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Journal of Archaeological Science 165 (2024) 105972
Available online 6 April 2024
0305-4403/© 2024 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
Prehistoric ornaments in a changing environment. An integrated approach
to the Late Upper Palaeolithic and Mesolithic Columbella rustica shells from
the Vlakno cave, Croatia
Barbara Cvitkuˇ
si´
c
a
,
b
,
*
, Emanuela Cristiani
b
,
**
, Andrea Zupancich
b
, Dario Vujevi´
c
c
a
Institute for Anthropological Research, Center for Applied Bioanthropology, Zagreb, Croatia
b
DANTE - Diet and Ancient Technology Laboratory, Sapienza University of Rome, Rome, Italy
c
Department of Archaeology, University of Zadar, Zadar, Croatia
ARTICLE INFO
Keywords:
Columbella rustica
chaîne op´
eratoire
Use-wear
Geometric morphometric
Experimental activity
Late Upper Palaeolithic
Mesolithic
Eastern Adriatic
ABSTRACT
This paper advances knowledge of human behavioural and adaptational strategies in coastal areas related to
acquiring, producing and distributing ornaments, specically, the omnipresent marine gastropod Columbella
rustica. By applying quantitative and qualitative approaches to the most extensive collection of Columbella rustica
shells in the Eastern Adriatic region discovered in the Late Upper Palaeolithic and Mesolithic levels of Vlakno
cave in Croatia, we have determined the complete step-by-step life cycle of this bead type, in particular, where
and how shells were collected, produced, used, distributed and discarded. By integrating different methodolo-
gies, our data revealed changes in the collection strategies, reduction of the shell size during the Mesolithic
period, and standardisation and continuity in production techniques. Detailed analyses of broken shells in the
archaeological assemblage identied the presence of technological traces resulting from processing mistakes,
supporting our hypothesis of on-site production. A signicant share of used and unused standardised beads
points that bead production at this site was for personal use but also likely for the exchange and distribution
systems. Standardised, systematic and long-lasting activity related to the ornaments places Vlakno cave as one of
the leading centres for maintaining regional exchange and communication networks in the Eastern Adriatic
region during signicant climatic and environmental changes happening in this region in the Late Pleniglacial
and the early Holocene. Detecting on-site activities related to the ornaments in Vlakno cave has extended our
understanding of how symbolic motives inuenced the settlement model of the Late Pleniglacial and Early
Holocene hunter-gatherers in Eastern Adriatic region and overall contributed to fundamental questions about the
complexity of ancient human societies’ adaptation strategies.
1. Introduction
Pigments and ornaments have attracted humans since the early
phases of the Palaeolithic and, from then, remained a part of human
material culture in almost all human societies (Sehasseh et al., 2021;
Radovˇ
ci´
c et al., 2015). Ornaments and pigments testify that our ances-
tors could not only "recognise" the potential to use and modify raw
material, e.g., ocher or animal tooth or shell, in bead but, consequently,
to create and share symbols and their meanings. Subsequently, these
specic archaeological remains point out how the prehistoric mind was
aware of aesthetics and, more importantly, the symbolic dimension
originating from wearing and exposing these creations to be visible to
others. Accordingly, prehistoric beads, the fundamental units of pre-
historic ornamental sets, are, together with pigments, one of the earliest
material proxies proving our ancestors’ ability to operate symbolically,
marked as a critical development point in human cognition
(´
Alvarez-Fern´
andez and J¨
oris, 2007; Bednarik, 2008; White, 1992). In
1992, R. White introduced the social and symbolic roots of the origin of
prehistoric body adornments in the anthropological debate. Since then,
they have played an essential role in our endeavours to understand our
cultural evolution better.
Displayed in their multiformity and worn in varied ways, personal
ornaments have become polysemic objects carrying diverse denotations
through their evolutionary history. In prehistoric contexts, beads are
* Corresponding author. Institute for Anthropological Research, Center for Applied Bioanthropology, Gajeva 32, 10 000, Zagreb, Croatia.
** Corresponding author. Sapienza University of Rome, Department of Oral and Maxillo-facial Sciences, Via Caserta 6, 00161, Rome, Italy.
E-mail addresses: barbara.cvitkusic@inantro.hr (B. Cvitkuˇ
si´
c), emanuela.cristiani@uniroma1.it (E. Cristiani).
Contents lists available at ScienceDirect
Journal of Archaeological Science
journal homepage: www.elsevier.com/locate/jas
https://doi.org/10.1016/j.jas.2024.105972
Received 27 October 2023; Received in revised form 6 March 2024; Accepted 27 March 2024
Journal of Archaeological Science 165 (2024) 105972
2
only rarely found in situ as part of a completely preserved individual
ornamental composition, i.e., in burials (Alarashi et al., 2023; Gravel--
Miguel et al., 2022; Orschiedt, 2016; Vanhaeren and d’Errico, 2001;
White, 1999). Beads are found scattered throughout the deposits or
recovered during the subsequent sieving of the sediment. Despite the
challenging circumstances of their recovery, analysis of these
small-sized artefacts can still provide important information about
prehistoric choices, lifestyles and symbolic behaviour. Preserved mate-
rials purposefully collected and used for bead production in early pre-
history include, amongst others, various types of mollusc shells, bones of
different animal species, stones (clay and soft ones), and wood (e.g.,
Taborin, 2004; White, 2007; Werker, 1988; Vanhaeren and d’Errico,
2006). We can only use our imagination while referring to ethno-
graphical records to conditionally complement our ancestors’ list of
other potentially perishable raw materials used for ornamentation. Until
now, prehistoric ornaments’ role was attributed to the aesthetic, pro-
tection, and construction of personhood (e.g., Bori´
c and Cristiani, 2019)
or as a marker of age, gender, power, or marital status (e.g., Bar-Yosef
Mayer; 2015; Kuhn and Stiner; 2007). Also, throughout varied qualita-
tive and quantitative analyses supported by ethnographical records of
modern forager populations, ancient personal ornaments are commonly
interpreted as indicators of social status and identity (Stiner, 2014;
Vanhaeren and d’Errico, 2011; Rigaud, 2011). Accordingly, ornaments
were essential "technology" for transmitting messages characterised by
the standardisation of beads’ form and formal redundancy, fundamental
features of any communicational system (Kuhn and Stiner, 2007).
Standardisation, necessary for the consistency of symbolic meaning, can
be observed in the prehistoric beads’ shapes, i.e., in the selectivity of
certain natural forms (shells, teeth) and production technology. The
small size and durability of the beads favour the redundancy supporting
the successful transmission of messages/information. In that regard,
beads have been one of the most signicant objects in studying accul-
turation or diffusion systems (Bourdieu, 1977) and are commonly part of
hunter-gatherers exchanging systems aiming at building and maintain-
ing social networks (Wiessner, 1982; Rigaud et al., 2019; Cvitkuˇ
si´
c,
2015). Still, we do not have enough knowledge and insights about the
acquisition and distribution routes of raw materials used for ornaments.
