Article pp.103-117 du Vol.25 n°3-4 (2013)

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ARTICLE /ARTICLE

Late Upper Palaeolithic human diet: first stable isotope evidence from Riparo Tagliente (Verona, Italy)

Alimentation humaine au Paléolithique supérieur : premières données isotopiques du site de Riparo Tagliente (Verone, Italie)

V. Gazzoni · G. Goude · E. Herrscher · A. Guerreschi · F. Antonioli · F. Fontana

Revised: 12 January 2012, Accepted: 12 October 2012

AbstractThis article reports results of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope analysis performed on the bone collagen of a Late Epigravettian human individual and 11 faunal remains from the Upper Palaeolithic deposits of Riparo Tagliente (Verona, Italy). Riparo Tagliente is located in Valpantena on the pre-alpine massif of Monti Lessini, at 250 m a.s.l. Its strategic position, about halfway from the plain and the top of the limestone plateau, has allowed the groups that occupied the site to exploit different ecosystems.

The human skeleton comes from an incomplete burial exca- vated in 1973 and belongs to a young adult male. It is dated between 16,634 and 15,286 cal BP (OxA-10672). Theδ¹³C (−18.4‰) andδ¹⁵N (13.0‰) values of the human individual are enriched compared to those of herbivores on average by +1.2‰in¹³C and +10.0‰in¹⁵N, and to the omnivores on average by +1.4‰in¹³C and +7.2‰in¹⁵N. Isotopic values would indicate an origin of proteins from terrestrial herbivores and high trophic level species. These data match with the results of taphonomic analyses carried out on the bone remains of herbivores, while the study of other species as fish is still in progress.

Keywords Epigravettian · North-eastern Italy · Carbon · Nitrogen · Subsistence strategies

RésuméDans l’objectif d’étudier les modes de subsistance à l’Épigravettien récent, des analyses isotopiques (carbone et azote) ont été réalisées sur le collagène osseux d’un sujet humain et de plusieurs espèces animales de la série Paléo- lithique supérieur de Riparo Tagliente (Vérone, Italie). Le site est localisé dans le fond de vallée du Valpantena, sur le massif préalpin des Monts Lessini, à 250 m au-dessus du niveau marin. Ce site occupe une position qui favorise l’accès à différents écosystèmes, environ à mi-chemin entre la plaine et le sommet du plateau calcaire. Les restes humains sont ceux d’un jeune adulte de sexe masculin, datés entre 16634–15286 cal BP. Les valeurs isotopiques du sujet humain (δ¹³C :–18,4‰;δ¹⁵N : 13,0‰) montrent un enrichissement important comparativement à la faune herbivore (+1,2‰en¹³C et +10,0‰en¹⁵N) et par rapport à la faune omnivore du même site (+1,4‰en¹³C et +7,2‰ en ¹⁵N). Les valeurs isotopiques (δ¹³C,δ¹⁵N) enregistrées sur le sujet indiquent que les protéines consommées ont plusieurs origines impliquant des espèces de niveaux trophiques différents. L’étude des traces de boucherie sur la faune herbivore terrestre confirme une partie de ces résultats. Malgré la présence de poisson sur le site, aucune information n’est disponible à ce jour pour discuter des espèces concernées et de leur rôle dans l’alimentation humaine.

Mots clésÉpigravettien · Italie nord-orientale · Carbone · Azote · Stratégies de subsistance

Introduction

After over 30 years of intensive research and study on the Late Upper Palaeolithic occupation of the south-eastern Alps, several data on settlement dynamics and landscape

V. Gazzoni (*)

via Ariosto 26, 46100 Mantova, Italy e-mail : valentina.gazzoni@unife.it G. Goude · E. Herrscher

LAMPEA–Université de Provence, CNRS, MCC (UMR 7269), Maison méditerranéenne des sciences de l’Homme,

5, rue du Château de l’Horloge, BP 647, F-13094 Aix-en-Provence cedex 02, France A. Guerreschi · F. Fontana

Dipartimento di Biologia ed Evoluzione, Università degli Studi di Ferrara,

Corso Ercole I d’Este 32, 44121 Ferrara, Italy F. Antonioli

ENEA, Department of Environment, Global Change and Sustainable Development, via Anguillarese 301, 00123 Roma, Italy DOI 10.1007/s13219-012-0079-x

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use of the hunter-gatherers are now available [1]. In particular, zooarchaeological researches have allowed the primary role played by hunting in human subsistence strategies to be highlighted [2]. Ibex, red deer and chamois were the most hunted species while elk, auroch, roe deer and wild boar occupy a secondary role. Among carnivores, fox, wolf,mus- telidaeand bear were hunted frequently [1,2]. Exploitation of birds is also documented at some sites such as Riparo Dalmeri (Trento) where Galliformes bones show numerous butchering marks [1]. Compared to hunting practices, gath- ering and fishing leave less visible traces in the archaeological record. As far as bone remains are concerned, fishing is testified only at a few sites including Riparo Dalmeri (Trento) [3], Villabruna (Belluno) [4] and Riparo Soman (Verona) with the presence of some vertebras ofCyprinidae, Salmo trutta,Thymallus thymallusandExos lucius. Impor- tant traces of fish consumption also come from Riparo Tagliente [5], where the identification of species is still in progress. By contrast, fish remains from Riparo Cogola (Trento) are probably the result of carnivores’activities [6].

In spite of the discovery of freshwater fish from these sites, archaeological data do not allow to understand the role of this activity in human subsistence in relation to hunting practices [3–6]. We should also consider the possibility that fish could be consumed in a place other than the excavation site [7]. Therefore, the comparison between archaeological and isotopic data is applied here in order to get a better insight into the different resources exploited by human groups.

