A systematic review of natural processes in creating pierced shells ...

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Kubicka AM, Rosin ZM, Tryjanowski P, Nelson E. (2017) A systematic review ofanimal predation creating pierced shells: implications for the archaeologicalrecord of the Old World. PeerJ 5:e2903 https://doi.org/10.7717/peerj.2903

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https://doi.org/10.7717/peerj.2903

A systematic review of natural processes in creating piercedshells: implications for the archaeological recordAnna Maria Kubicka, Zuzanna M Rosin, Piotr Tryjanowski, Emma Nelson

Background. The shells of molluscs survive well in most sedimentary contexts and yieldinformation about the diet of prehistoric humans. They also yield evidence of symbolicbehaviours, through their use as beads for body adornments. Researchers often analysethe location of perforations in shells to make judgements about their use as symbolicobjects (i.e., beads), the assumption being, that holes attributable to deliberate humanbehaviour are more likely to exhibit low variability in their anatomical locations, whileholes attributable to natural processes yield more random perforations. However, thereare non-anthropogenic factors that can cause perforations in shells and these may not berandom. The aim of the study is to look at the association between variation of holes inshell beads from archaeological sites and shells pierced by natural (non-human) processes.Methods. Two hundred and sixty scientific papers retrieved from online databases byusing keywords, (e.g., ‘shell beads’; ‘pierced shells’); 77 of these publications enabled usto conduct a systematic review and assess the location of the hole in the shell beads in thepublished articles. Results. Almost all archaeological sites described shells beads withholes in a variety of anatomical locations. High variation of hole-placement was also foundwithin the same species from the same site, as well as among sites. In contrast, predatorswere more specific in where they attacked molluscs; birds often select the thinnest part ofthe shell, while molluscs and cephalopods target thicker parts. Discussion. These resultsindicate that variation in hole-location on shells pierced by humans is greater thanvariation in the placement of holes created by natural processes. Consequently, thesepatterns are opposite to those expected. We also found that Gastropod and Bivalvepredators choose similar hole locations to humans. Research into human shell-beadsrecovered from archaeological contexts should take into account non-anthropogenicfactors, which can lead to more realistic scenarios of the cultural behaviours of prehistoricpeople.

PeerJ PrePrints | https://doi.org/10.7287/peerj.preprints.1710v1 | CC-BY 4.0 Open Access | rec: 4 Feb 2016, publ: 4 Feb 2016

1 A systematic review of natural processes in creating pierced shells: implications for

2 the archaeological record

3 Anna Maria Kubicka1, Zuzanna M. Rosin2, Piotr Tryjanowski3, Emma Nelson4,5

4

5 1Adam Mickiewicz University in Poznań, Faculty of Biology, Department of Human

6 Evolutionary Biology, Poznań, Poland.

7 2Adam Mickiewicz University in Poznań, Institute of Experimental Biology, Department of

8 Cell Biology, Poznań, Poland.

9 3Poznań University of Life Sciences, Institute of Zoology, Poznań, Poland.

10 4School of Medicine, University of Liverpool, Liverpool, UK

11 5Archeology, Classics and Egyptology, University of Liverpool, Liverpool, UK.

12

13 Corresponding author:

14 Anna Maria Kubicka

15 Umultowska 89, Poznań, PL61-614, Poland

16 e-mail address: [email protected]


17 ABSTRACT

18 Background. The shells of molluscs survive well in most sedimentary contexts and yield

19 information about the diet of prehistoric humans. They also yield evidence of symbolic

20 behaviours, through their use as beads for body adornments. Researchers often analyse the

21 location of perforations in shells to make judgements about their use as symbolic objects

22 (i.e., beads), the assumption being, that holes attributable to deliberate human behaviour

23 are more likely to exhibit low variability in their anatomical locations, while holes

24 attributable to natural processes yield more random perforations. However, there are non-

25 anthropogenic factors that can cause perforations in shells and these may not be random.

26 The aim of the study is to look at the association between variation of holes in shell beads

27 from archaeological sites and shells pierced by natural (non-human) processes.

28 Methods. Two hundred and sixty scientific papers retrieved from online databases by

29 using keywords, (e.g., ‘shell beads’; ‘pierced shells’); 77 of these publications enabled us to

30 conduct a systematic review and assess the location of the hole in the shell beads in the

31 published articles.

32 Results. Almost all archaeological sites described shells beads with holes in a variety of

33 anatomical locations. High variation of hole-placement was also found within the same

34 species from the same site, as well as among sites. In contrast, predators were more

35 specific in where they attacked molluscs; birds often select the thinnest part of the shell,

36 while molluscs and cephalopods target thicker parts.

37 Discussion. These results indicate that variation in hole-location on shells pierced by

38 humans is greater than variation in the placement of holes created by natural processes.

39 Consequently, these patterns are opposite to those expected. We also found that Gastropod


40 and Bivalve predators choose similar hole locations to humans. Research into human shell-

41 beads recovered from archaeological contexts should take into account non-anthropogenic

42 factors, which can lead to more realistic scenarios of the cultural behaviours of prehistoric

43 people.

44 Keywords: Gastropoda, Scaphopoda, Bivalvia, shell beads, interspecies interactions,

45 jewellery, predators.

46 INTRODUCTION

47 The adornments of prehistoric people play an important role in our understanding

48 of the evolution of human behaviour (Bednarik, 2001; Gutiérrez-Zugasti et al., 2013)

49 because they can indicate evolutionary changes in the cognitive and linguistic abilities of

50 early humans (Vanhaeren & d’Errico, 2006; Schick & Toth, 2013; Stiner, 2014). These

51 findings help anthropologists to construct a picture of the life of prehistoric human groups,

52 and can give insights into their social status (Bednarik, 1998; Stiner, 1999; Vanhaeren &

53 d’Errico, 2005), group membership, age or marital status (Kuhn et al., 2001). Personal

54 adornments made from molluscs survive well in most sedimentary contexts (Bar-Yosef-

55 Mayer & Beyin, 2009) and can be interpreted in a various ways, depending on the context

56 of the find. Usually the deposits are associated with graves (Vanhaeren et al., 2004;

57 Vanhaeren & d’Errico, 2005), human made-shelters (Kuhn et al., 2001) and hearths (Douka

58 et al., 2014). Some of the earliest forms of body adornments are in the form of shells beads

59 and date back to ~75 Kya (Henshilwood et al., 2004)and ~82 Kya (Bouzouggar et al.,

60 2007), possibly even 100-130 Kya (Vanhaereny et al., 2006)or earlier (Bednarik, 2015).

