Journal of Archaeological Science: Reports 25 (2019) 129–143

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Journal of Archaeological Science: Reports

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Large bovids on the Neanderthal menu: Exploitation of Bison priscus and Bos T
primigenius in northeastern Italy
⁎
Gabriele Terlatoa,b, Alessandra Livraghia,b, Matteo Romandinia,b,c, Marco Peresania,
a
Università degli Studi di Ferrara, Dipartimento degli Studi Umanistici, Sezione di Scienze Preistoriche e Antropologiche, Corso Ercole I d'Este 32, 44121 Ferrara, Italy
b
Area de Prehistoria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, 43002 Tarragona, Spain
c
Dipartimento di Beni Culturali, Università di Bologna, Via degli Ariani 1, 48121 Ravenna, Italy

A R T I C LE I N FO A B S T R A C T

Keywords: In northern Italy, Fumane Cave (Lessini Mountains -Verona), San Bernardino Cave, and De Nadale Cave (Berici
Bison priscus Hills - Vicenza) provide data to interpret the exploitation dynamics of the Pleistocene's large bovids Bos primi-
Bos primigenius genius and Bison priscus between 70 and 42 ky BP. Through the taphonomic study of bone assemblages, we have
Middle Palaeolithic attempted to reconstruct the strategies, methods, and butchery practices in the exploitation of these game an-
Taphonomy
imals adopted by different cultural groups of Middle Palaeolithic hunters. Therefore, Neanderthal hunting be-
Hunting
Italy
haviour has been examined using different proxies such as the choice of anatomical parts, selective transport of
elements, prey selection, and age estimation. Results suggest bovines were an important subsistence resource in
some cases, even if they were not the most exploited taxa in others, indicating differences and similarities
depending on the context across the Italian peninsula. These results highlight additional aspects of Neanderthal
landscape use and hunting strategies.

1. Introduction 2012; Blasco et al., 2013; Costamagno et al., 2006; Finlayson et al.,
2012; Fiore et al., 2016; Gaudzinski, 2006; Romandini et al., 2018a,
One of the most controversial issues in the understanding of human 2018b). In general, the core meat sources of Neanderthals were
evolution is the debate over Neanderthal ecology, subsistence strate- medium-sized and large herbivores like cervids, equids and large bo-
gies, and diet (Gaudzinski-Windheuser and Roebroeks, 2011; Morin vids. Hunting practices focused on large-sized prey increased with the
et al., 2016; Ready, 2010, 2013). In recent years, several approaches, introduction of Levallois technology, a new knapping method that was
among them faunal and taphonomic analysis (Brugal et al., 1999; not a simple modification in flake production, but was part of relevant
Gaudzinski-Windheuser and Kindler, 2012; Germonpré et al., 2014; behavioural changes (Gaudzinski, 2006). Neanderthal subsistence tac-
Jaubert and Brugal, 1990; Patou-Mathis, 2000; Romandini et al., tics inferred from many archaeological sites in Europe comprised long-
2018a, 2018b), dental microwear patterns (Harvati et al., 2013; Pérez- term and seasonal exploitation strategies. Within cave and open-air
Pérez et al., 2003), tooth calculus analysis (Henry et al., 2014; Weyrich sites, which were regularly used for their abundance of biotic resources,
et al., 2017), lithic use-wear and residue analysis (Hardy and Moncel, some hunting locations were selected to take advantage of animal
2011), and the investigation of stable carbon and nitrogen isotopes of routes or seasonal migration, which indicates planning for the future
bone and tooth collagen (Bocherens et al., 2005; Bocherens, 2009; (Mellars, 1996).
Richards and Trinkaus, 2009; Wißing et al., 2016), as well as the Some of the best evidence of Neanderthal subsistence tactics and
contextualisation of the overall archaeological record have vastly im- hunting practices of large herbivores come from the zooarchaeological
proved our knowledge of food acquisition and exploitation and thus studies of bovines: Bison priscus and Bos primigenius. The study of several
subsistence strategies by Neanderthals. Among all the different bone assemblages suggests that bovines were an important source of
methods, there is common agreement that Neanderthal subsistence subsistence for Middle Palaeolithic hunters. Large accumulations of
strategies show great flexibility in foraging systems, with adaptations to butchered carcasses have been recovered in western-central Europe, at
different environments and variation across the faunal spectrum de- La Quina (Chase, 1999; Rendu and Armand, 2009), Mauran (David
pending on species availability (Blasco and Fernández Peris, 2009, et al., 1994; David and Fosse, 1999; Rendu et al., 2012), Coudoulous

⁎
Corresponding author.
E-mail addresses: gabriele.terlato@unife.it (G. Terlato), ales.livraghi@gmail.com (A. Livraghi), matteo.romandini@unibo.it (M. Romandini),
marco.peresani@unife.it (M. Peresani).

https://doi.org/10.1016/j.jasrep.2019.04.006
Received 31 July 2018; Received in revised form 26 February 2019; Accepted 9 April 2019
2352-409X/ © 2019 Elsevier Ltd. All rights reserved.
G. Terlato, et al. Journal of Archaeological Science: Reports 25 (2019) 129–143

