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Quaternary International xxx (2014) 1e17

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Quaternary International

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The ungulate assemblage from layer A9 at Grotta di Fumane, Italy: Azooarchaeological contribution to the reconstruction of Neanderthalecology

Matteo Romandini a, Nicola Nannini a, Antonio Tagliacozzo b, Marco Peresani a,*aUniversità di Ferrara, Dipartimento di Studi Umanistici, Sezione di Scienze Preistoriche e Antropologiche, Corso Ercole I d’Este 32, I-44100 Ferrara, Italyb Soprintendenza al Museo Nazionale Preistorico Etnografico “L. Pigorini”, Sezione di Paleontologia del Quaternario e Archeozoologia, P. le Marconi 14,Roma, Italy

a r t i c l e i n f o

Article history:Available online xxx

Keywords:UngulatesZooarchaeologyMiddle PalaeolithicMIS3Italy

* Corresponding author. Università di Ferrara, Stud’Este 32, I-44100 Ferrara, Italy.

E-mail addresses: matteo.romandini@unife.it (M.libero.it (N. Nannini), antonio.tagliacozzo@beniculturperesani@unife.it (M. Peresani).

http://dx.doi.org/10.1016/j.quaint.2014.03.0271040-6182/� 2014 Elsevier Ltd and INQUA. All rights

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a b s t r a c t

One of the most widespread flaking methods in Europe was the Discoidal technique. However, tech-nological analyses of these lithic artifacts are not yet sufficiently integrated into a corpus of zooarch-aeological indicators that outline an ecological profile of Neanderthal mobility. To address this issue, thisstudy presents evidence from Grotta di Fumane in northern Italy, where the exclusive use of the Discoidmanufacturing technology is embedded in a Late Mousterian sequence with Levallois industries. Thepaper begins with a presentation of the regional ecological and contextual setting, and then explains thetaphonomic and zooarchaeological data from the large and varied ungulate assemblage. Results showthat hunting activity was shaped by the availability of game and that well-established, cost-effectivepatterns were used in carcass processing. Compared on a broader scale with other contexts whereDiscoid implements have been taken into account in relation to faunal assemblages, these foragingpractices show that a common model for Neanderthal subsistence strategy cannot be applied.

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1. Introduction

Reconstructing Neanderthal foraging behavior is one of themost challenging areas in anthropological research in westernEurasia, before and at the dawn of the MiddleeUpper Palaeolithictransition. During this period, and especially in the Late MiddlePalaeolithic, extensive cultural replacements occurred, which areindicated by many diverse archaeological signatures. For example,technological innovations are the product of adaptive strategiesand eventual changes in dietary habits, as inferred throughzooarchaeological and isotopic data (Stiner, 2001; Marean, 2005).In the past three decades, the technological variability in theMousterian has generated debate over its role in subsistence stra-tegies and in mobility, which requires assessment on differentscales (Kuhn, 1995; Mellars, 1996; Delagnes et al., 2007).

The focus of this study grew out of the analysis of a faunalassemblage that dates to the first half of the MIS3 and is associatedwith a lithic assemblage produced by means of Discoid technology.

di Umanistici, Corso Ercole I

Romandini), nicola.nannini@ali.it (A. Tagliacozzo), marco.

reserved.

i, M., et al., The ungulate asseerthal ecology, Quaternary In

Although noted in many zones of continental Europe, this method(Boëda, 1993; papers in Peresani, 2003) is not yet sufficiently in-tegrated into a corpus of data that would permit the evaluation ofthe technology’s impact on the adaptive strategy of the ancienthumans who employed it. While the production of Discoid lithicimplements is associated with a few zooarchaeological assem-blages in sparse contexts starting in MIS 6, this pattern becomesmore frequent in isotopic stages 3 and 4, especially in some areaswhere different technologies are also attested (Peresani, 2003;Delagnes and Rendu, 2011). It is perhaps even more interesting tonote that some assemblages document technological replacementswithin short chronological intervals. For this reason, their associa-tion with economic and subsistence indicators becomes funda-mental for outlining Neanderthal ecological and mobility profiles.

In order to contribute further information to these topics, thisstudy presents zooarchaeological evidence from the Grotta diFumane in northern Italy, a cave which preserves a late Middle andEarly Upper Palaeolithic sequence finely layered and extensivelyinvestigated for two decades. It records abrupt climatic, environ-mental and cultural changes. This paper considers unit A9, inwhichthe exclusive use of discoid manufacturing technology replaces theLevallois technology employed in the underlying layer A10(Peresani, 2012).

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

M. Romandini et al. / Quaternary International xxx (2014) 1e172

Given this particular cultural context, the related aspects ofNeanderthal behavior are here examined. Additionally, the dataprovide a baseline for assessing changes in foraging practicesaround the transition from the Middle to Upper Palaeolithic in theAlpine foreland. The paper begins with a presentation of theregional ecological setting and context of Grotta di Fumane, andthen explains the zooarchaeological data from the large faunalassemblage recovered there. The final section of the paper em-phasizes broader scale comparisons with other contexts wherediscoid lithic implements have been considered in relation tofaunal assemblages.

2. Regional setting

The central-eastern pre-Alps are formed by successivemountainranges that constitute discontinuous series of short chains withsummits above 2000 m.a.s.l., limestone massifs and karst plateausat 1000e1500 m.a.s.l., as well as gorges, deep valleys and wide

Fig. 1. The northern Adriatic region with the location of Grotta di Fumane (a, F), Riparo Tagliedi Manune view (b); the site (c); unit A9 during the 2009 and 2010 excavations (d, e); sketch(A4-A3) and earliest Aurignacian layers (A2) and of their variable content in archaeologicaPeresani, redrawn by S. Muratori) (f), plan of the cave with zone of origin of the materials

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basins. These are crossed by rivers and are the setting for glacialAlpine lakes such as Garda Lake.

During MIS3 until HE4, the Upper Po Plain and the Alpinefoothills saw persistent afforestation with some temperate trees,such as Tilia and Abies. Moderate forest reduction occurred duringearly DO events that supported the expansion of steppe environ-ments in combination with mixed conifers (Pinus and Picea) andBetula forests (Ravazzi, 2002; Pini et al., 2010). On the basis of somepollen sequences located in the southern Alpine foreland, it hasbeen argued that Middle Würm stadials experienced summertemperatures very close to the growth limit of oak (Fimon, AzzanoDecimo and Notranje Gorice; Pini et al., 2009; 2010).

Grotta di Fumane is located at 350 m of elevation in the westernpart of theMonti Lessini, a fan-shaped plateau dipping gently to thesouth towards the alluvial plain of the Adige River, which is char-acterized to the north by summits 1500e1600m.a.s.l. (Fig.1aec). Tothewest of Fumane, the plateau ends at the Adige Valley, a long anddeep cut connecting the inner Alpine region with the Po Plain. The

nte (a, RT), Mezzena (a, M), San Bernardino (a, SB) and Grotta del Rio Secco (a, RS); Vajoof sagittal section of the cave with evidence of the late Mousterian (A11-A5), Uluzzianl remains (increasing from light gray to dark gray and black) (by M. Cremaschi & M.in gray and position of the section above (g).

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

M. Romandini et al. / Quaternary International xxx (2014) 1e17 3

immediate surroundings of the cave are composed of severalmorpho-tectonic terraces connected to the bottom of the Fumanevalley by steep slopes and rock walls that include many caves andshelters. The cave is strategically placed due to its geographicsetting, which may have facilitated human penetration to thehighlands.