61 However, others researchers argue that this “modern behaviour” was probably established

62 much earlier and taphonomic processes mean that shell beads are simply the earliest


63 material associated with body adornment that have survived (Bowdler & Mellars, 1990;

64 Noble & Davidson, 1996; Botha, 2008, 2010).

65 Generally shell artefacts are limited to marine molluscs found in habitation levels

66 (Vanhaeren & d’Errico, 2006), while evidence of land and freshwater gastropods are more

67 rare [e.g., El Cuco, 23-34 Kya(Gutiérrez-Zugasti et al., 2013)]. The majority of shell artefacts

68 are known from the Palaeolithic and Neolithic sites in Levant, South Africa, Europe, North

69 America and Asia. Interestingly, while early dated shell remains from occupation layers in

70 Africa are only associated with anatomically modern humans (AMHs), molluscs recovered

71 from Middle Palaeolithic (MP) layers in Europe are mainly related to Neanderthal

72 occupations(Gutiérrez-Zugasti et al., 2013). Although some researchers have argued that

73 strategies for exploiting coastal resources do not differ between European Neanderthals

74 and AMHs in Africa during the MP and MSA (Stringer & Barton, 2008), others interpret the

75 evidence differently, with Neanderthals exploiting marine environments infrequently

76 compared to AMH (Shipman, 2015).

77 Researchers make detailed analyses of adornments, radiometric dates and

78 stratigraphic information to explain innovations in shell beads, as well as the spread of

79 cultural traditions (Kuhn et al., 2001). Piercings in shells carry indications as to the

80 placement, the rigidity or flexibility of the ornament within a larger assemblage and, in

81 relation to the substrate or cord, the direction the traction was exerted (Cristiani, 2012).

82 Based upon this kind of painstaking evidence-gathering, experts make judgements as to

83 whether perforations in shells from archaeological sites are anthropogenic in origin or

84 formed by natural processes (d’Errico et al., 2005), such as those made by hole-drilling

85 animals (predators, parasites; (Kowalewski, 2004; Li, Young & Zhan, 2011)). While the


86 location of the perforation is only part of raft of evidence that indicates an operational

87 chain (starting with the collection of the raw material, followed by the manufacture and

88 use, and ending with its discard), it is subject to bias. For example, it is believed that the

89 anatomical locations of holes drilled in shells made by humans exhibit low variability

90 whereas holes made by non-human animals yield more random perforations (Bouzouggar

91 et al., 2007).

92 In the Palaeolithic, beads made from molluscs were desirable for making larger

93 compositions, which vary due to decorative traditions of prehistoric human groups (Stiner,

94 2014). Although there is only rare evidence of arrangements of Palaeolithic ornaments, we

95 might expect beads strung in different arrangements to require differently placed piercings

96 for the shells to hang correctly (according to design). The evidence indicates that

97 prehistoric people were adepts at drilling holes in shells, but also made use of natural

98 perforations when possible(Bar-Yosef Mayer, Vandermeersch & Bar-Yosef, 2009).

99 Preferences for shell beads appear to transcend temporal and regional variations in the

100 Palaeolithic andpeople appear to have preferred shells with vivid markings (e.g., beads

101 made from different mollusc taxa which vary due to shape, size, shell thickness and

102 ornament placement (Stiner, 2014)). All these factors may influenced the attraction of

103 humans to the particularly species of mollusc and likely increased the variability of hole

104 placement in shells, hence, any assumption that hole location in shells made by humans

105 should be consistent, may not be borne out (d’Errico et al., 2005; Kuhn et al., 2009; Stiner,

106 Kuhn & Güleç, 2013).

107 Experiments are used to understand the shell anatomy (i.e., mineralogy and

108 structure) and the processes involved in the production of piercings. The microscopy can


109 provide evidence of the shape of the tools used for piercing shells, as well as other tell-tale

110 signs of human activity(d’Errico et al., 2005; Nigra & Arnold, 2013). For example, piercings

111 will be examined for presence of residues, such as ochre, and any polishing or other

112 indications of use wear, such as striations and notches close to the perforation, that might

113 indicate the presence of a suspension system. Similarly, microscopic analyses of naturally

114 made holes in molluscs are insightful (e.g.,(Li, Young & Zhan, 2011; Gorzelak et al., 2013)).

115 Animal predation on mollusc populations is a widespread phenomenon

116 (e.g.(Quensen & DS, 1997; Rosin et al., 2011)), suggesting that predator-prey interaction

117 may be underestimated when analysing holes in shells (Kowalewski, 2004). Such

118 behaviour has been observed for many drilling predators, such as naticids, muricids,

119 octopuses, crabs and birds (Grey, Lelievre & Boulding, 2005; GREY, 2005; Rosin et al., 2011;

120 Li, Young & Zhan, 2011). Moreover, predators can be specific in where they attack molluscs

121 because shell strength and location of internal organs can be important traits used in prey

122 selection (Hagadorn & Boyajian, 1997; Dodge & D, 1999; Rosin et al., 2013). For instance,

123 birds usually choose the part of the shell near the apex, which is less resistant to crushing

124 than, for example, the labium (Rosin et al., 2013). In contrast, octopuses and predatory

125 snails choose areas close to the umbo, which tends to be thicker than other areas of the

126 shell, but is near the heart. This strategy appears to be a compromise between drilling time

127 and effectiveness of the injected toxin (Dodge & D, 1999).

128 Furthermore, recent evidence indicates that holes in shells can be made without the

129 action of predators or humans. In a set of shell-rolling experiments that mimic the action of

130 the waves and tides, Gorzelak and co-workers (Gorzelak et al., 2013) showed that abrasive


131 action of rolling shells together can create holes in predictable locations that mimic the

132 holes of predators. It is therefore possible that abrasion may also mimic human actions.

133 Considering that pierced beads can not only be produced by humans for making

134 Palaeolithic jewellery, but also by natural processes (e.g., predator-prey interaction, natural

135 erosion), we decided to examine following two assumptions: 1) variability of hole location

136 in shell beads made by human is underestimated; 2) variation of hole location in shells

137 made by non-human animals or natural processes is overestimated. Evidence that outlines

138 the contribution by natural processes to shell piercing could provide a valuable resource

139 for future research on human behavioural evolution.

140 MATERIALS & METHODS

141 To assess shell and hole placement variation in archaeological findings we analysed

142 material within 260 publications such as articles, PhD theses and chapters in books about

143 malacological findings in archaeological context. We searched for these using Google

144 Scholar and SCOPUS, and using keywords such as: shell beads, pierced shells, beads, shells,

145 mollusc, gastropoda, bivalves, pendant, shell midden, ornaments, shell ornaments,

146 predator. Once the publications were selected, their references (backward search) and

147 citation records (forward search) were analysed to find other articles that could provide

148 relevant data (Fig 1). Gathered literature was published between the 1966 and first quarter

149 of the 2015. All papers were analysed in respect of searching for the following information:

150 1) mollusc species from which the shell beads were made; 2) name and country of the

151 archaeological site where shell beads were founded; 3) from which period the shell beads

152 come from; 4) hole location in shell beads; 5) predators who made holes in shells; 6)

153 gastropods habitats. We used the identifications made by the authors of the published


154 papers as to whether the remains were culturally significant (i.e., shell beads) or were

155 naturally perforated holes.

