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Home , Aquatic ape hypothesis, Melanopsis, Mytilus (bivalve), Otala lactea, Phorcus turbinatus, Semicassis

Science Reviews 217 (2019) 284e296

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Quaternary Science Reviews

journal homepage: www.elsevier.com/locate/quascirev

Shellfish collection on the westernmost Mediterranean, Bajondillo cave (~160-35 cal kyr BP): A case of behavioral convergence?

Miguel Cortes-S anchez a, b, c, María D. Simon-Vallejo a, b, c, * Francisco J. Jimenez-Espejo d, e, , María del Carmen Lozano Francisco b, Jose Luis Vera-Pelaez b, Adolfo Maestro Gonzalez f, Arturo Morales-Muniz~ g a Departamento de Prehistoria y Arqueología, Facultad de Geografía e Historia, Universidad de Sevilla, C/. Dona~ María de Padilla, S/n, 41004, Sevilla, Spain b HUM-949 Group, Universidad de Sevilla, Spain c ICArEHB-Interdisciplinary Center for Archaeology and Evolution of Behaviour, Universidade Do Algarve, Portugal d Instituto Andaluz de Ciencias de La Tierra, CSIC-Univ. de Granada, Armilla, 18100, Spain e Japan Agency for Marine-Earth Science and Technology, Yokosuka, Japan f Instituto Geologico y Minero de Espana,~ Madrid, 28003, Spain g Laboratorio de Zooarqueología, Departamento de Biología, Universidad Autonoma de Madrid, Madrid, E-28049, Spain article info abstract

Article history: The Middle (MP) and Early Upper Paleolithic (EUP) evidences of shellfish collection on the southern Received 28 November 2018 Iberian site of Bajondillo cave are presented and compared with Westernmost Mediterranean archaeo- Received in revised form logical sites. The main feature is stasis for Mytilus galloprovincialis represents the dominant taxon during 4 February 2019 a ~120kyr temporal sequence. The second feature is the decrease of the shellfishing signal when site Accepted 4 February 2019 distance to the coast increases. The data reveal that shellfish collection was practiced during Marine Available online 19 February 2019 Isotopic Stage 4, a poorly documented stage in terms of aquatic . Striking is also that mollusc assemblages evidence an uninterrupted decreasing trend in terms of remains from the earliest to the Keywords: Pleistocene latest levels, in particular when H. sapiens replaced H. neanderthalensis. Although taxa of secondary Western Europe importance are too scarce to make reliable inferences, another difference between the MP and EUP Malacology collections is the substantial increase of infaunal bivalves in the latter cultural period. Warm and cold Shellfish harvesting water mollusc records match temperature rises and drops although the scarcity of data do not allow one Middle Paleolithic to proceed beyond qualitative statements. Likewise, the prevalence of fresh and brackish water mollusc Upper Paleolithic hint at a permanent presence of freshwater around the site at all times. When compared with assem- Geomorphology blages from the Alboran sea region (Westernmost Mediterranean Sea), the Bajondillo cave collections are Coastal remarkable for their abundance of . Comparison between Bajondillo cave and Pinnacle Point reveal that infaunal bivalve abundances in the South African site are far higher than those recorded in the MP levels, though not those from the EUP. Whether this feature hints at subtle differences existing between the collection of shellfish by H. sapiens and H. neanderthalensis,reflects a behavioral conver- gence between the two hominine lineages or represents an inherited cognitive trait from a common ancestor is an issue in need of further analysis. © 2019 Elsevier Ltd. All rights reserved.

1. Introduction the association of shellfish collection with a putative cognitive development taking place in anatomically modern (AMH) The origin and evolutionary implications of marine resource but presumably not in other hominines is one major reason that exploitation in hominins has been hotly debated for nearly three accounts for this (vid. e.g. Cunnane et al., 1993; Parkington, 2001; decades. Although the reasons for this controversy are manifold, Langdon, 2006; Foley and Lahr, 2014, Erlandson, 2017).

* Corresponding author. Instituto Andaluz de Ciencias de la Tierra, CSIC-Univ. de Granada, Avda. de las Palmeras nº4, Armilla, 18100, Spain. E-mail addresses: [email protected] (M. Cortes-S anchez), [email protected] (M.D. Simon-Vallejo), [email protected] (F.J. Jimenez-Espejo), gaiadidactica@gmail. com (M.C. Lozano Francisco), [email protected] (J.L. Vera-Pelaez), [email protected] (A. Maestro Gonzalez), [email protected] (A. Morales-Muniz).~ https://doi.org/10.1016/j.quascirev.2019.02.007 0277-3791/© 2019 Elsevier Ltd. All rights reserved. M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 285