Nowadays, the reconstruction of prehistoric diets is based on the application of different approaches. Among these, stable carbon and nitrogen isotope analysis provides individual dietary information [8]. The study of animal and vegetal remains tends to be affected by several cultural, environmental and taphonomic factors; whereas the assumption that the isotopic composition of an animal tissue is a direct constant function of the diet makes stable isotope analysis a relevant tool to document the consumption of proteins [9]. Compared to zoological and botanical data, it helps to reconstruct past human and animal diets and to detect changes in subsistence strategies, often linked to environmental and cultural trans- formations [10,11]. As collagen turnover is continuous and slow in bones, it provides information on the proteins consumed over the last years of the individual’s life (ca. last decade) [8]. In particular, carbon isotope ratios (¹³C/¹²C) indicate the environmental origin of proteins (terrestrial vs.

marine) [12]. Theδ¹³C values from plants depend on their photosynthetic pathway: in the terrestrial ecosystems C3-, C4

- and CAM-type plants are present [13]. Considering the quasi-absence of C4plants in Pleistocene Europe, theδ¹³C values of animal bone collagen allow an estimation of the consumption of marine proteins, the latter being more enriched in¹³C than terrestrial proteins [12,14,15]. Nitrogen isotope ratios (¹⁵N/¹⁴N) indicate the trophic level at which

the individual is operating (herbivore vs. carnivore): the δ¹⁵N value detected in the bone collagen of the consumer is enriched by 3–5‰in comparison with the δ¹⁵N values of the prey [16,17]. An example of marine protein consumption in human diet is provided by the Gravettian burial, known as“Il Principe”, coming from Arene Candide (Finale Ligure, Savona) [18]. Considering Holocene end-point values of −20 ± 1‰for a 100% terrestrial diet and−12 ± 1‰ for a 100% marine diet, the authors suppose that the values detected in this individual (δ¹³C:−17.6‰andδ¹⁵N:

12.6‰) indicate a 20–25% consumption of marine protein probably coming from the Mediterranean Sea [18]. Isotopic analyses in Neolithic contexts have also underlined δ¹³C value of −18‰ relating to human and animal individuals that did not consume marine foods [19]. We may thus assume that the recognition of the consumption of marine resources in bone collagen is possible only if it is superior to 20–25% of the total proteins taken by the individual.

In Italy, the first stable carbon and nitrogen isotope analysis on bone collagen from a Palaeolithic context was applied by Francalacci [20] to two Epigravettian skeletons from Arene Candide (Finale Ligure, Savona). Isotopic data from Arene Candide 17 and 18 did not show a significant consumption of marine resources, which represents an unexpected result for a coastal site (Table 1). These results are in contrast with the clear contribution of marine foods documented in the diet of the Gravettian skeleton from the same site (Table 1)– “Il Principe” – which was studied by Pettitt et al [18], as just reported. More recently, new analyses have been conducted on the Villabruna skeleton (Sovramonte, Belluno) [21], the nine burials from Grotta del Romito (Papasidero, Cosenza) [22,23], a sample of human remains (n = 5, NI = 4: San Teo- doro 1, 2, 7 and Addaura 1) from Grotta di San Teodoro (Acquedolci, Messina) and Grotta dell’Addaura (Palermo) (Table 1). Particularly, the Villabruna hunter (KIA-27004:

12,140 ± 70 BP; 14,160−13,820 cal BP) shows a diet mainly based on the consumption of terrestrial proteins coming from herbivores [21]. At Grotta del Romito, most individuals had a similar terrestrial diet with the exception of Romito 9 dating to several millennia earlier than the others (LTL-3034A:

13,915 ± 70 BP; 17,000–16,150 cal BP) for whom a more varied diet based on both freshwater and/or marine fish in addition to terrestrial animals has been detected [22]. A difference in the diet of the individuals from the two sicilian sites has been noted by Mannino et al [23]. According to these authors, the hunter-gatherers from Grotta di San Teodoro had access to resources such as anadromous brown trout (Salmo trutta) which might not have been similarly available in the north western part of the island at the Addaura caves. In general, most authors argue that the limited evidence of the consumption of marine resources in the Italian peninsula could be a consequence of the low productivity of the Medi- terranean Sea compared to the Atlantic Ocean as previously

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Table1CarbonandnitrogenvaluesofhumanremainsfromUpperPalaeolithicsitesinItaly.CalibrateddatawereobtainedusingCalibversion6.0/Valeursisotopiques(CetN) dessujetshumainsdesitespaléolithiquessupérieursenItalie.Lesdatationssontcalibréessuivant«Calib6.0». SiteLocation ma.s.l.

BurialSexAgeLabCode14 CBPcalBP(2σ)PeriodC:Nδ13 C (‰)δ15 N (‰)

References Arene Candide, FinaleLigure (SV)

Coastal, 90m IlPrincipeMAdolescentOxA-1070023,440±19028,626−27,834Gravettian3.2−17.612.6Pettittetal2003 Arene Candide17

IndtYoung3.2−20.08.9Francalacci1988 Arene Candide18

MIndt3.4−18.99.1Francalacci1988 Grottadel Romito, Papasidero (CS)

Inland, 275m

Romito2FAdolescent3.5−20.010.3Craigetal2010 Romito3MYoungadult3.3−19.310.1Craigetal2010 Romito4FYoungLTL-3032A11,340±9013,417−12,984LateEpigravettian (Bølling–Allerød) 3.3−19.610.0Craigetal2010 Romito5FYoung-adultLTL-3033A10,862±7012,918−12,597LateEpigravettian (DryasIII)

3.3−19.79.3Craigetal2010 Romito6MAdult3.4−19.58.9Craigetal2010 Romito7MYoung3.2−19.19.7Craigetal2010 Romito8MAdult3.3−19.59.7Craigetal2010 Romito9IndtAdultLTL-3034A13,915±7017,208−16,776LateEpigravettian (Firstpart Lateglacial)

3.2−18.912.4Craigetal2010 Villabruna, Sovramonte (BL)

Inland, 515m

Villabruna1MYoungadultKIA-2700412,140±7014,183–13,797LateEpigravettian (Bølling–Allerød)

3.6−19.78.0Vercellottietal 2008 Addaura, Palermo (PA)

Coastal, 14m

Addaura1KIA-3605512,890±6015,950–15,007LateEpigravettian (Firstpart Lateglacial)

3.3−19.79.6Manninoetal. 2011 SanTeodoro, Acquedolci (ME)

Coastal, 135m S.Teodoro1FYoungadultETH-3445112,580±13015,232–14,126LateEpigravettian (Bølling–Allerød)

3.5−20.012.5Manninoetal. 2011 S.Teodoro2IndtAdultLateEpigravettian3.6−20.012.0Manninoetal. 2011 S.Teodoro2IndtAdultLateEpigravettian3.9−20.211.4Manninoetal. 2011 S.Teodoro7IndtAdultLateEpigravettian3.3−19.111.5Manninoetal. 2011

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assumed for other Mediterranean contexts (e.g. Balma Guila- nyà) [24]. By contrast, a different explanation to this situation could be connected to the absence of an adequate technology, which may have prevented Italian Late Epigravettian groups to carry out intensive fishing [23].