Ethnographically, connections and networks can be traced amongst
ethnographic groups if this information is available, such as in North
America (Dubin, 1999).
Despite the wealthy opus on the ornamental research in prehistory,
to date, we still miss more information on the human activities related to
the complete process of marine shell ornaments’ life cycle, i.e. where
shells were collected and selected, where and how they were produced,
used, distributed and nally discarded (Baysal and Yel¨
ozer, 2023;
Rigaud et al., 2019; White et al., 2007; Cristiani et al., 2014).
This paper aims to complement our knowledge of activities related to
acquiring, producing, using, and discarding ornaments in prehistoric
hunter-gatherer societies by integrating quantitative and qualitative
analysis and experimental activity by exploring the most extensive
collection of Columbella rustica shells in the Eastern Adriatic region
discovered in the Late Upper Palaeolithic and Mesolithic deposits of
Vlakno cave, characterised by multiform states of integrity and use.
During the Late Pleniglacial and early Holocene, the distribution of or-
naments is well documented in the Eastern Adriatic region and its hin-
terland (Cristiani et al., 2014; Cvitkuˇ
si´
c and Komˇ
so, 2015; Cvitkuˇ
si´
c,
2017; Cvitkuˇ
si´
c and Vujevi´
c, 2021; Komˇ
so, 2007; Komˇ
so and
Vukosavljevi´
c, 2011). The rich ornamental assemblage from the Late
Upper Paleolithic (LUP) and the Mesolithic deposits of Vlakno cave,
characterised by continuity in the representation throughout the se-
quences, variety of the bead types, the multiform outcome of production
activity (i.e. technological mistakes, perforated beads) and modalities of
use (i.e., used, unused) (Cvitkuˇ
si´
c et al., 2018), point that this site likely
served as a specialised workshop for the ornaments in the broader region
during indicated periods. The dynamic and activity of ornament pro-
duction at the site stayed undisturbed even with the profound Holocene
change of Vlakno becoming part of the island. The rich assemblage of
Columbella rustica in Vlakno cave allowed us to study and gain better
insight into behavioural and adaptational strategies in coastal and
inland areas and the connectivity of our ancestors related to symbolic
behaviour and mobility patterns associated with raw materials collec-
tion and chaîne op´
eratoire for ornaments.
2.Columbella rustica shells
Two species of Columbella (Lamarck, 1799) inhabit the Mediterra-
nean Sea and nearest parts of the Atlantic Ocean: (1) Columbella rustica
(Linn´
e, 1758) present in the entire Mediterranean Sea and extending
into the neighbouring Atlantic southward to Senegal and northward to
Portugal, and (2) Columbella adansoni (Menke, 1853) recorded in Cape
Verde islands, and assumed to occur across Macaronesia, from the
Azores to the Canary Islands, and along the West African coasts from
Ghana to Angola (Russini et al., 2017). Columbella rustica is a small
marine gastropod inhabiting warm waters’ rocky or sandy bottoms
(Poppe and Goto, 1991). It is well-adjusted to external environmental
alterations, such as temperature and pH changes (Wahl et al., 2016). The
natural colour varies from white to red and brown with irregularly
speckled patterns. The compact solid shell is oval-shaped, with the spire
not too high. The thickest parts of the C. rustica shell are (i) the area
along the top of the body whorl attaching to the spire, (ii) the siphonal
canal, and (iii) the outer lip of the aperture, while the thinnest part is the
apex, followed by the periphery part of the body whorl (Bosch et al.,
2023). This shell taxon is considered non-edible (Syrides, 2019).
Since the Upper Palaeolithic, C.rustica was collected and used
exclusively for ornamental purposes in European hunter-gatherers’ so-
cieties. The oldest C. rustica beads are discovered in Bizmoune Cave in
southwest Morocco in deposits dated ≥142 thousand years (Sehasseh
et al., 2021). With the beginning of the Mesolithic, C. rustica became
omnipresent in the archaeological record (´
Alvarez Fern´
andez, 2008) as a
unique element of a shared symbolic vocabulary of forager groups. In
later periods, it was not exclusively related to hunter-gatherer pop-
ulations (´
Alvarez Fern´
andez, 2003, 2008; Karali, 1999). C. rustica has a
wide spatiotemporal distribution in the Mediterranean area (´
Alvarez
Fern´
andez, 2003, 2008; Mussi, 2002; Stiner, 1999; Benghiat et al., 2009;
Cristiani, 2012; Kuhn et al., 2001; Bar-Yosef Mayer, 2005). It is present
in the continental part of Europe as well (e.g., Eriksen, 2002; ´
Alvarez
Fern´
andez, 2003; ´
Alvarez Fern´
andez, 2008) and is one of the few bead
types distributed over more extensive distances (Cucart-Mora et al.,
2022).
3. Regional and site background
Vlakno cave is located on the inner side of Dugi otok island, centrally
positioned within the Eastern Adriatic area (Fig. 1, nr. 9). The cave is a
simple speleological object with a small inner space of ~40 m
2
and a
wide opening facing southeast. There is a source of fresh water in the
vicinity, less than 150 m away (Brusi´
c, 2005:198; Vujevi´
c, 2021:35).
With these characteristics, it was almost ideal for a campsite and shelter
for small groups of foragers since prehistoric times (Vujevi´
c and Parica,
2011; Vujevi´
c and Bodruˇ
zi´
c, 2014; Cvitkuˇ
si´
c et al., 2018; Vukosavljevi´
c
et al., 2014). At the beginning of the Holocene, a profound environ-
mental change encompassed the Adriatic region, with a rapid sea level
rise (Suri´
c, 2006:182). Nine thousand years ago, the Adriatic plain
amounted to only 17% of its former area and was reduced to a narrow
strip around the Gulf of Trieste (Miracle 1995:117–118). The Adriatic
area’s geography was fundamentally reshaped and transformed from the
Adriatic Plain into the Adriatic Sea as we know it today (Peresani et al.,
2021; Sikora et al., 2014). During lower sea level, the cave was located
far from the sea, and the entire island was part of the mainland and
formed a 100–400 m high ridge rising above the valleys (Vujevi´
c and
Parica, 2011:23).
Due to sea level rise and ooding of the Adriatic Plain, Dugi otok and
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
3
the surrounding islands got separated from the mainland, forming one
much larger island, and Vlakno became a coastal site (Brusi´
c, 2005:198).