Bajondillo cave has provided the earliest evidences of shellfish In terms of shellfishing activities, except for the temporal depth collection in the northern Hemisphere at ~160kyr BP, demon- that improved dating methods permitted, no attempt has been strating that a coastal was present in the Alboran Sea made to provide a unitary interpretative frame for the origins of (Westernmost Mediterranean sea, Fig. 1) at essentially the same shellfish collection in the Alboran sea region. For such reason, time that shellfish collection is documented in South Africa certain particulars of the shellfish collection practice, including (Pinnacle Point: ~164kyr BP) (Cortes-S anchez et al., 2008). A changes in sea level and temperature dictating accessibility to the remarkable difference between these two sites is that at Bajondillo resource, foraging range within tidal zones, substrate type where cave it was , not Anatomically Modern Humans shellfish are documented or dietary preferences will be attempted (AMH), the hominines carrying out such practice (Marean et al., in this paper. 2007; Cortes-S anchez et al., 2008; Jerardino and Marean, 2010). Cortes-S anchez et al. (2011) published the taxonomic compo- In view that systematic collection of shellfish has been increasingly sition of the mollusc assemblage from Marine Isotope Stage (MIS6) recognized as a crucial element of human adaptation to aquatic at Bajondillo cave (archaeological level Bj/19). In this new study we ecosystems and evidence associated with the appearance of include the mollusc assemblages from eight levels (Bj/19 to Bj/11) sapiens, the synchronicity of these findings raise the possibility of a that range from the Middle Paleolithic (MP) to the Early Upper behavioral convergence existing in the two hominine lineages Paleolithic (EUP). One major aim of our study is to contrast how, in triggered by causes yet to be determined. terms of taxa and taxonomic abundances, the shellfish assemblages Bajondillo cave harbors the most important record of shellfish of six levels compare with those generated by AMH (2 collection to date in the Alboran Sea but not the only one. The first levels). In so doing, data will be provided for shellfish collection evidences of shellfish collection in the area (Fig. 1) were reported in during MIS4 (~51e74kyr BP), a poorly documented isotopic stage in the 1924e1925 excavations of the Mousterian rock-shelter of terms of aquatic adaptations that corresponds with a marine Devil's Tower in Gibraltar on the basis of (sic.) “large quantities” of regression episode that lowered sea levels worldwide by marine shells that, unfortunately, were never described in detail some 70 m (vide Erlandson, 2017). (Fischer, 1928). The 1951e1954 excavations on the dune (outer zone) of Gorham's cave (Gibraltar) also reported an MIS5-MIS3 2. Geographic setting sequence featuring substantial numbers of molluscs (Baden- Powell, 1964). More recently, in Gibraltar, marine molluscs have Bajondillo cave is found on a sheltered where the rivers been reported on Vanguard's Cave (MIS3) and the inner portion of from the Sierra de Mijas drain (Figs. 2 and 3). As such, it qualifies as Gorham's Cave (MIS3), and also on the MIS8-MIS5 North African an inlet shielded from the open sea by a . The cave rock shelter at Benzú (Ceuta, Spain) and in two MIS3 sites on the features a sequence with 20 archaeological levels that range from coast of Malaga (Abrigo 3 and Nerja Cave) (Barton, 2000; Cortes- an advanced Middle Pleistocene to the early . Based on Sanchez et al., 2008; Ramos Fernandez et al., 2012; Fa et al., techno-cultural and chronological data (i.e. 48 14C/AMS, TL and U/ fi 2016). Hints of shell shing by Iberian Neanderthals do not seem Th dates from 6 different laboratories), this sequence documents to be restricted to the Alboran sea, as both in Murcia (SE Spain) and four chrono-cultural stages (Cortes-S anchez, 2007; Cortes-S anchez the Algarve (S. Portugal) sites exist that report evidences of such et al., 2011, 2019). These include a Middle Palaeolithic (MP: Bj/19- activity (see Cortes-Sanchez et al. (2011) with references therein). Bj/14, <162 and >43.1 cal kyr BP) that is the main focus in this

Fig. 1. Site location and Northern Westernmost Mediterranean region and archaeological sites mentioned in the text. P: location of Padul Lake (Camuera et al., 2018). O: location ODP site 977 (Martrat et al., 2007) 286 M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296

Fig. 2. Overview of the travertine perimeter within the Sierra de Mijas, Torremolinos springwater and Bajondillo cave. paper, along with an Upper Palaeolithic (Bj/13-Bj/6: ca. 43- increased aridity during past glacial periods (Camuera et al., 2018), 16 cal kyr BP), an Epipalaeolithic (Bj/5-Bj/4, ca. 16e7.5 cal kyr BP) freshwater availability would have been the major agent dictating and a Neolithic (Bj/3- Bj/0, ca. 7.5e4 cal kyr BP) (Fig. 4). the permanent occupation of any given site and, for coastal sites, The coastlines of the Alboran Sea feature a far from homo- the year-round availability of molluscs. geneous and quite discontinuous archaeological record. The The main hydrogeological feature of the western sector of the reason for this is a geo-morphological dynamism due, among Bay of Malaga is the aquifer associated with the marble deposits of others, to the pushing of the African plate against the European the Sierra de Mijas (Martín-Arias et al., 2018). The discharge of this plate (Hernandez-Molina et al., 2014). From a strictly tectonic system occurs at the zone of contact between the carbonate series standpoint, even within the northern Alboran Sea coast, one and the impermeable metapelitic rocks of the Malaguide forma- must set apart its most active sector, located on the Strait of tion (Figs. 2 and 3). Aquifer configuration prevents major in- Gibraltar, undergoing uplifting during most of the Pleistocene ferences from sea level fluctuations taking place during at least from the Western sector of the Bay of Malaga that has remained the last 200,000 years because these do not affect the location stable for the past 200,000 years (Zazo et al., 1999). Such stability between these two lithologies (Fig. 3). Aquifer water level evolu- allows one to not only properly analyze the effects of glacioeu- tion is determined by the quantity and distribution of rainfall static dynamics (Zazo et al., 2003) but also to expect the presence during the hydrological year (Martín-Arias et al., 2018). Travertine of essentially continuous, long-term archaeological sequences in deposits (50 m deep, Figs. 2 and 3)indicatethatfromthe this area, of which Bajondillo cave is a paradigmatic case, to advanced Middle Pleistocene onwards, the springs of Torremoli- emerge (Cortes-S anchez and Simon-Vallejo, 2007). Tectonic sta- nos are documented to have provided water all around Bajondillo bility further allows one to apply the coastal level global model to cave. These lithochemical deposits underwent karstification Bajondillo cave in order to determine the mean distance to the events, of which Bajondillo cave constitutes one of the most shore for any of its MP/UP levels (Siddall et al., 2003; Bintanja spectacular cases (Cortes-S anchez and Simon-Vallejo, 2007). At et al., 2005). present, the travertine building lies below the piezometric level as Freshwater availability is a major agent in order to allow per- a result of the urban overexploitation of the aquifer although it manent site occupation and Homo expansion (e.g., Cuthbert et al., still features numerous seeps and streams that emerge in the area 2017). This critical resource may also have significantly deter- during the rainy season, along with a seasonal network of streams. mined the qualitative and quantitative of shellfish collection The nearest river, the Guadalhorce River, flows into the sea in prehistoric times. For regions such as the southern Iberian ~6.5 km away from the cave during high or low sea level periods Peninsula, that suffer prolonged episodes of summer droughts and (Fig. 5). M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 287

Fig. 3. Geologic section (A'-A in Fig. 2) of the zone where Bajondillo cave is located with a projection of glacio-eustatic changes from the Middle Pleistocene to the Holocene.