Based on the analysis of isotopic data from human collagen and for comparison on animal collagen, this study attempts to reconstruct the ecosystems exploited by the Late Epigravettian individual buried at Riparo Tagliente (Stallavena di Grezzana, Verona) (Fig. 1). In particular, it aims at understanding the role of different species, as terrestrial and aquatic ones, in human diet. Even if stable isotope analyses allow distinguishing fish from other food resources, the range of values they exhibit is wide due to local conditions [7]. Therefore, in order to make inferences on resources consumed, the establishment of a local baseline value is required. Unfortunately, due to the lack of determined fish remains available for isotopic data at Riparo Tagliente, bib- liographical data have been considered [25–27]. Further- more, this article attempts to infer hypotheses about the sea-

sonal mobility routes of hunter-gatherers and to establish if these involved or did not involve the Adriatic coast. Con- tacts with this region are testified by the discovery of marine shells in some inland sites, i.e. Riparo Tagliente (see below) [28]. Nevertheless, the origin of these shells is unknown and most probably they represent the results of exchanges with other groups. With regards to settlement strategies after the LGM (50,000-10,000 BP) Broglio (1980) has pro- posed a “vertical seasonal nomadic system”[29]. Hunter- gatherers groups moved seasonally and locally from residen- tial winter sites located on the Adige valley floor to high altitude summer [30].

Background: the site of Riparo Tagliente The Epigravettian occupation

The site of Riparo Tagliente is located in the village of Stal- lavena di Grezzana (Verona) on the left slope of Valpantena, one of the main valley bottoms of the pre-alpine massif of Monti Lessini (Fig. 1). The shelter opens at the base of Monte Tregnago, in the Lessini Mountains, under a bank of oolitic limestones at an altitude of 250 m a.s.l. [31,32].

The rock-shelter occupies a strategic position, about halfway both from the plain (ca. 100 m a.s.l.) and the top of the limestone plateau (ca. 2000 m a.s.l.) and at the cross-way between different topographic situations: the plain, the valley bottom, the rocky slopes and the top of the massif. Such a variety of situations matches with a rich mosaic of land- scapes with different faunal and vegetal resources, which varied in their distribution along time [5].

The site was discovered in 1958 by Francesco Tagliente.

Archaeological investigations were carried out by the Museo Civico di Storia Naturale of Verona from 1962 to 1964 and resumed after 1967 by the University of Ferrara. Excavations have brought to light a stratigraphic series over 4.50 m deep.

It is formed by two main deposits separated by a river erosion:

the lower deposit contains Mousterian and Aurignacian industries, while the upper one is characterized by a Late Epi- gravettian record (Table 2) [31,33]. According to radiocarbon dates which range from 16,932–15,495 cal BP (layer 15) to 14,572–13,430 cal BP (levels 10–8)¹[33–35], the Epigravet- tian series is one of the most complete in northern Italy spanning between the first part of the Lateglacial and the beginning of the Bølling–Allerød interstadial (Table 2) [28].

During the first part of the Lateglacial, an Epigravettian individual was buried in the area protected by the over- hang of the shelter. The body was discovered during archaeological investigation in 1973 [36–38]. Unfortunately, a Fig. 1 Map of Italy showing the location of Riparo Tagliente. Dur-

ing the first part of the Late Glacial the coastline was located about 360 km from the site (see the coastlines shift between 20 ka, 17 ka and 14 ka cal BP) /Localisation du site de Riparo Tagliente (Italie).

Au cours de la première partie du dernier glaciaire le littoral se situait à environ 360 km du site

1All the samples were calibrated using the software Calib version 6.0 [26].

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Medieval pit had removed the upper part of the burial (Fig. 2). Only a few ribs and vertebrae, the distal fragments of the radius and ulna and some phalanxes were discovered on the trampling floor, while the lower part of the skeleton– from the pelvis to the feet–was well preserved in the grave [36]. The pit had an oval shape and was about 60 cm wide

and 60 cm deep with a concave section. The bottom was higher at the southern edge and the feet were located at a higher position than the pelvis. The same situation probably occurred at the opposite edge where the skull was originally located [36]. The skeleton had been laid in a supine position with outstretched arms–the left arm was slightly bent–and Table 2 Radiocarbon dates of human remain and charcoals from the Epigrattian layers of Riparo Tagliente. Data were calibrated using Calib version 6.0 /Datations radiométriques du sujet humain et de charbons des couches épigravettiennes de Riparo Tagliente.

Les données ont été calibrées suivant « Calib 6.0 ».

Level Sample Method Lab Code ¹⁴C BP cal BP (2σ) Period

8–10 Charcoal CONV R-371 12,040 ± 170 14,572–13,430 Bølling–Allerød

10a Charcoal AMS OxA-3530 12,650 ± 160 15,632–14,111 First part Lateglacial, Bølling–Allerød 10c Charcoal AMS OxA-3531 13,070 ± 170 16,596–15,117 First part Lateglacial

10e Charcoal AMS OxA-3532 13,270 ± 170 16,785–15,273 First part Lateglacial 15 Charcoal CONV R-605 13,330 ± 160 16,851–15,297 First part Lateglacial 15–16 Charcoal CONV R-605α 13,430 ± 180 16,932–15,495 First part Lateglacial Burial Human bone AMS OxA-10672 13,190 ± 90 16,634–15,286 First part Lateglacial

Fig. 2 View of the incomplete human skeleton after the excavation of the Epigravettian burial from Riparo Tagliente /Vue du squelette incomplet après la fouille de la sépulture épigravettienne de Riparo Tagliente

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the head facing north. Due to the incompleteness of the burial no information is available on the original composition of the grave assemblage. Only a limestone pebble covered with some traces of ochre was found between the feet and a fragment of bovid horn near the right femur (Fig. 2).