According to some studies (Miracle, 2007; Boschian 2003:99),
hunter-gatherer groups from the central and southern parts of the
Eastern Adriatic region were forced to settle in the hilly hinterland and
change their subsistence strategies. Reduced territory decreased con-
nections with the western Adriatic coast and directed groups to more
intensive use of local resources and movements towards the eastern
Adriatic hinterland. Nevertheless, many cave sites at the Adriatic Plain’s
edge attest to the landscape’s viability during that period (Pilaar Birch
and Miracle, 2017:87), and the rich archaeological assemblage from the
Vlakno cave further supports that notion (Vujevi´
c, 2016; Vujevi´
c and
Bodruˇ
zi´
c, 2021).
The Vlakno deposits yielded rich evidence of the LUP and Mesolithic
occupation without a visible hiatus in the stratigraphy, making Vlakno
an ideal place for studying the transition from the Pleistocene to the
Holocene and the adaptations of the Epigravettian communities to sig-
nicant climatic and environmental changes happening in this region.
Despite this profound change, the dynamic and activity traced in the
site’s entire sequence stayed consistent. Archaeological investigations in
the Vlakno cave started in 2004 (Brusi´
c, 2005), while systematic exca-
vations have been ongoing since 2011 (Vujevi´
c, 2011; Vujevi´
c and
Parica, 2011). In the test trench, a depth of almost 5 m was reached, with
cultural layers that can be traced back to the last 19.5 kya (Beta302247,
16330 ±70 BP, 17600-17450 cal BC (19550-19400 cal BP), taken from
Stratum 10). Systematic research is currently stopped at a depth of 3.7
m, i.e. Stratum 8 (Beta607870, 13640 ±40 BP, 14685 - 14372 cal BC
(16634 - 16321 cal BP). The stratigraphic sequence of Vlakno is well
documented in the test trench and conrmed with absolute dates, rep-
resenting a long-lasting occupation dated from early Epigravettian
throughout late Epigravettian and early Mesolithic in cultural
terms—absolute dates acquired from the site range from c. 19.5 ky cal
BP to c. 8.6 ky cal BP (Fig. 2.). The excavated area of a test trench is
divided into ten primary cultural strata with subdivisions, with many
layers without a visible hiatus separated only by thin layers of ash and
burnt soil representing occupation surfaces of the cave use (Vujevi´
c and
Bodruˇ
zi´
c, 2014; Cvitkuˇ
si´
c et al., 2018). At 2 m depth from the surface,
cultural layers are interrupted by a 10 cm thick layer of tephra, the
Neapolitan yellow tuff (dated to c. 14.9 ±0.4 ka cal BP; Deino et al.,
2004; Radi´
c et al., 2008). Stratum 1 represents the surface layer. It
displayed evidence of disturbance in stratigraphy and small nds
(several pieces of Neolithic and Roman-age pottery, along with some
recent archaeological material). Undisturbed Mesolithic layers below
the surface are phased in two: Stratum 2 and Stratum 3, corresponding
roughly to the Early Holocene. These upper strata consist of loose and
dusty sediments with high quantities of land snails and marine molluscs
in deposit. Preliminary sedimentology results indicate that most sedi-
ments are deposited through anthropogenic activities (G. Boschian, pers.
Fig. 1. Central position of Vlakno cave site (9) in the Eastern Adriatic region with the indicated coastline from Late Pleistocene to Holocene. Sites with ornaments: 1.
Nugljanska cave, 2.Abri ˇ
Sebrn, 3. Pupi´
cina cave, 4. Lim 001, 5. Romualdova cave, 6. ˇ
Sandalja II, 7. Ljubi´
ceva cave, 8. Zala cave, 9. Vlakno cave, 10. Vela spila.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
4
comm). The division is based primarily on 14C dating, although there is
also evidence of temporal changes in the faunal and lithic assemblages
from these two strata (Vujevi´
c and Bodruˇ
zi´
c, 2021; Radovi´
c et al., 2021).
Chronologically, the Mesolithic occupation started ca. 8000 cal BC
(Beta-677951, 9300 ±30 BP). The archaeological assemblage of
Mesolithic Strata 2 and 3 are typical Mesolithic but with pronounced
Epigravettian tradition, which can especially be observed in ornaments,
lithics, and subsistence strategies (Cvitkuˇ
si´
c et al., 2018). Stratum 4
marks the end of the LUP, while Stratum 5 represents the rst cave
settlement phase after the eruption in the Phlegraean elds. The tephra
layer, Neapolitan yellow tuff from the mentioned eruption, served as a
stratigraphic boundary between Stratum 5 and Stratum 6 (Vujevi´
c and
Parica, 2011:26; Vujevi´
c and Bodruˇ
zi´
c, 2021).
Strata 6 to 10 are located below the tephra layer and represent
various chronological sections of life in the cave during the chronolog-
ical period from about 15 to 19.5 ky cal. BP. Stratum 10 includes the
lowest layers from the test trench, although since systematic excavations
have not yet reached these layers, the phasing could be somewhat
different in the future. Lithics, especially endscrapers, back bladelets,
and backed points, show that all strata can culturally be designated to
the Epigravetien, although characteristics of the lithic assemblage from
Stratum 10 point to its early phase.
Preliminary analysis of faunal remains from Mesolithic Strata 2 and 3
shows a predominance of mammals. However, there are also remains of
sh and birds, which are extremely rare in the strata below (Radovi´
c
et al., 2021). Within the taxonomically identied mammalian assem-
blage in Stratum 3, red deer (Cervus elaphus) was the most common
mammalian taxon, followed by a red fox (Vulpes vulpes). In contrast, in
the chronologically later Stratum 2, the red fox is the predominant
taxon, followed by red deer. Marine sh remains are present in the
faunal assemblage from the beginning of the Mesolithic layers, with a
signicant increase in Stratum 2 (Radovi´
c et al., 2021). A combination
of dental calculus and stable isotope analyses from a well-preserved
buried skeleton of a male aged between 35 and 40 from Stratum 2
also showed that the individual regularly consumed marine resources
with various plant foods (Cristiani et al., 2018).
The ornamental assemblage (N =692) from Vlakno cave is
comprised of nine different taxa discovered throughout LUP (Stratum 4
to 8) and Mesolithic (Stratum 2&3) deposits (Table 1). The most rep-
resented ornaments are from marine species, i.e. gastropods Columbella
rustica and Tritia neritea and scaphopods Antalis sp. (Table 1). Besides
modied, many whole unmodied specimens have been discovered
(Table 1, number in the brackets), particularly 105 C. rustica shells in
Stratum 2 and 3.
4. Material and methods
We have analysed 379 C. rustica shells discovered in the LUP (N =
23) and Mesolithic (N =356) deposits of Vlakno cave (Fig. 3) aiming to
reconstruct the chaîne op´
eratoire, i.e., the sequence of actions related to
their production, such as modalities of collection, production, use, and
discard by performing technological, use-wear, and residue analyses
supported by experimental activity. The experimental activity was car-
ried out on a modern sample of 160 C. rustica collected along the
Adriatic coast, aiming to reconstruct the specic technique applied to
perforate the archaeological shells from Vlakno cave.
