Quaternary International 421 (2016) 62e77

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Pigeons and choughs, a usual resource for the Neanderthals in Gibraltar

* Ruth Blasco a, , Jordi Rosell b, c, Anna Rufa b, c, Antonio Sanchez Marco d, Clive Finlayson e, f a Centro Nacional de Investigacion sobre la Evolucion Humana (CENIEH), Paseo Sierra de Atapuerca 3, 09002 Burgos, Spain b Area de Prehistoria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, 43002 Tarragona, Spain c IPHES, Institut Catala de Paleoecologia Humana i Evolucio Social, C/ Marcel.lí Domingo s/n e Campus Sescelades URV (Edifici W3), 43007 Tarragona, Spain d Area of Neogene and Quaternary Faunas, Institut Catala de Paleontologia, Campus de la UAB, Cerdanyola del Valles, Barcelona, Spain e The Gibraltar Museum, 18-20 Bomb House Lane, P. O. Box 939, Gibraltar f Institute of Life and Earth Sciences, University of Gibraltar, Gibraltar article info abstract

Article history: An ecological perspective is essential to our understanding of variations in the faunal communities and Available online 12 November 2015 their relationships with human groups, as well as the foraging strategies that hominids practised in Pleistocene environments. Using the Middle Palaeolithic avifauna, the ecological quality of the Strait of Keywords: Gibraltar shows a complex scenario in which cliff-dwelling species represented one of the most Middle Palaeolithic important and abundant taxa. Pigeons (Columba) are a species that favour rocky habitats, typically Neanderthals nesting on cliff ledges and at the entrances to large caves, while corvids are forest birds and only a few Taphonomy Palaearctic species, including colonial ones such as choughs (Pyrrhocorax), can also breed on rocky Birds Gorham's Cave outcrops or cliffs. At Gorham's Cave, Gibraltar, the Neanderthals exploited pigeons and choughs for a Gibraltar period of more than 40 ka, with the earliest evidence dating from at least 67 ka. We show that such exploitation was not occasional, having found repeated evidence of the practice in different layers within the cave. The Gibraltar sites seem to provide ideal conditions for broadening the spectrum of prey during MIS 3. The high diversity of avian species identified in the faunal assemblages could be related to the location of the caves, which enables the exploitation of coastal and rocky habitats. This fact seems to point to the unique circumstances and foraging opportunities at this particular locality, where the available prey and environmental conditions seem to be influencing, among other factors such as site functionality and socio-cultural variables, the Neanderthal diet. © 2015 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction ecological data has not gone unnoticed by many researchers who study the past, using birds to aid them in making more accurate Birds are one of the most successful biological groups of the palaeoclimatic and palaeoecological reconstructions (e.g., Finlayson biosphere in terms of evolution. Their extensive history, which et al., 2011; Stewart and Jacobi, 2015). dates back to the Mesozoic period, has enabled them to develop a Recently, birds have become increasingly important in studies large number of forms and taxa (about 10,000 species currently related to the behaviour of human groups in the Pleistocene, listed according to Clements [2011] and Clements et al. [2014]) and especially relating to hominids that preceded anatomically modern maintain stable populations in virtually all climatic regions and ones. In this latter case, their contribution is not solely limited to environments on the planet. This makes them authentic bio- providing ecological details, but significantly contributes to the markers for research related to life sciences, not only because of the clarification of fundamental aspects of behaviour and subsistence. information they provide on the specific characteristics of ecosys- One of the best examples can be found in the ornamental use of tems, but also because of their ecological relationships with other feathers and talons of raptors by the Neanderthals of the Late biological entities. Therefore, their potential as a source of Pleistocene (Peresani et al., 2011; Finlayson et al., 2012; Morin and Laroulandie, 2012; Romandini et al., 2014; Radovcic et al., 2015). However, the debate on the interaction between hominids and * Corresponding author. birds in this period goes beyond its possible symbolic capabilities E-mail address: [email protected] (R. Blasco). http://dx.doi.org/10.1016/j.quaint.2015.10.040 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved. R. Blasco et al. / Quaternary International 421 (2016) 62e77 63 and should be considered within the complex world of the diversity et al., 2009; Codding et al., 2010; Cochard et al., 2012; Morin, of human behaviour and subsistence strategies. 2012). Perhaps, from this perspective, it would be possible to According to some studies, the hominids of this period were explain a growing body of data, but occasionally, birds are being sufficiently able to adapt to very different climates and to exploit considered within the faunal spectrum of some human groups in the wide range of animal resources depending on the eco- the European Middle Palaeolithic and previous periods (e.g., Roger, geographical area (see examples in Gaudzinski, 2006; Stringer 2004; Gaudzinski-Windheuser and Niven, 2009; Hardy and et al., 2008; Cochard et al., 2012; Yravedra et al., 2012; Yravedra Moncel, 2011; Morin and Laroulandie, 2012; Blasco et al., 2013, and Cobo-Sanchez, 2015; Sacca, 2012; Blasco et al., 2013). Against 2014; Gerbe et al., 2014; Fiore et al., 2016). this backdrop, birds seem to have always played a secondary role as Gibraltar is an ideal setting in which to study the possible ex- a source of meat, which could be due to their characteristics (small istence of such strategies among Neanderthals. From an ecological and elusive prey eespecially flying birds) and be linked directly to point of view, the Rock is located in a prime area where the Med- the tenets of the Optimal Foraging Theory ([OFT] e.g., Smith, 1983) iterranean Sea to the east connects with the Atlantic Ocean to the e that is, the imbalance between the energy invested in capturing west and the Iberian plate to the north comes into contact with the quick-flying animals and the energy return that they supply makes African plate to the south. This encourages the existence of a wide their regular catching unprofitable without the appropriate tech- variety of avian species, among which there are many migratory nology. The blanket application of OFT to fossil material is, however, birds that move between Africa and Europe on a regular basis, and highly questionable (Finlayson, 2004). Several observations made resident birds, which are mainly marine birds that usually nest in in present-day hunteregatherer populations seem to show the ease the cliffs. The sites associated with Middle Palaeolithic in this lo- with which some species can be captured with relatively simple cality tend to have an abundant number of recorded birds, which methods if prior knowledge of their behavioural patterns is present indicates that this ecological diversity has remained unchanged (e.g., Levi-Strauss et al., 1992; Juste et al., 1995; Emperaire, 2002; over time (Finlayson et al., 2016). These species have natural hab- Negro et al., 2016). These captures often coincide with 1) incuba- itats associated with crags and currently remain numerous in the tion periods of ground-nesting species, in which the cryptically- area of Gibraltar, forming resident colonies and arriving to settle coloured females of some species remain fairly static for long pe- there at various times in the year. This study focuses on the most riods (e.g., waterfowl such as pochards); 2) capturing hatched in- represented taxa in the Mousterian levels of the Gorham's Cave site, dividuals of altricial species which usually remain in the nest for which are pigeons and corvids (especially choughs). Our intention several weeks depending on the species; and 3) birds forming here is to find out the extent of Neanderthal predation on these colonies of hundreds of individuals, as is the case with many sea- cliff-dwelling avian species in Gorham's Cave, in order to expand birds. Many of these animals (or their eggs) may be obtained and complement the previously published data on pigeons (Blasco directly by hand without much difficulty. An example of this is the et al., 2014). Furthermore, the data obtained will be compared with case of Cory's shearwaters (Calonectris borealis) in the Canary the finds from level III ewhich is associated with anatomically Islands, which were heavily hunted by the Guanches before the modern humans (Finlayson et al., 2006), allowing us to contribute arrival of Europeans in the fifteenth century (Rando and Perera, to the debate on the exploitation of birds in Middle Palaeolithic 1994), or the case of fulmars (Fulmarus glacialis) in the Faroe contexts. Islands, whose chicks were taken by Inuits who climbed the cliffs to collect them (Mallory, 2006). Therefore, these cases (and other, 2. The archaeological setting: Gorham's Cave similar ones) suggest some degree of profitability in the capture of some species depending on the time of year and the surrounding Gibraltar is located on the southern tip of the Iberian Peninsula environment. We should also bear in mind that birds contain (360701300N52003100W), at the eastern end of the bay of the same potentially useful non-edible products (Peresani et al., 2011; name. The peninsula is part of the northern shore of the Strait of Finlayson et al., 2012; Morin and Laroulandie, 2012; Blasco et al., Gibraltar, and is adjacent to the Mediterranean Sea and the Atlantic 2014; Romandini et al., 2014; Radovcic et al., 2015), and that fac- Ocean (Fig. 1). Currently, the Rock of Gibraltar contains 213 listed tors such as age composition, sex, and the physical-motor limita- caves, of which at least 26 feature archaeological remains. One such fl tions of some individuals of the group may also have in uenced the cave is Gorham's Cave, which was discovered by Captain A. Gorham inclusion of certain small animals in foraging strategies (e.g., Bird of the 2nd Battalion Royal Munster Fusiliers in 1907. The name

Fig. 1. Location of Gorham's Cave, Gibraltar, in the southern Iberian Peninsula. 64 R. Blasco et al. / Quaternary International 421 (2016) 62e77