The intentional deposition of a piercedCyclopenear the left knee is uncertain [36]. After the deposition of the body, the legs were covered with local stones of different dimensions.

A large stone located on the femurs was engraved with the figures of a lion and the horn of an auroch [36,37]. Based on pelvic and pubic surface morphology, anthropological studies support that the skeleton belonged to a young adult male aged 20–29 years [38]. The reason of his death is unknown.

His stature was about 163 cm, i.e. intermediate compared to other Upper Palaeolithic human individuals [38]. A radiocarbon date was obtained from the analysis of a bone fragment of the right femur giving an age of 13,190 ± 90 BP (OxA- 10672) and 16,634–15,286 cal BP (Table 2). This date confirms that the young hunter was buried in the shelter during the first part of the Lateglacial [37,39].

After the LGM, the major glaciers retreated from the Alpine pedemontane amphitheatres [40]. Pollen analyses carried out on the Epigravettian series of Riparo Tagliente indicate that during the first part of the Lateglacial, the landscape was dominated by herbaceous plants (Graminaceae, Artemisia) with some conifers and colonizing plants (Pinus sylvestris-mugo, Juniperus, Salix) (layers 16–15). In the Lateglacial interstadial, woodland expansion with conifers (Pinus sylvestrisemugo) and deciduous trees began (Quer- cussp.,Tiliasp.,Ulmussp.,Ostrya carpinifolia,Fraxinus, Corylus) (layers 14–5) [31]. The analysis of malacofauna

also testifies a progressive climate improvement along the Epigravettian series: xero-thermophilous species (layers 12–10) replace the most ancient association (layers 18–15).

The micromammal assemblage is mostly composed of fos- sorial species (Pytimys savii, Microtus nivalis, Arvicola, Microtus arvalis-agrestis) and is characteristic of a mountain steppe [31,32].

The distribution of ungulates and other mammals along the Late Epigravettian sequence highlights an opposition between the lower layers (17–14), characterized by species adapted to an open environment and the higher ones (12–5) which document the development of milder climatic conditions. Only in the layer which is coeval to the burial (layer 13) both associations are present and equivalent [28,41].

In particular, this assemblage is dominated by ibex (29%

Capra ibex), marmot (29% Marmota marmota) and red deer (20% Cervus elaphus) and to a lesser extent by roe deer (9% Capreolus capreolus), bison/aurochs (4% Bos/

Bison), wild boar (1.7% Sus scrofa), elk (1.6%Alces sp.), bear (1.5% Ursus arctos) and chamois (1.2% Rupicapra rupicapra) [41] (Fig. 3). The analysis of butchering marks confirms that herbivores represented one of the main food resources [28]. Fish and bird remains are also documented.

Nonetheless, a detailed analysis is still missing and no pre- cise information about their consumption is available from archaeological data. Data on the seasonality of occupation indicate that the site was occupied all year long with a greater emphasis on spring and summer time [28,41].

All Late Epigravettian layers are rich in lithic industry documenting an intense knapping activity and on-site blanks transformation. The identified chaînes opératoires

Fig. 3 Percentage of terrestrial faunal remains for species discovered in layers 13 from Riparo Tagliente /Distribution (%) des restes de faune terrestre de la couche 13 de Riparo Tagliente

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were aimed at producing a wide variety of rectilinear lam- inar blanks and small curved bladelets [28]. Raw materials are almost exclusively local and come from the Lessini area within a range of about 15 km from the shelter.

Bone and antler industries confirm the high technical abil- ity and good knowledge of animal hard tissues. The most frequent typological classes of tools are points and awls, followed by spatulas and tools with a distalbiseau. A further category is represented by ornamental elements mainly composed of perforated mammal teeth, namely red deer atrophic canines. Also shell ornaments are abun- dant: one third of the malacological collection is characterized by at least one intentional perforation. Almost all determined species are widely diffused in the Mediterra- nean Sea. The assemblage is dominated byCyclopeand it seems to be the result of human selection. A group of mobile art objects is also of great importance: most of them are engraved with both geometric and figurative representations. A high quantity of ochre nodules and fragments are documented in the Late Epigravettian series of Riparo Tagliente [28].

A note on the ancient coastline

A compilation of global sea-level estimates based on deep- sea oxygen isotope ratios at a millennial-scale resolution and even higher has been published since the 1970s. Nonethe- less, these global sea-level curves [42] do not take into consideration isostasy and vertical tectonics movements.

Sea level change is the sum of eustatic, glacio-hydro- isostatic and tectonic factors. The first is time-dependent while the latter two vary according to the location. This means that at the same time slices the relative sea level (the sum of the three different movements) should be different in different coastlines.

In order to reconstruct the Adriatic sea palaeocoastline in prehistoric times, we have used the predictions recently published for Italian sea since the LGM [43]. Lambeck’s model has been tested in the stable areas of Italy on hundreds of sites by using geological, geomorphological and archaeological markers; furthermore, on the basis of observed data this model has undergone little changes.

We have used these predicted sea level curves that take in account the eustatic and glacio-hydro-isostatic movements.

The Northern Adriatic sea suffers from tectonic and sedi- ment compaction [44,45]; these movements vary between

−1 and−0.1 mm/year. Central and southern Adriatic coast do not show significant vertical tectonic movements.