Taxonomical determination of archaeological C. rustica shells from
the Vlakno cave is based on the previous taxonomical identication
(Cvitkuˇ
si´
c, 2015) and strict geographic separation of two species
inhabiting the Mediterranean Sea.
Fig. 2. The stratigraphic sequence of Vlakno cave.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
5
The good state of preservation and the abundance of C.rustica shells
in the deposits above the tephra (from Stratum 5 to 2) allowed us to
apply a methodological approach of quantitative and qualitative
analysis based on integrated metrical, technological, and use-wear
studies, aided by taphonomical analysis. Furthermore, aiming to
detect specic production techniques, we have compared the
Table 1
Ornamental assemblage from Stratum 2 to Stratum 8, Vlakno cave. Numbers in the brackets indicate whole, unmodied specimens.
Stratum Marine gastropods, bivalve and scaphopods Freshwater gastropods Mammal
teeth
Columbella
rustica
Tritia
neritea
Tritia
nitida
Glycymeris
sp.
Acanthocardia
tuberculata
Antalis
sp.
Theodoxus
danubialis
Lythoglyphus
naticoides
Cervus
elaphus
2 222 (81) 3 2 (1) 1 2 (1)
3 29 (24) 22 (1) (3) 1 1 1 (1)
4 9 (1) 42 (1) 4 (5) 2 (1) 1 2 2 3
5 13 37 2 (6) 19 (2) (1) 7 1 1 2 (3)
TEPHRA ~ 14.9 ± 0.4 ka cal BP
6 45 (1) 1 7 174 2 1 18 (14)
7 7 1 4 (3)
8 1 2
Fig. 3. Selection of LUP and Mesolithic Columbella rustica shells from Vlakno cave.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
6
archaeological C. rustica with an experimental collection using Geo-
metric Morphometrics (Cristiani et al., 2020). All the analyses have been
performed at the DANTE—Diet and ANcient TEchnology laboratory
(Sapienza University of Rome, Italy).
4.1. Morphological and morphometric analyses
Before performing detailed analysis, LUP and Mesolithic C. rustica
shells were divided into three groups according to integrity: Group 1 –
whole specimens with perforation; Group 2 – whole specimens without
perforation; and Group 3 – damaged/broken specimens. Group 1 (G1) is
comprised of entire, undamaged perforated beads and beads with
missing small parts, i.e., the apex or small fragments of the outer lip (N
=223). Group 2 (G2) includes whole shells without modications (N =
106). Group 3 (G3) includes incomplete and/or damaged specimens, i.
e., shells with missing signicant parts of the body whorl or the entire lip
area with or without signs of perforation hole (N =50).
For all G1, G2, and experimental specimens, metric data were
recorded with a digital calliper: (1) dimensions of the bead (maximum
length and width in mm) and for the G1 (2) dimensions of the perfo-
rations (maximum length and width in mm).
The taphonomic study was carried out on all specimens with a focus
on pre-depositional alterations (predator drilling, bioerosion), anthro-
pogenic modications (perforations, ocher residues, thermic alter-
ations), and post-depositional alterations (fragmentations, de-
calcications) (Driscoll and Weltin, 1973; Claassen, 1998; Crothers,
2004; d’Errico et al., 2005).
4.2. Experimental activity
C. rustica shells used in the experimental activity were collected on
the shore of Zadar County, Croatia. Experimental shells (N =160)
divided into four groups (N =40) and perforated by testing two direct
and two indirect percussion techniques already documented in the
archaeological record (´
Alvarez Fern´
andez, 2006; Benghiat et al., 2009;
Rodríguez-Hidalgo et al., 2010; Cristiani, 2012; M˘
arg˘
arit, 2016;
M˘
arg˘
arit et al., 2018). Techniques of perforation applied in experi-
mental activities included the production of holes on the central body
whorl of the shell with 1. direct percussion using a pebble; 2. direct
percussion using a int core; 3. indirect percussion using int ake and
pebble; 4. indirect percussion using retouched point and pebble (Cris-
tiani et al., 2020). Attempts to perforate C. rustica using (1) a bone tool
and (2) a wooden stick have been unsuccessful, resulting in damaged
and broken points of tools made of this compared to shell, softer, organic
material.
Described and recorded criteria on the experimental C. rustica shells
comprehend perforation shape, section morphology of the walls, per-
cussion ake, micro-aking, compressions, crushing and notching
marks, striations, and cracks. In particular, the outline of the perforation
(circular, oval, sub-regular, and irregular), the section morphology of
the perforation walls (straight, internally bevelled or jagged), the pres-
ence/absence of the percussion ake, the position (internal or external)
and the invasiveness of the micro-aking, the presence/absence and the
organisation (isolated or bands) of striations, the invasiveness of com-
pressions marks, as well as the presence/absence of crushing and
notching, were recorded together with the presence/absence of cracks
starting from the perforation rim for each perforated shell.
4.3. Microscopic analysis
Specimens from G1 and G3 were analysed by low and high magni-
cation by using a Zeiss Axio Zoom V16 binocular stereo microscope
with progressive magnications ranging between ×10 and ×112 and
equipped with a Zeiss Axiocam 305/506 colour camera. For the G1
specimens, the use-wear and technological analysis aimed to identify
functional modications such as rounding of the perforation, faceting of
the prole, changes of colour, striations, and residues in order to discern
patterns of acquisition, manufacturing, wear, use, and nally, deposi-
tion/discard. The type and distribution of use-wear traces and residues
related to the perforation hole, the lip, and dorsal and ventral surfaces
have been recorded. For the G3, the use-wear and technological analysis
aimed to identify specimens that were discarded as a mistake during
perforation activity, i.e., technological accidents based on technological
marks such as striations together with attening or compression from
counterblows, crushing and notching marks, percussion ake in the
perforation area, and to detect possible used shells among broken
specimens. All technological traces on the archaeological specimens
were observed according to the traces recorded on the experimental
sample.
4.4. Geometric morphometric
A 2D geometric morphometric approach was applied to test whether
it could distinguish between perforation holes on C. rustica shells on the
base of the perforation shape. Zenithal pictures of experimental and
archaeological shells were taken using a Canon EOS 100D with a xed
50 mm macro lens.