Gorham was initially associated with a fissure at the rear of the middle and lower part of the stratigraphic sequence. The knap- cavern, but later both the cavern and the fissure system became ping technique mainly follows the discoid reduction sequences, known as Gorham's Cave. although a very significant increase in lithic flakes originating John D'Arcy Waechter was the first to perform large-scale from Levallois bipolar cores in the unit SSLm.5-6 is observed. The archaeological excavations in Gorham's Cave and the first to last moments of the Middle Palaeolithic are represented in this establish the stratigraphic sequence, which stretched from the area of the cave by a Levallois point from UBSm.4, with the CHm.5 Middle Palaeolithic to the Holocene, with occupation for more than unit being the first attributed to the Upper Palaeolithic (Barton 100 ka (Waechter, 1951, 1964). The stratigraphic profile described and Jennings, 2013). The inner area gathers the same techno- by Waechter consisted of several horizontal levels from east to complexes from the Middle Palaeolithic at level IV, which are west. However, this has proved to be considerably more straight- produced on autochthonous raw materials (mainly flint, sand- forward than the complex stratigraphic reality that more recent stone and limestone) originating from fossil deposits found at the interventions have shown (Barton et al., 2013). beach near the caves and from the immersed flint levels in the The second phase of the systematic excavations was developed Jurassic units of the Rock (Giles Pacheco et al., 2012; Shipton et al., by the Natural History Museum of London, directed by Dr Chris- 2013). The characteristics of the cave indicate the presence of topher Stringer, and the British Museum of London, led by Jill Cook. discoid reduction and Levallois methods, with some cores dis- After the first excavations, work was then carried out as part of the playing unipolar orthogonal and opposite bipolar reductions. Gibraltar Caves Project, which was jointly operated by the Gibraltar From a typological point of view, level IV shows a predominance Museum and the Natural History Museum of London. The work of scrapers and denticulates, with indentations and abrupt focused on the outer part of the cave, where Waechter (1951, 1964) retouching. The length of the flint flakes seems to be conditioned carried out his first interventions, and from 1997 the project by the size of the pebbles, which are of small dimensions in the incorporated teams from the Museum of El Puerto de Santa María current breccia on the beach. The archaeological horizon imme- and the University of Huelva, moving the target area to within the diately above records a characteristic Upper Palaeolithic technol- cave (Finlayson et al., 2006). ogy with diagnostic pieces attributed to both Solutrean (IIIb) and Excavations of the outer part of the cave have exposed a strat- Magdalenian (IIIa) (Giles Pacheco et al., 2012). igraphic sequence of more than 16 m (Barton et al., 2013), mainly In Gorham's Cave, the anthropogenic evidence goes beyond the composed of earthy materials covering a cemented beach-rock lithic industry, hearths and human processing marks, as this de- deposit, which presumably accumulated during MIS 5 (Fig. 2). posit documents the first known example of an abstract pattern Jimenez-Espejo et al. (2013) argue that the sedimentary structures engraved by Neanderthals, which consists of a deeply impressed appear to show a massive aeolian accumulation related to trans- cross-hatching carved into the bedrock of the cave that remained gressive coastal dunes, which seem to have migrated during MIS 3 covered by sediments from the archaeological level IV (Rodríguez- highstand substages and/or cold, arid periods. In general, the Vidal et al., 2014). stratigraphic series of Gorham's Cave includes silty dark-brown Paleo-environmental studies based on pollen and charcoal clays, which are rich in organic material, grey sand, irregularly samples reveal a varied Mediterranean landscape in both the bedded yellowish-brown sand, clay rich in organic matter with an Middle and Upper Palaeolithic (Carrion et al., 2008). Native vege- almost black colour and with phosphate crystals of whitish sand tation includes oaks, junipers and pine forests mixed with intercalated with massive, coarse and homogeneously brown sand savannah, grasslands with heaths, heliophysical scrub, phreato- (Collcutt, 2013). The radiocarbon dates give a chronological range phytic formations (such as wetlands and riparian forests) and between ca. 29 and 51 ka in the UBSm.7 and BeSm.1 units. However, Mediterranean coastal scrubland. the dates from the underlying LBSmff.1-5 (ca. 42 and 56 ka BP) The stratigraphic sequence shows no significant variations in indicate that most charcoal fragments could have been derived the taxonomic representation of macro-fauna, with the constant from lower down the sequence (Higham et al., 2013). The single- presence of the same species of ungulates and two predominant grain (SG) optically stimulated luminescence (OSL) age chronol- taxa, Cervus elaphus and Capra pyrenaica (Currant et al., 2013). Only ogy and the Bayesian model provided an age of MIS 5 near the base the presence of the grey seal (Halichoerus grypus) could constitute of the stratigraphy (119,300 ± 14,800 ka for CSM; Rhodes, 2013). evidence of a cool phase in the CHm unit (unit D in Waechter, 1951, Archaeological interventions inside the cave (inner sector) 1964). Regarding micro-vertebrates, at least 33 taxa of amphibians covered an area of ~29 m2, unearthing bedrock in the innermost and reptiles have been recovered, including 24 in the inner area, part and determining a stratigraphy composed of four archaeo- such as newts, toads, frogs, tortoises, turtles, lizards, geckos and logical horizons (IeIV from top to bottom; Fig. 2). The stratigraphic various snakes (Blain et al., 2013). However, the most significant composition differs greatly from that of the external sector as it groups in terms of quantity come from the LBSmcf.11 unit where a shows falls of autochthonous rocks, airborne dust and, principally, total of 21 species have been found, of which the main one is the karstic clay (Finlayson et al., 2006). Similarly, the sedimentary western spadefoot (Pelobates cultripes)(Gleed-Owen and Price, strength is lower, with a thickness of barely 2 m, mainly due to the 2013). Small mammals are present throughout the stratigraphic higher position of the substrate of the cave. Levels I and II corre- sequence, with a predominance of Oryctolagus cuniculus, Apodemus spond to Phoenician and Neolithic horizons, respectively. Level III sylvaticus, Eliomys quercinus, Microtus brecciencis and Terricola shows an average depth of ~60 cm and, archaeologically, is divided (Microtus) duodecimcostatus (Lopez-García et al., 2011; Price, 2013). into two sub-levels, corresponding to Solutrean (sublevel IIIb) at In the innermost area, there is a marked presence of Myotis myotis the bottom and Magdalenian (sublevel IIIa) on top. It is also (Lopez-García et al., 2011). possible to observe certain differences on a sedimentary level in the The outer area has contributed an important selection of birds, middle part of level III given the high proportion of fallen fragments with at least 90 registered species (seabirds, ducks, birds of prey, of angular limestone and speleothem. Levels III and IV differ in their partridges, waders, pigeons, common swifts, crows and small textural composition, with level III being sandier and mixed with passerines; Cooper, 2013). Excavations carried out in the inner area dark brown clay, while level IV is more clayey and characterised by of the cave complete the taxonomic list above and increase the a pure beige-coloured clay (Finlayson et al., 2006). number of taxa to 142 (Finlayson et al., 2014). Among the pre- Lithic industry studies conducted in the outer sector reveal a dominant species are pigeons and corvids, which are the focus of consistent set with Middle Palaeolithic techno-complexes for the the present study. R. Blasco et al. / Quaternary International 421 (2016) 62e77 65

Fig. 2. Geological sequence of Gorham's Cave e left: schematic profile of the outer sector (middle area of the cave) modified from Collcutt (2013); right: stratigraphic profile of the inner sector together to the general geological interpretative section of the site (NWeSE section) based on Jimenez-Espejo et al. (2013) and previous publications (e.g., Waechter, 1951, 1964; Goldberg and Macphail, 2000; Finlayson et al., 2006) showing the location of the excavated sectors (outer sector including the entrance and middle area of the cave, and inner sector [back of the cave]). Red boxes mark the archaeological levels/units presented here. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

3. Methods The proximal/distal ratio was used to assess the integrity of the sample according to their portions by dividing the total proximal The analysis of avian bones from Gorham's Cave was conducted elements (coracoids, scapula, humerus, femur, tibiotarsus) by the according to the published standards for taphonomy (e.g., sum of the proximal and distal elements (coracoid, scapula, hu- Laroulandie, 2000; Lyman, 2008). The NISP (Number of Identified merus, femur, tibiotarsus, radius, ulna, carpometacarpus, tarso- Specimens), MNE (Minimum Number of Elements), MNI (Minimum metatarsus), expressed as a percentage (*100) (Bochenski and Number of Individuals) and MAU (Minimum Animal Units) were Nekrasov, 2001). In addition, we performed a Kolmogor- calculated to estimate the relative abundance of various species, as oveSmirnov test (KS-test) of one sample to test if the series of ratio- well as the percentage of survival of skeletal elements (Lyman, values come from a uniform distribution in all the three anatomical 1994). The MAU percentage was used to standardise anatomical calculated ratios. profiles and compare the frequencies of several elements by taxa. Bone surfaces were macro- and microscopically inspected under Apart from the MNE and MAU, the wing to leg bone ratio was also a stereo light microscope (with a magnification of up to 60) with computed by calculating the percentage resulting from dividing the an oblique cold light source in order to document damage produced total number of wing remains (humerus, ulna and carpometa- by abiotic processes as well as both non-human and human agents. carpus) by the sum of the wing and leg (femur, tibiotarsus and The study was completed with the observation of selected speci- tarsometatarsus) remains (Ericson, 1987; Livingston, 1989). The mens at a magnification of up to 300 using an analytical FEI core to limb ratio was calculated by dividing the total core elements QUANTA 600 Environmental Scanning Electron Microscope (ESEM) (sternum, coracoid, scapula and pelvis) by the sum of the core and operated at low vacuum mode (LV). limb bones (humerus, femur, radius, ulna, tibiotarsus, carpometa- Fresh and dry fractures were analysed (Steadman et al., 2002; carpus, tarsometatarsus) (Bramwell et al., 1987; Bochenski, 2005). Laroulandie and Lefevre, 2014) to differentiate between possible 66 R. Blasco et al. / Quaternary International 421 (2016) 62e77 human/carnivore inputs and post-depositional fragmentation. 4. Results Fresh breakages were determined by bearing oblique angles and smooth edges, while the dry fractures were usually more trans- We examined a total of 2662 Columbidae and Corvidae speci- versal with straight, rough edges. In addition, the new breaks mens from the inner [levels III, IV] and outer areas [levels from produced during or after excavation were also registered and BeSm (Ossm).1 to SSLm (Usm).5] of Gorham's Cave. This sample defined by colour changes in the bone breakage planes. spans the temporal range from 67 ka to 28 ka (Finlayson et al., Fresh or green bone breakage on avian specimens can result 2006; Barton et al., 2013), coinciding with the occupation of the from several processes associated with both human activity and cave by Neanderthals and, subsequently, by modern humans. In carnivore actions. Despite the methodological framework now total, 21 discrete archaeological units were analysed taphonomi- being developed, the information currently available is still too cally, among which 20 were associated with Neanderthals and one scarce to completely isolate the breakages resulting from one agent with modern humans (level III from the inner area). Despite the from those generated by another (hominins versus carnivores). In remains being distributed along the sequence, the majority come this situation, the location of the fracture plane on the bone and its from the inner part (NISP ¼ 650 for level III and NISP ¼ 800 for level co-occurrence with other alterations (e.g., breaks associated with IV). Levels BeSm (PLSsm).3 (NISP ¼ 102), LBSmcf.2 (NISP ¼ 145), peeling) become relevant to identify its genesis and, therefore, the LBSmcf.4 (NISP ¼ 274), LBSmcf.9 (NISP ¼ 210) and SSLm (Usm).5 responsible agent. Several human activities on birds can produce (NISP ¼ 102) are also important in terms of number of specimens. fresh bone breakage, such as disarticulation or the removal of fat The remaining levels contain from 2 to 81 bones at most (Table 1). and/or cartilage. The experimental and archaeological cases pro- The Columba genus is the most abundant (NISP ¼ 1724), inte- vided by Laroulandie et al. (2008) are an example of how some grating 1364 specimens from Mousterian contexts and 360 from breakages are produced during the dismembering of the forelimbs the Upper Palaeolithic. Despite some remains of Columba palumbus by using overextension of the elbow. This activity usually produces being identified, the majority of the Columba bones correspond to a breakdown of the olecranon fossa of the humerus, with a medial Columba livia/oenas and, for this reason, they are all grouped as wrench of the distal part, and a fracture of the proximal joints of Columba in the data analysis. The same is true of the Pyrrhocorax the radius and ulna. In addition, other types of damage can appear group. Among corvids, the genus Pyrrhocorax stands out with 804 associated, for example peeling, which was defined by White remains (518 from Mousterian contexts). Other specimens from the (1992) as roughened surfaces with parallel grooves and a fibrous Corvidae family are present, but their representation is low when texture. This co-occurrence of alterations on the zones adjacent to compared with the others. Regarding the total MNI, levels III, IV and the area of breakage of the articular ends was grouped here as LBSmcf.4 accumulate 53.22% of the MNI (66, 82 and 42 MNI, over-extending damage and listed in Table 1 as BBr (Bone respectively). The Columba genus is still the best represented in all Breakage). the studied levels, reaching more than 50% of the MNI in the main Humans can also produce tooth marks on avian bones as a assemblages. Among corvids, the Pyrrhocorax genus represents consequence of direct consumption of meat, fat or cartilage (e.g., more than 65% of the total corvid MNI, if all the levels are consid- Laroulandie, 2001, 2005a; Blasco and Fernandez Peris, 2009; Blasco ered together. In the inner part, Pyrrhocorax represents nearly 100% et al., 2014). Given the potential risk of confusion with carnivore of all corvids (93.10% of corvids at level III and 92.59% at level IV). gnawing, human tooth-marks were identified using the criteria The sum of the MNE is 2376, roughly half of which is Columba in proposed by Laroulandie (2005a) and following a systematic all the levels. They are closely followed by the Pyrrhocorax genus. In comparison with bone damage generated by mammal carnivores general, Columba and Pyrrhocorax have representation of all the and raptors (e.g., Bochenski, 1997; Bochenski et al., 1998, 2009; body elements, with the exception of pelvises. Nonetheless, if the Laroulandie, 2000; Mallye et al., 2008; Monchot and Gendron, percentage of the MAU is considered, the representation changes 2013; Rodríguez-Hidalgo et al., 2016). Carnivore modifications slightly. As a general trend, cranial and axial skeletons, as well as were recorded in the form of pits, punctures, scores, crenulated acropodials, are under-represented. Nevertheless, it should be edges and beak-marks, alongside their quantification and location taken into account that the pedal phalanges are difficult to assign to on the skeleton and anatomical region. Chemical alterations pro- one or another species and this circumstance may have caused an duced by gastric acids were also listed as a result of carnivore ac- under-representation here. Despite the absence of pelvises, girdles tivities on a scale of five degrees of corrosion: degree 0 refers to a present important percentages of representation in the levels with complete lack of alteration and degree 4 refers to bones with a higher number of NISP. This is due to the predominance of cor- extreme corrosion undergoing a severe degree of destruction acoids and scapulae. In the inner part of the cave, (MAU) coracoids (Andrews, 1990). and humeri present higher percentages in the Columba genus: We also searched for other human evidence, such as cut marks, 90.54% (MNE ¼ 67) and 68.18% (MNE ¼ 75) of coracoids at levels III according to the criteria described by Shipman and Rose (1983), and IV, respectively; and 98.65% (MNE ¼ 73, level III) and 89.09% Lyman (2008) and Domínguez-Rodrigo et al. (2009). These au- (MNE ¼ 98 level IV) of humeri. Conversely, ulnae and carpometa- thors define cut marks as striations showing a V-shaped section, carpi are more important in the Pyrrhocorax genus. Ulnae of Pyr- internal microstriation and a linear outline of variable length, rhocorax are present in 94.44% (MNE ¼ 51, level III) and 82% width, and depth. In some cases, incisions show Hertzian cones (MNE ¼ 41, level IV) of cases, while carpometacarpi exist in 70.37% (Bromage and Boyde, 1984), shoulder effects, barbs and narrowing (MNE ¼ 38, level III) to 82% (MNE ¼ 41, level IV). The outer area that indicates directionality (Shipman and Rose, 1983). The char- shows an important number of coracoids at levels LBSmcf.2 (83.3% acteristics of cut marks such as type, morphology, number of stri- or MNE ¼ 20) and LBSmcf.9 (80% or MNE ¼ 24) in Columbiformes, ations, location and orientation regarding the longitudinal axis of although the sternum is more or equally important at levels the bone (oblique, longitudinal, transverse) were also recorded. LBSmcf.2 (83.3% or MNE ¼ 10) and SSLm (Usm).5 (60% or MNE ¼ 6) Thermal-alterations were analysed in terms of presence/ in this group, as well as scapulae at LBSmcf.4 (72.73% or MNE ¼ 32 absence and based mainly on colour changes and other physical represented in the Columba group). On the other hand, the Pyr- alterations produced during exposure to fire, such as deformation, rhocorax genus, which is the second most important group repre- cracks and/or fissures (e.g., Laroulandie, 2000, 2005a, 2005b). Any sented, exhibits high values of carpometacarpi at LBSmcf.4 (68.75% burning damage identified was arranged on a scale of six degrees or MNE ¼ 11) and tarsometatarsi at level LBSmcf.9 (71.43% or (from degree 0 [unburned] to degree 5 [calcined]). MNE ¼ 10) (see Supplementary data for details). Table 1 Number of specimens attributed to genus Columba (cf. Columba livia/oenas and Columba palumbus), Corvus (Corvus monedula, Corvus corone/frugilegus and Corvus corax), Pica (Pica pica) and Pyrrhocorax (P. pyrrhocorax and P. graculus) from Gorham's Cave, Gibraltar (see Supplementary data for details).