(Jaubert et al., 2005), La Borde (Jaubert et al., 1990), Biache-Saint- et al., 2015; Massilani et al., 2016). The diet of B. priscus included ty-
Vaast (Auguste, 1995), Hénin-sur-Cojeul (Marcy et al., 1993), Wal- pical steppe and grassland (C3) vegetation and lichens (Bocherens et al.,
lertheim (Gaudzinski, 1995, 1996), Il'skaja (Hoffecker et al., 1991), and 2015; Julien et al., 2012) but probably did not totally exclude woody
Mezmaiskaja Cave (Baryshnikov et al., 1996). A group of these sites vegetation, since some individuals, such as the bison bull mummy “Blue
provides evidence that Neanderthal groups used favourable landscape Babe” from Alaska, had 7% of woody material among the plant frag-
topography, such as cliffs, swamps or gorges to drive and trap the herds. ments trapped in its dentition (Guthrie, 1990). However, the structure
The planned and systematic exploitation and in some cases the selection of the neck and hump of the fossil species indicates that they held their
of individuals, also suggest the existence of repeated collaboration and heads higher, which might be a result of the adaptation to grazing taller
organization between hunters (Rendu et al., 2012). and sparser grasses (Guthrie, 1990). All living bisons are gregarious, the
The situation differs in southern Europe, where the faunal spectrum size and structure of the herds varies according to age, sex, season,
is more varied (e.g. Brugal and Valente, 2007; Garralda et al., 2014; resources, habitat and environment (Meagher, 1986, 1989; Plumb
Miracle, 2005; Morin and Soulier, 2017a; Panagopoulou et al., 2004; et al., 2009; Krasińska and Krasiński, 2013). Available resources con-
Rosell et al., 2012; Starkovich, 2017; Starkovich et al., 2018; Zilhão dition herd size; for example, North American average group size can be
et al., 2010) and specialized hunting behaviours that focus on particular as low as ~11–20 bison during winter, increases to ~200 in summer,
species such as large bovids are variable, as noted in Italian peninsula. and reaches a maximum of ~1000 during the rut season in June and
Faunal assemblages from sites in southern Italy (Grotta di Santa Croce, September (White et al., 2016). The B. bonasus average group size is
Riparo l'Oscurusciuto, and Grotta del Cavallo) are characterized by the environment-dependent. Typically, groups consist on average of ~8–13
predominance of Bos primigenius (Boscato and Crezzini, 2006, 2012; individuals in different populations. Sometimes, European bison forage
Boscato et al., 2011; Villa et al., 2009). in open areas (mown or mountain meadows and deforested grassland
In northern Italy, Neanderthals exploited large bovids as well glades) and form larger groups, of ~23 individuals. In all populations
(Romandini et al., 2014), but not as intensively. Cervids and caprinae bulls are often found in pairs, though more than half of males lead a
are commonly well-represented, while bovines are scarcely docu- solitary life (Krasińska and Krasiński, 2013). Bison herds may be se-
mented (Cassoli and Tagliacozzo, 1994; Fiore et al., 2004; Moroni et al., dentary or seasonally migratory; movements are altitudinal or direc-
2018; Peresani et al., 2014; Sala, 1990; Thun-Hohenstein and Peretto, tional. There are no historical data on seasonal migrations of lowland
2005; Valensi and Psathi, 2004). Thus, several questions revolve around European bison; however, seasonal movements are observed in the
the exploitation of bovines in the Middle Palaeolithic. Why were Bison forest-field landscape in Poland and also altitudinal seasonal move-
priscus and Bos primigenius not frequently exploited? Was the exploita- ments related to weather condition and snow cover are observed in
tion of these animals influenced by site topography or, rather, by the mountain areas (Krasińska and Krasiński, 2013). In the closely related
animals' seasonality? Could the climate have affected the availability of American plains, seasonal movements of bison were observed with
these resources? Could the technological systems adopted by different distances as long as 250 km (Meagher, 1986). Bison forage dis-
Neanderthal groups (e.g. Quina or Levallois) have influenced the ex- continuously on open bottomlands and lower adjacent slopes, but their
ploitation of these animals? In order to answer these questions, the routes sometimes traverse forest areas and steep slopes (Meagher,
analysis presented here focuses on three sites in northern Italy, Fumane 1989). Bison have poor vision but can detect movement up to 1.5 km.
cave (Grotta di Fumane), San Bernardino cave (Grotta Maggiore di San Their hearing and olfactory senses are highly developed, and they rely
Bernardino) and De Nadale cave (Grotta De Nadale), which preserve on them heavily to detect danger. They can smell humans 500 m away,
taxonomic and taphonomic data on several distinct human occupations. and upon sensing danger, can run at speeds reaching up to 60 km/h
The aim of this work is to assess the condition that might have affected (Fuller, 1960; Krasnokutsky, 1996). Juveniles and young bisons may act
the presence or absence of Bison priscus and Bos primigenius, and to unpredictably and run in other directions, but the adults will try to
better understand their exploitation in the late Middle Palaeolithic, adjust their path accordingly. If the herd encounters an obstacle, the
during Marine Isotope Stages (MIS) 4-3, on the southern foothill of the lead animals will veer sharply, but if the momentum of the herd is too
Eastern Alps. To this end, we use ethological information (ecology and great, it might plough forward into further danger and over cliffs
behaviour) from fossil evidence and through analogy of living re- (Krasnokutsky, 1996).
presentatives to infer the size and social composition of the herds. With
such information, we attempt to contribute to interpretations of 1.2. Aurochs ethology
Neanderthal hunting and subsistence strategies.
The aurochs, Bos primigenius (Bojanus, 1827), became extinct during
1.1. Bison ethology the 17th century. Any estimate of its social, behavioural ecology and
morphology must therefore rely on historical reports, baseline data
The Eurasian steppe bison, Bison priscus (Bojanus, 1827), dis- from extant species of wild cattle, and images from Lascaux, Chauvet,
appeared from the fossil record around 10–12 ky ago, although it seems Altamira, and Romito caves (van Vuure, 2002). From these sources, the
to have survived until the mid-Holocene in Siberia (Boeskorov et al., height of the withers has been estimated between 160 and 180 cm for
2016; Kirillova et al., 2015). It was a formidable animal with long horns the bull, and around 150 cm for the cow. Their horns grew outside from
and robust legs over 2 meter long at the withers and reached a total the skull then curved forward (in an angle of ca. 60° with the forehead)
length of > 2.7 m. The reconstruction of its ecology and ethology must and inward (towards each other) (van Vuure, 2002, 2005).
be inferred from fossil evidence and through analogy with its closest As studies indicate (Von Koenigswald, 2007), Bos primigenius was an
living relatives, the American bison, Bison bison (Linnaeus, 1758), and interglacial animal. European specimens would withdraw to the Med-
European bison or wisent, Bison bonasus (Linnaeus, 1758). As a genus, iterranean area during cold periods and expand to the north during
they are ecologically flexible: American bison are primarily grazers of warmer periods (Von Koenigswald, 2007). Fossil records, in western
wooded steppe mosaics, while wisents now occupy deciduous and Europe (Brugal, 1985) and in some parts of Asia, which show the
mixed forest, the areas to which they were introduced. However, recent presence of both Bos primigenius and Bison priscus in the same sites,
studies suggest that the European bison was not necessarily a forest suggest that aurochs were probably adapted to colder and drier climatic
specialist (Bocherens et al., 2015; Kerley et al., 2012; Kowalczyk et al., conditions, at least to a certain extent (van Vuure, 2002). B. primigenius
2015). B. priscus was adapted to forest-steppe and steppe ecosystems was a grazer or intermediate feeder, feeding mainly on grasses and
and is considered indicative of open environments (Brugal et al., 1999). graminoids, supplemented with forbs, leaves and branches of trees and
Paleoenvironmental evidence suggests that, like B. bison, B. priscus was bushes (van Vuure, 2002, 2005). According to palynological data, re-
more of a grazer than B. bonasus, which is a mixed feeder (Bocherens search on fossil insects, and historical accounts, the habitat that aurochs