3. Context

At Fumane, the Middle e Upper Palaeolithic deposits consist ofnumerous thin to very thin parallel levels and lenses that aregrouped into stratigraphic units labelled, frombottom to top, A13 toA1 (Fig.1). Mousterian industries are found in A12-A11, A10, A9, A6-A5, followed by the Uluzzian in A4 and A3, and the Proto-Aurignacian in A2 and A1 (Broglio et al., 2006; Peresani, 2008,2012; Peresani et al., 2011). The Mousterian levels, which wereinvestigated over areas varying from six to 75 sqm in size, arecharacterized by lithic and faunal remains that are densely scat-tered on the living floors, as is the case in units A11, A10, A9 and A6.Flints in A12-A11, A10V, A10, and A6-A5 were exploited mostlythrough Levallois technology (Peresani, 2012; Peresani et al., 2013).

Unit A9, the focus of this paper, is a complex of layers with a totalthickness of 15e20 cm, due to the overlapping of thin layers andlenses. These are composed of frost-shattered stones, aeolian siltand sands, and dark sediments, which are widely distributed as aresult of intense anthropic accumulation. These sediments havebeen removed at various intervals through trenches and limitedexcavated sectors, while more extensive excavations were under-taken over a 68 sqm area during 2009e2012 campaigns at the caveentrance (Fig. 1d and e). Intact or fragmented bones, teeth, anddeterminable fragments�5 cm have beenmapped topographically,while smaller fragments, micromammal and small avifaunal boneswere recovered through wet sieving.

Dwelling structures with hearths and possible toss-zones wereidentified within the area of investigation. To date, unit A9 has pro-duced 50 structures, most of which are hearths distributed at thecave entrancedmore so in the western than in the easternzonednear the present-day drip-line. Considering the 14C datesobtained from overlying units (Higham et al., 2009) and the knownbiases due to sample contamination in this age range (Higham, 2011),it is probable that the chronometric position at 47.6e45.0 ka cal BP ofA9 as provided by the oldest (42,750 � 700 BP) of six 14C dates(Peresani et al., 2008) is the more reliable minimum age.

The flint used in the Discoid lithic industry covers the litho-logical variability of the Valpolicella, an area rich in lithics within arange of 5e10 km from the site, where primary exposures and loosedeposits along stream beds, slopes and soils can be easily exploited.The use of rawmaterials in this industry does not vary in respect tothat of the Levallois assemblages, showing that the most abundanttypes in the site surroundingsweremore exploited than others. Theindustry is typically represented by thick flakes, pseudo-Levalloispoints, backed flakes with a thin opposite edge, polygonal andtriangular flakes, as well as some scrapers, points and denticulates(Peresani, 1998). Functional analyses have shown that brute flakesand retouched tools were used for scraping or cutting fresh and dryskin as well as wood. With some exceptions, the types of motionsand actions performed are randomly spread across the studiedsample, with transversal motions being more frequent than lon-gitudinal, unidirectional and bidirectional. Furthermore, while theuse of certain artefacts for specific functions has not been observed,different kinds of flakes were used in similar ways, and supportinterpretation of the discoid artefacts as being “strong” tools(Lemorini et al., 2003).

Previous osteological analyses carried out on the faunal assem-blage from the 1988e1991 fieldwork have shown that the sample of

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identifiable bones (2.2% of the total) is dominated by cervids (80%,red deer, giant deer and roe deer), followed by bovids and caprids(15%, ibex and chamois) and other species (5%, marmot, fox, bear,hyena, wild boar) (Cassoli and Tagliacozzo, 1991). Very preliminaryobservations on the ungulate bone surfaces identified anthropicactions ascribable to different stages of the butchering process likeskinning, dismembering and filleting. Several tiny fragments haveimpact flakes, and diaphyseal fragments show typical stigmata oftheir use as retouchers forflint tools. Integrating thesedatawith thatof the lithic industry suggests that through Discoid technology,Neanderthalswere able toaccomplish all the tasks involvedwith theprocessing of animal prey (Lemorini et al., 2003).

4. Materials and methods

The material under consideration here represents about 80% ofthe total zooarchaeological assemblage recovered (Nannini, 2012).The remainder (20%) of the assemblage comes from the innercavity, which is a zone not yet extensively explored. The sample isconsidered representative, as it comes from an area of about 58 m2

(Fig. 1g) that is characterized by variations in the spatial distribu-tion of bones and lithic tools, and includes hearths and possibletoss-zones.

Taxonomic and skeletal identifications made in this study arebased on the complete Alpine fauna reference collection of theDepartment of Humanities at the University of Ferrara and, formore problematic cases, on the collection of the Quaternary Pale-ontology and Zooarchaeology Laboratory of the Ethnographic-Prehistoric National Museum “L. Pigorini” (Rome, Italy). Micro-scopic analyses of bone surfaces were carried out using a Leica S6DGreenough stereomicroscope with 0.75e70� magnification range.In order to identify the nature of bone fractures and surface alter-ations, and to distinguish human from animal traces (pits, punc-tures, scores, furrowing, scooping-out, etc.), trampling abrasion,and modern mechanical modifications produced by excavationtools, reference was made to well-established taphonomic litera-ture (Binford, 1981; Brain, 1981; Shipman, 1981; Blumenschine andSelvaggio, 1988; Capaldo and Blumenschine, 1994; Blumenshine,1995; Fisher, 1995; Domínguez-Rodrigo and Piqueras, 2003). Evi-dence of anthropic modification has been achieved by taking intoaccount the type, position and orientation of the cut-marks, in or-der to reconstruct the performed action (skinning, defleshing,periosteum removal, etc.) and the intentional bone breakage toaccess marrow (Binford, 1981; Potts and Shipman, 1981; Shipman,1981; Shipman and Rose, 1984; Blumenschine and Selvaggio, 1988;Villa and Mahieu, 1991; Capaldo and Blumenschine, 1994; Lyman,1994; Blumenshine, 1995; Fisher, 1995). According to excavationprotocol, natural post-depositional degradation was identified onlyon those objects longer than 3 cm or on those taxonomically rele-vant. The degree of combustion was evaluated by employing themethodology developed by Stiner et al. (1995). Sex and age at thetime of death were determined by dental wear and by the state offusion of epiphyses (Aitken, 1974; Mariezkurrena, 1983; Vigal andMachordom, 1985; D’Errico and Vanhaeren, 2002; Fiore andTagliacozzo, 2006). Measurements were taken following Drieschand von den (1976). In order to evaluate species abundance, thefollowing methods were used: number of remains (NISP) (Grayson,1984), minimum number of elements (MNE) (Binford, 1981; Kleinand Cruz-Uribe, 1984; Stiner, 1994), and the estimate of the mini-mum number of individuals (MNI) (Bökönyi, 1970). On the basis ofmorphological or size characteristics, unidentifiable remains weresorted (by weight) into mammal body-size classes: small (mus-telids, marmot, lagomorphs, fox, etc.), medium (roe-buck, roe-deer,caprids, wolf, hyena, etc.) and large (giant-deer, bison, bear, elk,deer, ursids, lion, etc.). Estimation of deer MNI is based on dental

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

Table 1 (continued )

Taxa NISP NISP% NMI by age Total NMI

Infant Young y-ad Adult Senile

Ursus spelaeus 8 0.6 1 1 1 3Ursus sp. 5 0.4 1 2 1 4Crocuta crocuta

spelaea3 0.2 1 1

Panthera leospelaea

1 0.1 1 1

Sus scrofa 2 0.2 1 1 2Alces alces 17 1.4 2 1 3Megaloceros

giganteus79 6.2 6 6

Cervus elaphus 495 39.3 1 4 5 4 14Capreolus

capreolus281 22.3 2 1 7 1 11

Cervidaelarge size

166 13.2

Bos primigenius 6 0.5 2 2Bison priscus 6 0.5 1 2 3Bos/Bison 29 2.3Capra ibex 46 3.7 1 2 1 4Rupicapra

rupicapra68 5.4 1 4 1 6

Caprinae 19 1.5Total NISP 1259 100 1 14 5 44 8 72

Table 2General anatomy and taphonomy detail of bone remains; CS ¼ combustion struc-tures; GM¼ gnawing marks; B¼ burned¼ black/brown bones; C¼ calcined¼ grey/white bones.