3. Materials and methods remains were found below that sieve threshold. Screened materials were packed in plastic bags and transported to the laboratory for All materials were retrieved during field seasons taking place in sorting and labelling. 1989, 2000 and 2002. The stratigraphic sequence of Bajondillo cave Specimens were identified with the aid of the reference col- is well stratified and chrono-culturally defined featuring 19 levels lections of one of the authors (AMM) housed at the Laboratorio de that have been recognized in terms of artifacts and absolute dates Arqueozoología, Universidad Autonoma de Madrid (LAZ-UAM), (Fig. 4). Six of these levels (Bj/19-Bj/14) correspond to the Mous- although occasionally identifications had to remain at a higher terian culture (MP) and three, of which only two are reported here taxonomic level (-Phyllum). (Bj/13 and Bj/11) date back to the Aurignacian (EUP) (Cortes- To estimate minimum number of individuals (MNI), size (all Sanchez et al., 2019). Sediment volumes were measured during groups) and side (bivalves) were taken into account (size is excavation and bulk samples of sediment taken from every strati- extremely difficult to estimate when fragmentation is extensive graphic unit. A total of 20.5 m3, representing ~5% from an estimated and no diagnostic elements of the shell are found (see Moreno 400 m3 of sediment, have been excavated. Of these, and except for Nuno,~ 1994; Gutierrez Zugasti, 2009). For any given size, the small volume of sediment from Bj/19 whose marine in- gastropod MNIs were quantified through columellar counts or the vertebrates have been thoroughly studied, the samples from the number of apexes (). For bivalves, the highest number of remaining levels reported in this study represent ~30e40% of the either right or left umbos (hinges) was used to determine MNI. In a material pending analysis. All observed finds were located in three few instances, an MNI ¼ 1 was established when no countable shell dimensions and sequentially captured through nested 10 mm/ parts were encountered yet taxonomically identifiable fragments 5mm / 2mm / 1mm/ 0.5 mm/ 0.1 mm dry-sieving. This were found. Although specimens were measured, metrical data will study does not report finds below 0.5 mm as only unidentifiable be included in a future publication. Densities have been estimated 288 M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296

permanently above water (supratidal) and a low-shore (subtidal) permanently submerged are set apart from the intertidal (eulit- toral) zone with its divisions into upper, medium and lower sectors (Little and Kitching, 1996). Preservation features have been established through a qualita- tive evaluation of the degree of burning (charred or carbonized), type of fragmentation (natural vs. intentional fractures), water erosion, and presence of epibionts (i.e. organisms attached to the shell) (for definitions of thermoalterations and fractures, see Moreno Nuno,~ 1994, Gutierrez Zugasti, 2009). On account of these evidences, three taphonomical groups, sensu Gautier (1987) are recognized: a) reflecting human consumption, b) non-food species that, although potentially edible, reflect gathering for non- dietary purposes, and c) intrusive/incidental species. Included in the latter one finds penecontemporaneous intrussives (so-called “background faunas”) that are essentially synchronous with the anthropic faunas from a given level and late intrusive, the two groups reaching the deposits through natural processes including the activities of alternative accumulating agents such as predators.

4. Results

The marine assemblages from Bajondillo cave total 2768 mol- luscs (2607 of them marine) plus 11 crustacean remains, the latter being acorn barnacles (Balanus) that probably reached the deposits as epibionts. A total of 27 categories of were identified (Tables 1 and 2). These include 17 marine molluscs to species level (Mytilus galloprovincialis, Perna perna, , Cerastoderma edule, Acantocardia tuberculata, Callista chione, Venus verrucosa, Lutraria angustior, Donacilla cornea, Panopea glycymeris, Stramonita haemastoma, Patella caerulea, Patella vulgata, Patella ferruginea, Cymbula nigra, Charonia lampas and Glycymeris bima- culata), another 4 to Genus level (Mytilus sp., Cerastoderma sp., Thracia sp. and Glycymeris sp.), one mollusc Family (), two mollusc Classes ( and ) and one remain only to the level of Phyllum (). Crustacean remains were identified to species (Balanus trigonus) and Genus levels (Balanus sp.). In addition, 3 species of freshwater gastropods (, M. laevigata and Pissium cassertatum) and 7 terrestrial gastropods (Rumina decollata, Theba pisana, Cecilioides acicula, Bithynia tenta- culata, , Iberus marmoratus and lactea) plus 2 genera of terrestrial molluscs (Xerotricha sp., Vitrea sp.) and one terrestrial Family () were identified. In terms of taphonomic groups, as can be seen in Table 1, the majority of the assemblage is represented by what on account of preservation features, in particular fragmentation and occasionally thermo-alterations, constitute food taxa. In fact, most of the taxa from Bajondillo cave eexcept for the smallest species-are edible yet Fig. 4. Bajondillo cave: Overview of the chronostratigraphic sequence and techno- the essentially intact shells and hibernating habits of the pulmo- typological sequence. nate snails suggest that these most probably represent intrussives. Although not qualifying as such for obvious reasons, four additional through number of identified remains (NISP) and MNI counts per marine taxa seem devoid of culinary connotations. Dog cockles (G. cubic meter. Fragmentation has been calibrated through NISP:MNI Glycymeris) have striking shell colorations and are essentially ratios. subtidal taxa that appear on beaches as specimens rolled post- Behavioral inferences, in particular the coastal zone where mortem. In the case of the pearl (Fig. 6), its minuscule size in- molluscs were collected, are based on actualistic studies referring dicates that it derives from a but the possibility that it was to the biology of the species, the biocenoses in which they are found kept after eating the mollusc cannot be ruled out. Lastly, the small and their abundance within each coastal community (Per es and size of the shells from the Genus Thracia rules them out as edible Picard, 1964; Hayward et al., 1998; Poppe and Goto, 1991, 1993; items. Although one cannot decide whether these four groups Rolan, 1984, 1993; Rolan et al., 1990; Hawkins and Jones,1992; Little represent ornamental items as reported in SE Iberian Neanderthal and Kitching, 1996). In our case, were evaluated from three deposits, this is, however tentative, a hypothesis to contend with in ~ standpoints: a) coastal exposure (i.e. open shore vs. estuary), b) future investigations (Zilhao et al., 2010). substrate type [rocky, sandy, muddy and mixed (sand/mud)], and c) Mussels (M. galloprovincialis) dominate the Bajondillo cave vertical zonation. In the latter case an upper zone almost mollusc assemblages, their highest abundances recorded on the oldest levels. This in itself is striking since the fragility and dome- M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 289