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occupied in Europe was characterized by closed forests, fluvio-lacus- well as the Alpine and Berici foothills (Monegato et al., 2011). In the
trine environments, and marshy areas (van Vuure, 2002, 2005). In 65,000 years that span from MIS5d to MIS3, no signs of fluvioglacial or
general, Bos primigenius's behaviour was similar to the rest of the Bovini significant aggradation have been recorded: rivers flowed within stable
tribe; aurochs tended to separate into cow (or mixed) herds, bull herds, trenched paths, and relatively low sedimentation rates (Monegato et al.,
and solitary bulls. Bulls only joined cow herds during the rut season 2011) in both lacustrine and alluvial successions indicate water table
(between August–September), and most calves were born at the end of stability. Persistent afforestation with some temperate trees, notably
spring. Tilia and Abies, have been recorded throughout the entire early and
middle part of the last glaciation, with only moderate forest with-
2. The north Italian context drawals during early Dansgaard-Oeschger events (Pini et al., 2010). The
first part of MIS3 records a prevailing zonal vegetation on the plain that
2.1. The pre-alpine fringe and subalpine area of the southeastern Alps. includes open birch-conifer forests, xerophytic scrubs and steppe, and
Physical and ecological setting from MIS 4 to MIS 3 phases of contracting conifer forests and expanding steppic commu-
nities alternating with mixed conifer (Pinus and Picea) – Betula forests
Along the southern slope of the Alps, the pre-alpine mountain range (Pini et al., 2009). Soil formation affected aeolian deposits (Zerboni
and the sub-alpine zones succeed one another, forming a discontinuous et al., 2015). During a late phase of this period (about 38.2 ± 1.45 ky
series of short chains and mountain groups from the Lago Maggiore to cal BP according to Pini et al., 2010) aggradation of the alluvial fans in
Istria. This region encompasses a 40 km-wide mountain belt that is the western Venetian Plain occurred, in coincidence with the estab-
oriented west to east in its western-middle sector and progressively lishment of long-lasting marshes in the Friulian-Venetian Plain
turns to the north in the eastern transect. The landscape of the Eastern (Fontana et al., 2008).
Pre-Alps is composed of limestone massifs and high, karstic plateaus
1000–1200 m above sea level (a.s.l.), with summits that rise above 2.2. Fumane Cave
2000 m. This is interspersed with gorges, large river valleys, and wide
basins that host important glacial alpine lakes. Of these high plateaus, Fumane Cave is located at 350 m a.s.l. (Fig. 1) and has provided a
the Monti Lessini is a fan-shaped karstic plateau that dips gently to the dated sequence spanning from MIS 5 to MIS 2 (López-García et al.,
south towards the alluvial plain of the Adige River, and is characterized 2015). The Middle and Upper Palaeolithic deposits consist of numerous
to the north by summits that reach 1500–1600 m. It is bounded to the thin to very thin parallel levels and lenses grouped into stratigraphic
west by the Adige Valley, a long and deep cut that connects the inner units labelled from A13 to A1 (from bottom to top). The Mousterian
Alpine region with the Po Plain. The Monti Lessini is characterized by levels are characterized by lithic and faunal remains densely scattered
tectonic terraces connected to the bottom of the stream valleys by steep on the living floors, as is the case in units A11, A10, A9 and A6
slopes and rock walls that include caves and shelters. At the eastern (Peresani, 2012; Peresani et al., 2011b). Cherts in A12-A11, A10V, A10,
end, the Trieste karstic plateau forms a typical flat landscape that ex- and A6–A5 were exploited mostly through Levallois technology
tends at a low elevation to the Sava and Danube basins. The subalpine (Peresani, 2012). The focus of this paper is the stratigraphic complexes
zone of the Eastern Pre-Alps comprises hills of different origins, such as unit A8–A9 and unit A5–A6, dated to 47.6–45 and 44.8–42.2 ky cal BP,
the pre-MIS2 glacial moraines south of Garda Lake, the Monti Berici respectively (Higham et al., 2009; Peresani et al., 2008). Of these units,
plateau, and the Colli Euganei cone-shaped hills. The latter two groups layer A6 in particular has provided evidence of a structured use of the
are isolated from the alluvial plain. The Alpine foreland is a large al- living space, covering the entire entrance area (Peresani et al., 2011a).
luvial plain that mostly originated during the Middle and Late Over 40 combustion features and dumps of combustion debris are as-
Pleistocene from the major rivers, including the Po, Adige and the rivers sociated with zones used for Levallois manufacture, tool shaping and
of the Friulian-Venetian Plain (Mozzi, 2005). The Berici Hills are a curation, ungulate butchery, and the treatment of hides and furs. The
karstic plateau of special interest for our investigation. At an average faunal assemblage includes a rich association of ungulates, carnivores,
elevation of 250 m a.s.l., the plateau contains a honeycomb of sink- and birds from diverse environments and climates. Notably, unit A9
holes and various depressions that delineate an extremely uneven to- with Discoid technology is ecologically analogous to A5–A6
pography with peaks and karstic blocks that are affected by surface (Romandini et al., 2014), where the most abundant faunal remains are
dissolution. The plateau is dissected by depressed systems (e.g., the red deer, ibex, and roe deer, whereas chamois, bison, and giant deer are
Fimon, Liona, and Calto valleys) with pocket-valleys where ephemeral less frequent. Moose, horse, and wild boar are rare (Table 1). All un-
streams produced swampy environments and fed historic mills. The gulate species, with the exception of horse and wild boar, bear traces of
slopes are steep all around. To the east, a steep slope with rock cliffs human exploitation. Tibias, femurs and metapodials, radii and humeri
connects the plateau to the alluvial lowlands that were occupied by from Cervids (red deer, roe deer, and very large specimens of giant deer
marshes and swamps during the Pleistocene and earlier phases of the or moose), and to a lesser extent from other ungulates, were used as
Holocene. In the southern eastern area, the Pozzolo depression is a wide retouchers (Jéquier et al., 2012, 2018). Fox, wolf, brown bear, and cave
trench cutting through the plateau in a NW-SE direction and with an bear are the most numerous carnivores (Table 1). Except for wolves,
elevation of 150 m at both the SE and NW ends. It is the relict segment carnivore remains show butchery marks from skinning and defleshing
of an old valley that was cut by a river during the first uplifting phases (Romandini et al., 2018a, 2018b). The elevated number of cut marks,
of this morpho-unit. The ancient karst surface is covered with palaeo- compared to the scarcity of carnivore gnawing on bones, supports the
sols and thick red clayey residual deposits (Sauro, 2002). anthropogenic nature of bone accumulation in these units. In addition
For long parts of the Late Pleistocene, the eastern Italian Alps were a to the butchery marks on ungulate and carnivore bones, we also report
physical and ecological barrier due to the presence of ice fields and on the discovery of human modifications to some anatomical elements
alpine glaciers formed during the cold stages, while temperate vege- of large raptors and Passeriformes (Peresani et al., 2011a; Fiore et al.,
tation was only partially restored during the warmest interstadials. The 2016; Romandini et al., 2016).
morphological and sedimentary evolution of this part of the Po Valley
and the shoreline displacements that occurred during this period have 2.3. San Bernardino Cave
been assessed in several works (Antonioli, 2012; Fontana et al., 2008;
Mozzi, 2005; Zecchin et al., 2008). After the last interglacial, the The cave opens on the eastern slope of the Berici plateau (Vicenza),
sudden lowering of sea-levels at the beginning of the glaciation trig- 135 m a.s.l., west of the alluvial plain of the Bacchiglione River (Fig. 1).
gered erosion of the near-shore sediments and the incision of the lower In a 4 m thick stratigraphic sequence, eight lithological units and three
reaches of the rivers, probably affecting the entire Venetian foreland as main paleoclimatic cycles have been identified, shifting from temperate