NISP %

Total remains 111,841 100Total diaphysis 50,113 44.8Total epiphysis þ spongy bones 9036 8.1Other elements þ unidentified 52,692 47.1Total Ungulata remains (NISP taxa þ Ungulata size) 2345 2.3Total diaphysis 1871 7.8Total epiphysis þ spongy bones 122 5.2

M. Romandini et al. / Quaternary International xxx (2014) 1e174

eruption and wear. Fragmentation indexes (Binford, 1981; Brain,1981; Lyman, 1994) were calculated to evaluate the skeletal rep-resentation of different animals and the skeletal survival rate.

According to their ecological characteristics, the ungulate spe-cies in the assemblage are assumed to be representative of twohabitat types (Mysterud, 2000; Miller, 2003). Lowland speciesinclude elk Alces alces, giant deer Megaloceros giganteus, red deerCervus elaphus, roe-deer Capreolus capreolus, and wild-boar Susscrofa that would have been found at lower elevations in valleys,grasslands, wetlands, and forests. Mountain species include preytypes like ibex Capra ibex and chamois Rupicapra rupicapra athigher elevations or in rocky habitats. The distinction betweenthese habitat types is coarse, and it may obscure more refined preydistribution patterns within each type (Ready, 2013).

In order to consider how Neanderthals at Fumane used thehabitats described above, prey types within each patch are rankedhere using the body size rule (Bayham, 1979; Broughton, 2002).This rule assumes that the post-encounter return rates of prey in-crease with body size (i.e., that a larger animal provides more cal-ories per unit of foraging time) and is valid within the ensemble oflarge mammals in our assemblage. Mobility patterns among mid-to large-sized artiodactyls show decreasing maximal velocity withincreasing body mass; this in particular suggests that the body sizerule may be relatively strong for this group (Morin, 2012).

5. Results

5.1. First taphonomical layout

Within the analyzed sample of 111,841 bone fragments, it waspossible to identify 1259 remains of large mammal species(Table 1). The number of identified bones is low due to the highrates of bone breakage (more than 92.5% are smaller than 2 cm) andcombustion (50%). These conditions are due to a set of depositionaland post-depositional processes also noted in other fully anthropiccontexts (Morin, 2012). The number of bones altered by mecha-nisms of mobilization is very low (2%). The most common cause ofdegradation is root marks (38%), while other remains showmicrofissures from weathering and/or weathering cracks (0.8%),manganese stains (18%) and exfoliations (2.2%). Some concretedfragments (21.4%) were concentrated near the external area at thelimit of the drip-line, while the few with trampling abrasions(15.9%) in the east part of the cave are associated with greaterstoniness. There is high incidence of burned remains that couldsuggest the use of butchering refuse, rich in fats, as fuel for fires.Within this body of burned fragments, 92.6% were subject tomoderate combustion (200e500 �C, black/brown) while 7.4% werecalcined (>700 �C, grey/white). At a general level, these patternsand phenomena are similar to evidence from the final Mousterianlevels A5-A6, and to other anthropogenic deposits in the region(Fiore et al., 2004).

Table 1Number of bones, NISP and NMI by age of animals.

Taxa NISP NISP% NMI by age Total NMI

Infant Young y-ad Adult Senile

Erinaceuseuropaeus

1 0.1 1 1

Marmotamarmota

8 0.6 3 3

Mustela nivalis 4 0.3 1 1 2Mustelidae 1 0.1Canis lupus 4 0.3 2 2Vulpes vulpes 6 0.5 1 1 2Ursus arctos 4 0.3 2 2

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Low-density bone portions (ribs, spurious ribs, sternum, verte-brae fragments, shoulder blades) are rare in the assemblage (8.1%),as demonstrated by the extremely poor representation of long boneepiphyses (4%) relative to shaft fragments, which play a significantrole both in number and inweight (Table 2). This marked imbalancein low/high density rate of bone portions indicates several factors,such as the aforementioned post-depositional processes, the exis-tence of equifinality phenomena including the use of spongyepiphyses as fuel (which generally burn much better than compactdiaphyses, as suggested from experimental data (Costamagno et al.,2005, 2010; Morin, 2010)), bone grease production, and the activ-ities generated by carnivores during secondary access to the re-mains scattered on the surface.

Other elements þ unidentified 352 15Number combustion structures 26Total burned remains 54,411 48.7B diaphysis 23,915 44C diaphysis 2170 4B epiphysis þ spongy bones 2336 4.3C epiphysis þ spongy bones 92 0.1Other elements þ unidentified (B þ C) 25,898 47.6Total remains combustion structures 10,543 9.4CS diaphysis 6000 56.9CS epiphysis þ spongy bones 918 8.7CS other elements þ unidentified 3625 34.4Total gnawing marks remains þ digested 101 0.1Diaphysis GM 79 78.2Epiphysis þ spongy bones GM 7 6.9Other elements þ unidentified GM 15 14.9

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M. Romandini et al. / Quaternary International xxx (2014) 1e17 5

Carnivores are common agents of density-mediated destructionas well as of bone accumulation in Palaeolithic assemblages. Thedegree and the variability of these interventions can be assessedfrom examining prey type, skeletal representation, age at death,bone surface and structural modifications, and bone spatial distri-bution, and is also achieved from ethological studies (e.g., Binford,1982; Blumenschine, 1988; Selvaggio, 1994; Capaldo, 1997;Domínguez-Rodrigo, 2001; Pickering, 2002; Domínguez-Rodrigoand Piqueras, 2003; Camarós et al., 2013). This degree seems todepend on the use that these predators make of the site, especially

Table 3Ungulates with a list of butchering marks, thermal-alterations and carnivore marks. CM: cut-marks; PM: percussion marks; CM þ PM: cut-marks þ percussion marks; IF:impact flake; CM þ IF: cut-marks þ impact flake; Tot. BM: Total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/brown bones; C: calcined-grey/white bones; GM: carnivore gnawing marks.