156 We created a classification of hole location in shell beads which helped us analyse

157 gathered literature (Fig 2). As an example of shell shape we used two species from genus

158 Nassarius and Glycymeris, which are relatively common in the archaeological findings.

159 Then, we assessed hole location for shell beads found in the literature, based on figures

160 attached to each article. For this purpose we used 77 papers from 260 gathered scientific

161 articles. Articles were rejected from the study if they did not contain figures of shell beads.

162 If shells were described in more than one scientific publication, they were assessed only

163 once. Our estimation was based on the figures within the publications, thus the analysis is

164 not quantitative, but qualitative. We then searched for information on hole-making

165 predators for each mollusc species recovered from each archaeological site with shell

166 beads.

167 Next, we analysed the types of hole location in shell beads for normality by using the

168 Shapiro Wilk test, and for homogeneity of variances by using Levene’s test (all P>0.05).

169 Analysed material did not fulfil the criterion of normality, thus data were log or square root

170 transformed and tested again for normality(Sokal & Rohlf, 1981). After these

171 transformations the data were still not normally distributed; we therefore used

172 nonparametric Kruskal-Wallis tests with post-hoc Dunn-Bonferroni to analyse variation

173 and differences in number of hole location types among molluscs classes within

174 archaeological sites.

175 In order to analyse the strength of the differences between groups we calculated

176 size of the effect using the following equation: d=(M1-M2)/SDpooled, where d is the Cohen-d


177 index, M1 is the mean of the first group, M2 is the mean of the second group and SDpooled is

178 the pooled standard deviation (Cohen, 1988). To interpret d values we used the following

179 criteria for effect sizes: d≥0.1, small; d≥0.3, medium; d≥0.5, large(Cohen, 1988).

180 RESULTS

181 We found 46 taxa of Mollusca in the gathered literature (Table 1). The most

182 numerous and diverse class was Gastropoda (41 taxa). Only two taxa belong to the class of

183 Scaphopoda and four taxa to Bivalvia. Usually, a genus is represented by one species except

184 for the genera Littorina, Nassarius and Neritina. Usually, a genus was represented by only

185 one species. The number of hole location types was diverse amongst molluscs species (Fig

186 3). All shells with holes of Antalis sp. and Dentalium sp. (Scaphopoda class) were classified

187 to the same type (number 5; Fig 2). Bivalvia was the most diverse class in terms of hole

188 location. Only taxa Glycymeris sp. and Chlamys sp. had one type of hole location (number 9),

189 however, Chlamys sp. was only noted at one archaeological site. Shells of Acanthocardia

190 tuberculata (9 and 10 type of hole location) and Pecten sp. (4 and 10 type of hole location)

191 had more than one type of hole location.

192 Columbella sp. had seven types of hole location (2, 3, 4, 6, 7, 8 and 11 type of hole

193 location) and was the most variated taxa from the Gastropoda class among all

194 archaeological sites, while shells of Nassarius gibbosulus from Üçağızlı (Turkey) was the

195 most diverse in terms of hole location in shells from one archaeological site (5 types of hole

196 location: 1, 2, 3, 4 and 7). Twenty-five taxa exhibited more than one type of hole location in

197 their shells, while shells from 21 taxa were classified to only one type. However, 16 taxa

198 from the latter group were recovered at one archaeological site alone.

199 All taxa from the class Bivalvia have predators that make holes in shells (Table 2).


200 Species from this mollusc class are usually associated with more than one predator species

201 (except Acanthocardia tuberculata which has only one predator, Naticarius hebraeus).

202 Gastropoda, is the class with the most numerous predators (seven species), then the

203 Asteroidea (three species) and then Malacostraca with only one predator (Table 2). In the

204 Scaphopoda class only one taxon has predators that are able to bore holes in the shells. The

205 Dentalium sp. can be attacked by five species of the Gastropoda. Almost all taxa from the

206 Gastropoda class have non-human predators that attack the prey by drilling holes in the

207 mollusc’s shell. Gastropod predators belong to classes such as: Gastropoda (23 taxa),

208 Malacostraca (13 taxa), Cephalopoda (4 taxa), Asteroidea (4 taxa), Aves (5 taxa) and

209 Polychaeta (1 taxon). The Patella vulgata is the most diverse and numerous taxon in terms

210 of drilling predators (Table 2).

211 There were significant differences in number of hole location types among molluscs

212 classes within archaeological sites (Table 3). Gastropoda showed significantly higher mean

213 number of hole types than Scaphopoda and Bivalvia (Table 4).

214 DISCUSSION

215 Based on classifications of shell piercings in the literature, our results indicate that

216 variation in hole-location on shells reported to have been pierced by humans is greater

217 than variation in the placement of holes reported to have been created by natural

218 processes. Consequently, these patterns are opposite to those that might be expected, and

219 emphasise the importance of the battery of tests currently used to identify whether a

220 piercing in a shell is made naturally or an anthropogenic modification. Moreover, we found

221 that from among molluscs, Gastropoda was the most numerous and varied class in terms of

222 hole location types and species used as shell beads.


223 Some researchers claim that shell beads from the Middle and Lower Palaeolithic

224 could be perforated by natural process. For example, Bednarik(Bednarik, 2015) proposed

225 that predators and parasitic organisms commonly perforate mollusc shells (e.g. (Kelley &

226 Hansen, 1993; Li, Young & Zhan, 2011; Gorzelak et al., 2013)). Hahn (Hahn, 1972)

227 emphasized that indicators that signal human-made holes in shells from Aurignacian sites

228 such as Krems-Hundssteig, Willendorf, Kostienki 1 and Sjuren, are not always present and

229 could be made by predators. Beads made from shells with holes made by natural processes

230 are known, for example, the perforated bead of Antalis sp. from the Early Upper Paleolithic

231 site in El Cuco (Spain; (Gutiérrez-Zugasti et al., 2013)).

232 Researchers are at risk of discarding the possibility that perforations in shells are

233 made by predators due to the decreased likelihood of holes in suitable locations for

234 threating (d’Errico et al., 2005; Bouzouggar et al., 2007). There is evidence that coastal

235 environments were richer in prehistoric times and molluscs population more resistant to

236 overexploitation by humans (Bicho & Haws, 2008). Bicho and Haws (Bicho & Haws, 2008)

237 have suggested that the larger biomass of molluscs in the Palaeolithic likely meant that

238 mollusc gathering formed part of hunter-gatherers’ regular foraging behaviour in Portugal.