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Fig. 1. Geographical location of cave sites.

Table 1
Mammal NISP and NISP% from Fumane Cave (FC – unit A8–A9 and A5–A6), San Bernardino Cave (SB – unit II) and De Nadale Cave (CN – units 6–8).
Taxa FC – unit A8–A9 FC – unit A5–A6 SB – unit II CN – units 6–8

NISP NISP% NISP NISP% NISP NISP% NISP NISP%

Lepus sp. 1 0.1 3 0.3

Marmota marmota 4 0.2 1 0.1 16 1.8
Castor fiber 25 3
Total Lagomorpha and Rodentia 4 0.2 2 0.1 44 5.1
Canis lupus 4 0.2 11 0.3 3 0.3 3 0.2
Vulpes vulpes 4 0.2 46 1.5 7 0.8 2 0.1
Vulpes/Alopex 3 0.1
Ursus arctos 2 0.1 9 0.3 1 0.1
Ursus spelaeus 5 0.3 4 0.1 84 9.7 11 0.6
Ursus sp. 3 0.2 22 0.7 38 4.5 16 0.9
Martes sp. 1 0.1
Mustela putorius 2 0.2
Mustela nivalis 1 0.1
Crocuta crocuta 1 0.1
Felix silvestris 1 0.1
Linx linx 6 0.7
Panthera pardus 1 0.1
Panthera leo 1 0.1
Carnivora indet. 13 0.7 17 0.5 3 0.2
Total Carnivora 33 1.9 113 3.6 144 16.6 35 1.9
Sus scrofa 2 0.1 2 0.1 29 3.4 1 0.1
Megaloceros giganteus 65 3.9 38 1.3 4 0.4 105 5.6
Alces alces 11 0.6 5 0.2 24 2.8
Alces/Megaloceros 31 3.6
Cervus elaphus 391 23.3 1392 44.8 81 9.3 105 5.6
Capreolus capreolus 215 12.9 230 7.4 149 17.2 17 0.9
Cervidae indet. 134 8 167 5.3 56 6.5 54 2.9
Bison priscus 6 0.3 4 0.1 10 0.5
Bos primigenius 6 0.3 1 0.1 1 0.1
Bos/Bison 29 1.7 28 0.9 20 2.3 70 3.8
Capra ibex 42 2.6 84 2.7 1 0.1 1 0.1
Rupicapra rupicapra 54 3.2 87 2.8 24 2.8 3 0.2
Caprinae indet. 15 1 19 0.7 8 1 2 0.1
Stephanorinus sp. 1 0.1
Ungulata indet. 671 40 932 30 248 28.7 1455 78.2
Total Ungulata 1641 97.9 2988 96.3 677 78.3 1824 98.1
Total NISP 1678 3103 865 1859
Total Indeterminate 81,090 168,161 3354 22,288
Total NR 82,768 171,264 4219 24,147

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to dry cool conditions and dating from the Middle-Late Pleistocene For the identification of the remains described here, the publications of
(cycle 1) to the Late Pleistocene (cycles 2–3) (Cassoli and Tagliacozzo, Sala (1986) and Martin (1987) were used. Potentially burnt bones were
1994; López-García et al., 2017; Peresani, 2001). Unit II, the focus of determined by combustion grade level following the criteria described
this paper, was dug over 20 sqm and its thickness ranges up to 40 cm. It by Stiner et al. (1995). Microscopic analyses of bone surfaces were
records humid climatic conditions, the expansion of woodlands (Cassoli carried out using a Leica S6D Greenough stereomicroscope with
and Tagliacozzo, 1994; López-García et al., 2017), and an increase in 0.75–70× magnification range for capturing images. In order to iden-
the rate of accumulation of anthropogenic remains with respect to the tify the nature of surface alterations and to distinguish human from
underling units, as suggested from the frequency of faunal remains, animal damage, trampling abrasion, and modern mechanical mod-
lithic artefacts, and hearths. The most abundant ungulate remains be- ifications produced by excavation tools, reference was made to well-
long to roe deer, red deer, moose, wild boar, and bovines (Table 1), established taphonomic literature (Fernández-Jalvo and Andrews,
while cave bear is the most numerous carnivore (Romandini et al., 2016). Sex and age at time of death were determined by dental wear
2018b). Taphonomic analyses on large ungulate bone surfaces from the and by the state of epiphyseal fusion (Habermehl, 1975; Bunn and
main part of the sequence have revealed traces of disarticulation, de- Pickering, 2010; Silver, 1969). In order to evaluate species abundance,
fleshing, intentional shaft fracturing, and the use of bone fragments as the following methods were used: number of identified specimens
retouchers. Evidence of carnivore activity is scarce. (NISP) (Grayson, 1984), minimum number of elements (MNE) (Klein
and Cruz-Uribe, 1984; Stiner, 1994), and minimum number of in-
2.4. De Nadale Cave dividuals (MNI) (Bökönyi, 1970). Estimating MNI was difficult due to
the high degree fragmentation, which mostly involves the long bones
De Nadale Cave is a small cavity located at 80 m a.s.l., in the and some teeth. As those bones and teeth were frequently cracked into
southern slope of the Berici (Fig. 1). It opens on the Calto Valley, a several fragments, it was not always possible to compare the different
narrow V-shaped fluvio-karstic canyon-type valley with active water elements. In order to avoid distortion in the proportions of age groups
springs in its inner zones. The hydrographic drainage system of this in the various units, estimations were also calculated for those remains
area features the initial segment of the Liona Valley with the confluence generally determined as Bos/Bison by considering only the age or the
of other incisions currently supplied by springs from the north and the size of the animal not represented in the two categories of the de-