Taxa NISP CM PM CM þ PM IF CM þ IF Tot. BM % BM R B C GM

Sus scrofa 2 1 1 e 1Capreolus capreolus 281 44 2 3 49 17.4 1 63 5 9Cervus elaphus 495 116 26 38 6 3 189 38.2 28 97 4 17Alces alces 17 8 2 10 58.8 1Megaloceros giganteus 79 28 1 14 43 54.4 6 18Cervidae large size 166 30 7 6 2 45 27.1 8 48 4 4Bos primigenius 6 4 1 5 83.3 1Bison priscus 6 2 4 6 100 2Bos/Bison 29 7 2 3 1 13 44.8 3 3Capra ibex 46 7 1 1 1 10 21.7 6 3Rupicapra rupicapra 68 12 1 13 19.1 2 5 1 4Caprinae 19 2 1 3 15.8 1 1Ungulata large size 705 108 25 17 170 9 329 46.7 14 144 5 11Ungulata medium-large size 142 20 5 1 27 53 37.3 28 5Ungulata medium size 290 51 6 5 57 1 120 41.4 4 56 4 5IndeterminateLarge size 385 49 6 4 73 4 136 35.3 2 68 5 9Medium-large size 474 77 6 1 66 3 153 32.3 1 95 1 10Medium size 736 210 3 118 2 333 45.2 3 144 4 15Medium-small size 20 16 1 17 85 7 1Small size 16 5 5 31.2 2Unidentified 107,263 49 9 58 0.1 49,576 3979 4

in Pleistocene caves (Skinner, 2012), causing problems in recon-structing the formation processes of the assemblage or detectingspecific episodes within a given stratigraphic unit. Different ob-servations and experimental reproductions have been made, withboth wild animals and animals in captivity (e.g., Sutcliffe, 1970;Blumenschine, 1986a, 1986b, 1988; Bunn et al., 1988; Mareanet al., 1992; Camarós et al., 2013), through reproducing archaeo-logical contexts. For instance, Blumenschine (1988) exposed freshlybroken bovid bones to hyenas, and observed that these scavengerspreferred the rich-in-fat epiphyses. These bones were stronglytooth-marked and destroyed, leaving the shafts in their originalposition, and displacing some partially or completely consumedepiphyses. At Olduvai, carnivores acted on the skeletal parts with ahigher proportion of fat: vertebrae, ribs and epiphyses, leaving anassemblage with predominantly limb bones, comparable to thosegenerated by the anthropogenic exploitation (Marean et al., 1992)or scavenging of carcasses abandoned by large felids(Blumenschine, 1991). In more recent and more complex Palae-olithic sites, the role of nonhuman scavengers on the abandonedcamps was also assessed and compared to some ethno-archaeological studies (Binford, 1978a, 1978b, 1982; Binford et al.,1988; Bartram et al., 1991; Yellen, 1991; O’Connell et al., 1992).These observations suggest that carnivores destroy or modify thebones and can also change the original pattern of the remains withsignificant alteration (Binford et al., 1988). Camarós et al. (2013)have experimentally recorded the magnitude at which large car-nivores modify abandoned hearth structures and related

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assemblages, which strongly attracted their attention due to thesmells of meat, organic tissues, or fat remains. Results show thatbears, hyenas, lions and wolves produce considerable change in theoriginal discard.

At Fumane the frequency and patterning of rodent and carnivoregnaw marks, such as scoring and pits (carnivore marks: NISP 90 onbone shafts and NISP 7 on epiphyses; NISP 4, gastric juice erosion;rodent marks: NISP 18) (Tables 2 and 3), suggest secondary accessto the ungulate remains, and that carnivores did not make a sig-nificant contribution to the bone accumulation.

Looking in detail at the NMI of the carnivores, only seven in-dividuals belonging to three diverse species would have the ca-pacity to shatter the diaphyses of the large ungulates present: alion, a hyena, and five bears (Table 1), which would have left,however, clear and macroscopic evidence if they would haveinteracted significantly with the bone assemblage (Fig. 2A). Addi-tionally, the complex of unit A9, similarly to other contexts alongthe same stratigraphic sequence (Peresani et al., 2011), recordsrepeated anthropogenic frequentations containing hundreds ofavifaunal remains, some still in anatomical connection (Fig 2G), afew fragile and complete anatomical elements of foetuses, parts ofhyoid bones, and telemetacarpals of ungulates, in addition tocombustion structures (Table 2), distributed mainly in the atrialarea of the cave, in proximity to the present-day drip line. Some ofthese are partially superimposed, the result of successive fre-quentations over a relatively short period of time. The faunal re-mains contained within these structures (number of remains10543; 9.4%) seem to be generally very fragmented, for the mostpart with dimensions <1 cm. For this reason, the determination oftaxa was only possible for a few specimens, most of which arerepresented by ungulates and birds. The most highly representedspecies in the structures are cervids, strongly represented bydiaphyseal fragments (56.9%), which are particularly prevalent inrespect to the epiphyses and spongy bones (8.7%). This dispro-portion demonstrates the mechanical fragility typical of the ex-tremities and the spongy bone tissue, which seems to recur withlimited changes, even within the different tapho-taxonomic

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

Fig. 2. Sketch of the sagittal section at Grotta di Fumane (see Fig. 1) showing some structures and traces of bones modified by humans or carnivores. A: Bos and Bison bonesmodified by hyenas and bears from the top levels of the inner cave. BeC: Aurignacian levels (A2 þ A2R), reject structure with bones, flint artifacts and charcoals, B; extremity limb inanatomical connection of Capra ibex with cut-marks and percussion marks, C; DeE: Mousterian units (A5-A6, with Levallois industry), combustion structure and associated cut-marked bone and radio of Bison priscus, D; metapodial of young Cervus elaphus with scrapes and percussion mark (indicator), E; FeG: Mousterian unit (A9, with discoid industry),Bison priscus tibia with cut-marks and percussion mark, F; connected wing bones (Humerus-radio-ulna) of Pyrrhocorax graculus discovered in sq. 87, G.

M. Romandini et al. / Quaternary International xxx (2014) 1e176

ensembles (Table 2). The number of deer shaft portions graphicallyexpressed with chromatic scale (Fig. 8A) as compared with theother ungulates (Fig. 8B) denotes the same representation of theepiphyses of the long bones, while demonstrating the exactopposite for the appendicular elements of the extremities (meta-podials and phalanges). The close relationships between thedensity of anthropogenic indicators and the state of conservationare found further expressed in the Levallois Mousterian level (A5-A6) and the Aurignacian level (A2) above it (Fig. 2BeE). By contrast,at the top of the sedimentary sequence, when access to the innercavity was feasible only by carnivores, the bone remains belongingto the ungulates of medium-large size found scattered on the

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surface present an elevated degree of destruction, with tooth-marks present (Fig. 2A).

In further support of a very low degree of alteration or loss ofspatial and behavioral information at unit A9 due to secondaryaccess by carnivores, these occurrences do not seem to haveincluded cases of hibernation, except perhaps on sporadic occa-sions, and in any case not within the investigated zone (four re-mains from individuals under two years of age and one fetal bearindividual) (Table 1). Moreover, traces of human modifications (seebelow) exclude the possibility that long bone shafts and epiphyseswere deliberately disassociated prior to transport. Potential ex-planations might be sought only by large-scale examination, by

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

Fig. 3. Representation of the frequence of anatomical elements for the main taxa of ungulates.

M. Romandini et al. / Quaternary International xxx (2014) 1e17 7

taking into account the spatial patterning of burnt and calcinedbones, and comparing this patterning with the distribution of thecombustion structures.

5.2. Ecology and spectra of predated animals

Red deer and roe deer, the two most commonly occurring ani-mals (Table 1), indicate forested environments interspersed withclearings. The presence of elk, aurochs, and wild boar suggests thatwetlands with water sources also existed in a general cold-temperate climatic context. Bison and giant deer would haverequired access to open, sparsely forested environments, whichcould be found on the plain or above the timber line (7000e1000 m a.s.l.) not far from the site. This latter environment was alsovital to chamois and ibex. The high incidence of cervids is consis-tent with the faunal records from the Mousterian stratigraphicsequence (Cassoli and Tagliacozzo, 1991; Fiore et al., 2004), which

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from units A11-A10 shows a progressive increase of these species tothe detriment of those favoring open environments.