239 Therefore, the likelihood of finding Gastropods and Bivalves shells with naturally made

240 holes in suitable locations in the Palaeolithic could have been higher, possibly ranging from

241 2.8% to 50% (Taraschewski & Paperna, 1982; Hagadorn & Boyajian, 1997; Zagyvai &

242 Demeter, 2008; Sawyer, 2010). Importantly, all the Bivalves and almost all the Gastropods

243 found at archaeological sites were associated with several drilling predators (e.g. Fig 4, Fig

244 5, Fig 6).

245 Researchers often use morphological differences to distinguishing holes made by


246 human and predators. Predator-made holes are mostly made by chemical processes and

247 tend to be round in outline, while shells pierced by humans have elliptical or irregular

248 outlines (Stiner, 1999; Komšo & Vukosavljević, 2011). However, many predators form

249 holes in their prey which range in shape from nearly perfect circles to ellipsoids (Zagyvai &

250 Demeter, 2008). For example, muricids, naticids and cephalopods use their radula to bore

251 into mollusc shells and they can adapt the size and shape of the drill hole to the

252 morphology of their prey, as a result, bore holes can differ in shape(Walker & Brett, 2002).

253 Birds often take less time make holes in shells, and holes are often ellipsoid because of the

254 morphology of their beaks (Rosin et al., 2013). Bird beaks can also cause cracks and chips

255 the shells that mimic stone tool use (Ingolfsson & Estrella, 1978; Shumaker, Walkup &

256 Beck, 2011). Thus, close scrutiny of the tell-tale signs of anthropogenic manipulation is

257 critical. Striations indicating the direction of rotary drilling can provide a clear indication of

258 human actions (Zilhão et al., 2010).

259 According to Stiner (Stiner, 1999) the anatomical location of holes drilled by

260 prehistoric human groups is quite consistent, while predator-made holes are randomly

261 placed. However, this statement can be questioned; according to our results, hole location

262 in shell beads reported to have been pierced by humans, can be highly variable. Many

263 archaeological sites such as: Pupićina Cave (Komšo & Vukosavljević, 2011), Zala cave

264 (Komšo & Vukosavljević, 2011), Hautes-Pyrénées(Cattelain, Bozet & Di Stazio, 2012), El

265 Cuco(Gutiérrez-Zugasti et al., 2013) or Grotte des Pigeons (Elias, 2012) contained shells

266 beads with holes in a variety anatomical locations. The most variable holes in shells were

267 found at the site of Üçağizli (Turkey) dated to the Upper Paleolithic(Stiner, Kuhn & Güleç,

268 2013). Moreover, variation is observed among sites and within the same snail species like


269 in Columbella or Cyclope sp. Our assessment of hole location was based on figures from the

270 published literature only, it is therefore possible that the true level of this variation from

271 beads recovered from archaeological sites might be bigger (i.e., in collections not described

272 in the literature).

273 The reason for non-random placement of drilled holes by predators can be

274 associated with shell thickness of prey which varies across the body due to differential

275 predatory pressure on snails (Rosin et al., 2013). Predatory gastropods spend three to

276 twenty minutes locating a drilling site on their prey’s shell surface and, once a location is

277 made, a hole may take from several hours to several days depending on shell-thickness

278 (Hagadorn & Boyajian, 1997). Additionally, in molluscs with ornamental shells, up to five

279 times more force can be required to make a hole (Dalziel & Boulding, 2005). The variation

280 in shell thickness among and within molluscs is likely to impact piercing behaviours in both

281 humans and predators, and may be linked to why piercings in shells most often occur near

282 the labium and in the centre. This hole placement correspond to type number such as: 1, 2,

283 3 and 4. To be clear, our results show that Gastropod and Bivalve predators choose similar

284 hole locations to humans. Consequently, statements that indicate human agencies are

285 largely responsible for perforation in shells because perforations are rare in nature, can be

286 questioned (d’Errico et al., 2005).

287 Scaphopoda was the least diverse molluscs class in terms of hole location diversity.

288 All shell beads belonging to this class have two anatomical holes, one in the apex and

289 second one in the base of the shell. The anterior (apex) hole is for the burrowing foot and

290 captacula to protrude, while the posterior (base) hole is responsible for respiratory

291 currents-pass (Reynolds, 2002). As such, prehistoric humans did not have to drill holes in


292 Scaphopoda’s shell to make beads due to the elongated shape of the shells and two natural

293 holes. Drillhole predation frequency in Dentalium sp. is very low (0.9%) (Yochelson,

294 Dockery & Wolf, 1983). According to Klompmaker (2011), completely drilled holes in the

295 shells of Scaphopoda from the Miocene were usually made by naticids (predatory sea

296 snails) and were not randomly placed, but were frequently located in the middle and the

297 thickest part of the shell.

298 Bivalvia was found to be slightly more numerous and diverse than Scaphopoda in

299 terms of hole location. Type number 9 was the most frequent hole location in this class

300 (hole placed in the middle of the umbo) and rare types were number 10 (hole near the

301 umbo but closer to the centre of the shell) and 4 (hole at the centre of the shell). Bivalve

302 predators are also diverse and belong to Asteroidea, Gastropoda or Malacostraca. This

303 variation of predators might have influenced the frequency of drillhole predation in the

304 Bivalvia from the Miocene, which is much greater than in the Scaphopoda, ranging from

305 8.6% to 34.1% depending on the region (Sawyer, 2010). For example, variation in hole

306 location is very low in Chlamys sp., with naticids and muricids usually choosing the region

307 near the adductor muscle. This strategy of selection of a drill location may facilitate access

308 to the viscera (Guerrero & Reyment, 1988). Similar results were obtained by Amano

309 (2006), who showed that in the early and middle Pleistocene Glycymeris sp. were drilled by

310 predators usually close to the umbo or near the centre. These locations correspond to hole

311 location type number 9, 10 and 4 on shells recovered at the archaeological sites and

312 proposed to have been drilled by humans. Gastropoda was the most numerous and varied

313 class in terms of species used as shell beads and hole location types. However, we assume

314 that actual variation may be greater than obtained in our study because, in our dataset, one


315 archaeological site (Riparo Mochi) yielded a greater variety of taxa (13 taxa), compared to

316 other sites.

317 Furthermore, evidence suggests that bivalves were rarely used as beads due to size

318 and weight and most perforations are attributed to predation or taphonomy(Carter, 2008).

319 In most Palaeolithic sites the presence of bivalves is attributed to utilitarian purposes, and

320 not use as beads (Harper, 2005; Rogalla & Amler, 2007; Douka et al., 2014). Zilhão et al.