Thirty-six (36) landmarks were manually positioned at every 10◦
along the perforation in a clockwise direction using the open-source
software tpsDIG2 (v.2.32). To facilitate this process, a digital protrac-
tor was placed on the specimen picture at the centre of the perforation,
vertically aligned with the siphon. The analysis of the perforation holes
was carried out in RStudio (v. 2023.06.1 +524) using the packages
Momocs (1.4.0), tidyverse (v.2) and kableExtra (1.3.4). Elliptic Fourier
analysis of the perforation shapes was performed in Momocs, where the
TPS data for each perforation was imported as coordinates and scaled to
centroid size. Following Bonhomme et al. (2014), Fourier transforms
were computed for each perforation hole. Twelve harmonics were
retained, allowing to gather 99% of the harmonic power. Results were
obtained by applying Principal Component Analysis to assess shape
variation and potential clustering based on the shape of the perforation
hole. Raw data and R code can be found in https://zenodo.org/records
/10786315.
5. Results
The Mesolithic specimens (N =356) are better preserved than the
LUP (N =23). Visual inspection of C. rustica shells revealed that a sig-
nicant share of Mesolithic shells display the characteristic alterations
of shells collected in thanatocenoses, such as bioerosion marks, sponge
marks, and/or breakage of the spire, compared to LUP specimens where
most of the shells have the appearance of freshly collected shells with
preserved apex. The whole sample has a signicant share of specimens
characterised by multiform outcomes, such as entirely perforated and
intact shells, whole perforated and intact shells with changed natural
colour to black, broken specimens, shells with the marks of the initial
but unnished perforation process, and manufacturing discard. We
hypothesise that the heterogeneous sample of C. rustica shells is related
to the on-site production activity of the ornaments, in particular, that
broken specimens are likely discarded as technological mistakes, intact
shells are a raw material supply, and that among perforated beads, there
are a signicant share of used but also unused ones.
The distribution of C. rustica according to the state of integrity is
Table 2
Distribution of C. rustica according to the state of integrity: G1 – whole speci-
mens with perforation; G2 – whole specimens without perforation; G3 –
damaged/broken specimens.
Columbella rustica G1 G2 G3
Late Upper Palaeolithic 21 1 1
Mesolithic 202 105 49
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
7
presented in Table 2. An increased number of specimens in divided
groups (G1, G2 and G3) characterise the Mesolithic sample.
Results of morphometric analyses revealed differences between the
size of Mesolithic and LUP C. rustica. The average length and width of
the C. rustica with intact apex from LUP deposits is 14.62 ×8.85 mm,
and from Mesolithic, 13.49 ×8.69 mm (Table 3; Fig. 4). The average
dimension of the experimental sample is 13.62 ×8.39 mm (Table 3).
The naturally present pattern on the shell’s surface is invisible in
most archaeological C. rustica. Loss of the natural pattern results from
the various chemical and mineralogical changes that occur due to long-
term residence in the sediment (Claassen, 1998). Sponge marks and
pitting forms are detected in almost half of the Mesolithic sample,
indicating that shells were likely collected on the beach. Calcication
and root damage are present in 12% of Mesolithic specimens. Pitting
forms and root damage are observed on two LUP specimens.
Almost 20% of perforated LUP and ~20% of G1, G2, and G3 Meso-
lithic C. rustica shells are black, characterised by the even dark colour of
the surface and core of the shell, likely resulting from controlled expo-
sition to the re (Perl`
es and Vanhaeren, 2010). In the Mesolithic sample,
the share of perforated (used ~36%, unused ~28%) and whole un-
modied black shells (~36%) is represented almost equally. LUP and
Mesolithic C. rustica shells in Vlakno cave appear as (1) non-burnt, i.e.
with the natural colour of the surface, or (2) burnt, i.e. black or very dark
grey colouring of the surface and core of the shell (Claassen, 1998). In
the Vlakno sample, we have not detected heated but not burnt shells
with light grey colour or surface patches (Claassen, 1998).
The use-wear analysis of G1 showed that ~60% of the LUP and
Mesolithic C. rustica beads were used (Fig. 5).
The analysis revealed that ~40% of G3 specimens were broken
during the perforation process. They all have one or a combination of
more common features visible in a perforation area on the body whorl in
the form of percussion aking and/or int striations marks (Fig. 6; e, f,
h, k, s) and very sharp perforation edges with a characteristic pattern of
missing outer lip area (Fig. 6; e, s). Hence, they were marked as tech-
nological mistakes. A small share of broken beads in G3 have use-wear
traces, but it cannot be distinguished if the fractures result from the
use or post-depositional factors. Moreover, the cause of shell damage of
G3 specimens without technological traces conditionally can be attrib-
uted to natural factors (predators, post-depositional) or anthropic fac-
tors (trampling, gathering debris) (e.g. Bosch et al., 2023; Stiner, 1999;
Perl`
es, 2016; Stafford et al., 2015).
Moreover, during analyses, ocher residues, in the form of small steins
or spots, have been detected in one-fourth of the Mesolithic beads,
localised along the edge of the perforation for the most (Fig. 7; a, b, c, e),
across the columella (Fig. 7; a, d, g), on the surface of the body whorl
(Fig. 7; a, d, e) and along the lip (Fig. 7; f, h). Spots of red residues have
been identied inside one LUP specimen. The type of ochre distribution
could be associated with using coloured threads to retain the ornaments
(Cristiani et al., 2014). On one C. rustica (Fig. 7; e), a big part of the shell
surface appears covered by a reddish compacted patina that distributes,
leaving the surface below the perforation rim clean. This would conrm
that the ornaments could have been retained with coloured threads or
that the shells had been in contact with an intensely red-coloured
surface.
The experimental activity resulted in the biggest share of broken
specimens in the experimental samples perforated by indirect percussion
using int ake and pebble and retouched point and pebble. Techno-
logical mistakes are present, but less, in the experimental samples
perforated by direct percussion using a pebble and a int core. Also, we
have noted that production mistakes depend on the expertise of the
person performing the experimental activity, mainly that a person with
more experience had fewer mistakes.
When analysing the shapes of the experimental perforation holes, it
is possible to differentiate between perforations produced by the
different tested techniques. As seen in Fig. 8 the perforation holes made
through various techniques and tools form three distinct clusters. In
particular, the PCA results highlight the apparent difference in the shape
of holes produced through direct percussion using a pebble and the ones
made through indirect percussion using a ake edge. The former are
circular, while the latter are thinner and more elliptical; nally, the
perforation holes produced using a int point through indirect percus-
sion result in a more oval shape (Fig. 8).
Furthermore, the results of LDA on the PCA scores show that in 70%
of the cases, it is possible to discriminate between the tested perforation
techniques appropriately. In 90% of the cases, holes produced using a
pebble through direct percussion are correctly identied. A similar high
successful discrimination rate (80%) is also achieved for perforation
holes made using a ake edge adopting an indirect percussion. A much
lower correct identication rate (40%) characterised the holes produced
using a int point through indirect percussion.
Finally, the results of MANOVA performed on the PCA scores show a
signicant difference between the perforation produced by indirect
percussion with a pebble and indirect percussion with a ake edge (p =
<0.001) as well as in the case when the perforation produced by direct
percussion with a pebble and indirect percussion with a point are
compared (p =0.0165) (Table 4). On the other hand, no signicant
difference is recorded when the perforations produced through indirect
percussion using a ake and a point are compared (p =0.078) (Table 4).