As these prey species inhabited an environmental range thatmay have occurred around the cave, between the foothills of theplains and the highlands, the site’s occupants may not have spentequal time foraging in these different patches. The taxonomic data(see below) demonstrate that the majority of animal remainsbrought back to the site were from prey types inhabiting lowlandsand valley bottoms. Larger prey types, relatively infrequent in theassemblage, may have had lower encounter rates than red deer, orperhaps were disregarded on occasion if the hunters’ success ratesfor these were lower than for smaller prey species (Bird et al.,2009). Conversely, even if larger ungulates were moderatelyabundant and always pursued upon encounter, actual capture ratesmay have been so low that few attempts were successful, resultingin low representation of these animals in the assemblage (Coddinget al., 2010; Ready, 2013). The available data from Fumane layer A9,

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Fig. 4. Alces alces: tibia (a) and proximal rib (c) with defleshing cut-marks and respective close-ups (a1, c1); Megaloceros giganteus: cut-marked humerus (b) and calcaneum (d) andrespective close-ups (b1, d1).

M. Romandini et al. / Quaternary International xxx (2014) 1e178

which includes the lowest-ranked ungulate species in the localenvironment, suggests maximum exploitation of the ecologicalrange (Grayson and Delpech, 1998).

Several alternative explanations for this pattern can be sug-gested. First, while small ungulates like chamois and ibex mayhave had a high rate of capture upon encounter because of theirliving in groups, more so than wild boar or roe deer, for example,these may have been infrequently encountered compared to reddeer and roe deer. Mountain habitat areas may not have beenextensive near Fumane, but this is only partially confirmed by thecharcoal assemblage of unit A9, which suggests that the cavesurroundings were ecologically characterized by the presence oflarch, showing that the tree-line was positioned at an elevationnot far from the cave, with denser forests of spruce in the valleybottom (Basile, 2011e12). Another possibility is that roe deer andchamois were naturally mutually exclusive (Herrero et al., 1996),which would have led to a lower incidence of chamois

Fig. 5. Bos-Bison: metapodial (a) with skinning cut-marks (a1, close-up); Bison pris

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encounters. Larger species such as red deer and giant deer tend tofeed in open areas, and therefore may have been easier to accessand locate for the occupants at Fumane. Roe deer are elusive,territorial, and tend to live alone or in pairs (Geist, 1998), but mayhave been abundant in the environment in question. The patch-iness of ungulate prey distributions, as well as the variableencounter rates within these patches, may have been importantin determining the opportunity costs of different hunting envi-ronments. Finally, it is also possible that ungulates from higherelevations were generally excluded from the diet, but served as analternate food source at times when higher-ranked prey (such asred deer) was less abundant.

Age ranges determined by dental wear analyses and the state ofepiphyseal fusion reveal a predominance of adults (Table 1) andmature adults, while relatively few are prime individuals. Thescarcity of fetal and neonatal individuals (NISP 5) makes it impos-sible to identify a preferential seasonal use of the site. A mandible

cus: tibia (b) with defleshing cut-marks (b1, close-up) and impact notch (b2).

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Fig. 6. Sus scrofa: burned metapodial with cut-marks (a and a1 close-up); Rupicapra rupicapra: second phalanx with skinning cut-marks (b and b1 close-up); Capra ibex: carpal withcut-marks. (c and c1 close-up).

M. Romandini et al. / Quaternary International xxx (2014) 1e17 9

fragment from a young bison with traces of skinning is the singlesuggestion that capture took place in the spring.

A variety of species are identifiable amongst the carnivore re-mains (Table 1). The presence of wolf, hyena and lion, together withblack vulture (Peresani et al., 2011), confirm the existence of openspaces and an abundant ungulate biomass; meanwhile, ursids, redfox and weasel are suited to a predominantly forested landscape. Ahuman subsistence or utilitarian (furs) interest in these types ofanimals is attested by two rib fragments with skinning marksattributable to one mid-sized carnivore (wolf or hyena) and onelarge carnivore (bear or lion).

5.3. Exploitation of carcasses

The surfaces and the fresh fracture edges of bones that preserveanthropic marks produced during the butchering of carcasses areobserved on all types of ungulates (Table 3). In particular, from thefew cranial elements to the more abundant limbs (especially hindlimbs), the large cervids (red deer, giant deer and elk) and largebovids (bison, aurochs) show different phases of carcass processing,which were carried out in a more thorough way compared tosmaller animals (ibex, chamois, roe deer and wild boar) (Fig. 3).These data might be biased due to the overall larger bone size forthese ungulates, which offers greater surfaces for analysis. Amongthe ungulate remains, at the level of body size, these values aremore proportionate (Table 3).

Cuts made by lithic tools constitute the most common mark(Table 2), and are often repeated, especially on long bones andin places corresponding to muscle and tendon insertions(Figs. 4e6). Striae on cranial or appendicular elements suggestskinning. Longitudinal scrapings on shafts suggest that theperiosteum was removed for cleaning the bone of meat residuesand preparing it for deliberate breakage. Traces in articulated orepiphyseal zones are few (2.6%), probably due to the lownumber of bones.

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Overall, the distribution of marks can further be conditioned byvariations in the conservation of remains and their morphologies. Itis important to bear in mind that the high number of remains withanthropic modifications, and particularly fragments identified asmedium-sized ungulates and unidentified individuals, can becontextualized in the scarcity of certain types of markings on roedeer, chamois and ibex (Table 3).

Single or multiple percussion marks, percussion notches, andspiral fractures that are typical of fresh bone were observed taxo-nomically, at the level of size, and on unidentified bones. Impactpoints are found primarily on bones that are long, thick, andresistant to trauma, such as distal tibias and humeri, metapodialsand, in some cases, phalanges. Impacts are repeated and wide onbones of large ungulates, while impact flakes are well documentedfor both medium-sized and large ungulates (Table 3).

Numerous shaft fragments (total NISP 72) were used as ham-mers for retouching flint artifacts. Stigmata-like punctiform im-pressions, linear impressions, retouch-induced striae, and notchesrelated to this purpose were found not only on remains with sub-stantial weight and bone thickness, such as femur and tibia frag-ments of red deer and other large-size animals, like bison andmegaceros, but also on roe deer and chamois tibia fragments(Jéquier et al., in press).

5.4. Carcass representation index

The ungulates, as a whole, broken down and subdivided by theirrespective anatomical elements in cranio-caudal order, show aprevalence of fragments of metapodials (29.1%), zeugopods (24.2%),and stylopods (13%), followed by elements of extremities likephalanges and sesamoids (14.7%). Cranial remains are scarce andare predominantly represented by isolated teeth (11.5%) and partsof the hemimandible and jaw (4.4%). The trunk and spinal columnare nearly absent (1.8%) (Fig. 3) and, even considering that parts ofthe vertebrae and ribs could be present in a fragmentary and

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

Fig. 7. 1: Distribution of the butchering marks on the skeletal elements of the Cervus elaphus and skeletal survival rate (in red-scale). Details of cut-marks on distal humerus (a),radius (b), proximal femur (c) and metatarsal (d). 2: distribution of butchering marks on the skeletal elements of Capreolus capreolus and representation of the skeletal survival rate(in red-scale). Details of cut-marks on humerus (a), radius (b), tibia (c) and metatarsal (d).

M. Romandini et al. / Quaternary International xxx (2014) 1e1710

unidentified state or identified only on the level of size (Table 2),their scarcity is not attributable to differences in degree of post-depositional preservation. This bias becomes clear for the largecervids elk and giant deer, which are represented by shaft frag-ments, phalanges, sesamoids and isolated teeth. This becomes evenmore pronounced in the case of bison, aurochs, and remainsidentified as bos/bison, which are represented only by hind limbs. Asimilarly differential representation was also noted for small ani-mals such as ibex and chamois (Fig. 3), which are missing ribs,vertebrae, and fragments of spongy tissue.