a Units Columba Corvus/Pica Pyrrhocorax Total n % NISP MNE MNI Cm BBr Burn Carniv NISP MNE MNI Cm BBr Burn Carniv NISP MNE MNI Cm BBr Burn Carniv NISP MNE MNI Cm BBr Burn Carniv Cm BBr Burn Carniv c,e III 360 335 37 5 8 21 7 4 4 2 1 286 261 27 8 12 13 650 600 66 13 9 33 20 2.00 1.38 5.08 3.08 IVc,e 481 426 55 16 6 43 8 2 2 2 317 287 25 15 4 27 18 800 715 82 31 10 70 26 3.88 1.25 8.75 3.25 BeSm (Ossm).1 2 2 1 221 62 (2016) 421 International Quaternary / al. et Blasco R. BeSm (PLSsm).3 81 70 8 2 9 2 1 1 1 20 18 3 1 1 102 89 12 3 10 2 2.94 9.80 1.96 LBSmff.1 10 9 3 5 5 2 1 2 15 14 5 1 2 6.67 13.33 c LBSmcf.1 3 2 1 1 1 1 1 4 3 2 1 25.00 LBSmcf.2b,c,d 116 93 12 5 2 27 13 11 4 4 16 16 5 1 1 145 120 21 5 2 32 1 3.45 1.38 22.07 0.69 LBSmcf.4b,c,d,f 195 157 22 2 5 58 30 29 12 1 12 49 46 8 1 11 274 232 42 4 5 81 1.46 1.82 29.56 LBSmcf.1-4gend 11 9 2 1 1 1 1 1 1 1 13 11 4 1 7.69 LBSmcf.5b,c,d,f 52 47 7 1 5 14 14 6 1 1 15 14 3 1 81 75 16 1 6 2 1.23 7.41 2.47 LBSmcf.6 17 17 3 4 4 2 1 5 5 2 1 26 26 7 1 1 3.85 3.85 LBSmcf.7 22 20 2 2 5 5 3 1 4 4 1 1 31 29 6 1 3 3.23 9.68 LBSmcf.8 47 41 3 1 3 3 3 2 6 6 2 56 50 7 1 3 1.79 5.36 b,c,d LBSmcf.9 160 130 15 1 4 16 14 3 1 34 32 7 1 1 210 176 25 1 1 5 1 0.48 0.48 2.38 0.48 LBSmcf.10d 3 3 1 3 3 3 5 5 2 1 11 11 6 1 9.09 LBSmcf.11b,c,d,f 58 53 8 1 8 7 4 1 14 13 4 80 73 16 2 2.50 LBSmcf.12c 29 27 4 4 4 4 2 6 6 3 1 39 37 9 5 12.82 LBSmcf.13b,c,f 8 8 3 1 1 1 1 6 6 3 15 15 7 1 6.67 SSLm (Usm).2 2 2 1 1 1 1 3 3 2 SSLm (Usm).3 2 2 1 1 1 1 1 1 3 3 2 1 1 33.33 33.33 b,c,d SSLm (Usm).5 67 58 10 2 2 2 18 18 4 1 1 17 16 5 2 102 92 19 3 5 2 2.94 4.90 1.96 e

n-Total 1724 1509 198 33 23 179 21 134 128 55 2 2 22 4 804 739 104 27 5 59 33 2662 2376 357 62 30 260 58 2.33 1.13 9.77 2.18 77 MP units 1364 1174 161 28 15 158 14 130 124 53 2 1 22 4 518 478 77 19 5 47 20 2012 1776 291 49 21 227 38 2.44 1.04 11.28 1.89 UP units 360 335 37 5 8 21 7 4 4 2 1 286 261 27 8 12 13 650 600 66 13 9 33 20 2.00 1.38 5.08 3.08

Note that some bones can show co-occurrence of modifications. a Stratigraphic units/levels taken from Finlayson et al. (2006) and Barton et al. (2013). b Units containing Columba palumbus specimens: LBSmcf.2 ¼ 2; LBSmcf.4 ¼ 2; LBSmcf.5 ¼ 3; LBSmcf.9 ¼ 1; LBSmcf.11 ¼ 3; LBSmcf.13 ¼ 1; SSLm (Usm).5 ¼ 4. c Units containing Pyrrhocorax graculus: III ¼ 81; IV ¼ 85; LBSmcf.1 ¼ 1; LBSmcf.2 ¼ 1; LBSmcf.4 ¼ 2; LBSmcf.5 ¼ 3; LBSmcf.9 ¼ 1; LBSmcf.11 ¼ 2; LBSmcf.12 ¼ 1; LBSmcf.13 ¼ 1; SSLm (Usm).5 ¼ 2. d Units containing Corvus corone/frugilegus: LBSmcf.2 ¼ 6; LBSmcf.4 ¼ 2; LBSmcf.1-4gen ¼ 1; LBSmcf.5 ¼ 2; LBSmcf.9 ¼ 1; LBSmcf.10 ¼ 1; LBSmcf.11 ¼ 1; SSLm (Usm).5 ¼ 3. e Units containing Corvus corax: III ¼ 2; IV ¼ 1. f Units containing Pica pica: LBSmcf.4 ¼ 4; LBSmcf.5 ¼ 1; LBSmcf.11 ¼ 2; LBSmcf.13 ¼ 1. Cm ¼ Cut marks; BBr ¼ Bone breakage by over-extension; Burn ¼ Burnt bones; Carniv ¼ Carnivore damage; MP ¼ Middle Palaeolithic; UP ¼ Upper Palaeolithic. 67 68 R. Blasco et al. / Quaternary International 421 (2016) 62e77

Table 2 Wing/leg, core/limb and proximal/distal elements ratios from stratigraphic levels with the highest quantity of avian specimens.

Units Genus Wings Legs Wings/legs Core Limbs Core/limb Prox elements Dist elements Prox/dist

III Columba 169 68 71.31 99 248 28.53 213 124 63.20 Corvus 1 1 50.00 e 2 e 2 1 66.67 Pyrrhocorax 138 73 65.40 43 216 16.60 123 145 45.90 IV Columba 219 103 68.01 129 340 27.51 287 164 63.64 Corvus 2 ee e 2 ee 2 e Pyrrhocorax 145 71 67.13 60 232 20.55 158 124 56.03 LBSmcf.2 Columba 33 25 56.90 51 58 46.79 73 26 73.74 Corvus 4 4 50.00 e 2 e 8 5 61.54 Pyrrhocorax 7 5 58.33 5 8 38.46 7 8 46.67 LBSmcf.4 Columba 56 40 58.33 76 101 42.94 106 63 62.72 Corvus/Pica 16 6 72.73 6 23 20.69 8 15 34.78 Pyrrhocorax 19 16 54.29 12 36 25.00 24 23 51.06 LBSmcf.9 Columba 56 31 64.37 54 92 36.99 84 55 60.43 Corvus 5 6 45.45 e 2 e 8 5 61.54 Pyrrhocorax 12 13 48.00 5 11 31.25 12 24 33.33 KeS test 0.985 0.963 0.385

Prox ¼ proximal; dist ¼ distal.

The wing to leg ratio was calculated in the levels that had suf- Human-induced damage is particularly important in Colum- ficient bones to record e that is, level III, level IV, LBSmcf.2, bidae, where it affects 9.44% (NISP ¼ 34) of Columba from level III, LBSmcf.4 and LBSmcf.9. Levels III and IV show a clear predominance 13.51% (NISP ¼ 65) at level IV, 29.31% (NISP ¼ 34) at LBSmcf.2, and of wings over legs in both genera: Columba (71.31% at III and 68.01% 33.33% (NISP ¼ 65) at LBSmcf.4. Pyrrhocorax exhibits high values of at IV) and Pyrrhocorax (65.4% at III and 67.13% at IV). Other corvids damage in the inner levels (6.99% at III and 14.51% at IV), but the are not present in sufficient numbers to compute a reliable ratio. In most significant percentages of human modifications are at the outer area (LBSmcf.2, LBSmcf.4 and LBSmcf.9), the Colum- LBSmfc.4, where anthropogenic damage can be observed on 24.49% biformes wing/leg ratio shows percentages of 56.9%, 58.33% and of the Pyrrhocorax remains (NISP ¼ 12), as well as in 43.33% of other 64.36%, respectively. Otherwise, Pyrrhocorax shows a predomi- corvid elements (NISP ¼ 13). On the other hand, level LBSmcf.9, nance of wings at LBSmcf.2 (58.33%) and LBSmcf.4 (54.29%), which was important in terms of the NISP, is one of the levels with although legs are more important in LBSmcf.9 (wing/leg ratio: 48%). less representation of human activity (3.33% or NISP ¼ 7). However, the values are near to 50% representation in the afore- Cut marks and thermo-alterations are the most frequent mod- mentioned levels. In the case of the core/limb ratio, there is always ifications on bones. In the case of the Neanderthal occupation units, a prevalence of limb elements over core fragments. The ratios we detected cut-marks on 49 bird bones (2.44%), 31 from level IV estimated never exceed 46.79% of core remains (Columba from and 18 from the outer area (Table 1; Table 3; Fig. 3). The specific LBSmcf.4), and they can be even lower in most of the levels (16.60% percentage of incisions can oscillate between 0.48% (NISP ¼ 1 for in Pyrrhocorax at level III). The proximal to distal elements ratio LBSmcf.9) and 3.88% (NISP ¼ 31 for IV). Despite being more shows a slight predominance of proximal elements in Columba of important in terms of the number of remains in Columba (NISP ¼ 28 all the levels. Values oscillate between 60.43% for LBSmcf.9 and or 2.05% of Columbiformes), cut-marked elements exist in higher 73.74% for LBSmcf.2. For Pyrrhocorax, the values of proximal ele- percentages on Pyrrhocorax (3.67% of total NISP; NISP ¼ 19), to the ments are near to a 1:1 proportion (from 45.9% for level III to 56.03% extent that in level LBSmcf.9 Columba specimens do not show cut for level IV). Only level LBSmcf.9 was characterised by the especial mark evidence. On the contrary, level LBSmcf.2 only shows cut- prevalence of distal elements, with a 33.3% representation. Never- marked bones on the Columbidae group. As previously discussed, theless, the number of remains in that group is too low (12 prox- level LBSmcf.4 is also relevant in terms of cut marks (1.46% or imal and 24 distal elements) to compute a reliable ratio. In addition, NISP ¼ 4). In this unit, the cut marks are distributed among the results of the KS-test show not significant differences in any of different taxa: 1.03% (NISP ¼ 2) of Columba, 3.33% (NISP ¼ 1) of the three ratios, suggesting homogeneity in the values series Corvus/Pica and 2.04% (NISP ¼ 1) of Pyrrhocorax. Regarding level III, (Table 2). 13 avian bones show cut marks (2%), with special predominance on We found evidence of human intervention on both corvid and Pyrrhocorax exhibiting 8 bones with incisions (1.23%) while pigeon bones in 16 (80%) of the Neanderthal units, as well as in the Columba shows 5 specimens (0.77%). In contrast, no cut marks are modern human context. There was no observable difference in the detected on bones attributed to Corvus or Pica. Wing elements are tendency of damaged bones between Neanderthals and modern the most cut-marked, with humeri being 38.46% affected at level III humans (Table 1). In total, anthropogenic damage affects 13.22% of (NISP ¼ 5) and 41.94% at level IV (NISP ¼ 13). Ulnae are also the remains analysed. It is especially important in the inner levels important at level IV (25.81% cut-marked, NISP ¼ 8). Other bones (8.46% and 13.88% for levels III and IV, respectively), as well as in the such as coracoids, femora, tibiotarsi, fibulae, carpometacarpi, tar- outer levels LBSmcf.2 (26.9%) and LBSmcf.4 (32.85%). The latest level sometatarsi or sterna show cut-marks less frequently. Butchery is the one with a significant number of altered remains, also marks usually present as oblique or transversal incisions on prox- considering the NISP. Levels LBSmff.1 (fine facies) and LBSmcf.1 imal, distal or mid-shafts. Occasionally, they can be found at the (coarse facies) from the outer area show a high degree of anthro- ends of long bones (Table 3). When the marks are located on leg or pogenic damage, but their representation, considering the general core bones, they are always associated with defleshing activities NISP of the level, is not significant (NISP ¼ 15 and NISP ¼ 4, due to their meat content. In the case of wing bones, feather respectively). The other levels [BeSm (Ossm).1, LBSmcf.11, LBSmcf.13, removal has been linked to cut marks on ulnae and carpometacarpi, and SSLm (Usm).2] do not show signs of hominid activities, although as well as humeri in the case of raptors. Nonetheless, defleshing most of them have a low NISP. Only LBSmcf.11 could be outlined activity in flapping birds can also be found on humeri and ulnae, as since, with an NISP of 80, any human activity has been recorded. a higher proportion of meat tends to cover these bones. R. Blasco et al. / Quaternary International 421 (2016) 62e77 69