321 (41) suggest that, especially for bivalves of the most common taxa in archaeological

322 assemblages (e.g., Cerastoderma, Acanthocardia and Glycymeris sp.,) natural, as opposed to

323 anthropogenic processes, should be the null hypothesis for the origin of the perforations.

324 This can only be challenged if evidence exists to support that (i) a tool was involved in the

325 perforation; (ii) the weathering stage and perforation patterns do not agree with those

326 seen in natural death assemblages, or (iii) the hole is associated with artificial modification

327 of the shell's geometry.

328 Almost all Gastropoda taxa are vulnerable to predator-prey interaction. In most

329 cases, the drilling predators belong to the class Gastropoda (23 taxa), or Malacostraca (13

330 taxa). The rest belong to Aves, Asteroidea, Cephalopoda and Polychaeta. Drill frequency in

331 Gastropods is slightly greater than in Bivalves and ranges from 2.8% to 50.0% depending

332 on the provenance (Taraschewski & Paperna, 1982; Hagadorn & Boyajian, 1997; Zagyvai &

333 Demeter, 2008; Sawyer, 2010). Moreover, results obtained by researchers show that the

334 drill site selected by predators of Gastropod are not randomly selected (Arpad, 1993;

335 Zagyvai & Demeter, 2008). For example, in Neritina picta, access to the apex of the shell

336 was a preferred as a strategic location by predators belonging to Asteroidea, Miricidae, or

337 Polychaeta(Zagyvai & Demeter, 2008). In turn, Theodoxus sp. usually exhibit a muricid


338 (predatory sea snail) borehole often located close to the umbilicus (Arpad, 1993).

339 Shell beads are usually limited to marine molluscs. Terrestrial snails are very rare in

340 archaeological findings and, when recovered, are usually used in paleoenvironmental

341 reconstruction (Christensen & Kirch, 1981). Marine molluscs generally occur in more dense

342 colonies than terrestrial snails (Barnes & Hughes, 1999) thus, gathering molluscs is likely

343 to have been more profitable. Although it has been proposed that some species of seawater

344 snails could have been collected for their rarity and bright colours(Gutiérrez-Zugasti et al.,

345 2013). However, coastal productivity seasonally fluctuates (Dusseldorp, 2013), making the

346 location of some marine species only accessible to human foragers during spring low tides

347 (Kyriacou et al., 2015). In contrast, terrestrial snails may have been easier to obtain, but

348 have thinner shell. This structural difference is a morphological adaptation which allows

349 land snails to inhabit dryer environments (Machin, 1967). Shell morphology of terrestrial

350 snails makes it more difficult to drill a hole in the shell without inflicting catastrophic

351 damage. So although it is possible that terrestrial snails could have been incorporated into

352 jewellery, the fragile nature of their shells, especially after being damaged by piercing, is

353 likely to have left them more susceptible to taphonomic processes and decay.

354 According to the very low drill frequency in Scaphopoda, and the fact that holes

355 made by predators do not correspond to locations chosen by humans, the likelihood of

356 using natural perforated shells from this class, as shell bead by prehistoric human groups,

357 is remote. In turn, natural perforated Bivalvia and Gastropoda seem to be a better source of

358 beads. Variation in hole location in bivalves and gastropods, reported to have been pierced

359 by non-human animals, is overestimated. Moreover, the likelihood of finding shells with

360 holes made by predators in suitable locations is probably higher than researchers believe.


361 Specifically, all Bivalves and almost all Gastropods, found at archaeological site are

362 associated with drilling predators. Additionally, the high drill frequency suggests that the

363 Gastropoda and the Bivalvia may have been a valuable source for jewellery production.

364 Shells with holes can be produced by human-action in order to make Palaeolithic

365 jewellery, but holes can also be created by natural processes, such as predator-prey

366 interactions. Previous researchers have argued that holes in shells made by predators

367 would vary more than holes made by humans, and use this as a means of identifying

368 natural-made from human-made piercings. Brumm and Moore’s (Brumm & Moore, 2005)

369 claim that, “the absence of evidence for repeated pattering in symbolic behaviour cannot be

370 itself taken as evidence for the absence of behavioural modernity among past people”. It is

371 possible that before tools were used to bore holes, finding shells with holes in favourable

372 positions for threading into ornate jewellery may have increased their importance/value.

373 Similar examples could have been fish vertebrae or crinoid discs with natural apertures for

374 threading (e.g., see (Bednarik, 2015)). This study highlights how natural piercings and their

375 placement on the shell can mimic human activity and, as such, emphasises the importance

376 of other methods for investigating the tell-tale signs of anthropogenic manipulation. This,

377 in turn, should help to engender more realistic scenarios of the social and cultural

378 expressions of prehistoric people.

379 ACKNOWLEDGMENTS

380 We thank Mick Vernon and David Bell for their careful reading and helpful

381 comments on the manuscript.

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549


1PRISMA flow diagram



2Graphical representation of hole location types in shells.


3Number of hole location types in shell beads for molluscs species.

Star indicates taxon which was found at one archaeological site.


4Shell from the genus Venus with hole made by predator.


5Shell from the genus Bolinus with hole made by marine predator.


6Helix pomatia with a hole made by a bird.


Table 1(on next page)

Hole assessment in shell beads from the archaeological sites.


Mollusc species Country Site Date References Hole type

Balauzerie 40.000-28.000 BP (BARGE, 1983)

9

Régismont 40.000-28.000 BP (Fiocchi, 1998)

9

France

Tournal 40.000-28.000 BP (Fiocchi, 1998)

9

Fanciulli 40.000-28.000 BP (BARGE, 1983)

9Italy

RiparoMochi 34.870-32.280 BP (BARGE, 1983)

9

Spain Cueva de los Aviones 50.000 BP (Zilhão et al., 2010; Vanhaeren & d’Errico, 2011)

9

Acanthocardia tuberculata

Turkey Üçağızlı 41.000-39.000 BC (Stiner, Kuhn & Güleç, 2013)

10

Portugal Vale Boi 20.570-18.859 BP (Tátá et al., 2014)

5Antalis sp.