Due to the high number of remains, it is possible to carry out amore detailed analysis for red deer and roe deer. For red deer, thecombination of dental wear data and the state of epiphyseal fusionin the post-cranial skeleton shows that nearly all age classes arerepresented. Of 14 individuals in total, one is young (18e22months), four are young adults (three in the 20e32 months rangeand one of 24e36 months), five are adults (one in the 5e6 yearsrange, one in the 7e8 range, and three in the 8e10 range), and four

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are mature adults (one 9e11 years and three >12 years) (Table 1).Considering that hinds give birth at the end of spring, it is possibleto estimate the capture period for the sole young individual. Mostof the bones are fragmented shafts and teeth; epiphyses are un-derrepresented, and phalanges are relatively numerous comparedto the carpal and tarsal bones, which are almost absent (Table 4).The elements of the trunk are very scarce as well (Fig. 7), as shownby their survival rate. The absence of the atlas and the axis isremarkable, because these bones may have remained with thecranium or were still connected to the other vertebrae when thehead was disarticulated from the body. Only four ribs have beenattributed to this species, but among the indeterminate ribs wecounted 94 fragments whose size attributes them to this ungulate.Also significant is the rarity of specimens identified as coxal, andthe absence of scapulae (Fig. 7). The most common long bones aretibia, followed bymetatarsals and femurs, while the front limbs arelower in number (Table 4). The patella is completely absent, andcarpal and tarsal bones are dramatically underrepresented. In light

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M. Romandini et al. / Quaternary International xxx (2014) 1e17 11

of this anatomical composition, it results that the deer trunk andgirdle were left at the killing site and that only selected portionswith meat were transported into the cave. Considering the highnumber of metapodials it remains difficult to interpret the scarcityof carpal and tarsal bones (usually very resistant to post-depositional phenomena), while phalanges were often discardedor left with the skin. Regarding the MNE, our considerations aboutthe anatomical composition of the deer are different (Table 6): themost common anatomical elements belong to the tibia, followed bythe humerus and radius. The survival rate reveals that the trunkand the extremities of the limbs are severely underrepresented. Afragmentation index (MNE/NISP) reveals a high incidence of frac-tured elements except for short and compact bones of the ex-tremities, which have values that tend to 1, and other elements,which may have overestimated values due to the small number ofremains (such as coxal vertebrae or ribs) (Table 6). Countless tracesof anthropic modifications affect over 38.2% of the red deer bones,while few bones show evidence of gnawing by carnivores, or tracesof digestion (3.4%) (Table 3). Butchering traces like cut-marks andpercussion marks were detected on fragments, and 80 flakes wereused as retouchers (Table 3, Fig. 7.1). Each type of long limb boneshows cut-marks: femurs, humeri, radii, tibia and metapodials areaffected in over the 49.6% for the frontal limb and 48.4% of the casesfor the hind limb, and the remaining bones have values not higherthan 11.4%. Cut-marks were also observed on all the short end-bones of the limbs (Table 4).

Table 4Number of anatomical elements of Cervus elaphus and with anthropic modificationsmarks þ percussion marks; IF: impact flake; CM þ IF: cut-marks þ impact flake; Tot. BMbrown bones; C: calcined-grey/white bones; GM: carnivore gnawing marks.

Cervus elaphus

NISP CM PM CM þ PM IF

Cranium 23Hemimandible 35 2 1 1Tooth indet. 14HyoidTotal cranium 72 2 1 1Atlas-axisVertebra 3Rib 4 2Total axial skeleton 7 2ScapulaHumerus 27 10 1 3 1Radius/UlnaRadius 29 12 3Ulna 4CarpalsMetacarpal 55 18 4 3 2Metacarpal rudim.Total frontal limb 115 40 5 9 3Coxal 4 1Femur 44 15 8 6PatellaTibia 96 27 8 14 2Malleolar BoneCalcaneum 1AstragalusTarsals 2Metatarsal 76 18 4 2 1Total hind limb 223 61 20 22 3Metapodial 25 5 3First phal. 11 1 3Second phal. 12 1Third phal. 4First phal. rudim. 3 2Second phal. rudim. 5Third phal. rudim. 4Sesamoid 14 2Total indet. limb 78 11 6

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The data for the representation of roe deer anatomical elementsare less clear, despite the clear imbalance between shaft fragmentsand the axial skeleton (Table 5, Figs 7.2 and 8). This discrepancyseems evenmore acute whenwe consider that 37 rib fragments andeight vertebra fragments can be attributed to the small cervid on thebasis of size. The absence of skeletal elements from the girdles(scapular and coxal) could be due not only to anthropic selection, butalso to a difference in conservationof certain elements for amedium-sized animal. In fact, the same bias is visible among the indetermi-nate remains. On the other hand, unlike the red deer, rudimentalcarpal andmetacarpal bones have been preserved (Table 5). Analysisof age groups demonstrated that nearly all groups are represented.Among the 11 total individuals, there are twoyoung individuals (3e9months and >1 year), one young adult (c. 22 months), seven adults(from 4 to 8 years) and one mature adult (>8 years) (Table 1). Therepresentation of anatomical elements of all ungulates suggests thatthe treatment of the carcass took place at the kill site and that onlyselected parts of the animal were transported to the cave, increasingthe disparity between useful anatomical elements and those of “lowcaloric utility” (Binford,1978b;Metcalfe and Jones,1988; BarlowandMetcalfe, 1996; Bettinger et al., 1997).

6. Discussion

The archaeozoological data acquired here allow us to discussvarious topics related to the ecology and economy of subsistence

and carnivore marks. CM: cut-marks; PM: percussion marks; CM þ PM: cut-: Total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/

CM þ IF Tot. BM % BM R B C GM

4 11.4 2 3 3

4 5.6 2 3 3

2 22 28.6 2

15 55.6 3 6

15 51.7 4 31

27 49.1 4 20 4

57 49.6 11 30 41

29 65.9 4 5 2

1 52 54.2 8 17 2

1 26 34.2 3 25 2 42 108 48.4 15 47 2 81 9 36 4 1 1

4 36.4 3 11 8.3 1

2 66.731

2 14.3 3 11 18 23.1 15 2 2

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Table 5Number of anatomical elements of Capreolus capreolus with anthropic modifications and carnivore-marks. CM: cut-marks; PM: percussion marks; CM þ PM:cut-marks þ percussion marks; Tot. BM: Total butchering marks; % BM: % butchering marks; R: retoucher; B: burned; C: calcined; GM: carnivore gnawing marks.

Capreolus capreolus

NISP CM PM CM þ PM Tot. BM % BM R B C GM

Cranium 7Hemimandible 16 1 1 6.3 1 1Tooth indet. 6HyoidTotal cranium 29 1 1 3.4 1 1Atlas-axisVertebra 2Rib 2Total axial skeleton 4ScapulaHumerus 12 2 2 16.6Radius/UlnaRadius 10 4 4 40 1 2Ulna 1 1 1Carpals 6 1 1 1Metacarpal 32 3 1 4 12.5 9 1Metacarpal rudim. 4 1 1 e

Total frontal limb 65 11 1 12 18.5 11 1 4CoxalFemur 17 4 4 23.5 2 1Patella 1Tibia 38 11 1 12 31.6 1 6 1Malleolar BoneCalcaneumAstragalusTarsals 1 1Metatarsal 59 10 1 1 12 20.3 21 1 1Total hind limb 116 25 1 2 28 24.1 1 29 1 4Metapodial 11 2 2 18.2 5 1First phal. 21 1 1 2 9.5 4Second phal. 9 2 2 22.2 2 1Third phal. 4 3First phal. rudim. 4 1 1 e

Second phal. rudim. 4 3Third phal. rudim. 7 4Sesamoid 7 1 1 14.3 1 1Total indet. limb 67 7 1 8 11.9 22 3

Table 6MNE (Minimum Number Elements), NEE (Number Expected Elements), skeletal survival rate and fragmentation index of Cervus elaphus and Capreolus capreolus.