The calibrated radiocarbon ages of Table 2 range between 14.0 and 17.1 ka cal BP. Considering the Adriatic sea bathi- metry (Fig. 1), the altitude of sea level predicted by Lambeck et al (2011) [43] and the radiocarbon data obtained for

Riparo Tagliente (−81 m at 14 ka cal BP and −102 m at 17 ka cal BP), the prehistoric palaeocoastlines were located within the latitude of the central-southern Adriatic sea. These data have been obtained without applying tectonic correc- tions. In Fig. 1, we have drawn the LGM [46] and the 14 and 17 ka cal BP coastlines.

Materials and methods

Samples selected for stable isotope analysis

Twelve samples have been analysed for stable carbon (¹³C/

12C) and nitrogen isotope ratios (¹⁵N/¹⁴N) at Riparo Tagliente². Fragments of thick and compact bones (i.e. diaphysis) were selected avoiding spongy tissue, burnt bones and remains with cut-marks on the surface.

The human bone sample was taken from a broken fragment of a rib at the Museo Civico di Storia Naturale of Ver- ona where the skeleton is currently on display. In order to provide comparative data to interpret human isotope values, 11 adult bone samples were selected from the faunal assemblage stratigraphically associated with the burial (Table 3).

The well-represented faunal assemblage has given us the possibility to study different species with a known diet (Table 3). The samples of seven herbivores–two red deer (Cervus elaphus), one roe deer (Capreolus capreolus), two ibex (Capra ibex), one chamois (Rupicapra rupicapra), one bovid (Bos/Bison) – and four omnivores – two marmots (Marmota marmota) [47], one fox (Vulpes vulpes), one wild boar (Sus scrofa)–were analysed in order to interpret the values of the Late Epigravettian human individual. No fish remains were sampled since no detailed analysis is available yet. The collagen extraction was conducted in the

“Laboratoire de biochimie de l’Unité d’anthropologie de Marseille”(UMR CNRS 6578 UAAB) and stable isotopic analyses were performed at Iso-Analytical Ltd (Crewe, UK).

Collagen extraction and isotope ratio mass spectrometry

The collagen extraction from the 12 archaeological samples (1 human and 11 faunal bones) was performed according to the procedure described by Longin (1971) and Bocherens et al (1997) [48,49]. Each bone was cleaned by abrasion and washed with distilled water. Acetone was used when traces of consolidants were present on the samples. All the cleaned samples were ground into a powder (0.7 mm sieve).

2The sampling authorization was delivered by the Soprintendenza per i Beni Archeologici del Veneto and carried out with the collaboration of Dr. Aspes (Museo Civico di Storia Naturale di Verona) and Dr.

Giacobini (Università degli Studi di Torino).

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Approximately 600 mg of archaeological bone powder was used, as well as modern bovine bone powder as a bug sample to make sure that the extraction procedure had been carried out correctly. Bone powder is first demineralized in HCl solution (1 M, 20 min, room temperature), then the residue is soaked in NaOH solution (0.125 M, 20 h, room temperature) and solubilized in a weak acid solution (HCl, 0.01 M, 17 h, 100°C). The dissolved collagen solution is filtered and freeze-dried for 48 h to be ready for analysis.

Elemental compositions (C, N) and stable isotope ratios (¹³C/¹²C and ¹⁵N/¹⁴N) of the extracted collagen were performed using an Elemental Analyser (Europa Scientific) connected to an Isotope Ratio Mass Spectrometry (Europa Scien- tific 20-20 IRMS). IAEA standards used during the analysis of samples are the following: bovine liver (IA-R042), ammonium sulphate (IA-R045), beet sugar (IA-R005) and sugar cane (IA- R006)³. Stable isotope ratios are expressed with delta notation (δ¹³C,δ¹⁵N) in parts per thousand (‰) relative to international standards, respectively VPDB for C and AIR for N. Collagen preservation was tested according to DeNiro (1985), Ambrose (1990) and van Klinken (1999) criteria thus providing a satis- factory quality and quantity control. These criteria can be sum- marized as follows: yield≥10 mg/g [50], %C≥30% and %N≥ 11% [51], C:N ratio within the range of 2.9 to 3.6 [52]. Repro- ducibility of results is inferior to 0.1‰for bothδ¹³C andδ¹⁵N.

Results and discussion

Stable isotope results, collagen quality indicators and other details obtained after human and faunal bone samples analyses are presented in Table 3. Carbon (δ¹³C) and nitrogen (δ¹⁵N) values of human and faunal bone collagen are plotted in Figs 4, 5.

Bone collagen preservation

All the analysed samples satisfied the standard protocol for the control of collagen quality and quantity (Table 3) [50–52]. The collagen yield of each sample was clearly higher than 10 mg/g as established by Ambrose (1990) [50] and with an average value (n = 12) of 59.0 ± 32.9 mg/

g. Collagen was well preserved with C and N weight percentage values respectively included between 28.8% and 36.9% and between 11.2% and 13.1%. The isotopic integrity of collagen is also demonstrated by the atomic C:N ratios, which range between 3.1 and 3.3 [52].

Carbon and nitrogen isotope ratios from herbivores’ specimens

At Riparo Tagliente, theδ¹³C values of herbivores (n = 7) range between −20.5‰ and −18.4‰ (Table 3) and are therefore consistent with the consumption of C3 plants [53]. The carbon stable isotope value (−19.6 ± 0.8‰) falls on the least negative range of herbivore values coming from modern contexts of western Europe which range between −25.3‰ and −18.5‰ [54]. The same results were obtained from other isotopic studies: general ¹³C depletion was observed at the beginning of the Bølling– Allerød interstadial until the Atlantic when values stopped on the current average [54,55]. Authors agree that the decreasedδ¹³C values of herbivores during the Lateglacial and the Early Holocene transition are related to a decrease inδ¹³C value in the consumed plants [54–57]. At the origin of this depletion there were probably global and local factors such as the spread of forest vegetation [54,56,57]. As a result of global climatic warming after the LGM, the vegetation composition shifted from a steppe-tundra dominated by grasses and herbaceous species to a temperate dense Table 3 Isotopic results for human and animal remains from Riparo Tagliente /Résultats isotopiques du sujet humains et de la faune de Riparo Tagliente.