Spain El Cuco 29.000-22.000 ka (Gutiérrez-Zugasti et al., 2013)

5

Buccinum undatum

Italy Riparo Tagliente 14.600-11.5000 BC (Fontana et al., 2009)

4

Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998)

7Cerithiumsp Italy

Riparo Tagliente 14.600-11.5000 BC (Fontana et al., 2009)

6

Chlamyssp Italy Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998)

10

Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 6Clanculus corralinus Italy Cala 40.000-28.000 BP (Fiocchi,

1998)4

Austria Krems-Hundsteig 40,000-28,000 BP (Fiocchi, 1998; Wild et al., 2008)

2,3

Pupićina Cave 11.070-10.500 BP (Komšo & Vukosavljević, 2011)

3, 4, 7Croatia

Zala cave 11.070-10.500 BP (Komšo & Vukosavljević, 2011)

3, 4, 7

Greece Klisoura 41.000-38.000 BP (Kozłowski, 1996)

4

Cala 40.000-28.000 BP (Fiocchi, 1998)

2

Grotta di Pozzo 85.000-60.000 BP (Mussi et al., 2000)

4

Riparo Biarzo 12.000-5.600 BP (Cristiani, 2012)

3, 4, 6

Columbella

Italy

Riparo Tagliente 14.600-11.5000 BC (Fontana et 4


al., 2009)Ksar Akil 41.000-39.000 BC (Inizan, 1978;

Douka, 2013)7, 4, 3Near East

Sefunim 41.000-15.000 BP (Bar-Yosef, 1996)

4

Russia Kostienki 1 36.500-32.600 BP (Sinitsyn, 2003)

2

Spain Botiquería de Los Moros

6.000-4.000 BP (Álvarez-Fernández, 2010)

4, 8, 11

Pınarbaşı 8.5000-8.000 BC (Baysal, 2013)

8Turkey

Boncuklu Höyük 9.000-8.000 BC (Baysal, 2013)

8

Australia Mandu Mandu Creek rock-shelter

35.200-30.900 BP (Kate Morse, 1993; Balme & Morse, 2006)

8

Cala 40.000-28.000 BP (Fiocchi, 1998)

4Italy

Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998; Stiner, 1999)

2, 8

Conus sp


8

France Abri Peyrony 40.000-28.000 BP (Vanhaeren & d’Errico, 2011)

1, 3

Germany Andernach-Martinsberg

13.200-12.820 BP (Langley & Street, 2013)

1, 3

Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 6Riparo Biarzo 9.000-7.000 BP (Cristiani,

2012)1

Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998; Stiner, 1999)

1

Cyclope sp

Italy


1

Krems-Hundsteig 40.000-28.000 BP (Fiocchi, 1998; Neugebauer-Maresch, 1999)

5

Langmannersdorf 40.000-28.000 BP (Hahn, 1972) 5Senftenberg 40.000-28.000 BP (Hahn, 1972) 5

Austria

Willendorf 28.000-22.000 BP (Kozłowski, 1996)

5

Abri Peyrony 40.000-28.000 BP (Vanhaeren & d’Errico, 2011)

5

Blanchard 34.000-32.000 BP (Taborin, 1993)

5

Dentalium sp.

France

Caminade Est 37.200-32.140 BP (Taborin, 5


1993)Castanet 34.000-32.000 BP (Taborin,

1993)5

Cellier 40.000-28.000 BP (Taborin, 1993)

5

Laouza 40.000-28.000 BP (Fiocchi, 1998)

5

Pecheurs 28.000-22.000 BP (BARGE, 1983)

5

Rochette 40.000-28.000 BP (Movius, 1995)

5

Rothschild 40.000-28.000 BP (Zilhão, 2011) 5Saint-Césare 40.900-36.300 BP (d’Errico et

al., 1998)5

Saint-Germain-la-Rivière

15.570 BP (Vanhaeren & d’Errico, 2005)

5

Salpetriere 22.000-18.000 BP (BARGE, 1983)

5

Tournal 40.000-28.000 BP (Fiocchi, 1998)

5

Tuto de Camalhot 40.000-28.000 BP (Taborin, 1993)

5

Vachons 40.000-28.000 BP (Taborin, 1993)

5

Greece Klisoura 41.000-38.000 BP (Kozłowski, 1996)

5

Cala 40.000-28.000 BP (Fiocchi, 1998)

5

Grotta del Cavallo 31.000-21.000 BP (Cesnola & Mallegni, 1996)

5

Grotta di Pozzo 85.000-60.000 BP (Mussi et al., 2000)

5

Fanciulli 40.000-28.000 BP (BARGE, 1983)

5

Fumane 41.000-38.000 BP (Fiocchi, 1998)

5


5

Italy


5

Hayonim (Belfer-Cohen & Hovers, 2010)

5

KsarAkil 41.000-39.000 BP (Inizan, 1978) 5

Near East

Yabrud (Bar-Yosef, 1996)

5

Beneito 40.000-28.000 BP (Soler-Major, 2001)

5

Cova del Parco 13.175-12.460 BP (Mangado et al., 2010)

5

Spain

L’Arbreda 37.340-35.480 BP (Carbonell, 1996)

5


Çatalhöyük 7.200-6.000 BP (Bar-Yosef Mayer, Gümüş & Islamoǧlu, 2010)

5

Pınarbaşı 8.5000-8.000 BC (Baysal, 2013)

5

Turkey

Boncuklu Höyük 9.000-8.000 BC (Baysal, 2013)

5

Engina mendicaria

Eritrea Red Sea Coast 7.330-5.385 BP (Bar-Yosef-Mayer & Beyin, 2009)

4

Euthria cornea Turkey Üçağızlı 41.000-39.000 BC (Stiner, Kuhn & Güleç, 2013)

4

Gibbula sp. Turkey Üçağızlı 41.000-39.000 BC (Stiner, Kuhn & Güleç, 2013)

4

France Figuier 40.000-28.000 BP (Taborin, 1993)

9

Israel Qafzeh cave 90.000 y BP (Taborin, 1993)

9

Fumane 41.000-38.000 BP (Vanhaeren & d’Errico, 2011)

9Italy

Grotta del Cavallo 31.000-21.000 BP (Cesnola & Mallegni, 1996)

9

Cueva de los Aviones 50.000 BP (Zilhão et al., 2010)

9

Glycymeris sp.

Spain

Cueva Morín 18.000-10.000 ka (Álvarez Fernández, 2008)

9

Germany Andernach-Martinsberg

13.200-12.820 BP (Álvarez-Fernandez, 2009; Langley & Street, 2013)

1, 3, 4

Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 6

Homalopoma sanguineum

Spain Cova del Parco 13.175-12.460 BP (Mangado et al., 2010)

11

Pupićina Cave 11.070-10.500 BP (Komšo & Vukosavljević, 2011)

6Croatia

Zala cave 11.070-10.500 BP (Komšo & Vukosavljević, 2011)

6

Lithoglyphus sp.