Cervus elaphus Capreolus capreolus

NISP MNE NEE Skeletal surv.Rate(%)

Fragm. IndexMNE/NISP

NISP MNE NEE Skeletal surv.Rate(%)

Fragm. IndexMNE/NISP

Cranium 23 9 14 6.3 0.4 7 3 11 27.3 0.4Hemimandible 35 9 28 32.1 0.3 16 5 22 22.8 0.3Tooth indet. 14 6HyoidTotal cranium 72 18 42 42.9 0.3 29 8 33 24.2 0.3Atlas-axisVertebra 3 3 378 0.8 1 2 2 297 0.7 1Rib 4 4 364 1.1 1 2 2 286 0.7 1Total axial skeleton 7 7 742 0.9 1 4 4 583 0.7 1ScapulaHumerus 27 15 28 53.6 0.6 12 7 22 31.8 0.6Radius/UlnaRadius 29 14 28 50 0.5 10 7 22 31.8 0.7Ulna 4 4 28 14.3 1 1 1 22 4.5 1Carpals 6 6 132 4.5 1Metacarpal 55 10 28 35.7 0.2 32 9 22 40.9 0.3Metacarpal rudim. 4 4 44 9 1Total frontal limb 115 43 112 38.4 0.4 65 34 264 12.9 0.5Coxal 4 4 28 14.3 1Femur 44 13 28 46.4 0.3 17 8 22 36.4 0.5Patella 1 1 22 4.5 1Tibia 96 20 28 71.4 0.2 38 13 22 59.1 0.3Malleolar boneCalcaneum 1 1 28 3.6 1Astragalus

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Table 6 (continued )

Cervus elaphus Capreolus capreolus

NISP MNE NEE Skeletal surv.Rate(%)

Fragm. IndexMNE/NISP

NISP MNE NEE Skeletal surv.Rate(%)

Fragm. IndexMNE/NISP

Tarsals 2 2 84 2.4 1 1 1 66 1.5 1Metatarsal 76 13 28 46.4 0.2 59 13 22 59.1 0,2Total hind limb 223 53 224 23.7 0.2 116 36 154 23.4 0.3Metapodial 25 6 0.2 11 4 0.4First phal. 11 8 112 7.1 0.7 21 15 88 17 0.7Second phal. 12 7 112 6.3 0.6 9 8 88 9.1 0.9Third phal. 4 4 112 3.6 1 4 2 88 2.3 0.5First phal. rudim. 3 3 112 2.7 1 4 4 88 4.5 1Second phal. rudim. 5 5 112 4.5 1 4 3 88 3.4 0.8Third phal. rudim. 4 4 112 3.6 1 7 7 88 8 1Sesamoid 14 14 224 6.3 1 7 7 176 4 1Total indet. limb 78 51 896 7 0.7 67 50 704 8.7 0.7

M. Romandini et al. / Quaternary International xxx (2014) 1e17 13

for the Neanderthals of unit A9 at Fumane, and to propose aframework for models of mobility from comparisons with few butsignificant indicators from Western Europe.

As concerns the ecological context, the faunal spectrum noted inA9 is consistent with a picture of persistent afforestationwith sometemperate trees (Pini et al., 2010). However, due to chronometricuncertainty, we cannot exclude the occurrence of DO events thatsupported moderate mixed conifer (Pinus, Picea and Betula) forestcontraction with the expansion of steppe communities (Ravazzi,2002). Fumane Cave could thus well be inserted in a context notfar from open-spaced forests, in conditions of transitive todiscontinuous Alpine grasslands or pioneer vegetation on carbon-ate rocks. This picture therefore confirms that hunting activity wasnot specialized to target one or several selected taxa, but was rathershaped by the game availability in the western Lessini.

Fumane is not an isolated Late Middle Palaeolithic settlement inthe pre-Alpine fringe and sub-Alpine area. Evidence of humanfrequentation has been identified at sites that are concentratedalong a longitudinally oriented belt that begins in the LessiniMountains, and progressively curves north to the east, ending atthe Trieste Karst where it arcs towards the plain. Sites such asRiparo Tagliente and Riparo Mezzena in the Lessini Mountains, andGrotta San Bernardino and Grotta Rio Secco far to the east, werelocated in different ecological contexts. Their faunal assemblagesindicate that the prevalent association of ungulates e mostly reddeer and roe deer with fewer chamois and ibex, with limitedexploitation of Bos/Bison, giant deer, elk and wild boar e fit theecological conditions around each site and show shifts correlatedwith the most relevant climatic oscillations (Fiore et al., 2004).Nevertheless, each of these sites provides the possibility for asso-ciation between Levallois implements and faunal assemblages.

Some information comes from Riparo Tagliente and RiparoMezzena, positioned in contexts similar to that of Fumane, Grotta S.Bernardino, and Grotta Rio Secco, which are each at elevationsbetween 250 and 500 m.a.s.l. At Riparo Tagliente deposits from thefinal Mousterian still have not been dated. At Riparo Mezzena thereis a deposit that was partially disturbed and excavated in the 1960s,which was provisionally dated to at the latest 41,270e38,040 cal BP(Longo et al., 2012). Grotta San Bernardino in the Berici Hills, situ-ated on a steep eastern slope where the low karstic plateauabruptly connected the alluvial plain, was occupied by humans inthe Mousterian (unit II) in an environment that was forested bytemperate-cold woods with wetlands, as suggested by wild boar,elk and beaver findings (Cassoli and Tagliacozzo, 1994). Grotta RioSecco, on the Pradis plateau in the eastern part of the Carnic pre-Alps, preserves layers with a Mousterian industry that usedDiscoid and Levallois technology (Peresani et al., in press). Thefaunal association is from two different environments: dense forest

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and sparse trees, such as those typical of Alpine grasslands or high-elevation plateaus.

Quantitative and qualitative data available from these sitesreveal that the types of hunted ungulates at each site conform totheir respective specific environmental and climatic conditions. AtRiparo Tagliente, in layers 35e37, the most common species to havebeen brought into the cave are roe deer and red deer, with feweribex and chamois. Hunting at San Bernardino targeted differentspecies, primarily young adult and adult cervids. Among the fewcaprids, chamois was more commonly found than ibex. Cut markson some beaver bones indicate the recovery of pelts (Malerba andGiacobini, 1998). At Grotta Rio Secco the hunters killed, butch-ered, and consumed large cervids, bovids and caprids. San Ber-nardino, Riparo Tagliente, Fumane, and probably Grotta Rio Seccotherefore share a model of exploitation centered on the selection ofyoung, young adult, and adult prey (Fiore et al., 2004; Thun-Hohenstein and Peretto, 2005; Romandini, 2012), with in-dications of seasonality pointing primarily to spring and summermonths. Moreover, hunting of rodents and some carnivores for fursis documented. Nevertheless, besides this environmental vari-ability that has supported the hunting of many different ungulatespecies, differences in human group behavior or specific economicrequirements should be considered to explain the issue ofremarkable evidence, as for instance from Rio Secco, where theexploitation of bears and cave bears marks a particular signature ofhuman occupation (Peresani et al., in press).