west, where they are limited by the Pozzolo suspended valley. The termined taxa (Bison priscus and Bos primigenius). To estimate age at
geometry and nature of the lowest Calto Valley deposits are unknown, death, we used dental wear and the state of epiphyses fusion, as well as
but it is probable that this depressed area (about 55 m a.s.l.) was the proportions between thickness and growth of crests of femur and
characterized by moist or swampy environments with peaty deposits in tibia. Combustion degree was distinguished between moderate
the past, comparable to what is known along the Val Liona. (200–500 °C, black/brown) and elevated with calcination (> 700 °C,
Field investigations started in 2013 and are still in progress. Under a grey/white). Cut-marks were classified as incisions, like skinning
superficial and reworked layer, a 2 m thick stratigraphic sequence has marks, defleshing marks, and scrapes (Potts and Shipman, 1981;
been unearthed during the last three campaigns. Although ten strati- Shipman and Rose, 1984). Intentional bone breakage to access marrow
graphic units have been identified in this sequence, only one (unit 7) was documented by diagnostic criteria like percussion marks/notches
can be recognized as archaeological. This unit contains several osteo- and impact flakes, and positive flakes of the percussion marks
logical remains and lithic implements attributed to the Quina method. (Blumenschine and Selvaggio, 1988; Capaldo and Blumenschine, 1994;
Uranium-series (UeTh) dating, performed on an herbivore maxillary Pickering and Egeland, 2006; Villa and Mahieu, 1991). Traces left by
fragment, provided a minimum age of 70.2 + 1/−0.9 ky BP (Jéquier carnivores were identified as deep punctures in the cortical bone sur-
et al., 2015). The faunal assemblage (Table 1) reveals that the most face, concentrated on the articular ends of the bones, as well as pits and
abundant species are the giant deer and the red deer, followed by large- scores with U-shaped striations. These results were compared to data
sized bovines. Chamois and roe deer have also been identified, but in from Fisher (1995), Selvaggio and Wilder (2001), Domínguez-Rodrigo
lower numbers. Carnivores are scarcely present: among them, the most and Piqueras (2003).
common is the bear (Ursus sp.; Ursus spelaeus), while wolf and fox are
represented by 3 fragments each. A taphonomic analysis indicates an
excellent preservation of the osteological remains: nearly half of the 4. Results
total amount of bones preserves butchering marks, including scraping
and cut marks, impact notches and spiral fractures on the surface of The bovine bone assemblage, subdivided by anatomical element in
several bone shafts. Furthermore, the large number of pieces with re- craniocaudal order, shows that entire carcasses are not completely re-
touch damage is exceptional. This association of taxa, in addition to the presented. In particular, the cranial remains (maxilla, hemimandible,
ecological data based on the micromammal assemblage, reflects a cold horn, and teeth) and the axial skeleton (ribs and vertebra) are rarely
climatic context in a landscape dominated by open woodlands and open represented. The stilopodium (humerus/femur), zeugopodium (radius-
dry meadows, which, taking into account the chronology of the site, ulna/tibia-fibula), and metapodium (metacarpal/metatarsal) are well
could be related to the onset of MIS 4 (López-García et al., 2018). represented in all three sites, which contrasts with the low number of
Moreover, the importance of large-sized cervids and aurochs in the limb extremities (Table 2; Fig. 2).
faunal assemblage supports this interpretation and suggests the pre- In an osteological context it is difficult to distinguish between the
sence of a swampy area with water sources (Jéquier et al., 2015). two genera Bos and Bison, due to the great ambiguity of many of their
anatomical elements as well as the high fragmentation of the remains.
3. Materials and methods Due to these factors, most of the remains from the three sites were
generically assigned to the category Bos/Bison, which is also the case at
The bones examined in this study were discovered in the last many other contemporaneous sites. Overall, despite the small number
30 years of excavations using dry and wet sieving. Zooarchaeological of diagnostic remains, the Bison priscus was found at Fumane and De
and taphonomic analyses were performed on 175 bovine bones. Nadale (n = 10 respectively) and Bos primigenius was recovered from all
Taxonomic and skeletal identifications are based on the complete alpine three sites (n = 6 in Fumane Cave, n = 1 in San Bernardino and n = 1
fauna reference collection of the Department of Humanities at the in De Nadale).
University of Ferrara and, for more problematic cases, in the
Quaternary Paleontology and Zooarchaeology Laboratory of the
Ethnographic-Prehistoric National Museum “L. Pigorini” (Rome, Italy).