Lab code Taxon Bone sample Yield (mg/g) %C %N C:N δ¹³C (‰) δ¹⁵N (‰)

RT1 Homo sapiens Rib 105.9 36.1 12.8 3.3 −18.4 13.0

RT2 Capra ibex Humerus 16.8 28.8 11.2 3.0 −18.9 1.2

RT3 Capra ibex Humerus 68.7 34.8 12.3 3.3 −19.1 2.5

RT4 Cervus elaphus Metatarsal 52.3 34.2 12.7 3.1 −20.0 2.1

RT5 Cervus elaphus Tibia 37.1 32.2 11.5 3.2 −20.5 2.0

RT6 Capreolus capreolus Metacarpal 45.0 33.5 11.9 3.3 −20.5 5.0

RT7 Rupicapra rupicapra Metatarsal 85.6 34.0 12.4 3.2 −18.4 2.7

RT8 Bos/Bison Metacarpal 49.1 34.4 12.2 3.3 −19.6 5.8

RT9 Marmota marmota Ulna 23.5 31.7 11.2 3.3 −19.8 4.8

RT10 Marmota marmota Radius 31.3 31.9 11.2 3.3 −20.6 4.2

RT11 Vulpes vulpes Metatarsal 124.0 36.9 13.1 3.3 −19.3 9.4

RT12 Sus scrofa Mandible 68.5 34.0 12.0 3.3 −19.4 5.0

3http://www.iso-analytical.co.uk/standards.html

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deciduous forest affecting the carbon isotopic composition of plants and consequently the isotopic signature of herbivores [40]. Dated to the first part of the Lateglacial (Table 2), the high δ¹³C values of the faunal samples

from Riparo Tagliente can be explained both by climatic factors and the landscape features that characterized the area surrounding the site which was dominated by a wooded grassland [31,55,58].

Fig. 5 δ¹³C andδ¹⁵N mean and standard deviation of herbivores and omnivores remains from Riparo Tagliente, as well as human values / Moyenne et écart-type des valeurs isotopiques des herbivores, des omnivores et du sujet humain de Riparo Tagliente

Fig. 4 δ¹³C andδ¹⁵N values for human and faunal bone collagen from Riparo Tagliente / Valeurs deδ¹³C etδ¹⁵N des collagènes osseux du sujet humain et de la faune de Riparo Tagliente

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The range ofδ¹³C values (2.1‰) reflects ungulates carbon isotopic variability, and thus habitat variability. Even if these carbon values would indicate open habitats, the fact that some ungulates exhibit lowerδ¹³C values–i.e. red deer and roe deer (n = 3:−20.3 ± 0.3‰) than others, i.e. ibex and chamois (n = 3:−18.8 ± 0.4‰)–would suggest a consumption of food items isotopically different, as different type of trees or grasses according to species⁴. Moreover, chamois and ibex, which feed mostly on lichens in winter time and during per- iods of food shortages, display higherδ¹³C values than the

coeval ruminants [59]. Lichens exhibit more positive δ¹³C values than their coeval vascular plants and their consumption affects the isotopic value of the animals that feed on them as already observed for reindeers [58,60]. In addition, this difference could also underline that, in this study, ungulates could occupy different ecosystems at diverse altitudes: red deer and roe deer lived in the sparse forest on the plain and the valley bottom [40] while ibex and chamois could occupy the open steppe environments of the mid-altitudes [31] having as their main food resource vascular plants enriched in ¹³C compared to plants from under more closed environment like undergrowth [54,58]. Theδ¹³C values of the herbivores Table 4 Stable isotope values (δ¹³C, δ¹⁵N) of freshwater fish remains dating at the Early Holocene from European sites [62–64] / Données isotopiques (δ¹³C,δ¹⁵N) de poissons d’eau douce de site européens datés du début de l’Holocène [62–64].

Taxon Site Period Habitat δ¹³C δ¹⁵N References

Acipenseridae Vlasac (Danube) Pre-Neolithic Fluvial or lacustrine −19.7 7.8 Nehlich et al 2010

Anguilla anguilla North France ~8,000 BP Fluvial −23.8 8.3 Bocherens et al 2007

Anguilla anguilla SW France ~10,000 BP Fluvial −23.7 8.0 Bocherens et al 2007

Anguilla anguilla SW France ~12,500 BP Fluvial −20.8 8.5 Bocherens et al 2007

Cyprinidae Vlasac (Danube) Pre-Neolithic Fluvial or lacustrine −18.9 6.8 Nehlich et al 2010 Cyprinidae Vlasac (Danube) Pre-Neolithic Fluvial or lacustrine −20.1 7.4 Nehlich et al 2010

Cyprinidae SW France ~9,500 BP Fluvial −21.5 9.4 Bocherens et al 2007

Exos lucius SW France ~9,500 BP Fluvial −22.2 9.5 Bocherens et al 2007

Exos lucius Holmegard I (Denmark) Mesolithic Fluvial or lacustrine −15.4 7.8 Fischer et al 2007 Exos lucius Holmegard IV (Denmark) Mesolithic Fluvial or lacustrine −22.8 10.0 Fischer et al 2007 Exos lucius Mullerup (Denmark) Mesolithic Fluvial or lacustrine −9.3 9.4 Fischer et al 2007 Exos lucius Mullerup (Denmark) Mesolithic Fluvial or lacustrine −8.0 9.2 Fischer et al 2007 Exos lucius Mullerup (Denmark) Mesolithic Fluvial or lacustrine −9.5 8.7 Fischer et al 2007 Exos lucius Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −22.6 7.9 Fischer et al 2007 Exos lucius Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.3 8.0 Fischer et al 2007 Exos lucius Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −22.8 7.6 Fischer et al 2007 Exos lucius Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −24.0 8.1 Fischer et al 2007 Exos lucius Storelyng VI (Denmark) Kongemose/