Fig. 5. Reconstructed landscape around Bajondillo cave in a situation of marine regression (black line: present day coast line; yellow line: 90 mbsl: 120 m bsl. Blue line: tentative fluvial network during marine lowstand. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.) shaped morphology of mussel shells render them highly suscepti- is probably a methodological construct since these are very small ble to fragmentation, thus one would assume mussel remains to collections and, by virtue of design, NISP / MNI when NISP / 1 decrease from the top to the bottom of the stratigraphic column (Reitz and Wing, 2008)(Table 3)]. (see discussion section). That valve thickness also influences frag- mentation and associated loss of mussel shells at Bajondillo cave 5. Discussion one can infer from the shifting NISP and MNI ratios of the two Mytilid species [the thin-shelled M. galloprovincialis (Mga) and the 5.1. Bajondillo mollusc record and the MP-EUP transition thicker-shelled P. perna (Pp)] from two contiguous levels (Table 3). When NISPs are considered, Mga happens to be more than twice as For the purposes of comparing molluscs from different levels frequent as Pp in the younger level as opposed to the older one (Bj/ one needs to stress that geomorphological and taphonomic dif- 15: 10 Mga/1 Pp vs. Bj/16: 4 Mga/1 Pp). Although less marked, this ferences exist among them. Trampling and thermoalterations, for same phenomenon is documented when MNIs are considered (Bj/ example, are most frequent in levels Bj/15 and Bj/16, documenting 15: 1.5 Mga/1 Pp vs. Bj/16: 1 Mga/1 Pp)(Table 3). This suggests that a more intensive occupation also in terms of lithic industries and although shell loss due to fragmentation increases with depth, it is hearths (Cortes-S anchez, 2007, Table 3). Differential granulometry also related to the thickness of the shell given that these shifting of the sediment must have also affected the degree of conservation, NISP:MNI ratios most parsimoniously correlate with shell density levels Bj/14 and Bj/17 incorporating a significantly larger amount of (Mga ≪ Pp). Notwithstanding anthropogenic causes, such time- gravel than the remaining MP levels that could have produced a mediated, density-dependent, fragmentation suggests that the different amount of damage to the shells. Weight of the sediment is increasing abundance of mussels as one moves back in time is real also a factor to consider when fragmentation is evaluated as this is and was probably more marked in the original thanatocoenoses proportional to the depth a level is found. Such contingencies than in the taphocoenoses that were later excavated. In other would not allow for a straightforward comparison of NISPs, shell words, during Middle Palaeolithic times, shellfish collection at densities or fragmentation among levels to be established and this Bajondillo cave centered upon a single species. need to be kept in mind. Partly for this reason, NISP/MNI ratios may Humans undoubtedly generated shell fragmentation but this is turn out to be the best estimators for establishing comparisons far from straightforward to quantify. Anthropic activity one may amongst levels. infer from NISP:MNI ratios revealing a much more intensive frag- Shellfish collection is documented in all the studied levels, mentation of the hard-shelled marine taxa (i.e. molluscs that were including Bj/17 (MIS4) (Tables 1 and 3). Although the occupation consumed) as opposed to the thin-shelled, presumably intrusive, must have featured periods of abandonment, one can assume that, pulmonates. Despite terrestrial snail shells being the most fragile of in a more or less continuous manner, the activity was maintained all molluscs, Bajondillo cave pulmonates are mostly represented by for those 120kyr. In terms of both NISP and MNI the EUP samples complete or almost complete shells (i.e. NISP-MNI ratios of 1:1). In are orders of magnitude below those from the MP. Density values, the MP levels, marine mollusc NISP:MNI ratios increase steadily taken as Ʃ marine mollusc NISP/m3 and Ʃ marine mollusc MNI/m3, with depth [i.e. from 5.4 (Bj/14) to 45 (Bj/19)] but it is difficult to tell the same story (Table 3). This hints that the harvesting of decide what amount of that fragmentation is due to natural or molluscs decreased dramatically from the earliest to the youngest anthropic causes [a 1:1 NISP:MNI ratio is also recorded in mussel levels of the occupation (Table 3). Although fragmentation is a samples from the Upper Palaeolithic levels (Bj/11 and Bj/13) but this factor to take into consideration (see above) and the densities of 290 M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296

Table 1 Bajondillo invertebrate assemblages: Mousterian. NR and MNI per level (NR: number of remains, MNI: minimum number of individuals).