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Table 2 4.1. Taphonomy

Anatomical elements of Bovinae (Bison priscus, Bos primigenius, and Bos/Bison);
NISP and MNI. 4.1.1. Fumane Cave
FC – unit FC – unitA5–A6 SB – unit CN – units Among the natural taphonomic processes that have affected the
A8–A9 II 6–8 surface of the studied bovine bones from units A8–A9 and A5–A6 (NISP
73), root modifications were the most common, affecting 30% of the
Cranium 2 6 1 2
bones. In some cases, root etching was associated with the presence of
Hemimandible 2 2 2 4
Tooth indet. 2 manganese stains. Concretions were found on many bones (26%),
Hyoid probably due to their immediate proximity to the cave wall. Carnivore
Total cranium 6 8 3 6 damage is very rare, found on only 3 Bos/Bison bones. In both units,
Atlas-axis
several remains (56% and 46.8%, respectively) show traces of butchery
Vertebra 1
Rib 1 1
(Table 3; Figs. 2, 3). These are observed as isolated cuts or cuts in a
Sternum 2 series, testifying to short- and medium-length gestures. Percussion
Total trunk 1 2 2 marks (PM) on long bones for the extraction of marrow account for 13%
Scapula 1 2 and 46.6% of the total bovine remains with modifications (Table 3),
Humerus 2 6
and are in some cases associated with cut-marks (CM). It should,
Radius 1 2 9
Ulna 1 1 2 however, be clarified that for certain anatomical elements where the
Carpals bone is less protected from the muscular tissue, cut-marks may indicate
Metacarpal 2 2 the recovery of skin or disarticulation (Table 3; Fig. 3). Cut-marks found
Total frontal limb 1 6 3 21
on a fragmentary left mandible with deciduous dentition, as well as a
Coxal 1 1 1
Femur 3 3 2 4
few phalanges, sesamoids, and metapodials, demonstrate defleshing
Tibia 14 8 3 28 and in some cases, disarticulation. The defleshing and detachment of
Tarsals 2 muscle mass is evident: sixteen tibiae from different individuals, three
Metatarsal 7 1 9 femurs, two humerus and two radii preserve cut marks. Five bones from
Total hind limb 25 11 9 42
A8–A9 (12.1%) and one from A5–A6 show traces of thermal alteration.
Metapodial 5
First phalanx 2 3 In two cases, a Bos primigenius metatarsal (A9 SVI.2) and a radius of
Second phalanx 2 3 3 Bison priscus (A5 SIII) were associated with combustion features
Third phalanx 2 (Fig. 3F). So far, the only seasonal indication of human use of the site is
Sesamoid 3 1 limited to the A8–A9 unit, where a fragment of an infant Bison priscus
Total indet. limb 5 5 4 10
Indet. 3 2
(right mandible with attached premolar deciduous teeth) had evidence
Total 41 32 21 81 of skinning (Fig. 3A_1). This suggests that the animal was hunted be-
MNI 6 4 2 7 tween spring and summer. Five bone shafts (2 Bison priscus and 3 Bos/
Bison) were used as hammers for retouching flint artifacts. The re-
touchers were manufactured from metapodials, femurs, and tibia, most
of them cut-marked (Table 4; Figs. 2, 6A_1).