Fig. 3. Cut-marked bones of pigeons and corvid specimens from Gorham's Cave: proximal sternum of Columba livia/oenas from level IV (A); distal ulna of Corvidae from SSLm (Usm).5 (B) and ESEM view of the cut mark (B1); distal femur of Pyrrhocorax pyrrhocorax from level IV (C); distal humerus of Corvidae from LBSmcf.4 (D); ulna of Columba livia/oenas from level IV (E).

Burnt bones represent 9.77% of the total assemblages at Gor- Regarding the most significant levels, grade 2 coloration stands out ham's Cave (NISP ¼ 260): 227 come from Mousterian contexts in the inner sequence with percentages from 71.43% (NISP ¼ 15) for (11.3%) and 33 from level III (5.08%) (Table 1; Table 4). In the case of level III to 41.86% (NISP ¼ 18) for level IV in Columbidae. Similar the Neanderthal occupation units, burning affects 11.58% values are presented for Pyrrhocorax. At level LBSmcf.4, for (NISP ¼ 158) of Columba remains, 16.9% (NISP ¼ 22) of Corvus/Pica, example, 2e3 double colorations reach 13.79% (NISP ¼ 8) in and 9.07% (NISP ¼ 47) of Pyrrhocorax. Meanwhile, level III shows Columba and 9.09% (NISP ¼ 1) in Pyrrhocorax. At level LBSmcf.2, thermal alterations in 5.83% (NISP ¼ 21) of Columba remains and double colorations are important among Columba elements rising 4.2% (NISP ¼ 12) of Pyrrhocorax. In contrast, Corvus/Pica bones show to 18.5% of the total Columba burnt bones (Table 4). no alteration due to exposure to fire in this unit. In the most Apart from butchery marks and burning, bone breakage can be representative levels, the percentage of burnt bones oscillates be- interpreted as a consequence of human activity when associated tween 2.38% (NISP ¼ 5, LBSmcf.9) and 22.07% (NISP ¼ 32, LBSmcf.2) with human tooth marks or damage related to disarticulation or 29.56% (NISP ¼ 81, LBSmcf.4). At levels LBSmcf.2 and LBSmcf.4, processes, such as peeling on the ulna and holes by wrenching on the burning values are homogeneously distributed among all the the distal end of the humerus as a result of elbow over-extension taxa analysed. Colorations due to fire exposure are present on long (Laroulandie et al., 2008)(Table 1; Fig. 4). In that sense, bone bones, as well as girdles and phalanges, with especial incidence on breakage has been observed in 30 elements (1.13% of the total humeri, ulnae, carpometacarpi and femora. Different degrees of studied specimens). At levels III, IV and LBSmcf.4, 1.38% (NISP ¼ 9), coloration can be observed, although degrees 2 and 3 stand out in 1.25% (NISP ¼ 10) and 1.82% (NISP ¼ 5) of the remains show this all the units. Some 38.33% (NISP ¼ 87) of the altered remains pre- modification. Humeri (NISP ¼ 9) and ulnae (NISP ¼ 17) are the sent degree 2 of coloration, while 37.44% (NISP ¼ 85) exhibit degree most frequently affected. Level IV stands out in terms of the 3, all of them from Mousterian contexts. Level III shows 69.70% number of humeri (NISP ¼ 4) and ulnae (NISP ¼ 5) with this (NISP ¼ 23) with degree 2 and 18.18% (NISP ¼ 6) with degree 3. modification. Such a modification is also present on ulnae at level Double colorations are also important (NISP ¼ 42 or 16.15% of III (NISP ¼ 7). Level LBSmcf.4 exhibits bone breakage for humeri charred bones), which mainly stem from the outer area and level IV (NISP ¼ 2) and ulnae (NISP ¼ 2), as well as other levels, but is less (NISP ¼ 39 or 92.86% of bones bearing double colourations). representative. 70 R. Blasco et al. / Quaternary International 421 (2016) 62e77

Table 3 Cut-marked pigeon and corvid specimens from Gorham's Cave.

Units Genus NISP total NISP Cm Skeletal element [NISP] No striae by group Distribution Type Region location Orientation Activity

III Columba 360 5 Humerus [4] 1e7 isol, cl inc mid-distal shaft obl, trans Fr, Df Tibiotarsus [1] 3 cl inc mid-shaft obl Df Pyrrhocorax 286 8 Femur [2] 1 isol inc mid-distal shaft obl, trans Df Fibula [1] 5 cl inc distal shaft trans Df Humerus [1] 1 isol inc mid-shaft trans Fr, Df Tarsometatarsus [1] 2 cl inc distal shaft trans Df Tibiotarsus [1] 8 cl inc mid-shaft obl Df Ulna [2] 2e3 cl inc prox shaft trans Fr IV Columba 481 16 Coracoides [1] 2 cl inc body trans Df Humerus [10] 1e9 isol, cl inc prox, mid-distal shaft obl, trans, long Fr, Df Sternum [1] 4 isol inc prox area trans Df Tibiotarsus [1] 5 cl inc mid-shaft trans Df Ulna [3] 2e10 cl inc prox-mid shaft obl, trans Fr Pyrrhocorax 317 15 Carpometacarpus [1] 1 isol chop distal shaft trans Fr Coracoides [2] 5e8 cl inc prox-mid shaft obl, trans Df Femur [1] 2 cl inc distal shaft trans(obl) Df Fibula [2] 3 cl inc prox shaft obl Df Humerus [3] 3e5 cl inc mid-distal shaft obl, trans Fr, Df Tarsometatarsus [1] 2e3 cl inc prox-mid shaft trans Df Ulna [5] 1e7 isol, cl inc prox-mid shaft obl, trans Fr BeSm (PLSsm).3 Columba 81 2 Humerus [1] 2 cl inc distal shaft obl Fr Femur [1] 3 cl inc mid-shaft trans Df Pyrrhocorax 20 1 Carpometacarpus [1] 1 isol inc mid-shaft obl Fr LBSmcf.2 Columba 116 5 Carpometacarpus [1] 1 isol inc mid-shaft trans Fr Coracoides [1] 2 cl inc distal shaft (distal end) obl Df Humerus [2] 1e3 isol, cl inc prox end, mid-shaft obl Fr Tibiotarsus [1] 5 cl inc mid-shaft obl Df LBSmcf.4 Columba 195 2 Humerus [1] 2 cl inc mid-shaft obl Fr Tarsometatarsus [1] 5 cl inc prox-mid shaft trans Df Corvus/Pica 13 1 Humerus [1] 2 cl inc distal shaft obl Fr Pyrrhocorax 16 1 Carpometacarpus [1] 1 isol inc mid-shaft trans Fr LBSmcf.8 Columba 47 1 Ulna [1] 1 isol inc distal shaft trans Fr LBSmcf.9 Pyrrhocorax 34 1 Ulna [1] 3 cl inc prox shaft obl Fr SSLm (Usm).3 Pyrrhocorax 1 1 Humerus [1] 3 cl inc prox-mid shaft trans Fr SSLm (Usm).5 Columba 67 2 Femur [1] 1 isol chop distal shaft trans Df Humerus [1] 1 isol inc distal shaft obl Fr Corvidae 18 1 Ulna [1] 1 isol inc distal end obl Fr

Cm ¼ cut marks; cl ¼ clustered; cr ¼ crossed; isol ¼ isolated; inc ¼ incisions; chop ¼ chop marks; obl ¼ oblique; long ¼ longitudinal; tr ¼ transverse; Df ¼ Defleshing; Fr ¼ Feather removal.

The activity of non-human predators has been documented on 5. Discussion 58 avian bones (2.18%) at Gorham's Cave, including pits, punctures, scores, crenulated edges and digestive damage (Table 1; Table 5; As previous studies have pointed out (Finlayson et al., 2012; Fig. 5). The incidence is higher in the inner area, with 3.08% Blasco et al., 2014), the processing of birds by Neanderthals at the (NISP ¼ 20) and 3.25% (NISP ¼ 26) at levels III and IV, respectively. Gibraltar sites is a widespread phenomenon. The present study Regarding the outer area, those levels with significant NISP do not demonstrates that humans did not only focus their food activities on show more than 2% of damage on bones. Only level LBSmcf.11 can pigeons at Gorham's Cave, but also on corvids from those levels with be particularly noted as it has more carnivore incidence (2.5%, or abundant specimens. The presence of cut marks on lower and fore NISP ¼ 2) and no evidence of anthropogenic activity, but the total limb bones, and burning patterns, as well as over-extension of the NISP of the level is still low (NISP ¼ 80). The Pyrrhocorax genus is elbow, are direct evidence of that. Nevertheless, hominid activity on the one bearing more modifications, 4.55% (NISP ¼ 13) of Pyr- bones is not the same in all the levels and some appreciations should rhocorax remains at level III and 5.68% (NISP ¼ 18) at level IV. Form be done, because several taphonomic agents other than humans the outer area, level LBSmcf.4 is important because any modifica- may have also contributed to the Gorham's Cave bird accumulations. tion related to carnivores (mammalian carnivores or raptors) has Anatomical representation could provide us with clues about been documented. It is also a level with an important proportion of how to interpret archaeological assemblages and that is why anthropogenic activity. In the other units, mechanical modifica- several researchers have proposed methods to differentiate human tions (pits, punctures, scores) are normally found isolated, either on from non-human accumulations based on skeletal ratio (e.g., the epiphyses or on proximal, distal or mid-shafts. They mainly Mourer-Chauvire, 1983; Ericson, 1987; Bochenski, 2005). For affect long bones, but occasionally coracoids and sternums. example, Mourer-Chauvire (1983) suggested that in archaeological Although digestive damage has been found in some Mousterian contexts where human activity is attested on bird remains, ele- contexts, its rate of occurrence is low (NISP ¼ 13; 0.65% of the total ments such as coracoids, humeri and femora have higher fre- analysed remains), showing only slight and moderate grades of quencies of representation. This is because, when roasting, these corrosion (Table 5). elements are covered by meat, facilitating their preservation. On Beyond carnivore marks, rodent gnawing was also detected, the contrary, those bones directly in contact with the fire may although its presence is testimonial with only 3 damaged bones disappear as a result of fire exposure. This may be the case for (0.11%), all of them come from the inner area (1 from level III and 2 Gorham's Cave. Coracoids and humeri are the most abundant re- from level IV) (Fig. 5). mains at units III and IV, especially in the Columba group. In the R. Blasco et al. / Quaternary International 421 (2016) 62e77 71

Table 4 Burnt pigeon and corvid specimens from Gorham's Cave.