Level MIS3 MIS4 MIS5 MIS6

Bj/14 Bj/15 Bj/16 Bj/17 Bj/18 Bj/19

Abundance estimator NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI

Food taxa Mytilus galloprovincialis 4 2 99 5 191 17 186 10 497 11 1305 29 Mytilus sp. 12 2 3 1 60 2 28 3 15 1 ee Perna perna ee8 8 56 17 eeeeee Mytilidae sp. 38 6 12 4 1 1 ee11ee Pecten maximus ee11222111ee Cerastoderma edule 11eeeeeeeeee Cerastoderma sp. 1 1 eeeeeeeeee Acanthocardia tuberculata eeeeee11eeee Callista chione ee112153eeee Venus verrucosa eeee11eeeeee Lutraria angustior eeee33eeeeee Donacilla cornea eeeeeeeeee11 Panopea glycymeris ee1142332111 Stramonita (Thais) haemastoma eeeeeeeeee11 Patella cf. vulgata eeeeee11eeee Patella caerulea 11ee33ee22ee Patella cf. caerulea ee11eeeeeeee Patella ferruginea ee33eeeeeeee Cymbula nigra eeeeeeee11ee Charonia lampas eeeeee22eeee Non-food taxa Pearl eeeeeeeeee1 e Glycymeris bimaculata ee66eeeeeeee Glycymeris sp. ee51eeeeee33 Thracia sp.? eeeeeeeeee11 Taphonomically undefined marine Bivalvia 2 2 1 1 2 2 eeeeee Gastropoda eeeeeeeeee22 Mollusca sp. 1 1 eeeeeeeeee Intrussives/incidental molluscs Melanopsis praemorsa 11663322eeee Melanopsis laevigata eeeeee5 5 11 11 11 11 Pisium cassertatum eeeeeeeeee11 Rumina decollata eeeeee6 6 10 10 10 10 Theba pisana eeee33553333 Xerotricha sp. eeeeee11eeee Vitrea sp. eeeeee111111 Cecilioides acicula eeeeee223311 Bithynia tentaculata eeeeee6 6 15 15 ee Succinea putris eeeeee111122 Iberus marmoratus eeeeee11ee11 Otala lactea 113311112211 Helicidae 1 1 1 ee e526677 Crustacea Balanus trigonus eeeeeeeeee22 Balanus sp. e 3 ee44ee4433 TOTAL 62 19 151 42 336 62 264 57 575 74 1358 80

Table 2 Bajondillo mollusc assemblages from Aurignacian levels (Bj/13-Bj/11). NR and MNI per level (NR: number of remains, MNI: minimum number of individuals).

Level Bj/11 Bj/13

NISP MNI NISP MNI

Food taxa Mytilus galloprovincialis 54ee Mytilus sp. ee22 Mytilidae sp. 4 4 4 4 Fig. 6. Pearl from Bj/19. Cerastoderma glaucum 11ee Bivalvia ee41 Intrussives/incidental molluscs lithics do hint at a less intensive occupation of the cave by AMH Melanopsis praemorsa 3311 Otala lactea 1111(Table 3), the differences between the MP and the EUP are striking. Helicidae 1 1 6 4 These differences hold when Bj/19, the only level whose molluscs TOTAL 15 14 18 13 have been totally studied, were to be removed since Bj/18 also M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 291

Table 3 Selected parameters for the Bajondillo levels (Bj). (mm ¼ marine molluscs) (NISP: number of individual specimens).

Bj/ ~kyr NISP (total) Mussel (NISP) Mussel m3 m3 Lithic ind. Lithics ind./m3 mm/m3 (NISP) mm/m3 (MNI) mm (NISP) /m3 /kyr

11 8.9 10 60 1.9 4.6 e 1845 401 2.2 1.95 9 13 4.0 10 33 7.5 0.8 e 353 441 12.5 8.75 7 14 7.0 59 87 12.8 4.2 0.6 1999 476 14.3 3.8 16 15 5.0 141 75 36.9 3.3 0.82 5062 1534 42.7 10.0 33 16 6.0 325 74 158.0 1.95 0.65 1593 817 166.6 26.1 51 17 13.0 228 81 38.2 5.6 0.4 4682 835 40.7 4.28 24 18 [54.0] 519 89 68,266 0.0075 e 91 12,133 69,200 2400 18 19 e 1315 96 76,764 0.017 e 73 4291 77,352 2235 38

features dozens of thousands of mollusc remains per m3. The In terms of taxonomic groups, mussels at large (i.e. Mytilus sp., outlier of this trend is Bj/17 whose drastic lowering of the shellfish M. galloprovincialis, Perna perna and Mytilidae) constitute the signal may reflect a collection having taken place farther away from dominant taxon at all times, NISP values being also significantly the site during MIS4 due to the 70 m low-stand the marine higher during the MP (74e96%) than in the EUP (46e57%) (Tables 1 regression fostered (Fig. 7). and 3). Although abundance values decrease in the case of MNI If shellfish collection is taken to be the hallmark of H. sapiens (18.5e63.8%) their overall dominance holds at all times. If a aquatic adaptations, then the data from Bajondillo cave undoubt- restricted species spectrum is taken as the hallmark of a “special- edly call for a reconsideration of such paradigm. ized” exploitation strategy, then the ca. 120kyr mussel dominance Food items constitute the bulk of all samples when NISPs and at Bajondillo cave probably qualifies as the longest one on record. It MNIs are considered. Dominance of edible molluscs is more marked is also relevant from a biogeographical standpoint that archaeo- during the MP (Bj/14e19: 86e96%) than the EUP (Bj/11, Bj/13: logical level Bj/16 (MIS3) is the earliest record for P. perna in the 45.5e66%). Non-food species are marginal items documented only Mediterranean Sea. in half of the MP levels and none of the EUP levels. In contrast, Mussel dominances at Bajondillo cave imply that the almost intrussives are far more abundant in the EUP (33e44%) than in the anecdotic presence of the remaining taxa, including those of the MP (2e13.5%). second most abundant group (limpets), cannot be expected to yield