Fig. 2. Frequency of skeletal elements of the Bovinae (Bison priscus, Bos primigenius, and Bos/Bison) and distribution of butchery marks.

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Table 3
Number of anatomical elements of Bovinae with anthropogenic modifications and carnivore marks; CM cut-marks, PM percussion marks, CM + PM cut-marks + percussion marks, BM butchery marks, R retoucher, B
burned, GM gnaw marks.
Bovinae - FC unit A8–A9 Bovinae - FC unit A5–A6 Bovinae - SB unit II Bovinae - CN units 6–8

NISP CM PM CM + PM TOT. BM R B GM NISP CM PM CM + PM TOT. BM R B GM NISP CM PM CM + PM TOT. BM R B GM NISP CM PM CM + PM TOT. BM R B GM

Cranium 2 1 1 6 1 2
Hemimandible 2 1 1 2 2 4 2 2 1 1
Tooth indet. 2
Total cranium 6 2 2 8 3 6 2 2 1 1
Vertebra 1
Rib 1 1 1
Clavicle 2
Total trunk 1 1 2 2
Scapula 1 2 1
Humerus 2 1 1 2 6 1 1 2 4 2
Radius 1 1 1 1 2 1 1 1 9 4 2 6 5 1

135
Ulna 1 1 2 1 1
Carpals
Metacarpal 2 2 2 1 2
Total frontal limb 1 6 3 2 5 1 1 3 1 1 1 21 6 1 4 11 8 1
Coxal 1 1 1 1 1 1
Femur 3 1 2 3 1 1 3 1 1 2 1 1 4 1 1 2
Tibia 14 4 2 5 11 2 1 8 1 2 2 5 2 3 1 1 1 28 9 5 4 18 3 3
Tarsals 2 1
Metatarsal 7 2 1 1 4 1 1 1 1 9 1 2 2 5 2 1
Total hind limb 25 7 3 8 18 2 3 3 11 1 3 2 6 2 9 1 1 2 1 1 42 11 8 7 26 5 5
Metapodial 5 2 1 3 1
First phalanx 2 1 1 3
Second phalanx 2 1 1 3 1 1 3 1 1 2 1
Third phalanx 2
Sesamoid 3 1 1 1
Total indet. limb 5 2 2 5 2 2 4 10 3 1 1 5 1 1
Indet. 3 1 1 1 2 2 2
Total 41 12 3 8 23 2 5 3 32 6 7 2 15 3 1 21 2 1 3 2 1 81 22 10 12 44 15 8
MNI 6 4 2 7
Journal of Archaeological Science: Reports 25 (2019) 129–143
G. Terlato, et al. Journal of Archaeological Science: Reports 25 (2019) 129–143

Fig. 3. Fumane Cave (unit A8–A9 and unit A5–A6),

localization and details of anthropogenic traces: A)
right mandible with premolar deciduous teeth of
Bison priscus, with traces on buccal side (1 – close-up
of striae); B) distal portion of second phalanx of Bos/
Bison with traces (2); C) right metacarpal of Bos/
Bison with traces on lateral portion (3); D) right tibia
of Bos primigenius with traces on diaphysis (4); F)
Combustion structure (A5 SIII) and associated radio
of Bison priscus.