Several points can be raised by comparing the perforation holes of
C. rustica shells from Vlakno with the experimental ones (Fig. 9).
First, no difference in shape is observed between the LUP and the
Mesolithic C. rustica from the Vlakno assemblage. As shown in Fig. 9, the
PCA shows that the shape of the archaeological perforations falls to-
wards the more circular ones experimentally produced by direct per-
cussion using a pebble. These results are conrmed when a MANOVA
Pairwise is run to compare the perforation shapes of the C. rustica from
Vlakno with the ones produced experimentally (Table 5).
No signicant differences are recorded in perforation shapes among
the archaeological specimens, suggesting that a similar technique was
employed to perforate C. rustica shells during both periods. In this sense,
using a pebble through direct percussion was the technique most likely
employed, given the signicant differences in shape between the
archaeological specimens and the holes produced through indirect
Table 3
Descriptive statistics for Late Upper Palaeolithic, Mesolithic and experimental
C. rustica with preserved apex.
N Minimum
(mm)
Maximum
(mm)
Mean
(mm)
Std.
Deviation
Late Upper
Palaeolithic
18 12,37 16,09 14,6433 1,00009
Mesolithic 207 10,05 16,82 13,4109 1,31726
Experimental
sample
136 11,00 17,00 13,6213 1,10094
Fig. 4. The difference in the size between Late Upper Palaeolithic and Meso-
lithic C. rustica from Vlakno cave.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
8
percussion using a int point.
6. Discussion
By integrating quantitative and qualitative analyses of C. rustica
shells from LUP and Mesolithic deposits of Vlakno cave, we have
determined its anthropogenic collection, production, and use, indicating
that Vlakno cave was a specialised workshop for ornaments. Further-
more, the conducted study allowed us to elaborate on the on-site ac-
tivities related to ornaments, including complete chaîne op´
eratoire for
this widespread taxon for shell ornaments omnipresent in the Mediter-
ranean region ( ´
Alvarez Fern´
andez, 2008, 2010; Cristiani et al., 2014;
Cvitkuˇ
si´
c, 2017). Moreover, we have detected a reduction in the size of
the Mesolithic specimens. A signicant difference in size, i.e., length
(Perl`
es, 2016), of Vlakno’s Late Pleistocene and Holocene C. rustica
shells can be interpreted in favour of three possible intertwined factors:
(1) environmental changes; (2) human pressure on coastal resources;
and (3) changes in collection strategies. The environmental change in
the central Eastern Adriatic caused by the rise of the sea level, separating
Dugi otok and Vlakno cave from the mainland, has consequently
directed inhabitants to exploit mainly local coastal resources. Studies
have shown that these two factors can impact the decrease in shell size
(García-Esc´
arzaga et al., 2022; Klein and Steele, 2013). The decrease of
the shell size and signicant increase in the Mesolithic sample perfectly
t in the period of Dugi otok becoming an island induced by sea level
rise, where changes in the surface temperature of the Adriatic Sea were
undoubtedly present. However, research showed that this small-sized
gastropod is well-adjusted to environmental changes, i.e., temperature
and pH alterations (Wahl et al., 2016), an occurrence common in the
eastern Adriatic area (Miliˇ
si´
c, 2007). The increasing prominence of
shing activities during the Early Mesolithic period is likely connected
with an environmental change and rising sea levels, with Dugi otok
becoming separated from the mainland already before ca. 10,194–8351
cal BC at 95% condence (Z-3660: 9760 ±280 BP) (Suri´
c, 2006) and
Vlakno cave becoming a coastal site. Dental calculus and stable isotope
analyses (human burial, Stratum 2) showed that the inhabitants regu-
larly consumed marine resources (Cristiani et al., 2018), and the same is
recorded for early Holocene layers of the nearest site of Vela Spila
(Rainsford et al., 2014). Likely, environmental changes directed the
human population inhabiting Vlakno cave to the intensive exploitation
of coastal resources, a behaviour that consequently impacted the
C. rustica shell size.
Besides the intensive exploitation of coastal resources caused by
changed geography, the reduction in the size of the Holocene shell can
also point to modications in collection strategies. The exact size dif-
ference of C. rustica from the Late Upper Palaeolithic and Mesolithic
periods is recorded in Franchti cave (Perl`
es, 2016). Perl`
es’ experimental
activity (2016) revealed that the mode of collection (thanatocoenoses vs.
underwater) could be a signicant factor in size variation. C. rustica
collected underwater, on average, was bigger. Compared to the one
collected from the beach, the only recorded lack was the dull colour of
the shell. The signicant share of black shells, with the shell’s integrity
and signicant height of the LUP C. rustica, can indicate that shells were
collected alive from the sea during this period. It is rather challenging to
prove if hunter-gatherers collected them in the shallow waters or on
rocky shores since the only indicators are signicant mean height and
integrity of the shell with preserved apex, but it is something that should
be considered. ~30% of the Mesolithic sample is characterised by
sponge marks, pitting, and missing apex, indicating that shells were at
least partially gathered on the nearby coast. Interestingly, more than
80% of Mesolithic black C. rustica have an intact apex. By all means, a
two-mode gathering - thanatocoenoses and underwater - can explain
characteristics present in the Vlakno Mesolithic sample. On the contrary,
in the LUP sample, we found only two shells collected in thanatocenoses.
Hence, we can consider that the beginning of the Holocene brought
modications in gathering modes in the Vlakno cave.
Regarding black C. rustica shells, our hypothesis on the deliberate
burning, especially in Holocene layers, is supported by the following
facts: (1) Among nine different taxa used for beads, changes in natural
colour due to thermic alteration are limited exclusively to C. rustica and
Fig. 5. Selection of used LUP (a, b) and Mesolithic (c – l) C.rustica from Vlakno cave presented by sample number: a) 78.4; b) 56.6; c) 14.96; d) 14.206; e) 24.7; f)
14.76; g) 14.98; h) 14.4; i) 24.6; j) 14.78; k) 14.32; l) 14.78.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
9
Tritia neritea taxa; (2) In comparison to the 20% of burnt C. rustica, the
proportion of burning or heating traces on Mesolithic lithic artefacts is
~3,5% (Z. Perhoˇ
c and M. Bodruˇ
zi´
c; pers. comm. 2024); (3) The pro-
portion of intact, used and unused perforated black C. rustica is nearly
equal in the Mesolithic sample.