Stratigraphic units Genus NISP total NISP burnt Degree 1 Degree 2 Degree 3 Degree 4 Degree 5 Double colouration %NISP

0e20e32e3

III Columba 360 21 1 15 4 1 5.83 Pyrrhocorax 286 12 8 2 2 4.20 IV Columba 481 43 5 18 14 2 1 3 8.94 Pyrrhocorax 317 27 3 15 5 1 3 8.52 BeSm (PLSsm).3 Columba 81 9 2 5 1 1 11.11 Pyrrhocorax 20 1 1 5.00 LBSmff.1 (fine facies) Corvus 5 2 1 1 40.00 LBSmcf.1 (coarse facies) Columba 31 1 33.33 LBSmcf.2 Columba 116 27 5 15 1 1 5 23.28 Corvus 13 4 1 2 1 30.77 Pyrrhocorax 16 1 1 6.25 LBSmcf.4 Columba 195 58 27 21 2 8 29.74 Corvus/Pica 30 12 2 3 7 40.00 Pyrrhocorax 49 11 3 5 1 1 1 22.45 LBSmcf.1e4 general Corvus/Pica 1 1 1 100.00 LBSmcf.5 Columba 52 5 1 1 1 1 1 9.62 Corvus 14 1 1 7.14 LBSmcf.6 Pyrrhocorax 5 1 1 20.00 LBSmcf.7 Columba 22 2 2 9.09 Corvus 5 1 1 20.00 LBSmcf.8 Columba 47 3 2 1 6.38 LBSmcf.9 Columba 160 4 1 1 2 2.50 Pyrrhocorax 34 1 1 2.94 LBSmcf.10 Pyrrhocorax 5 1 1 20.00 LBSmcf.12 Columba 29 4 4 13.79 Pyrrhocorax 61 1 16.67 SSLm (Usm).3 Pyrrhocorax 11 1 100.00 SSLm (Usm).5 Columba 67 2 1 1 2.99 Corvus 18 1 1 5.56 Pyrrhocorax 17 2 1 1 11.76 260 13 110 91 3 1 5 8 29 other units where the NISP is sufficiently significant, coracoids are representation is normally low, which does not fit with Mourer- still important, but other elements such as ulnae, carpometacarpi Chauvire's hypothesis for anthropogenic accumulations. It also or scapulae are also significant (LBSmcf.2, LBSmcf.4, LBSmcf.9). does not fit with Ericson's (1987) proposals, which emphasise the Femora never stand out over other elements and their importance of leg elements in the human assemblages. However, it

Fig. 4. Co-occurrence between break down and peeling on the proximal part of ulnae of Columba livia/oenas from LBSmcf.4 (A) and Pyrrhocorax pyrrhocorax from level IV (B) probably produced by over-extension of the elbow. 72 R. Blasco et al. / Quaternary International 421 (2016) 62e77

Table 5 Carnivore damage on pigeon and corvid specimens from Gorham's Cave.

Units Genus NISP NISP Skeletal element Type No. tooth-marks Distribution Region location total Carniv [NISP]

III Columba 360 7 Coracoides [1] Crenulated edge 1 notch isol body Humerus [3] Pit, puncture, score 1e10 isol, cl prox and mid-shaft, prox end Sternum [1] Pit 1 isol apophysis Tmt [1] Pit 1 isol distal end Ulna [1] Pit 1 isol prox end Pyrrhocorax 286 13 Cmt [2] Score 2e12 cl, gen prox and mid-shaft Femur [1] Crenulated edge 2 notches cl mid-shaft Humerus [5] Crenulated edge, pit, puncture 1e2, 1 notch isol, cl prox and distal end, mid-shaft Ulna [5] Crenulated edge, pit, score 1e10 isol, cl, disp prox, distal and mid-shaft IV Columba 481 8 Coracoides [1] Pit 1 isol distal end Humerus [5] Crenulated edge, pit, puncture 1e3 isol, cl prox and distal end, mid-shaft Sternum [1] Crenulated edge 1 isol body Ulna [1] Score 11 cl mid-shaft Pyrrhocorax 317 18 Cmt [2] Pit, digestion [degree 1] 1 isol mid-shaft Coracoides [1] Score 10 cl distal shaft Femur [2] Pit 1 isol prox shaft, distal end Humerus [5] Crenulated edge, pit, score 1e6 isol, cl prox, distal and mid-shaft Tmt [1] Pit 1 isol distal shaft Tbt [1] Pit 7 gen distal shaft Ulna [6] Pit, score, puncture, 1e12 isol, disp, cl distal and mid-shaft, distal end digestion [degree 2] BeSm (PLSsm).3 Columba 81 2 Humerus [1] Puncture 1 isol prox end Coracoides [1] Crenulated edge 1 notch isol body LBSmcf.2 Pyrrhocorax 16 1 Ulna [1] Puncture 1 isol distal shaft LBSmcf.5 Corvus 14 1 Femur [1] Digestion [degree 1] e cl distal end Pyrrhocorax 15 1 Ulna [1] Digestion [degree 1] e cl prox end LBSmcf.6 Corvus 4 1 Ulna [1] Puncture 2 opp prox shaft LBSmcf.9 Corvus 16 1 Ulna [1] Digestion [degree 2] e cl prox end LBSmcf.11 Columba 58 1 Coracoides [1] Digestion [degree 1] e cl end Corvus/Pica 8 1 Coracoides [1] Crenulated edge 1 isol body LBSmcf.13 Columba 8 1 Ulna [1] Pit, score 2e9 cl mid and distal shaft SSLm (Usm).5 Columba 67 2 Tbt [1] Digestion [degree 2] e cl prox end Femur [1] Pit 2 cl mid-shaft

Cl ¼ clustered; isol ¼ isolated; gen ¼ generalize; disp ¼ dispersed; opp ¼ opposite; Cmc ¼ Carpometacarpus; Tbt ¼ Tibiotarsus; Tmt ¼ Tarsometatarsus.

should be taken into account that although the methods based on of this is the study conducted by Monchot and Gendron (2013) on skeletal ratio have been demonstrated to be relevant in some bird bones from an active fox den in the Hudson Strait, in which the archaeological contexts, they are usually subject to criticism and wing elements predominated over legs in the surroundings of the should be used with caution, always as a supplementary approach den. In contrast, Rodríguez-Hidalgo et al. (2016) observed an in- (e.g., Livingston, 1989; Laroulandie, 2000, 2010). Nevertheless, in crease of leg bones in a non-ingested red-legged partridge (Alectoris our case, the results from the skeletal profile could lead us to rufa) accumulation generated by the Iberian Lynx under captive propose that, apart from humans, other predators could have breeding programmes; however, given the risk of establishing introduced elements into the assemblage. analogies between captive carnivores and wild ones (because of Despite long bones being the best represented elements in all differences in environmental contexts and possible alteration of archaeological levels at Gorham's Cave, the importance of wing behaviour patterns and their resulting bone damage; see a dis- elements over leg elements must be noted. Apart from the studies cussion on large carnivores in Gidna et al. [2013] and Arilla et al. mentioned above (Mourer-Chauvire, 1983; Ericson, 1987), the [2014]), their results should be used with caution. Mallye et al. proportion of wing elements to leg ones is usually related to ac- (2008) also attempted to provide data by studying accumulations cumulations produced by birds of prey (e.g., Bochenski et al., 1993, generated by foxes and badgers, which included leporid and avian 1997, 1998, 1999, 2009; Bochenski and Tomek, 1994; Bochenski, bones, among other animals. These authors were not able to 1997; Bochenski and Nekrasov, 2001; Bochenski and Tornberg, differentiate the bones modified by one predator from those altered 2003). Considering the low incidence of digestive corrosion at by the other, and so the information provided is of limited useful- Gorham's Cave (0.65% at Mousterian units and no alteration at level ness when isolating a specific predator. The fact is that few detailed III), diurnal raptors, such as eagles and falcons, may be potential studies have been conducted on mammalian carnivore accumula- candidates (Bochenski et al., 1997, 1999; Bochenski and Tornberg, tions, making identifying them as a possible generating agent of 2003; Bochenski, 2005). Despite the Golden Eagle having cora- avian assemblages complicated (Laroulandie, 2000; Mallye et al., coids and humeri as dominant elements, it has been discarded as a 2008; Monchot and Gendron, 2013). In any case, the mammalian possible predator in the cave because the values it presents for carnivore presence is confirmed at Gorham's Cave by recording a anatomical representation are even more extreme (Bochenski et al., few modifications on bones in the form of tooth marks, especially 1999). Cooper (2013) also observed the probable action of Bubo pits, scores and crenulated edges (between 0.69% at LBSmcf.2 and bubo at the site and, therefore, sporadic inputs of this raptor cannot 6.67% at LBSmcf.13), especially in the inner area where no differ- be totally discarded, despite the low incidence of digested bones in ences in the percentages of carnivore alterations are observed be- the studied sample. Nonetheless, on some occasions, mammalian tween level IV and level III. However, the overall proportions are far carnivores such as foxes may present the same pattern as birds of from those recorded by Monchot and Gendron (2013), who docu- prey based on an over-representation of wing bones. One example mented values that reach 44% of the altered bones. On the contrary, R. Blasco et al. / Quaternary International 421 (2016) 62e77 73

Fig. 5. Examples of carnivore and rodent gnawing damage on pigeons and corvid specimens from Gorham's Cave: crenulated edge on humerus of Columba livia/oenas from level IV (A); digestion on distal end of a humerus of Corvidae (B); rodent gnawing on coracoid of Columba livia/oenas from level IV. the percentages extracted from lynx accumulations (4%; Rodríguez- and lower limb bones (2.15%), as well as on one sternum fragment Hidalgo et al., 2016) could fit better with some levels presented of Columba (16.67%). In the case of Pyrrhocorax, the presence of cut here (LBSmcf.6 and LBSmcf.13); nevertheless, the index of damage marks on femora and tibiotarsi suggests the use of this corvid as representation continues to be low as a tendency through the food, in addition to the exploitation of feathers proposed by stratigraphy. This fact leads us to suggest that some birds could Finlayson et al. (2012) on the basis of cut-marked wing bones. This have been provided by carnivores (regardless of whether they are type of alteration, however, is not represented in a huge quantity of raptors or mammal carnivores) when humans did not use the cave, bones. This is not a strange fact because the size of these birds probably making a differential use of the space with a tendency to makes the use of stone tools unnecessary for direct consumption. occupy the inner back areas or specific local zones (e.g., Binford, As in the case of leporids, birds can be easily consumed without the 1981; Brugal and Fosse, 2004; Rosell and Blasco, 2009). use of tools and so it is unusual to find a high number of anthro- Within this mixed picture provided by the skeletal ratio and pogenic marks on bones (Lefevre and Pasquet, 1994; Laroulandie, carnivore-induced damage, hominid inputs also have to be 2000, 2005; Steadman et al., 2002; Cruz, 2005; Laroulandie et al., considered as important because there also exist cases of anthro- 2008). As evidence of this, other modifications related to the pro- pogenic accumulations where wings predominate (e.g., Livingston, cessing of the prey can be observed, such as peeling, fresh breaks or 1989; Bovy, 2002, 2012; Laroulandie and Lefevre, 2014), as could holes and wrenching on the distal humerus, which is usually be the case at Gorham's Cave. In fact, the over-representation of associated with the over-extension of the elbow for disarticulation raptor and corvid wing bones, in addition to human-induced (Laroulandie, 2005a, 2005b; Laroulandie et al., 2008). Additionally, damage on fore limbs at this locality, was interpreted as being it is possible that some of the tooth marks assigned to carnivores related to the procurement of feathers for non-alimentary pur- might actually be produced by humans, because a potential risk of poses and, therefore, as a result of the human contribution confusion exits due to equifinality phenomena. Nevertheless, (Finlayson et al., 2012). In addition, the anthropic incidence on Laroulandie (2005a) tried to distinguish carnivore marks from corvid and pigeons is confirmed in 16 of the 20 analysed Nean- hominid ones, and described the latter as oval shaped marks with derthal units, as well as in the modern human context. Judging by associated crushing. But, in spite of this study, how to precisely the significant percentages of anthropogenic damage, the human identify human tooth marks is still a controversial issue. groups furthermore seem to play an important role in the most Concerning burning, it is the most widespread modification numerous units of Gorham's Cave (level III, level IV, LBSmcf.2, linked to hominid activities. LBSmcf.2, for example, is one of the LBSmcf.4). Level LBSmcf.9 is the one with a relatively high number units where burnt bones are substantially important and affect all of specimens where human activity is lower, but carnivore damage taxa (Columba, Corvus/Pica and Pyrrhocorax). At level LBSmcf.4, is even lower in this unit. The unique level that could be exclu- thermo-alterations represent the highest values in the whole sively attributed to mammal carnivores is LBSmcf.11, although this Gorham's sequence, with similar circumstances to those in data must be treated with caution as the total NISP of the level is LBSmcf.2. This, in addition to the fact that no signals of predation by not significant. carnivores have been observed in the case of unit LBSmcf.4, re- Among the anthropogenic activity, cut marks should be high- inforces the strong impact of Neanderthal activities during the lighted, as they are the clearest physical evidence left by humans. At period of its formation. It is worth noting the presence of double Mousterian levels, incisions tend to occur both on the wing (2.85%) colourations, which may have been generated during differential 74 R. Blasco et al. / Quaternary International 421 (2016) 62e77