Italy Riparo Biarzo 12.000-7.000 BP (Cristiani, 2012)

3, 7

Littorina fabalis Spain Los Canes Mesolithic (Álvarez Fernández, 2008)

1

Littorina littorea Hautes-Pyrénées 21.000-10.000 BP (Cattelain, Bozet & Di

1, 2, 4


Stazio, 2012)Spain El Cuco 29.000-22.000 ka (Gutiérrez-

Zugasti et al., 2013)

2

France Gargas Cave 26.910-23.590 BP (Juan-Foucher & Foucher, 2008)

11

Spain Tito Bustillo 18.000-10.000 ka (Álvarez Fernández, 2008)

11

Hautes-Pyrénées 10.000-6.000 BP (Cattelain, Bozet & Di Stazio, 2012)

1, 2, 4France

Rothschild 40.000-28.000 BP (Zilhão, 2011) 1El Cuco 29.000-22.000 ka (Gutiérrez-

Zugasti et al., 2013)

2, 3

La Garma 29.000-22.000 ka (Álvarez Fernández, 2008)

3, 4, 8

Littorina obtusata

Spain

Maltravieso cave 40.000-10.000 BP (Rodríguez Hidalgo et al., 2010)

1

Italy Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998)

8

France Tuto de Camalhot 40.000-28.000 BP (Vezian & Vezian, 1966)

1, 4

Littorina sp.

South Africa

Sibudu Cave Middle Stone

70.000-60.000 BP (D’Errico, Vanhaeren & Wadley, 2008)

1, 3

Melanopsis sp. Turkey Üçağızlı 41.000-39.000 BC (Kuhn et al., 2009)

3

Mitrella corniculata

Italy RiparoMochi 34.870-32.280 BP (Kuhn & Stiner, 1998)

4

Monodonta sp. Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 4Nassarius circumcintus

Spain Moroccan cave 83.000-60.000 BP (Elias, 2012) 4

Nassarius corniculus


4

Algeria Oued Djebbanna 35.000 BP (Vanhaereny et al., 2006)

2

Blanchard 34.000-32.000 BP (Taborin, 1993)

4France

Rothschild 40.000-28.000 BP (Zilhão, 2011) 4Israel Skhul 110.000 BP (Vanhaereny

et al., 2006)1, 2


4Italy

Riparo Mochi 40.000-28.000 BP (Stiner, 1999) 4

Nassarius gibbosulus

Morocco Grotte des Contrebandiers

40.000-12.500 BP (Vanhaeren & d’Errico,

4


2011)Grotte des Pigeons, Taforalt

83.000-81.000 BP (Vanhaeren & d’Errico, 2011; Elias, 2012)

1, 3, 4, 6

KsarAkil 41.000-39.000 BP (Douka et al., 2013)

1,4 Near East

Sefunim 41.000-15.000 BP (Bar-Yosef, 1996)

4

Spain Moroccan cave 83.000-60.000 BP (Elias, 2012) 2, 3, 4Turkey Üçağızlı 41.000-39.000 BC (Kuhn et al.,

2001b; Stiner, Kuhn & Güleç, 2013)

1, 2, 3, 4, 7


2Nassarius incrassatus

Italy


4


4

Blombos Cave 78.000-75.600 BP (d’Errico et al., 2005; Vanhaeren et al., 2013)

3South Africa

Border Cave 44.000-22.000 BP (d’Errico et al., 2012)

3, 4, 11

Nassarius kraussianus

Spain La Garma A 29.000-22.000 ka (Álvarez Fernández, 2008)

2

France Rothschild 40.000-28.000 BP (Zilhão, 2011) 4Italy Fumane 41.000-38.000 BP (Vanhaeren &

d’Errico, 2011)

4Nassarius mutabilis


3

France Rothschild 40.000-28.000 BP (Zilhão, 2011) 3Russia Mezmaiskaya Cave 36.000-28.510 BP (Golovanova,

Liubov, Doronichev, Vladimir & Cleghorn, Naomi, 2010)

2, 4Nassarius reticulates

Spain Tito Bustillo 18.000-10.000 ka (Álvarez Fernández, 2008)

2

Italy Riparo Mochi 34.870-32.280 BP (Kuhn & Stiner, 1998)

4Nassarius sp.

Morocco Contrebandiers 40.000-12.500 BP (d’Errico & Vanhaeren, 2009)

4, 7

Natica sp. Italy Fumane 41.000-38.000 BP (Vanhaeren & d’Errico,

3


2011)Spain Cova del Parco 13.175-12.460 BP (Mangado et

al., 2010)3

Naticarius sp. Turkey Üçağızlı 41.000-39.000 BC (Stiner, Kuhn & Güleç, 2013)

1


1, 4Neritina fluviatilis

France

Rothschild 40.000-28.000 BP (Zilhao, 2010) 1France Hautes-Pyrénées 21.000-10.000 BP (Cattelain,

Bozet & Di Stazio, 2012)

2, 4

Abri Peyrony 40.000-28.000 BP (Vanhaeren & d’Errico, 2011)

1

Neritina picta

Gargas Cave 26.910-23.590 BP (Juan-Foucher & Foucher, 2008)

1


1


1

Rothschild 40.000-28.000 BP (Zilhao, 2010) 3, 4

France

Saint-Germain-la-Rivière


4

Nucella lapillus

Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 4Cueto de la Mina 18.000-10.000 ka (Álvarez

Fernández, 2008)

2

El Horno 18.000-10.000 ka (Álvarez Fernández, 2008)

2

Spain

La Garma A 29.000-22.000 ka (Álvarez Fernández, 2008; Avezuela Aristu & Álvarez Fernández, 2012)

4, 11

Ocinebrina edwardsii

Italy Riparo Mochi 34.870-32.280 BP (Stiner, 1999) 4Olivella biplicata USA Irvine 9.420-7.780 BP (Erlandson et

al., 2005)4, 5

Patella vulgata Spain Maltravieso cave 40.000-10.000 BP (Rodríguez Hidalgo et al., 2010)

10

Riparo Mochi 40.000-28.000 BP (Stiner, 1999) 4Pecten sp. FranceGargas Cave 26.910-23.590 BP (Juan- 10


Foucher & Foucher, 2008)

Persicula terveriana

Eritrea Red Sea Coast 7.330-5.385 BP (Bar-Yosef-Mayer & Beyin, 2009)

4

Pirenella plicata France Hautes-Pyrénées 21.000-10.000 BP (Cattelain, Bozet & Di Stazio, 2012)

2

Greece Klisoura 41.000-38.000 BP (Stiner, 2010) 6Theodoxus sp.Turkey Üçağızlı 41.000-39.000 BC (Kuhn et al.,

2001b)6


3


1

Rothschild 40.000-28.000 BP (Zilhao, 2010) 3Saint-Germain-la-Rivière


5


3

France

RiparoMochi 34.870-32.280 BP (Kuhn & Stiner, 1998; Stiner, 1999)

2, 3

Los Azules I 10.000-8.000 ka (Álvarez Fernández, 2008)

2

Trivia sp.