Ertebølle

Fluvial or lacustrine −24.0 7.8 Fischer et al 2007

Exos lucius Storelyng VI (Denmark) Kongemose/

Ertebølle

Fluvial or lacustrine −25.9 6.6 Fischer et al 2007

Exos lucius Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −22.6 9.1 Fischer et al 2007 Exos lucius Akonge (Denmark) Neolithic Fluvial or lacustrine −22.3 7.9 Fischer et al 2007 Exos lucius Akonge (Denmark) Neolithic Fluvial or lacustrine −22.4 8.8 Fischer et al 2007 Exos lucius Akonge (Denmark) Neolithic Fluvial or lacustrine −21.9 7.3 Fischer et al 2007 Exos lucius Argus (Denmark) Kongemose Fluvial or lacustrine −13.3 11.8 Fischer et al 2007 Exos lucius Bøgebjerg (Denmark) Ertebølle Fluvial or lacustrine −23.1 12.5 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.6 5.0 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.0 5.5 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.7 5.4 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.4 5.5 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −25.9 5.6 Fischer et al 2007 Tinca tinca Storelyng VI (Denmark) Neolithic Fluvial or lacustrine −26.2 5.8 Fischer et al 2007 Tinca tinca Akonge (Denmark) Neolithic Fluvial or lacustrine −25.7 5.6 Fischer et al 2007 Tinca tinca Akonge (Denmark) Neolithic Fluvial or lacustrine −25.5 4.7 Fischer et al 2007

4Mann–Whitney U test, p≤0.05.

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from Riparo Tagliente compared with the same data from Grotta/Riparo del Romito are relatively more positive [22].

This difference may be due to the consumption of plants growing at different altitude and climatic conditions. In particular, north-eastern Italy is characterized by the presence of the Alps and therefore it has been more affected by the effects of glaciation processes than southern Italy [1].

By contrast, the herbivores from Riparo Tagliente show a wide range (4.6‰) ofδ¹⁵N values, ranging from 1.2‰to 5.8‰ (Table 3). The average value (3.0 ± 1.7‰) is much lower than the one recorded by Holocene European herbivores [61,62]. Nonetheless, it matches the values obtained for other European sites dated to the same period [55–57].

Low δ¹⁵N values can be explained by climate-driven edaphic effects and are particularly linked to the colder climatic conditions associated with the LGM, between 20,000 and 10,000 BP [55]. In Italy, lowδ¹⁵N values have already been recorded on a deer bone (1.6‰) from the Late Epigravettian site of Riparo Villabruna in Val Cismon (Belluno, Veneto, Italy) [21]. In this case, two different factors interact resulting in the value obtained: the chronologi- cal attribution of the burial to the first warm phase of the Late Glacial (Bølling–Allerød interstadial) and the geographical position of the site which is located in a narrow Alpine valley. On the other hand, no change has been identified in the bone collagen δ¹⁵N values of Upper Palaeolithic faunal remains coming from sites located in the southern part of the peninsula such as Grotta Paglicci (Rignano Garganico, Foggia, Puglia) [62], Arene Candide (Finale Ligure, Savona, Italy) [20] and Grotta del Romito (Papasidero, Cosenza, Italy) [22] possibily in relation to the more temperate climatic conditions of the areas where they are located.

At Riparo Tagliente, among herbivores the auroch (Bos/

Bison n = 1, δ¹⁵N: 5.8‰) shows the highest δ¹⁵N value.

This value is even higher than the one obtained on some omnivores (i.e. the marmot and the wild boar). A similar isotope value has been observed in the bovine collagen of other Euro- pean Upper Palaeolithic sites, e.g. Grotta Paglicci (Rignano Garganico, Foggia, Italy) [62], Grotta del Romito (Papasidero, Cosenza, Italy) [22] and Saint-Germain-La-Rivière (Dor- dogne, France) [65]. Craig et al [22] explain this result as a physiological difference or a dietary difference between forest or mountain living species (such as ibex, roe deer and wild boar) feeding on¹⁵N depleted woody shrubs and open living species (such as aurochs) on relatively¹⁵N enriched grasses [64]. It also seems to improve the idea that ungulates from Riparo Tagliente occupied different ecosystems, feeding different types of plants.

Carbon and nitrogen isotope ratio in omnivore’s species

The average isotopic values of the four analysed omnivores from Riparo Tagliente are as follows: 19.8 ± 0.6‰forδ¹³C

and 5.8 ± 2.4‰forδ¹⁵N. Theδ¹³C values range from−19.8 to−19.3‰with a narrow variation of 1.3‰(Table 3). They are included between the highest (ibex and chamois) and the lowest (red deer and roe deer)δ¹³C values of the herbivores.

This value may be related to the effect of the omnivores’ great adaptability (e.g. wild boar and fox) to different food resources from different habitats. On the other side, the wide range of δ¹⁵N values (5.2‰) showed by omnivores is included between 4.2‰ and 9.4‰ and it is related to the different trends in animal protein consumption that charac- terize the different omnivores species (Table 3, Figs 4, 5).

From this point of view, the δ¹⁵N values of marmot and wild boar range between the values of herbivores. This may be due to their diet which is mainly based on the consumption of plant proteins and only occasionally of bugs and little birds [66]. The fox shows isotopic values higher than those of herbivores respectively of +0.3‰ for δ¹³C and 2.8‰for δ¹⁵N (Fig. 4). The position of the fox at the top of the food chain is consistent with a great consumption of terrestrial animal proteins (small mammals and rodents) rather than of plant proteins (berries, fruits, herbs, etc.) on which it feeds only during summer and autumn [66,67].