Combined analyses of C. rustica allowed us to elaborate on step-by-
step activities related to the on-site production of ornaments, specif-
ically (i) modes of the acquisition and selection, (ii) colour modication
of the selected sample, (iii) perforation activity, and (iv) sorting of the
outcome (Fig. 11). With the high percentage of likely deliberately burnt
intact black C. rustica shells in the overall Mesolithic sample of intact
shells (~28%), we can suggest that the change of natural colour in
Vlakno cave preceded perforation activity. As for our experimental ac-
tivity, perforation actions in Vlakno cave resulted in two possible
outcomes: (1) successful perforation and (2) unsuccessful perforation, i.
e., technological mistake. Moreover, one specimen from Vlakno cave
exhibits an unnished perforation, indicated by a small round hole of
int trace mark characterised by a counterblow mark on the dorsal side
of the shell (Fig. 11, 3). Interestingly, the same case is recorded on one
C. rustica from the Ifri Oudadane site in NW Morocco (Hutterer et al.,
2021).
As illustrated in Fig. 10, our analysis revealed that following the
perforation activity, beads were sorted as (1) perforated shells (intended
for use (used ~60%; Figs. 5 and 7), or as a supply (unused ~40%; Fig. 6),
and (2) discard, i.e., technological mistakes (~40%; Fig. 6).
On-site bead production in Vlakno cave is strongly indicated by (i)
unused perforated LUP and Mesolithic C. rustica (~40%), (ii) unperfo-
rated black shells (~20%), and (iii) technological mistakes (~40%).
Fig. 6. Selection of LUP (a, b), Mesolithic (c, e, f, h, j, k, m, o, q, s) and experimental (d, g, I, n, p, r, t) C. rustica (framed in red) showing unused specimens,
technological mistakes and technological traces. Unused - a, b, c, e, h, j, m, q, s; Flint striations - c, d, e, f, g, h, k, n; Compression mark – I, m, n, o; Flake of production
– s, t; Fracture of the body whole – q, r; Technological mistakes and perforation akes– e, s.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
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Broken beads in archaeological contexts were generally not recognised
as technological mistakes, while whole and broken specimens were not
considered for the use-wear analysis. In the case of Vlakno cave,
microscopic analyses of the complete C. rustica assemblage, regardless of
the integrity state, have revealed important information related to the
production activity and the selection process of the nished products.
Broken shells with perforation holes could be considered accidently lost
or damaged during use. Experimental activity suggested that in Vlakno’s
case, a signicant portion of broken shells without use-wear were likely
fractured during production and subsequently discarded as waste. Use-
wear is present on only a few fractured specimens. All technologically
broken and unused C. rustica in Vlakno cave exhibit standardised fea-
tures, such as percussion aking in the hole area, int striations marks
on the body whorl, and a missing part of the outer lip of the aperture (see
Fig. 6). The last feature strengthens our hypothesis regarding techno-
logical mistakes, as this part is one of the thickest areas of C. rustica
shells and is rarely missing due to post-depositional processes. This is in
contrast to, for example, the apex (Bosch et al., 2023), which is missing
in 20% of the Vlakno archaeological sample. Moreover, it should be
mentioned that the breakage of the shell in gastropods can result from
predator attacks, such as crustaceans, which are responsible for much of
the apertural lip damage observed in modern systems (Walker and Brett,
2002). While a gastropod that survives an attack can repair its damaged
shell, the process will leave visible scars as disruptions in the growth
lines, surface ornament, or colour pattern of the shell (Stafford et al.,
2015). We have ruled out the possibility of a crustacean attack as the
cause of the breakage of C. rustica in Vlakno cave due to the presence of
technological traces. However, crustacean attack as a cause of breakage
Fig. 7. Selection of C. rustica with ocher residues presented by sample number: a) 14.102; b) 24.7; c) 14.13; d-f) 14.101; g-h) 14.211. The insert in gure "e" marks
the part of the shell surface where residues are less distributed.
Fig. 8. Morphometric analysis of the experimental perforation. a) PCA of the experimental perforation holes; b) perforation shape comparison.
Table 4
Results of the MANOVA pairwise performed on PCA scores of experimental perforation holes. Signicance codes:’***’ =0.001; ’**’ =0.01; ’*’ =0.05; ’-’ =0.1
MANOVA Pairwise – Perforation shape vs. Perforation technique
Df Pillai approx F num Df den Df Pr(>F) Signicance
Direct percussion with pebble - Indirect percussion with ake edge 1 0.9681009 27.313975 10 9 0.0000158 ***
Direct percussion with pebble - Indirect percussion with point 1 0.8338897 4.518087 10 9 0.0165126 *
Indirect percussion with ake edge - Indirect percussion with point 1 0.7467457 2.653741 10 9 0.0787874 –
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
11
cannot be ruled out for the specimens without technological marks
based solely on fracture patterns. Additionally, shells could have been
accidently brought to the cave already naturally broken as a part of
gathering activities (e.g., Bosch et al., 2023; Perl`
es, 2016), or broken in
the cave, for example, by trampling.
Regarding perforation techniques, they were simple and consistent
during both periods, most likely involving direct percussion with a
pebble. In general, the perforation of prehistoric beads involved simple
techniques (e.g. Cristiani et al., 2014; Perl`
es and Vanhaeren, 2010;
Fig. 9. Morphometric analysis of the archaeological and experimental perforation holes. a) PCA combining both archaeological and experimental specimens; b)
shape comparison between archaeological and experimental perforation holes.
Table 5
Results of the MANOVA pairwise performed on PCA scores of experimental and archaeological perforation holes. Signicance codes:’***’ =0.001; ’**’ =0.01; ’*’ =
0.05; ’-’ =0.1
MANOVA Pairwise of Archaeological Perforation vs. Experimental Perforation
Df Pillai approx F num Df den Df Pr(>F) Signicance
Late Upper Palaeolithic - Mesolithic 1 0.0838367 0.9791407 10 107 0.4659763 –
Late Upper Palaeolithic - Direct percussion with pebble 1 0.1889574 0.3960673 10 17 0.9305411 –
Late Upper Palaeolithic - Indirect percussion with ake 1 0.8261988 8.0812895 10 17 0.0001057 ***
Late Upper Palaeolithic - Indirect percussion with point 1 0.6402423 3.0254024 10 17 0.0216002 *
Mesolithic - Direct percussion with pebble 1 0.1040516 1.1497436 10 99 0.3339882 –
Mesolithic - Indirect percussion with ake 1 0.5726424 13.2656139 10 99 <0.001 ***
Mesolithic - Indirect percussion with point 1 0.2774711 3.8018733 10 99 0.0002308 ***
Fig. 10. Illustration of the step-by-step on-site activities related to the pro-
duction of C. rustica beads.
Fig. 11. On-site bead production in Vlakno cave is strongly indicated by: 1.
Intact shells with natural colour; 2. Intact shells with changed color; 3. Shell
with the mark of perforation punch; 4. Used black shells; 5. Unused black shells;
6. Used with natural color; 7. Unused with natural color; 8. Technolog-
ical mistakes.