Spain

Los Canes 7.930-7.580 BP (Álvarez-Fernández, 2010)

3

Trophon muricatus

Russia Mezmaiskaya Cave 36.000-28.510 BP (Golovanova, Liubov, Doronichev, Vladimir & Cleghorn, Naomi, 2010)

11

Italy Cauna de Belvis 40.000-30.000 BP (Taborin, 1993)

1, 2

France Abri Peyrony 40.000-28.000 BP (Vanhaeren & d’Errico, 2011)

2,3

Turritella sp

Spain El Horno 18.000-10.000 ka (Álvarez Fernández, 2008)

2

1



Drilling predators for molluscs species used as a shell bead by prehistoric humangroups.


Molluscs specie Predators class Predator ReferencesAcanthocardia tuberculata Gastropoda Naticarius hebraeus (Calvet et al., 1992)Antalis sp. No dataBuccinum undatum Gastropoda Euspira macilenta (Sawyer, 2010)

Gastropoda Euspira macilentaCerithium sp.Malacostraca Callinectes danae

Eriphia gonagraMenippe node frons Panopeus occidentalis

(Turra, Denadai & Leite, 2005; Sawyer & Zuschin, 2010; Coleman, 2010; Gorman, Sikinger & Turra, 2015)

Asteroidea Pycnopodia helianthoidesChlamys sp.

Gastropoda Murex sp.Naticidae

(Guerrero & Reyment, 1988; Farren & Donovan, 2007; Chattopadhyay & Dutta, 2013)

Clanculus corralinus No dataColumbella Cephalopoda Octopus vulgaris (Mather & O’Dor,

1991)Conus sp. Gastropoda Euspira macilenta (Sawyer & Zuschin,

2010)Cyclope sp. Asteroidea Astropecten spp. (Baeta & Ramón,

2013)Dentalium sp. Gastropoda Euspira macilenta

Euspira obliquataNatica canrenaNeverita duplicata, Oichnus spp.,

(Yochelson, Dockery & Wolf, 1983; Sawyer & Zuschin, 2010; Li, Young & Zhan, 2011)

Engina mendicaria No dataEuthria cornea Cephalopoda Octopus vulgaris (NIXON &

MACONNACHIE, 1988)

Gibbula sp. Malacostrata Carcinus maenas (Mowles, Rundle & Cotton, 2011)

Gastropoda Cryptonatica spp., Euspira spp. Glossaulax spp.,

Glycymeris sp.

Malacostraca Cancer pagurus

(Ramsay, Richardson & Kaiser, 2001; AMANO, 2006; Sawyer & Zuschin, 2010)

Homalopoma sanguineum No dataLithoglyphus No dataLittorina fabalis No dataLittorina littorea Asteroidea Pisastero straceaus

Pycnopodia helianthoides(Harley et al., 2013)

Aves Calidris canutus (Alerstam, Gudmundsson & Johannesson, 1992)

Littorina obtusata

Malacostraca Carcinus maenas (Edgell et al., 2008; Edgell & Rochette, 2009)

Littorina sp. Malacostraca Carcinus maenas (Reimchen, 1982)Melanopsis sp. Gastropoda Gastropoda (Kowalewski, Michał

Domenech, Rosa Martinell, Jordi


Mancheno, 2009)Mitrella cornicula Gastropoda Euspira macilenta (Sawyer & Zuschin,

2010)Monodonta sp. Aves Haematopodidae

Laridae(Harris, 1984)

Malacostraca Carcinus maenas, Nassarius circumcintusNassarius corniculusNassarius gibbosulusNassarius incrassatusNassarius kraussianusNassarius mutabilisNassarius reticulatesNassarius sp.

Gastropoda Euspira macilentaLunatiaheros, Natica tecta

(Stenzler & Atema, 1977; Sawyer & Zuschin, 2010)

Natica sp. Gastropoda Euspira macilentaNaticidae, Muricidae

(Arua, 1989; Zlotnik, 2001; Sawyer, 2010; Das, Mondal & Bardhan, 2013)

Asteroidea Asterina sarasiniNaticarius sp.Gastropoda Euspira macilenta

(Sawyer, 2010)

Aves Gallinula chloropusNeritina fluviatilisMalacostraca Macrobrachium spp.

(Blanco-Libreros & Arroyave-Rincón, 2009)

Gastropoda ActeocinaMiricidae

Neritina picta

Polychaeta Polychaeta

(Zagyvai & Demeter, 2008)

Nucella lapillus No dataOcinebrina edwardsii No dataOlivella biplicata Gastropoda Naticidae (Chojnacki & LR,

2013)Aves Haematopus ostralegus

Cephalopoda Octopus vulgaris

Gastropoda Euspira macilenta

Patella vulgata

Malacostraca Cancer pagurus,Carcinus maenas,Necora puber, Pachygrapsus marmoratus,

(Coleman et al., 1999; Smith, 2003; Silva et al., 2008, 2010; Sawyer, 2010)

Asteroidea Asteria srubensMarthasterias glacialis

Pecten sp.

Gastropoda Euspira macilenta

(Sawyer, 2010; Magnesen & Redmond, 2011)

Persicula terveriana Cephalopoda Octopus insularis (Leite, Haimovici & Mather, 2009)

Pirenella plicata Gastropoda Naticidae (Taraschewski & Paperna, 1982)

Theodoxus sp. Gastropoda Muricidae (Arpad, 1993)Trivia sp. Asteroidea Asterina sarasini (Sadhukhan &

Raghunathan, 2013)Trophon muricatus Gastropoda Naticidae (Delance & Emig,

2004)Turritella sp. Gastropoda Euspira macilenta

Naticidae(Hagadorn & Boyajian, 1997;


Muricidae,Odostomia sp.

Filipescu & Popa, 2001; Sawyer, 2010)

1



Descriptive statistics and results of Kruskal-Wallis test for molluscs classes.

N – number of archaeological site with assessed shell beads in context of hole location, Mean

– mean number of hole location types at archaeological sites, SD – standard deviation, Min –

minimal number of hole location types at archaeological site, Max – maximal number of hole

location types at archaeological site, Χ2 result of Kruskal-Wallis test. Bold indicates significant

difference between classes (P<0.05).


Class N Mean SD Min Max Χ2 PScaphopoda 11 1.00 0.00 1 1

Bivalvia 40 1.00 0.00 1 1Gastropoda 48 2.67 1.56 1 7

53.758 0.000

1



Summary class differences for hole location of Scaphopoda, Bivalvia and Gastropoda.

Effect sizes (d) is calculated for post-hoc Donn-Bonferroni test. Bold indicates significant

difference between classes (P<0.05).


N P Post-hoc d

Scaphopoda vs. Bivalvia 51 1.000 0.000 0.00Scaphopoda vs. Gastropoda 88 0.000 -6.887 1.53Gastropoda vs. Bivalvia 63 0.000 -4.411 1.53

1


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A systematic review of natural processes in creating pierced shells ...

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