Fumane Cave was used as a habitat place where the processingof carcasses was finalized after having been begun at the kill site.This is clearly indicated by the pattern of preservation of skeletalelements, at least for red-deer and roe-deer, from which it ispossible to infer that groups of humans transported to the siteselected anatomical parts with high nutritional value (bothmarrowand meat), such as limbs, especially hind, and to a lesser extent thecranium. Therefore, well-established, cost-effective patterns can beenvisaged in the selection of specific skeletal portions, as a functionof several factors including the weight of the red-deer and roe-deercarcass portions and the distance between the kill/butchering siteand the home base. The location and the ecological context ofFumane, a cave placed at low-mid altitude in proximity to the al-luvial plain and the mountain grassland belt, supported theexploitation of different prey, but required efforts to cover thedistance in a landscape dissected by deep valleys with cliffs andsteep slopes. Moreover, other social factors, such as the number ofparticipants in hunting parties, regulated by economic and socialcontexts, have to be considered. These variables will be better takeninto account in future works, when more detailed data from layersabove and belowA9 and fromother sites is achieved using the samelevel of investigation can assess whether the selection of specific

mblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeologicalternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.03.027

Fig. 8. A: graphic color representation of the some bone shaft portions based on a chromatic scale with distribution of the anthropic marks on humerus, femur and tibia (all rights)of Cervus elaphus. B: Frequency of bone shaft portions graphically expressed using a black-gray scale: femurs, tibias, metacarpals þmetatarsals, second phalanges, humeri and radii-ulne of ungulates (ungulate size þ ungulates NISP e Cervus elaphus on bones rights of cervus) (by M. Romandini).

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M. Romandini et al. / Quaternary International xxx (2014) 1e17 15

prey or parts could beworthwhile for human groups. Data is scarce,though, from the eastern Alps, where no faunal remains at any sitehave been found in context with a Discoid industry that wouldallow us to infer models of resource management. However, somesimilarities in physiographic and ecologic context can be found atMujina Pecina. This site is located along a potential migrationcorridor for hominin populations moving into Europe fromwesternAsia. The faunal composition is comparable to sites from north-eastern Italy (Fiore et al., 2004) due to the predominance of reddeer. Nothing about the composition of the assemblage indicatesthat hominins were targeting particular prey to the exclusion ofother species. Although non-hominin carnivores played a signifi-cant role in the modification of the faunal assemblages throughoutthe stratigraphic sequence, the Late Mousterian faunal assemblageshows hominins to be competent hunters (Miracle, 2005).

Outside the pre-Alpine and High Adriatic region, ecologicalcontexts with similar faunal spectra are found in the Ligurianlandscape, where archaeozoological data associatedwith industriesincluding Discoid flaking are known from Arma delle Manie, Ma-donna dell’Arma, and Santa Lucia Superiore in the MIS5 e MIS3interval. These sites have yielded faunal assemblages rich in cer-vids, and are generally dominated by red deer. The frequency ofother ungulate taxa, including roe deer, aurochs and caprids, isinextricably linked to the topography of each site, as well as toclimatic and environmental conditions. The accumulation of her-bivore remains could have been caused by carnivores, but evidencefor human activity involving the same remains is also documented,albeit marginally (Valensi and Psathi, 2004). Mortality and sea-sonality data show various patterns of age distribution of the un-gulate species, and suggest no relation between occupation seasonof these sites and the hunting of a particular species. The domi-nance of red deer can be explained by their natural abundance andby their high rendering in terms of hunting transport, meat quan-tity, and other useful materials including skins and tendons(Valensi and Psathi, 2004).

In general, there are few sites with Discoid industry that areassociated with faunal remains that have undergone archae-ozoological analysis at a level of detail that includes survival rates.In France and on the Iberian Peninsula, sites that fit this descriptionare diverse in terms of ecology, but at the same time are compa-rable from logistical and economical points of view. In the RhôneValley, the recurrence of similar occupation types along thesequence of Payre attests to the permanency of the Neanderthalmode of occupation through time in this area and is associated witha ‘‘logistical” type organization of territorial management(Daujeard and Moncel, 2010).

As at Fumane, a similar trend can be observed at Saint-Cézaire(layer Egpf) in the lateMousterian of MIS3. Morin (2012) has shownthat the site was used as a residential habitat, based on the con-sumption of horse and the selective introduction of the richestbody parts of bovines and reindeer. Clear analogies in the mode ofcarcass treatment also come from level J from Abric Romanì, whereit is hypothesized that the transport of selected parts (head andlimbs) to the site took place during the spring and fall (Rosell et al.,2012). The example from Fumane unit A9 distinguishes it fromassemblages attributed to the Discoidal lithic assemblages in someregions of western Europe (Mauran, La Rouquette 1, La Quina 6c)that confirm the existence of selective and seasonal hunting stra-tegies correlated with this specific technology, which becamedominant from MIS4 until the end of the Middle Palaeolithic(Delagnes and Rendu, 2011). The fact that at a given time of the yearsome sites were repeatedly used for the exploitation of a particulartaxon is evidence of a year-round pattern of scheduling the huntingof gregarious and migratory prey. This model does not seem validfor Grotta di Fumane, as the system of Discoid lithic production is

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associated with an exploitation not targeted at a specific type ofungulate. Rather, more opportunistically, this exploitation seems tofollow the natural abundance of ungulates and then determine theencounter rate in the environment.

Systematic practices in the faunal processing sequence shouldalso be assessed under the light of social and cultural factors, whichmight become identifiable in cases where the zooarchaeologicalrecord produces signatures of standardized actions on both soft andhard tissues (Yellen, 1977, 1991; Blasco et al., 2013). To evaluate ifeach limb bone was treated following a standardized process anddetermine if this process took different forms depending on thecultural group is an intriguing issue if it is related to the exclusiveuse of the Discoid lithic technology in A9. Nevertheless, this re-quires more integrated spatial, taphonomical and use-wear ana-lyses from A9 and other units across the late Mousterian sequenceof Fumane, where the Discoid technology alternates or associateswith others (Peresani, 2012).

7. Conclusion

A significant set of evidence shows that Neanderthals were ableto face environmental constraints and change over time, by puttinginto action different types of responses that are visible in techno-logical innovations, lithic procurement, and mobility. The dataobtained from Fumane fit this scenario, in that they show that thezooarchaeological assemblage derives from the economic activityperformed by groups employing a Discoid technology in a territoryrich in lithic raw material and variable enough to have multipleenvironments and ecological niches. In other systems, the pre-dictability of seasonal movement of game was significant and thusresulted in more highly structured and planned mobility patterns,such as, for instance, those associated with the Quina system(Delagnes and Rendu, 2011). By contrast, the Discoidal system re-flects more opportunistic hunting displacements, which is consis-tent with Fumane in the exploitation of lithic raw materials of lessthan top quality, contrary to what has been observed across the MPand the MPeUP transition sequence (Bertola et al., 2009; Peresani,2012).

Acknowledgments

Research at Fumane is coordinated by the Ferrara University inthe framework of a project supported by the Ministry of Culture e

Veneto Archaeological Superintendency, public institutions (Lessi-nia Mountain Community e Regional Natural Park, Fumane Mu-nicipality, Veneto Region e Department for Cultural Heritage), andprivate associations and companies (National Geographic Society,Cariverona Foundation, Valpolicella-Benaco BCC Bank, RobertoGardina & C., Albino Armani Winegrowers and others). The authorsare grateful to two anonymous reviewers for constructive sugges-tions and to K. Heasley for revision of the English text.

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