B. Cvitkuˇ
si´
c et al.
Journal of Archaeological Science 165 (2024) 105972
12
Vanhaeren and d’Errico, 2001; White, 2007), although few complex and
time-consuming processes have been detected in Palaeolithic assem-
blage (Heckle, 2018; Wei et al., 2017; White, 1989). Standardisation in
Vlakno cave is evident in both production technology and the preference
for raw material in favour of C. rustica shells during the Mesolithic. The
small size and durability of the C. rustica shell support the redundancy,
which is essential for successfully transmitting messages/information
((Kuhn and Stiner, 2007). Hence, standardisation and redundancy as
fundamental features of the communication system further support the
role of Vlakno cave as a relevant taskscape (Ingold, 1993), i.e., a speci-
alised workshop for producing and distributing ornaments in the Eastern
Adriatic region, and likely further. The presence of C. rustica beads at the
contemporaneous sites in the region (Fig. 1, sites 2–4, 7–10) supports the
notion of the shared symbolic vocabulary and aesthetic standards of the
foragers’ groups inhabiting the Eastern Adriatic region (Cvitkuˇ
si´
c and
Vujevi´
c, 2021; Cristiani et al., 2014; Cvitkuˇ
si´
c, 2017; Komˇ
so and
Vukosavljevi´
c, 2011). Conducted studies on the ornaments from this
area revealed no evidence of bead production (Cvitkuˇ
si´
c, 2017;
Cvitkuˇ
si´
c and Komˇ
so, 2015; Komˇ
so and Vukosavljevi´
c, 2011), except for
the Vela Spila site (Cristiani et al., 2014). In general, ornaments are
found in low numbers, with anthropogenic modications suggesting
they were most likely introduced, already used, and accidently lost at
those sites. A slightly different situation is recorded in the Holocene
levels of the Vela spila site, with the ornament assemblage comprised
exclusively of C. rustica characterised by mainly used beads and with a
small share of unused beads together with ~13% unperforated speci-
mens, pointing to on-site production (Cristiani et al., 2014). Compared
to Vela Spila, the Holocene ornamental assemblage from Vlakno cave is
more diverse concerning the type of raw materials and the state of the
beads’ integrity, with C. rustica as the dominant bead type. Furthermore,
the selection and use of C. rustica shells in Vlakno cave more straight-
forwardly reect modes of collection strategies and reveal stages of
production activity to the specic details (Fig. 11). Accordingly, the
Vlakno cave, with its central position in the Eastern Adriatic region,
seems to have represented a central point for the acquisition, produc-
tion, and distribution of the ornaments in the region. Extensive spatial
distribution of C. rustica from the Italian peninsula along the Adriatic
area to the south in Montenegro and Greece (e.g., Bori´
c et al., 2023;
´
Alvarez Fern´
andez, 2008; Mussi, 2002; Taborin, 1993; Cvitkuˇ
si´
c, 2017;
Cristiani et al., 2014; Cristiani and Bori´
c, 2017; Perl`
es, 2018), suggest
the existence of wide-ranging networks and communication paths in this
northern corner of Mediterranean. Shell ornaments are suitable for
tracking our ancestors’ symbolic behaviour and ideas (e.g. Cucart Mora
et al., 2022; Rigaud et al., 2022), and several studies have shown the
potential of isotope analysis of shells for tracing the movements (Colo-
nese et al., 2009; Milano et al., 2022). In future, this approach could
contribute to a more profound understanding of the role of the Vlakno
cave in the settlement patterns and mobility of prehistoric populations
in this region.
7. Conclusion
This paper has complemented our knowledge related to the symbolic
behaviour of our ancestors, how ornaments inuenced movements, and
how they contributed to the creation of ’the pattern of dwelling activ-
ities’ (Ingold, 1993: 153) and opened new directions for studying or-
naments. Since the Middle Palaeolithic, shells have been intentionally
collected and used as ornaments. Our results conrmed that the Vlakno
cave was no exception, with continuity of purposeful shell gathering
during the Late Upper Palaeolithic and Mesolithic and solid arguments
that it was used as a specialised workshop for ornaments where on-site
production was taking place. Furthermore, C. rustica from Vlakno cave
allowed us to perceive ornaments in a different light and better under-
stand our ancestors’ symbolic behaviour and mobility patterns related to
the collection of raw materials and chaîne op´
eratoire that can be applied
to the ornaments. Standardisation of production technology and a
signicant share of used and unused standardised beads of C. rustica
shells indicate that bead production in Vlakno cave was for personal use
and the exchange and distribution systems in the broader region. During
the environmental Pleistocene - Holocene and techno-cultural Palae-
olithic-Mesolithic transitions, the Vlakno cave kept a signicant role in
the Eastern Adriatic and most likely operated as one of the leading
taskscape locations for maintaining regional exchange and communica-
tion networks. Overall, identication of the raw material preferences,
technological choices, modes of use, and ornamental production
standardisations allowed us to understand better the symbolic behav-
iour of prehistoric hunter-gatherers hidden behind the scattered beads
throughout the Vlakno deposits. Finally, the Vlakno cave, as a signi-
cant location for activities related to ornaments, broadens our knowl-
edge of how symbolic motives inuenced the settlement model of Late
Upper Palaeolithic and Mesolithic foragers of the Eastern Adriatic area.
CRediT authorship contribution statement
Barbara Cvitkuˇ
si´
c: Writing – review & editing, Writing – original
draft, Methodology, Funding acquisition, Formal analysis, Data cura-
tion, Conceptualization. Emanuela Cristiani: Writing – review & edit-
ing, Supervision, Methodology, Funding acquisition, Formal analysis,
Conceptualization. Andrea Zupancich: Investigation, Formal analysis,
Data curation, Methodology, Writing – original draft, Writing – review &
editing. Dario Vujevi´
c: Data curation, Funding acquisition, Investiga-
tion, Writing – review & editing.
Declaration of competing interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
Acknowledgements
The research at Vlakno cave was conducted through the Croatian
Science Foundation projects IP-2019-04-6115
″
Epigravettian commu-
nities of Northern Dalmatia" and UIP-2014-09-1545
″
Transition and
tradition in Vlakno cave: Model of the transition from the Palaeolithic to
the Mesolithic in the Northern Dalmatia region“ by DV.
This research was supported by the European Research Council
under the European Union’s Horizon 2020 Research and Innovation
Program (grant agreement no. 639286 HIDDEN FOODS to EC; http://
www.hidden-foods.eu) to EC and by Postdoctoral SAPIExcellence
Fellowship (no. 616_22_SAPIEX ReDress) at Sapienza University of
Rome, Rome, Italy to BC. We thank the anonymous reviewers for their
insightful comments, which have greatly contributed to the improve-
ment of this paper.
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