Fig. 7. Multiproxy data comparison with Bajondillo cave selected archaeological levels and age (kyr cal BP). From top to bottom a) Reconstructed global sea level relative to present (blue) (Bintanja et al., 2005) b) Mean annual Sea Surface Temperature (ºC) (Uk'alkenones) from Alboran sea (Martrat et al., 2007) c) Arboreal Pollen concentration (%) from Padul Lake (Southern Spain) (Camuera et al., 2018). Dashed yellow line represents aprox. >5 km distance from Bajondillo Cave to coast line and SST ¼ 16 C. Dashed red line represents aprox. >6 km distance from Bajondillo Cave to coast line and arboreal Pollen ¼ 40% at Padul Lake. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.) 292 M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 reliable complementary information of any kind. Still, one note- In colder NW Atlantic waters, mussels in sheltered , as worthy difference between the MP and the EUP assemblages refers is the case at Bajondillo cave, often form extensive beds in the to the comparative abundance infaunal bivalves (i.e. buried intertidal zone their densities reaching up to 1000 individuals/m2 in the substrate) feature. This group, which includes all bivalves (Poppe and Goto, 1993). The presence at Bj/17 (MIS4) of P. vulgata, a with the exception of mussels and (Pecten sp.), represents cold water of mussel beds in sheltered estuaries with calm 1.6% of the identified marine molluscs in the Neanderthal levels yet waters, reinforces that scenario. All of the remaining marine bi- raises to 25% in the AMH levels. As will be seen, this feature may be valves, with the exception of scallops, bury in sandy or mixed a crucial clue for setting apart the shellfish collection of the two bottoms of sand and mud from the intertidal zone downwards. hominine lineages. Terrestrial pulmonates characteristic of limestones lying close to If the NE Atlantic biological preferences of M. galloprovincialis the coast seem restricted to the lowermost of the MP levels (i.e. Bj/ were to be taken as a reference, their shellfish collection at 17e19), whereas Otala lactea, which thrives on vegetation in areas Bajondillo cave would be taken to indicate a preferential foraging in with abundant water, is, along with mussels, the only species the mid- and lower intertidal zones of a rocky shore. Such hy- documented on all eight of the studied levels (Tables 1e3). pothesis would be reinforced by the prevalence of limpets (Patella) in all but the earliest of the MP levels. But in the Alboran Sea, as 5.2. Bajondillo mollusc assemblage: regional and global significance throughout the Mediterranean, M. galloprovincialis is a denizen of the shallow subtidal, a permanently submerged zone where the To evaluate the Bajondillo cave assemblages from a regional species defines a facies (Rodríguez, 1982). In this zone mussels are perspective, Table 4 summarizes the essentials of the mollusc col- accompanied by the limpet Patella caerulea, documented on Bj/14, lections thus far published from the Alboran sea region as well as Bj/15, Bj/16 and Bj/18, and the red-mouthed rock shell Stramonita the unpublished assemblages from Abrigo 3. Although the earliest haemastoma, documented on Bj/19. Although the limpet exhibits a of the assemblages from MIS8-MIS5 are poor both in terms of NISP wider distribution from the supralitoral to the subtidal, and taxonomic diversity, a pattern that seems clear is that limpets, S. haemastoma is only documented over permanently submerged not mussels, constitute the most important and prevalent group in rocks (MCLZ and JLVP, pers. obs.). Lack of coincidence of these the region as well as the only major group in the earliest stages of gastropods at any level may simply be a chance event due to low the temporal sequence [ƩNISP ¼ 1665 (47.8%)]. In contrast with sample yet the idea that mussels were systematically collected Bajondillo cave where mussel frequencies exhibit very high values underwater is striking. during MIS6 (Bj/19: 96% NISP), MIS5 (Bj/18: 89% NISP) and MIS4 (Bj/ From a paleoenviromental standpoint, the prevalence of fresh- 17: 81% NISP), mussels only reach substantial abundances in MIS3 water snails and a freshwater bivalve (Pisium cassertatum) sites and in the MIS5b þ to MIS5d þ collections from Gorham's throughout the sequence hints at the existence of freshwater levels K,R,S and T, mussels are secondary items (Baden-Powell, around the cave, a fact that the hydrogeological data of the area 1964, Table 4). Although both limpets and mussels evidence support (Fig. 5, Tables 1 and 3). Cockles (Cerastoderma sp., shellfish collection to have been carried out preferentially on rocky C. glaucum and C. edule) and wedge shells (Donacilla cornea) tell a shores, mussels found as aggregations must have been a more similar story in the case of brackish waters (C. edule can tolerate an productive resource in terms of total cropped/unit time up to 5% decrease in salinity levels) though the restriction of cockles but also as isolated individuals. For both reasons but also because to the levels Bj/14-Bj/11, along with the absence of another typical the earliest documented limpet species in the sequence (P. vulgata) estuarine species as is the (Ostrea edulis) seems intriguing. is often an accompanying species of mussels beds in the NE These data and geological evidences (travertine deposits) indicate Atlantic, such absence/scarcity of mussels on MIS8-5 sites other that, for the period our study covers, there must have existed no than Bajondillo cave seems counterintuitive and only explainable in freshwater shortages in the area around Bajondillo cave at no time terms of taphonomic loss or restricted accessibility to the resource. of the year. In other words, freshwater availability should not Taphonomic loss should never be ruled out since mussel shells are explain putative diachronic shifts taking place in the mollusc as- very fragile when compared with those of other marine molluscs. semblages in terms of diversities and abundances. This would also Also, although the pearly inside of the shell often suffices to explain why, with independence of the distance to the shore at determine a small fragment as a member of the family, if remains different moments, Bajondillo cave features an uninterrupted were retrieved by hand small fragments are likely to have been sequence from MIS6 to MIS3, being also one of the very few cases missed. Under such circumstances it is difficult to decide whether incorporating evidences of shellfish collection during the poorly absence of mussels in those samples would be genuine or not. documented MIS4 stage. Accessibility would be essentially determined by two factors: coast Regional warm/wet or cold/dry climate are represented by exposure and bathymetric range. The present day coastlines of the Arboreal Pollen (AP) percentages (%), with Pinus excluded, from the Strait of Gibraltar, where most of the sites appear, are dominated by near Padul lake terrestrial pollen sum (Camuera et al., 2018). It is rocks and quite exposed although beaches are occasionally found. significant that minor AP% values are reached during Bj-18 to Bj-17 Intense current velocities in Gibraltar Strait region and abrupt hiatus and during MP to UP transition (Fig. 7). Temperature fluc- continental shelf promote that beaches are poorly developed even tuations also seem coincident with the distribution of certain bio- during past glacial periods (Fernandez-Salas et al., 2015). These indicator species (Fig. 7b) as the subtropical limpet Cymbula nigra. problems would not be so acute in the more sheltered coast along Intertidal zone denizens such as P. perna (Bj/15 and Bj/16) and the Bay of Malaga thus the comparatively high frequencies of P. ferruginea (a SW Mediterranean endemism found in Bj/15), are mussels from these sites (Abrigo 3: 67% NISP, Bajondillo cave: documented when the coast was closer to the site (i.e. tempera- 57e96% NISP) and the slightly lower values from the more exposed tures were on the rise) (Fig. 7 and Tables 1e3). Both species thrive coast in front of Nerja cave (37% NISP) make sense. in the upper intertidal, where on exposed shores they overlap the But the really intriguing question in terms of mussel availability barnacle belt (Rodríguez, 1982, Gofas et al., 2011a,b). During MIS5, is whether or not during prehistoric times M. galloprovincialis was, Bj/18 documents the earliest record of Cymbula nigra in the Medi- as is the case today, found only in the permanently submerged terranean. This subtropical limpet occupies rocky surfaces from the subtidal zone or was instead an intertidal species, as is today the shallow subtidal to the lowermost reaches of the intertidal zone in case in the NE Atlantic waters down to the Strait of Gibraltar (Gofas areas of moderate hydrodynamics. et al., 2011a,b). This is a critical issue to explain former mussel M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 293