Human diet: terrestrial and aquatic resources consumption

Theδ¹³C (−18.4‰) andδ¹⁵N (13.0‰) values of the human individual buried at Riparo Tagliente are higher than +1.2‰ for carbon and +10.0‰for nitrogen compared to herbivores, and higher than +1.4‰for carbon and +7.2‰for nitrogen compared to omnivores (Table 3, Figs 4, 5). The isotopic signal therefore indicates an emphasis on the consumption of animal rather than plant resources and within the animal items those with a high trophic level as carnivore or fish species. Indeed, ungulates did not represent the main protein source of human diet. In fact, the δ¹⁵N value of 13.0‰ obtained from the analysis of the bone collagen of this individual is over 8‰ above the average value obtained on the coeval faunal remains analysed at Riparo Tagliente (4.1 ± 2.3‰) and much higher than the suggested range of 3–5‰indicated for a trophic level shift [16]. Considering theδ¹⁵N value of fox which is higher than the one of herbivores (on average +6.4‰) and lower than the value of the human individual (on average +3.6‰; Table 3, Figs 4, 5), a consumption of meat from fox and/or other carnivores could explain theδ¹⁵N value of the human individual. However, zooarchaeological and archaeological data do not show any evidence of a significant consumption of carnivores’ meat [41]. Indeed, fox (4%) and other carnivores (Canis lupus, Panthera leo spelaea,Linx linx) are poorly documented in the layers coeval to the burial [41]. Moreover, carnivores’ hunting would represent a more difficult task than that of

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herbivores, the carcasses of carnivores were probably more exploited for their fur rather than for meat.

The young individual from Riparo Tagliente could also have included another additional dietary source than carnivores’meat characterized by high δ¹⁵N values. Consider- ing the archaeological and geographical contexts and the expected isotopic distribution of values of dietary sources from marine and freshwater ecosystems (fish, shellfish and birds), the consumption of aquatic resources would be a relevant candidate to explain the elevated δ¹⁵N values of the hunter from Riparo Tagliente [7,26,68]. Information currently available on fish remains from the Late Epigra- vettian layers coeval to the burial does not allow so far either to identify the fish species that were consumed by the inhabitants of the shelter or to understand their origin– e.g. sea, lake or river. Nevertheless, the site location, far away from the Adriatic Sea (ca. 360 km), would suggest a preferential fishing activity in a local or circum-local environment. This hypothesis is also supported by the presence of different freshwater ecosystems in the area surrounding the site–including the Progno river flowing in front of the shelter, the Garda Lake (ca. 22 km) and the Adige River (ca. 15 km), although the possibility that the high humanδ¹⁵N value could derive from the consumption of marine resources cannot be totally excluded [28].

In order to detect the origin of the possible aquatic resources consumed, i.e. marine vs. freshwater, the isotopic literature provides several elements of comparison [7,68,69].

The relatively lowδ¹³C value of freshwater fish (<−17‰for modern samples)⁵[7] does not always allow to detect the potential incorporation of these aquatic proteins in human collagen [22,69]. If it is compared with δ¹⁵N values, it could give more information. After a review of the available literature, France (1995) asserts that marine fish is enriched in¹⁵N compared to freshwater specimens [68,70]. For estu- arine and anadromous fish, the author indicates intermediate δ¹⁵N values depending on the time spent feeding in either fresh or salted waters [68]. In such a context, freshwater fish could also be an excellent potential candidate to explain both δ¹³C (similar to terrestrial one) andδ¹⁵N (similar to aquatic resources eater) values of the human individual of Riparo Tagliente. In order to verify this hypothesis,δ¹³C andδ¹⁵N values available from the literature for Early Holocene⁶ archaeological remains of freshwater fish and from central Europe are considered [25–27] (Table 4). Bone collagen stable isotope values obtained from these remains and those from the human and faunal specimens of Riparo Tagliente

are plotted in Fig. 6. The human isotopic values are consistent with the consumption of freshwater species (Fig. 6).

Seemingly they are inconsistent with the range of values recently reported for archaeological and recent Mediterra- nean marine fish bone collagen [71,72]. In general, the bone collagen of marine consumers is definitively more enriched in¹³C compared to theδ¹³C value of the individual from Riparo Tagliente [17,18]. According to bibliographic data for freshwater fish species [62–64], the fish species which would display the values closer to those of the human specimen are pike (Exos lucius), sturgeon (Acipen- seridae) andCyprinidae, i.e. carps (Fig. 6). Pike and sturgeon are autochthonous species from the Po Valley and today they are widespread both in the Garda Lake and the Adige River. The ideal season to capture pikes is between spring and autumn during migrations from deep waters to the shore [73]. This feature of the pike ecology becomes more interesting if we consider that the main seasonal evidence of occupation at Riparo Tagliente is during spring and summer time [5]. As previously mentioned, remains of Cyprinidae, such as churbs (Squalius cephalus) and barbels (Barbus barbus), in addition to pikes (Exos lucius), trouts (Salmo trutta) and graylings (Thymallus thymallus), were discovered in some Late Epigravettian sites from Trentino and Veneto, i.e. Soman (Verona), Villabruna (Belluno), Dal- meri (Trento) and La Cogola (Trento) [3,4,6]. All these species live in the riverine and lacustrine biomes of north- eastern Italy and may have been captured by the Epigravet- tian groups during their seasonal migrations.

Conclusions

This article reports the first results of carbon and nitrogen stable isotope analysis on 1 human and 11 faunal bone specimens from the Late Epigravettian deposits of Riparo Tagliente (Stallavena di Grezzana, Verona) located in north-eastern Italy. As the stratigraphical position of the grave and a radiocarbon date (Table 2) undertaken on a bone sample taken from the skeleton of the individual indicate, the adult male was a member of one of the first groups of hunter-gatherers who had re-occupied the southern slope of the Alps after the LGM [6]. As documented by the faunal assemblage, the strategic position of this site, located about halfway from the plain and the top of the limestone plateau, had allowed these last Palaeolithic groups to exploit a wide variety of ecosystems (Fig. 3).

Isotopic data obtained on ungulates remains are consistent with the averageδ¹³C andδ¹⁵N values obtained on herbivores from mainland Europe dating from the cold period before the Bølling–Allerød interstadial [54–57]. In particular, theδ¹³C values of herbivores are probably strictly related to the consumption of plants growing in different ecosystem

5 Isotopic measurements were performed on bone collagen of teleostean fish from three European lakes (from Lake Geneva, Lake Constance, Lake Aiguebelette) and from Lake Baikal (Russia) by Dufour et al [59].

6No data for freshwater fish remains from Late Pleistocene contexts were discovered.