fire exposure of the skeleton when the prey (de-feathered or fixed position, the nest, where the nestlings grow until they reach otherwise) was placed on the embers for roasting. In this situation, full adult size (for altricial birds such as the pigeon). This makes the entire bone surface would not receive the same degree of eggs, nestlings and brooding adults relatively easy to catch by hand thermal intensity and only the areas without meat or those covered by a moderately skilful and silent climber (Blasco et al., 2014). In by a very thin soft tissue would be affected more intensely, devel- addition, bird colonies are predictable in time and space, leading to oping higher degrees of charring. In contrast, the bone areas this resource being easy to harvest; the only risk incurred would be covered with large muscle mass would remain unmodified or to descend or climb up to the nest-site if it is located in rocky change only slightly, thus presenting lower degrees of colouring outcrops, coastal cliffs or trees. Even flying species, which are (e.g., Cassoli and Tagliacozzo, 1997; Laroulandie, 2005b; Blasco and harder to catch during the day, spend the night forcibly at roost Fernandez Peris, 2009; Blasco et al., 2013, 2014). As a consequence, sites, where they become much more vulnerable (Negro et al., at Gorham's Cave, the highest degree of burning of the humerus is 2016). found on the proximal joint (head), on the distal end of the ulna and Since birds have traditionally been perceived as elusive prey distal portion of the palmar side of the radius, as well as on the (basically due to their flight capabilities), their systematic exploi- distal end of the tibiotarsus. This pattern is slightly higher on tation has been considered to be an exclusive and defining feature Columba bones (18.35% of burnt bones) than on corvids (17.39%). of modern human behaviour (e.g., Klein, 1989). However, growing Nevertheless, the high values of burning at LBSmcf.2 and LBSmcf.4 evidence points to the use of birds by Neanderthals for both in association with the low number of butchery marks, as well as nutritional and non-nutritional proposes (Fiore et al., 2004; Roger, the significant presence of completely burnt bones bearing degrees 2004; Soressi et al., 2008; Blasco and Fernandez Peris, 2009; Dibble 2 and 3 (brown and black colour, respectively; 77.31%), give rise to et al., 2009; Gaudzinski-Windheuser and Niven, 2009; Hardy and other questions about what may have happened at that site. One Moncel, 2011; Peresani et al., 2011; Finlayson et al., 2012; Morin and option is that part of the burning process occurred accidentally due Laroulandie, 2012; Blasco et al., 2013, 2014; Gerbe et al., 2014; to contact of bones with fire. This situation may occur if a fireplace Romandini et al., 2014; Radovcic et al., 2015), suggesting a more was set up on the ground where bird bones were left. It is known complex scenario as to subsistence strategies and foraging behav- that bones can be subjected to different coloration changes due to iour than was previously thought. The fact that some Mousterian thermal exposure, even if the bones are buried (e.g., Perez [2015] sites with detailed taphonomic studies do not show anthropogenic for lagomorph specimens; Bennett [1999] for larger fauna). damage on small prey (e.g., Lloveras et al., 2011; Rufa et al., 2016) Another scenario contemplates that burnt bones were intentionally encourages further research about what caused Neanderthals to thrown on the fire as a part of cleaning processes (removal of incorporate this type of game in some localities, with particular waste). From this perspective, some burnt specimens at Gorham's emphasis on the small-scale ecological aspects, as well as the socio- Cave may have resulted from non-nutritive episodes that occurred cultural variables and site functionality. after consumption and/or deposition. In either case, it should be taken into account that the complexity of the human occupations 6. Conclusions and the palimpsests formation processes may undergo the destruction or overlapping of bone alterations after the first human In this study, we have tried to contribute to the reassessment of activity; therefore, the roasting should not be discarded in latter the Neanderthal use of avian resources by taphonomically ana- cases because, previous to the secondary burning, the animals lysing 21 discrete archaeological units from Gorham's Cave, 20 could have been exposed to fire with a nutritional purpose. associated with Neanderthals and one with modern humans (level Our results demonstrate that Neanderthals, and later on modern III from the inner area). We propose mixed contributions where the humans, processed pigeons and choughs as part of their spectrum Neanderthals would have played a main role as accumulator agents of prey. Birds seem to have played a role as supplementary re- in the formation of the bird assemblages of Gorham's Cave. Among sources, since the contribution of meat coming from birds is the 16 levels showing anthropogenic evidence on avian bones (80% extremely low compared to the hunted mammals and the strati- of the total Middle Palaeolithic units), LBSmcf.4 shows exclusive graphic sequence of Gorham's Cave shows a constant presence of hominid inputs, in as far as any other modifications have proved the large game, mainly characterised by Cervus elaphus and Capra presence of non-human predators and the anthropogenic damage pyrenaica, among other ungulates (Currant et al., 2013). The is significant on pigeons and choughs. Mammal carnivores would important fact here is that the procurement of pigeons and choughs have contributed a minor input, as shown by the presence of pits, is not an isolated or casual behaviour as it affects a significant punctures and scores. Diurnal birds of prey would also have taken number of individuals over a long temporal span. The special part as another predator, if anatomical representation is ecological conditions of Gibraltar may have favoured the exploita- considered. tion of a broad spectrum of resources, among which birds would Gorham's Cave constitutes an ideal scenario to show formation occupy a special place. The Rock is a mountain in a lowland coastal dynamics based on a complex interplay of cultural and non-cultural setting, providing a catchment area with a diverse combination of factors. Predation on pigeons and choughs seems to be common environmental situations rarely encountered together in the fossil among the Neanderthals at Gorham's Cave, similar to that observed record (Cooper, 2013). A wide variety of cliff-dwelling avian species at level III in association with anatomically modern humans. The and carrion-feeding taxa, as well as wetland, coastal and marine special ecological conditions of this site, located in an area of high species are present, pointing towards an important zone of high biodiversity, seem to have led Neanderthals to the exploitation of a biodiversity within the Strait of Gibraltar (Finlayson et al., 2016). broad spectrum of resources, among which birds stand out as The unique aspects of this locality may have made other manners of backup sources of food. Some colonial and fast-reproducing spe- foraging feasible beyond ungulates. The continued observation of cies, such as pigeons, make them an ideal candidate for sustainable the landscape by Neanderthals could have led them to incorporate exploitation in this context. The knowledge of certain behavioural birds as a backup source of food. Historic and ethnographic ac- and ecological aspects resulting from the observation of the land- counts explain the relative simplicity of catching some species of scape may have favoured the inclusion of certain avian species in birds if certain behavioural and ecological aspects are known (e.g., the hominid socio-economic life. Our results, together with those Levi-Strauss et al., 1992; Juste et al., 1995; Emperaire, 2002; Negro from a growing body of evidence, reflect behavioural complexity et al., 2016). For example, birds are forced to incubate their eggs in a and large versatility among Neanderthal communities, as well as R. Blasco et al. / Quaternary International 421 (2016) 62e77 75 emphasising the importance of the ecological aspects of small- Bochenski, Z.M., Korovin, V.A., Nekrasov, A.E., Tomek, T., 1997. Fragmentation of bird scale, socio-cultural variables and site functionality when making bones in food remains of Imperial Eagles Aquila heliaca. International Journal of Osteoarchaeology 7 (2), 165e171. interpretations. Bochenski, Z.M., Huhtala, K., Jussila, P., Pulliainen, E., Tornberg, R., Tunkkari, P.S., 1998. Damage to bird bones in pellets of Gyrfalcon Falco rusticolus. Journal of Archaeological Science 25, 425e433. Acknowledgements Bochenski, Z.M., Huhtala, K., Sulkava, S., Tornberg, R., 1999. Fragmentation and preservation of bird bones in food remains of the Golden Eagle Aquila chrys- We are grateful to Juan Jose Negro, Geraldine Finlayson, Fran- aetos. Archaeofauna 8, 31e39. Bochenski, Z.M., Nekrasov, A.E., 2001. The taphonomy of Sub-Atlantic bird remains cisco Giles Pacheco, Joaquín Rodríguez-Vidal, Marco Antonio Bernal from Bazhukovo III, Ural Mountains, Russia. Acta Zoologica Cracoviensia 44 (2), and Stewart Finlayson for helpful discussion of the subject covered 93e106. in this paper. Special thanks to Carlos Lorenzo for his help with the Bochenski, Z.M., Tornberg, R., 2003. Fragmentation and preservation of bird bones statistical methods. This research is supported by funding from the in uneaten food remains of the Gyrfalcon Falco rusticolus. Journal of Archaeo- logical Science 30, 1665e1671. Spanish Ministry of Science and Innovation, project nos. CGL2012- Bochenski, Z.M., 2005. Owls, diurnal raptors and humans: signatures on avian 38434-C03-03, CGL2012-38358, HAR2013-48784-C3-1-P, bones. In: O'Connor, T. (Ed.), Biosphere to Lithosphere. New Studies in Verte- e CGL2011-28681 and CGL-BOS-2012-34717, from SeNeCa Founda- brate Taphonomy. Oxbow Books, Oxford, pp. 31 45. Bochenski, Z.M., Tomek, T., Tornberg, R., Wertz, K., 2009. Distinguishing nonhuman tion, project no. 19434/PI/14, and from Generalitat de Catalunya, predation on birds: pattern of damage done by the white-tailed eagle Haliaetus projects nos. 2014 SGR 900, 2014/100573, 2014 SGR 416 and 437 albicilla, with comments on the punctures made by the golden eagle Aquila K117. Anna Rufa has a predoctoral research fellow FPU by the chrysaetos. Journal of Archaeological Science 36, 122e129. Bovy, K.M., 2002. Differential avian skeletal part distribution: explaining the Spanish Ministry of Education, Culture and Sports. R. Blasco is a abundance of wings. Journal of Archaeological Science 29, 965e978. Beatriu de Pinos-A post-doctoral scholarship recipient (Generalitat Bovy, K.M., 2012. Why so many wings? A re-examination of avian skeletal part de Catalunya and COFUND Marie Curie Actions, EU-FP7). This study representation in the south-central Northwest Coast, USA. Journal of Archae- e “ ological Science 39, 2049 2059. was carried out in the framework of the session entitled Human- Bramwell, D., Yalden, W., Yalden, P.E., 1987. Black grouse as the prey of the golden bird interactions during the Prehistory” of the 17th UIPPS World eagle at an archaeological site. Journal of Archaeological Science 14, 195e200. Congress. Bromage, T.G., Boyde, A., 1984. Microscopic criteria for the determination of directionality cutmarks on bone. American Journal of Physical Anthropology 65, 359e366. Appendix A. Supplementary data Brugal, J.-Ph., Fosse, Ph., 2004. Carnivores et Hommes au Quaternaire en Europe de l'Ouest. Revue de Paleobiologie 23 (2), 575e595. Carrion, J.S., Finlay son, C., Fernandez, S., Finlayson, G., Allue, E., Lopez-S aez, J.A., Supplementary data related to this article can be found at http:// Lopez-García, P., Gil-Romera, G., Bailey, G., Gonzalez-Samp eriz, P., 2008. dx.doi.org/10.1016/j.quaint.2015.10.040. A coastal reservoir of biodiversity for Upper Pleistocene human populations: palaeoecological investigations in Gorham's Cave (Gibraltar) in the context of the Iberian Peninsula. Quaternary Science Reviews 27, 2118e2135. References Cassoli, P.F., Tagliacozzo, A., 1997. Butchering and cooking of birds in the palaeolithic site of Grotta Romanelli (Italy). International Journal of Osteoarchaeology 7, Andrews, P., 1990. Owls, Caves and Fossils. University of Chicago Press, London. 303e320. Arilla, M., Rosell, J., Blasco, R., Domínguez-Rodrigo, M., Pickering, T.R., 2014. The Clements, J.F., 2011. The Clements Checklist of Birds of the World. Cornell University “bear” essentials: actualistic research on Ursus arctos arctos in the Spanish Press, Ithaca. pyrenees and its implications for paleontology and archaeology. PLoS One 9 (7), Clements, J.F., Schulenberg, T.S., Iliff, M.J., Roberson, D., Fredericks, T.A., Sullivan, B.L., e102457. Wood, C.L., 2014. The eBird/Clements Checklist of Birds of the World: Version Barton, R.N.E., Jennings, R., 2013. The lithic artefacts assemblages of Gorham's Cave. 6.9. Downloaded from http://www.birds.cornell.edu/clementschecklist/ In: Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Neanderthals in Context: a download/. Report of the 1995-1998 Excavations at Gorham's and Vanguard Caves, Cochard, D., Brugal, J.-Ph., Morin, E., Meignen, L., 2012. Evidence of small fast game Gibraltar. Oxford University Press, Oxford, pp. 151e187. exploitation in the Middle Paleolithic of Les Canalettes (Aveyron, France). Barton, R.N.E., Stringer, C.B., Finlayson, J.C., 2013. Neanderthals in Context: a Report Quaternary International 264, 32e51. of the 1995-1998 Excavations at Gorham's and Vanguard Caves, Gibraltar. Ox- Codding, B.F., Porcasi, J.F., Jones, T.L., 2010. Explaining prehistoric variation in the ford University Press, Oxford. abundance of large prey: a zooarchaeological analysis of deer and rabbit Bennett, J.L., 1999. Thermal alteration of buried bone. Journal of Archaeological hunting along the Pecho Coast of Central California. Journal of Anthropological Science 26, 1e8. Archaeology 29, 47e61. Binford, L.R., 1981. Bones. Ancient Men and Modern Myths. Academic Press Inc, Collcutt, S.N., 2013. Gorham's Cave lithological and lithogenetic patterns. In: Orlando, Florida. Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Neanderthals in Context: a Bird, D.W., Bliege Bird, R., Codding, B.F., 2009. In pursuit of mobile prey: Martu Report of the 1995-1998 Excavations at Gorham's and Vanguard Caves, hunting strategies and archaeofaunal interpretation. American Antiquity 74, Gibraltar. Oxford University Press, Oxford, pp. 37e49. 3e29. Cooper, J.H., 2013. The late Pleistocene avifauna of Gorham's Cave and its envi- Blain, H.-A., Gleed-Owen, C., Lopez-García, J.M., Carrion, J.S., Jennings, R., ronmental correlates. In: Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Finlayson, G., Finlayson, C., Giles-Pacheco, F., 2013. Climatic conditions for the Neanderthals in Context: a Report of the 1995-1998 Excavations at Gorham's last Neanderthals: herpetofaunal record of Gorham's Cave, Gibraltar. Journal of and Vanguard Caves, Gibraltar. Oxford University Press, Oxford, pp. 112e127. Human Evolution 64, 289e299. Cruz, I., 2005. La representacion de partes esqueleticas de aves. Patrones naturales e Blasco, R., Fernandez Peris, J., 2009. Middle Pleistocene bird consumption at Level XI interpretacion arqueologica. Archaeofauna 14, 69e81. of Bolomor Cave (Valencia, Spain). Journal of Archaeological Science 36, Currant, A.P., Price, C., Sutcliffe, A.J., Stringer, C., 2013. The large mammal remains 2213e2223. from Gorham's Cave. In: Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Ne- Blasco, R., Rosell, J., Fernandez Peris, J., Arsuaga, J.L., Bermúdez de Castro, J.M., anderthals in Context: a Report of the 1995-1998 Excavations at Gorham's and Carbonell, E., 2013. Environmental availability, behavioural diversity and diet: a Vanguard Caves, Gibraltar. Oxford University Press, Oxford, pp. 141e150. zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra de Dibble, H.L., Berna, F., Goldberg, P., McPherron, S.P., Mentzer, S., Niven, L., Richter, D., Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain). Quaternary Sandgathe, D., Thery-Parisot, I., Turq, A., 2009. A preliminary report on Pech de Science Reviews 70, 124e144. l'Aze IV, layer 8 (Middle Paleolithic, France). PaleoAnthropology 2009, 182e219. Blasco, R., Finlayson, C., Rosell, J., Sanchez Marco, A., Finlayson, S., Finlayson, G., Domínguez-Rodrigo, M., Juana, S. d., Galan, A.B., Rodríguez, M., 2009. A new pro- Negro, J.J., Giles Pacheco, F., Rodríguez Vidal, J., 2014. The earliest pigeon fan- tocol to differentiate trampling marks from butchery cut marks. Journal of ciers. Scientific Reports 4, 5971. http://dx.doi.org/10.1038/srep05971. Archaeological Science 36, 2643e2654. Bochenski, Z.M., Boev, Z., Mitev, I., Tomek, T., 1993. Patterns of bird bone frag- Emperaire, J., 2002. Los nomades del mar. Lom Ediciones, Santiago. mentation in pellets of the Tawny Owl (Strix aluco) and the Eagle Owl (Bubo Ericson, G.P., 1987. Interpretations of archaeological bird remains: a taphponomic bubo) and their taphonomic implications. Acta Zoologica Cracoviensia 36 (2), approach. Journal of Archaeological Science 14, 64e75. 313e328. Finlayson, C., 2004. Neanderthals and Modern Humans. An Ecological and Evolu- Bochenski, Z.M., Tomek, T., 1994. Patterns of bird bone fragmentation in pellets of tionary Perspective. Cambridge University Press, Cambridge. the long-eared Owl Asio otus and its taphonomic implications. Acta Zoologica Finlayson, C., Giles Pacheco, F., Rodriguez-Vidal, J., Fa, D., Gutierrez-Lopez, J.M., et al., Cracoviensia 37 (1), 177e190. 2006. Late survival of Neanderthals at the southernmost extreme of Europe. Bochenski, Z.M., 1997. Preliminary taphonomic studies on damage to bird bones by Nature 443, 850e853. Snowy Owls Nyctea scandiaca, with comments on the survival of bones in Finlayson, C., Carrion, J., Brown, K., Finlayson, G., Sanchez-Marco, A., Fa, D., Rodrí- palaeontological sites. Acta Zoologica Cracoviensia 40 (2), 279e292. guez-Vidal, J., Fernandez, S., Fierro, E., Bernal-Gomez, M., Giles-Pacheco, F., 2011. 76 R. Blasco et al. / Quaternary International 421 (2016) 62e77