Table 4 Mollusc assemblages from prehistoric sites from the Alboran Sea. Data taken from Fischer, 1928), Baden-Powell (1964), Barton (2000), Lozano-Francisco et al. (2003), 2004, Cortes-S anchez et al. (2008), Ramos Fernandez et al., 2012, Fa et al., (2016), Ramos Munoz~ et al., 2016).

TAXON Site/Level

MIS 8 MIS 6 MIS 5 MIS 3 MIS 2

Benzú 1-5 Benzú 6 Benzú 7 Gorham KeU Vanguard Devil's Gorham Gorham Abrigo 3 Nerja Gorham B Tower D/E IV b

Antalis inaequicostatum e eee e eeee 5 e Antalis vulgare e eee e eeee 1 e Fissurella sp. eeee eee1 eee Patella vulgata 107 22 2 eeþþþ e 615ee Patella caerulea ee2 197 11 e 548 1 151 105 211 Patella ulyssiponensis eeee 2 ee e 20 27 e Patella ferruginea 1 ee64 e þ 2 e 73e Patella rustica eeee eeee45 9 e Patella depressa eeee eþ e 19 41e Patella sp. eeee eee11 e 25 e Cymbula nigra eeee eeee614e Throchidae sp. e eee e eeeeee Gibbula richardi eeee eeee171e Gibbula sp. eeee eeee91e lineatus eeee eeee8 ee Phorcus articulatus eeee ee76 ee 53 Phorcus turbinatus eeee ee9133ee Littorina sp. eeee eee2 ee2 Littorina littorea eeee eee1 e 1 e Littorina saxatilis eeee eee1 ee1 Nucella lapillus e eee e eeeeee Stramonita haemastoma e eee e eeeeee Cyclope donovani e eee e eeee 51 Erosaria spurca eeee ee1 ee 12 eeee ee1 ee 1 e Charonia lampas eee1 eeee1 ee Ocinebrina edwarsi eeee eeee1 ee Nassarius reticulatus eeee eeee11e eee2 e eeeeee 1 e 1 eeeee32e Mytilus galloprovincialis eee12 100 213 5 675 271 15 Perna perna e eee e eeeeee Modiolus modiolus eeee ee6 ee 5 e Ostrea edulis eeee eeee14e Spondilus gaederopus e eee e eeeeee Pecten maximus eeee ee50 2 5 21 10 e eee e eeee 1 Pecten sp. eeee eee2 eee Chlamys multistriata eeee eeee1 ee Lucina borealis e eee e eeeeee Acanthocardia tuberculata eeee 1 eeee 61 Cerastoderma edule e eee e eeee 88 e Cardium sp. e eee e eeeeee Venus verrucosa e eee e eeee 1 e Callista chione eeee 1 eeee 1 e Tapes decussatus e 1 ee e e2 ee 58 1 Veneridae sp. eeee eee3 eee Gastrochaena dubia eeee eeee1 ee Sepia nodifera e eee e eeeeee TOTAL 109 23 5 276 115 þ 908 37 993 774 247

presence and abundance that cannot be solved with the data at 45e80%, Jerardino and Marean, 2010, Table 5). In the case of the hand. South African site it appears that the fluctuation of infaunas was Equally interesting would be to explain the scarcity of infaunas determined by alternating marine transgression and regression (i.e. molluscs that bury in the substrate) at both Bajondillo cave and events that turned the shore into rocky or sandy. But these authors the remaining Alboran sea assemblages. Taken as a whole, this are keen to stress that whereas rocky intertidal species are “… group represents a mere 1.6% of the NISP in the MP levels at largely visible, fixed to rocks and their collection entails relatively Bajondillo cave (Table 3) and some 8% of the NISP in the Alboran sea shallow entry into the water during low tides …” the collection of collections. But in the EUP collections the abundance of infaunas clams buried in sand “… involves more than simple foraging … seems rises to 25% (Table 3). And although this figure may not be reliable to require a relatively more complex set of variables such as antici- due to the minimal sample size of the EUP levels, that value is very pation, planning, and possibly also additional help in the form of dilly similar to that of infauna on Nerja cave (Gravettian: 20%) (Cortes- bags and/or cooperation with other people” (Jerardino and Marean, Sanchez et al., 2008). In fact, when one reflects upon it, these 2010: 420). Although regional conditions and species spectra may values are more in concordance with those obtained on some of the have determined the course of various of the aquatic adaptations, samples studied on the South African site of Pinnacle Point (i.e. species living in sediments would have been largely invisible for 294 M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296

Table 5 Abundance, expressed in terms of NISP (Number of identified remains) and MNI (Minimum number of individuals), and their corresponding percentages over the assemblage's total, of selected mollusc taxa from Middle and Early Upper Paleolithic assemblages from the Alboran Sea. (*mentions Lucinia, Cardium, Pecten, Spondylus and Sepia) (Gorham's a-c are Waechter's excavations, see Baden-Powell, 1964; d is taken from Fa et al., 2016, other data taken from Ramos Munoz~ et al., 2016, Fisher, P., 1928, Barton, N. 2000, Cortes- Sanchez et al., 2008, Jerardino and Marean, 2010).