The Homo habitat niche: using the avian fossil record to depict ecological Peters, J. (Eds.), Feathers, Grid and Symbolism Birds and Humans in the Ancient characteristics of Palaeolithic Eurasian hominins. Quaternary Science Reviews Old and New Worlds. Proceedings of the 5th. Meeting of the ICAZ, Munich, 30, 1525e1532. pp. 165e178. Finlayson, C., Brown, K., Blasco, R., Rosell, J., Negro, J.J., Bortolotti, G.R., Finlayson, G., Laroulandie, V., Costamagno, S., Cochard, D., Mallye, J.-B., Beauval, C., Castel, J.C., Sanchez Marco, A., Giles Pacheco, F., Rodríguez Vidal, J., Carrion, J.S., Fa, D.A., Ferrie, J.G., Gourichon, L., Rendu, W., 2008. Quand desarticuler laisse des Rodríguez Llanes, J.M., 2012. Birds of a feather: Neanderthal exploitation of traces: le cas de l'hyperextension du coude. Annales de Paleontologie 94, raptors and corvids. Plos One 7 (9), e45927. 287e302. Finlayson, F., Blasco, R., Rodríguez-Vidal, J., Giles Pacheco, F., Finlayson, G., Laroulandie, V., 2010. Alpine chough Pyrrhocorax graculus from Pleistocene sites Gutierrez, J.M., Jennings, R., Fa, D.A., Rosell, J., Carrion, J.A., Sanchez Marco, A., between pyrenees and alps: natural versus cultural assemblages. In: Finlayson, S., Bernal, M.A., 2014. Gibraltar excavations with particular reference Prummel, W., Brinkhuizen, D., Zeiler, J. (Eds.), Birds in Archaeology. Groningen to Gorham's and Vanguard Caves. In: Sala, R. (Ed.), Pleistocene and Holocene Archaeological Studies, vol. 12, pp. 219e232. Hunter-Gatherers in Iberia and the Gibraltar Strait: the Current Archaeological Laroulandie, V., Lefevre, C., 2014. The use of avian resources by the forgotten slaves Record. Universidad de Burgos-Fundacion Atapuerca, ISBN 978-84-92681-87-7, of Tromelin Island (Indian Ocean). International Journal of Osteoarchaeology 24, pp. 506e514. 407e416. Finlayson, C., Finlayson, S., Giles Guzman, F., Sanchez Marco, A., Finlayson, G., Lefevre, C., Pasquet, E., 1994. Les modifications post-mortem chez les oiseaux: Jennings, R., Giles Pacheco, F., Rodriguez Vidal, J., 2016. Using birds as indicators l'exemple de l'avifaune holocene de Patagonie australe. Artefacts 9, 217e229. of Neanderthal environmental quality: Gibraltar and Zafarraya compared. Levi-Strauss, C., 1992. Tristes Tropicos. Ediciones Paidos Iberica S.A, Barcelona. Quaternary International 421, 32e45. Livingston, S.D., 1989. The taphonomic interpretation of avian skeletal part fre- Fiore, I., Gala, M., Tagliacozzo, A., 2004. Ecology and subsistence strategies in the quencies. Journal of Archaeological Science 16, 37e547. Eastern Italian Alps during the Middle Palaeolithic. International Journal of Lloveras, L., Moreno-Garcia, M., Nadal, J., Zilhao, J., 2011. Who brought in the rab- Osteoarchaeology 14, 73-286. bits? Taphonomical analysis of Mousterian and Solutrean leporid accumula- Fiore, I., Gala, M., Romandini, M., Cocca, E., Tagliacozzo, A., Peresani, M., 2016. From tions from Gruta do Caldeirao (Tomar, Portugal). Journal of Archaeological feathers to food: reconstructing the complete exploitation of avifaunal re- Science 38, 2434e2449. sources by Neanderthals at Grotta di Fumane, unit A9. Quaternary International Lopez-García, J.M., Cuenca-Bescos, G., Finlayson, C., Brown, K., Giles Pacheco, F., 421, 134e153. 2011. Palaeoenvironmental and palaeoclimatic proxies of the Gorham's cave Gaudzinski, S., 2006. Monospecific or species-dominated faunal assemblages dur- small mammal sequence, Gibraltar, southern Iberia. Quaternary International ing the Middle Palaeolithic in Europe. In: Hovers, E., Kuhn, S. (Eds.), Transitions 243, 137e142. before the Transition. Springer, New York, pp. 137e148. Lyman, R.J., 1994. Vertebrate Taphonomy. Cambridge University Press. Gaudzinski-Windheuser, S., Niven, L., 2009. Hominin subsistence patterns during Lyman, R.J., 2008. Quantitative Paleozoology. Cambridge University Press. the Middle and Late Pleistocene in northwestern Europe. In: Hublin, J.-J., Mallory, M.L., 2006. The Northern Fulmar (Fulmarus glacialis) in Arctic Canada: Richards, M.P. (Eds.), The Evolution of Hominin Diets: Integrating Approches to ecology, threats, and what it tells us about marine environmental conditions. the Study of Palaeolithic Subsistence. Springer, Dordrecht, pp. 99e111. Environmental Reviews 14, 187e216. Gerbe, M., Thiebaut, C., Mourre, V., Bruxelles, L., Coudenneau, A., Jeannet, M., Mallye, J.B., Cochard, D., Laroulandie, V., 2008. Accumulations osseuses en Laroulandie, V., 2014. Influence des facteurs environnementaux, economiques periph erie de terriers de petits carnivores: les stigmates de predation et de et culturels sur les modalites d'exploitation des ressources organiques et frequentation. Annales de Paleontologie 94 (3), 187e208. minerales par les Neandertaliens des Fieux (Miers, Lot). In: Jaubert, J., Monchot, H., Gendron, D., 2013. Les restes osseux d'une taniere de renard en milieu ^ Fourment, N., Depaepe, P. (Eds.), Transitions, ruptures et continuites en periglaciare (Ile Digges, rive sud du Detroit d'Hudson, Canada). In: Prehistoire, Actes du XXVIIeme Congres Prehistorique de France, Bordeaux-les Laroulandie, V., Mallye, J.-B., Denys, C. (Eds.), Taphonomie des petits vertebr es: Eyzies 31 mai-5 juin 2010, vol. 2, pp. 257e279. ref erentiels et transferts aux fossiles, BAR International Series 2269. Archaeo- Gidna, A., Yravedra, J., Domínguez-Rodrigo, M., 2013. A cautionary note on the use press, Oxford, pp. 65e76. of captive carnivores to model wild predator behaviour: a comparison of bone Morin, E., 2012. Reassessing Paleolithic Subsistence: the Neandertal and Modern modification patterns on long bones by captive and wild lions. Journal of Human Foragers of Saint-Cesaire. Cambridge University Press. Archaeological Science 40, 1903e1910. Morin, E., Laroulandie, V., 2012. Presumed symbolic use of diurnal raptors by ne- Giles Pacheco, F., Giles Guzman, F., Gutierrez Lopez, J.M., Santiago Perez, A., anderthals. Plos One 7 (3), e32856. Finlayson, C., Rodríguez Vidal, J., Finlayson, G., Fa, D.A., 2012. The tools of the Mourer-Chauvire, C., 1983. Les oiseaux dans les habitats paleolithiques : gibier des last Neanderthals: morphotechnical characterization of the lithic industry at hommes ou proies des rapaces? In: Grigson, C., Clutton-Brock, J. (Eds.), Animals level IV of Gorham's Cave, Gibraltar. Quaternary International 247, 151e161. and Archaeology: 2. Shell Middens, Fishes and Birds, BAR International Series Gleed-Owen, C.P., Price, C., 2013. Amphibians and reptiles from Gorham's Cave. In: 183, Oxford, pp. 111e124. Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Neanderthals in Context: a Negro, J.J., Blasco, R., Rosell, J., Finlayson, C., 2016. Potential exploitation of avian Report of the 1995-1998 Excavations at Gorham's and Vanguard Caves, resources by early humans: an overview from ethnographic and historical data. Gibraltar. Oxford University Press, Oxford, pp. 102e111. Quaternary International 421, 6e11. Goldberg, P., Macphail, R.I., 2000. Micromorphology of sediments from Gibraltar Peresani, M., Fiore, I., Gala, M., Romandini, M., Tagliacozzo, A., 2011. Late Nean- caves: some preliminary results from Gorham's Cave and Vanguard Cave. In: dertals and the intentional removal of feathers as evidenced from bird bone Finlayson, C., Finlayson, G., Fa, D. (Eds.), Gibraltar during the Quaternary: the taphonomy at Fumane Cave 44 ky B.P., Italy. Proceedings of the National Southernmost Part of Europe in the Last Two Million Years. Gibraltar Govern- Academy of Sciences 108 (10), 3888e3893. ment Heritage Publications, Gibraltar, pp. 93e108. Perez, L.J., 2015. Aproximacion experimental a los indicadores de desocupacion Hardy, B.L., Moncel, M.-H., 2011. Neanderthal use of fish, mammals, birds, starchy humana en yacimientos del Pleistoceno superior a partir de los restos ter- plants and wood 125e250,000 years ago. PLoS One 6 (8), e23768. moalterados de conejo. In: Sanchis, A., Pascual, J.L. (Eds.), Preses petites i grups Higham, T.F.G., Bronk Ramsey, C., Cheney, H., Brock, F., Douka, K., 2013. The radio- humans en el passat. II Jornades d'arqueozoologia. Museu de Prehistoria de carbon chronology of Gorham's Cave. In: Barton, R.N.E., Stringer, C.B., Valencia, Valencia, pp. 27e46. Finlayson, J.C. (Eds.), Neanderthals in Context: a Report of the 1995-1998 Ex- Price, C., 2013. The small mammal fauna of Gorham's Cave. In: Barton, R.N.E., cavations at Gorham's and Vanguard Caves, Gibraltar. Oxford University Press, Stringer, C.B., Finlayson, J.C. (Eds.), Neanderthals in Context: a Report of the Oxford, pp. 62e76. 1995-1998 Excavations at Gorham's and Vanguard Caves, Gibraltar. Oxford Jimenez-Espejo, F.J., Rodríguez-Vidal, J., Finlayson, C., Martínez-Ruiz, F., Carrion, J.S., University Press, Oxford, pp. 128e140. García-Alix, A., Paytan, A., Giles Pacheco, F., Fa, D.A., Finlayson, G., Miguel Radovcic, D., Srsen, A.O., Radovcic, J., Frayer, D.W., 2015. Evidence for Neandertal Cortes-S anchez, M., Rodrigo Gamiz, M., Gonzalez-Donoso, J.M., Linares, M.D., Jewelry: modified White-Tailed Eagle Claws at Krapina. PLoS One 10 (3), e0119802. Caceres, L.M., Fernandez, S., Iijima, K., Martínez Aguirre, A., 2013. Environmental Rando, J.C., Perera, M.A., 1994. Primeros datos de ornitofagia entre los aborígenes de conditions and geomorphologic changes during the Middle-Upper Paleolithic in Fuerteventura (Islas Canarias). Archeofauna 3, 13e19. the southern Iberian Peninsula. Geomorphology 180e181, 205e216. Rhodes, E.J., 2013. OSL dating of sediments from the lower part of Gorham's Cave. Juste, J., Fa, J.E., Perez del Val, J., Castroviejo, J., 1995. Market dynamics of bushmeat In: Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Neanderthals in Context: a species in Equatorial Guinea. Journal of Applied Ecology 32, 454e467. Report of the 1995-1998 Excavations at Gorham's and Vanguard Caves, Klein, R.G., 1989. The Human Career. University of Chicago Press, Chicago. Gibraltar. Oxford University Press, Oxford, pp. 77e88. Laroulandie, V., 2000. Taphonomie et archeozoologie des oiseaux en grotte: ap- Rodríguez-Hidalgo, A., Saladie, P., Marín, J., Canals, A., 2016. Bird-bones modifica- plications aux sites Paleolithiques du Bois-Ragot (Vienne), de Combe Sauniere tions by Iberian lynx: taphonomic analysis of non-ingested red-legged par- (Dordogne) et de la Vache (Ariege) (PhD dissertation). Universite de Bordeaux I. tridge remains. Quaternary International 421, 228e238. Laroulandie, V., 2001. Les traces liees a la boucherie, a la cuisson et a la con- Rodríguez-Vidal, J., d'Errico, F., Giles Pacheco, F., Blasco, R., Rosell, J., Jennings, R.P., sommation d'oiseaux: apport de l'expeerimentation. In: Bourguignon, L., Queffelec, A., Finlayson, G., Fa, A.D., Gutierrez Lopez, J.M., Carrion, J.S., Ortega, I., Frere-Sautot, M.-C. (Eds.), Prehistoire et approche experimentale. Negrok, J.J., Finlayson, S., Caceres, L.M., Bernal, M.A., Jimenez, S.F., Finlayson, C., Collection prehistoire 5. Monique Mergoual, Montagnac, pp. 97e108. 2014. A rock engraving made by Neanderthals in Gibraltar. Proceedings of the Laroulandie, V., 2005a. Anthropogenic versus non-anthropogenic bird bone as- National Academy of Sciences 111 (37), 13301e13306. semblages: new criteria for their distinction. In: O'Connor, T. (Ed.), Biosphere to Roger, T., 2004. L'avifaune du Pleistoc ene moyen et superieur au bord de la Lithosphere. New Studies in Vertebrate Taphonomy. Oxbow Books, Oxford, Mediterran ee europeenne : Orgnac 3, Lazaret (France), Caverna delle Fate, Arma pp. 25e30. delle Manie (Italie), Kalamakia (Grece), Karaïn E (Turquie), Paleontologie, Laroulandie, V., 2005b. Bird explotaition pattern: the case of Ptarningan Lagopus sp. taphonomie et paleo ecologie. Doctorat du Museum National d'Histoire In the Upper Magdalenian site of la Vache (Ardiege, France). In: Grupe, G., Naturelle, p. 388. R. Blasco et al. / Quaternary International 421 (2016) 62e77 77