MIS 3 Level Mussels Mussels Limpets Topshells Periwinkles Other NISP (%) MNI (%) MNI (%) MNI (%) MNI (%) MNI (%) MNI (%)

MIS 3 7 eee 4 (80.0%) ee 1 (20.0%) MIS 3 6e1 eee 130 (98.5%) ee 2 (1.5%) MIS 3 all “abundant” ee “abundant” ee * MIS 3 B 15 (6%) ee 211 (85.0%) 5 (2.0%) 1 (0.4%) 12 (4.8%) MIS 3 D-G 245 (26%) ee 550 (58.5%) 85 (9.0%) ee MIS 3 KeU 12 (4%) ee 261 (94.5%) ee 3 (1.0%) MIS 3 IVb 5 (13%) ee 19 (50.0%) 2 (5.2%) 3 (4.0%) 9 (23.6%) MIS 3 e z100 z50 e 13 (11.0%) ee 2 (1.7%) MIS 3 ee 276 (8%) 2626 (78.0%) 224 (6.5%) 77 (2.2%) 1 (0.02%) 211 (6.1%) MIS 3 11e13 15 (45.4%) 14 (51.8%) 11 (40.7%) eee1 (3.7%) MIS 3 14 54 (87%) 10 (52%) 2 (10.5%) 1 (5.2%) ee 4 (21.0%) MIS 3 15 114 (75.5%) 18 (42.8%) 4 (9.5%) 4 (9.5%) ee 4 (9.5%) MIS 3 16 317 (95.5%) 37 (63.8%) 4 (6.9%) 3 (5.1%) ee 11 (19%) MIS 3 17 214 (81%) 13 (22.8%) 26 (45.6%) 1 (1.7%) ee 10 (17.5%) MIS 3 18 512 (89%) 13 (18.5%) 41 (58.5%) 3 (4.3%) ee 2 (2.8%) MIS 3 19 1305 (96%) 29 (38.6%) 26 (34.6%) eee6 (8.0%) MIS 3 ee 198 (29%) e 6 (0.9%) 4 (0.6%) e 471 (69.0%)

coastal foragers. Could it be that the rise of infauna we witness in Mediterranean (Oguz et al., 2014; Yebra et al., 2018). Whether these the EUP collections from Bajondillo cave is reflecting a change in similar environments triggered changes in the behavioral reper- the shellfish collection strategy from Neanderthals to AMH? Or toires of the hominine populations stationed on their shores at a would perhaps such rise of infaunas merely reflect a preferential given time is something that needs to rank high in the priorities of collection of clams after heavy storms suck animals in the sedi- the research agenda of researchers. ments up to the surface and wash them ashore? Here lie probably the most challenging questions deriving from the analysis of the Bajondillo cave mollusc assemblages. 6. Conclusions On a global scale, coastal resource exploitation has been regar- ded as one of the hallmarks of AMH behavior and a contributing Evidences of shellfish collection at Bajondillo cave range from factor in the modern humans Out-of-Africa expansion although its the MP and EUP (~160e35kyr BP). These reveal that Neanderthals uniqueness and significance have been contested (Walter et al., had been harvesting mollusc well before the arrival of anatomically 2000; Finlayson, 2009). In looking at examples of prehistoric modern humans at the site. Comparative analyses additionally coastal resource exploitation, the question of whether such be- demonstrate that Bajondillo cave Neanderthals are not exceptional haviors are the product of long-established cultural traditions or by any means since the evidences of shellfishing during Mousterian independently derived is one of great importance. times are also documented at other Alboran Sea sites. In evolutionary in biology one traditional way to address such In contrast with Western Mediterranean sites mostly dominated kind of questions is through the comparative method (Harvey and by mobile and non-colonial gastropods, the dominant taxon of the Pagel, 1991). This can be done by comparing coastal resource molluscs assemblages from Bajondillo cave throughout the tem- exploitation between the two AMH and Neanderthal lineages poral sequence is the mussel Mytilus galloprovincialis. which separated around 500kyr BP checking for quantifiable co- Bajondillo cave shellfishing signals seem sensitive to the dis- incidences and differences (Green et al., 2010). The existence of a tance of the site to the coast and paleoenviromental conditions. shellfish collection behavior in two lineages one can interpret as: a) This may explain why the site, lying never too far away from the a behavioral repertoire that was present in the common ancestor shore, provides evidences of shellfishing during MIS4, a poorly and is therefore much more ancient than present day estimates documented stage in terms of aquatic adaptations that is associated suggest, b) a behavioral repertoire that was transmitted from one to a major sea level drop. lineage to another, and this would require contact (i.e. coincidence At Bajondillo cave, the transition from Neanderthals to AMH in space and time), and c) a behavioral repertoire that arose inde- features a sharp decline of molluscs yet a relative increase of pendently in different lineages thus constitutes a phenomenon of infaunal bivalves. An increase of infaunal bivalves also marks the evolutionary convergence. MP-EUP transition at Pinnacle Point although only AMH are present Convergence, also in terms of behavior, is expected in different at this site. Despite their temporal coincidence, differences in the lineages of organisms exploiting resources in a similar way or un- collection of shellfish in the Westernmost Mediterranean and der similar environmental conditions (pressures). In our case, the South Africa regions exist that suggest the practice to have arisen so-called Mediterranean (i.e. the Mediterranean proper, independently in the two hominine lineages. This hypothesis South Africa, south-west Australia, California and Chile) are well awaits confirmation once new collections become available for known for their numerous instances of convergent evolution study. (Fig. 8). In our case, Gibraltar-Bajondillo and Pinnacle Point not only In the light of our research, it appears that shellfish collection share similar climatic regimes but are also areas of high marine will have to join those signatures of human associated with specific currents and sys- considered as exclusive heritage of Homo sapiens. Our shellfishing tems: the Agulhas Current in the case of eastern South Africa data hint instead at parallel and gradual trajectories, stepped in (Pinnacle Point) (Lutjeharms, 2007, Goschenad et al., 2015) and the time and space, where other hominines, in this case Neanderthals, Malaga upwelling and Alboran Gyres in the Westernmost may have played a significant role. M. Cortes-Sanchez et al. / Quaternary Science Reviews 217 (2019) 284e296 295

Fig. 8. Mediterranean climate biogeographic lands (green) associated with zones of coastal (orange) showing the location of archaeological sites with shellfishing explotaition (Map: http://donationsworldmap.com/images/worldmapt.gif). (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)

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