Romandini, M., Peresani, M., Laroulandie, V., Metz, L., Pastoors, A., Vaquero, M., nouveaux gisements, nouveaux resultats, nouvelles methodes, Societ e Slimak, L., 2014. Convergent evidence of Eagle talons used by Late Neanderthals prehistorique française, Memoire XLVII, pp. 95e132. in Europe: a further assessment on symbolism. PLoS One 9 (7), e101278. Steadman, D.W., Plourde, A., Burley, D.V., 2002. Prehistoric butchery and con- Rosell, J., Blasco, R., 2009. Home sharing: carnivores in anthropogenic assemblages sumption of birds in the Kingdom of Tonga, South Pacific. Journal of Archaeo- of the Middle Pleistocene. Journal of Taphonomy 7 (4), 305e324. logical Science 29 (6), 571e584. Rufa, A., Blasco, R., Rivals, F., Rosell, J., 2016. Who eats whom? Taphonomic analysis Stewart, J.R., Jacobi, R.M., 2015. The long term response of birds to climate change: new of the avian record from the Middle Paleolithic site of Teixoneres Cave (Moia, results from a cold stage avifauna in northern England. PLoS One 10 (5), e0122617. Barcelona, Spain). Quaternary International 421, 103e115. Stringer, C., Finlayson, C., Barton, R.N.E., Fernandez-Jalvo, Y., Caceres, I., Sabin, R.C., Sacca, D., 2012. Taphonomy of Palaeloxodon antiquus at Castel di Guido (Rome, Rhodes, E.J., Currant, A.P., Rodríguez-Vidal, J., Giles-Pacheco, F., Riquelme- Italy): proboscidean carcass exploitation in the Lower Palaeolithic. Quaternary Cantal, J.A., 2008. Neanderthal exploitation of marine mammals in Gibraltar. International 276e277, 26e41. Proceedings of the National Academy of Sciences 105 (38), 14319e14324. Shipman, P., Rose, J., 1983. Early hominid hunting, butchering and carcassprocessing Waechter, J.D.'A.,1951. The excavations at Gorham's Cave, Gibraltar, preliminary report behaviors: approaches to the fossil record. Journal of Anthropological Archae- for the seasons 1948 and 1950. Proceedings of the Prehistoric Society NS 17,83e92. ology 2, 57e98. Waechter, J.D.'A., 1964. The excavations at Gorham's Cave, Gibraltar, 1951-1954. Shipton, C., Clarkson, C., Bernal, M.A., Boivin, N., Finlayson, C., Finlayson, G., Fa, D., Bulletin of the Institute of Archaeology of London 4, 189e221. Giles Pacheco, F., Petraglia, M., 2013. Variation in lithic technological strategies White, T.D., 1992. Prehistoric Cannibalism at Mancos 5M TURM-2346. Princeton among the Neanderthals of Gibraltar. PLoS One 8 (6), e65185. University Press. Smith, E.A., 1983. Anthropological applications of optimal foraging theory: a critical Yravedra, J., Rubio-Jara, S., Panera, J., Uribelarrea, D., Perez-Gonz alez, A., 2012. El- review. Current Anthropology 24, 625e651. ephants and subsistence. Evidence of the human exploitation of extremely Soressi, M., Rendu, W., Texier, J.-P., Daulny, L., d'Errico, F., et al., 2008. Pech-de-l'AzeI large mammal bones from the Middle Palaeolithic site of PRERESA (Madrid, (Dordogne, France) : nouveau regard sur un gisement mousterien de tradition Spain). Journal of Archaeological Science 39 (4), 1063e1071. acheuleenne connu depuis le 19eme siecle. In: Jaubert, J., Bordes, J.G., Ortega, I. Yravedra, J., Cobo-Sanchez, L., 2015. Neanderthal exploitation of ibex and chamois (Eds.), Les societ es du Paleolithique dans un Grand Sud-Ouest de la France : in southwestern Europe. Journal of Human Evolution 78, 12e32.