4.1.2. San Bernardino Cave remains are cut marked (27%), 10 have percussion marks (12.3%), and
Among the natural taphonomical processes that have affected the 12 bone fragments (14.8%) have both (Table 3; Figs. 2, 5). The presence
bovine remains (NISP 21), manganese coatings and roots had the big- of cut marks on a second phalanx and on two fragmented mandibles
gest impact, modifying the bone surface of 38% and 26% of the as- reflects skinning activities; the same traces detected on three fragments
semblage, respectively. No traces attributable to carnivores were found of humerus, six radii, one ulna, one coxal, one femur, thirteen tibias,
in this assemblage. The bovine remains with traces of butchering marks three metatarsals, and three generic metapodial, suggest defleshing and
are scarce (Table 3; Figs. 2, 4), and include only one radius and one muscle mass removal. Furthermore, there are some large, deep scrape
tibia. The documented activities probably relate to defleshing and the marks on two tibia shafts, on a metatarsal and on two mandibles,
detachment of muscle mass. Percussion marks are visible only on the pointing to the removal of the periosteum. On one of the mandible
diaphysis of a femur (Table 3; Fig. 4B), which presents two relatively fragments, these traces seem to be linked to the retouch damage. Per-
large percussion grooves, one of which covered most of the cortical cussion marks have been observed on three humerus diaphyses, two
surface. Two retouchers were present (Table 4, Figs. 2, 6B_2), made radii, one coxal bone, one femur, nine tibias, four metatarsals, one
from a radius and tibia of generic Bos/Bison. Cut-marks on the dorsal generic metapodials, and one second phalanx, attesting to the inten-
face of the radius (defleshing) were noted in association with the area tional fragmentation of the long bones to extract marrow (Fig. 5A and
used for retouching. D). In total, there are 15 bone retouchers (Figs. 2, 6C_3; D_4–5): one
mandible, one scapula, two humeri, five radii, three tibias, three me-
tatarsals and one generic metapodial. Of these 15 bone tools, 12 are
4.1.3. De Nadale Cave also cut-marked and 8 have percussion marks for marrow extraction.
Taphonomic analysis suggests that the most common bone surface
modifications are due to manganese staining and root etching.
Manganese stains have been detected on 47 bovine bones (56.6%), 4.2. Age and minimum number of individual estimation
while root modifications are present on 52 remains (62.6%). Trampling
also affected several remains, as it has been recognized on 38 bone Data from Fumane indicate the presence of individuals of almost all
shafts (47.8%). Carnivore gnawing is rare: 8 of the 81 (9.8%) bone ages, with a small preference towards adults (Table 4). In A8–A9 spe-
fragments show pits and tooth marks. Butchering marks are abundant: cifically, MNI indicates at least 6 bovines (2 Bison priscus; 2 Bos primi-
44 fragments (54.3%) show anthropogenic modifications related to genius and 2 Bos/Bison): Bison priscus-1 infant (> of 1 month); 1 generic
defleshing, skinning, and marrow extraction activities. Moreover, 22 adult; Bos primigenius-2 generic adults of different ages; Bos/Bison-1

Table 4
NISP and MNI divided by age of Bison priscus, Bos primigenius, and Bos/Bison.
FC – unit A8–A9 FC – unit A5–A6 SB – unit II CN – units 6–8

Taxa NISP MNI by age Total MNI NISP MNI by age Total MNI NISP MNI by age Total MNI NISP MNI by age Total MNI

I Y-A A S Y A S Y A I Y A

Bison priscus 6 1 1 2 4 2 1 3 10 4 4
Bos primigenius 6 2 2 1 1 1 1 1 1
Bos/Bison 29 1 1 1 2(1) 28 1 3 1 1(4) 20 1 2 1(2) 70 1 1 3 2(3)
TOT. 41 1 1 3 1 6 32 1 2 1 4 21 1 1 2 81 1 1 5 7

Italic numbers represent the MNI by age of Bos/Bison not considered in the total because already represented in the two categories of determinate taxa (Bison priscus
and Bos primigenius).

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Fig. 4. San Bernardino Cave, localization and details of anthropogenic traces: A) tibia of Bos/Bison with traces on diaphysis (1–2); B) femur of Bos/Bison with
percussion marks (3) and percussion notch.

young adult; 1 senile. To date, in A5–A6 an MNI of 4 bovines has been 1987; Sala, 1986; Sher, 1997) have shown that well-established char-
estimated (3 Bison priscus; 1 Bos/Bison): Bison priscus-2 generic adults; 1 acteristics can also be used to identify some of the postcranial skeletal
senile; Bos/Bison-1 young (< 24 months). elements to taxon. Taphonomic analyses have provided evidence of
At San Bernardino, a combination of young and adult individuals is human exploitation of large bovids for meat and other resources.
noted, while infant and senile individuals are absent. An MNI of 2 bo- Butchering marks are abundant: 38 NISP (52.0%) in Fumane A9–A5, 3
vines was estimated (1 Bos primigenius and 3 Bos/Bison): Bos primigenius- (14.3%) at San Bernardino and 44 (54.3%) at De Nadale provide evi-
1 generic adult; Bos/Bison-1 young (12–24 months). dence of defleshing and marrow extraction (Table 3). In most of the
All ages are present at the De Nadale, with a clear predominance of examined cases, the low number of axial skeleton elements induces us
adults. The bovine remains can be ascribed to seven individuals (4 Bison to hypothesize that Neanderthals intentionally preferred certain ana-
priscus, 1 Bos primigenius and 2 Bos/Bison), at least: Bison priscus-4 tomical elements (front and hind limb) over others (skulls and mandi-
generic adults; Bos primigenius-1 generic adult; Bos/Bison-1 infant and 1 bulae, which are represented in small numbers). This selection likely
young. The presence of a Bos/Bison fetus establishes that the occupation took place directly at the kill site in order to facilitate the transport of
of the site occurred mostly between spring and summer. the highly valued body elements to the sites, while body parts with low
economic value would have been left behind (Marín et al., 2017). Such
decisions are influenced by various factors including the body sizes of
5. Discussion prey, distance to the site, weather, topography, the number of hunters
involved, the cost of carcass processing and transporting, as well as the
In the all three bone assemblages, it was not possible to distinguish competition with other carnivores (Bunn et al., 1988; Monahan, 1998;
between Bison priscus or Bos primigenius for all remains, due to their Schoville and Otárola-Castillo, 2014). Furthermore, considering the
high fragmentation rate and similarities of many anatomical elements. potential weight of these animals, which in some cases were con-
The taxonomy of large bovids is traditionally based on cranial and horn siderably bigger than their modern descendants, it makes sense that the
characteristics (Breda et al., 2010), but a number of authors (Martin,

Fig. 5. De Nadale Cave, localization and details of

anthropogenic traces: A) right radius of Bos/Bison,
with percussion marks (white arrow); B) metacarpal
of Bos/Bison with traces on dorsal face (1 – close-up
of striae); C) rib of Bos/Bison with traces (2); D) left
tibia of Bison priscus with traces on diaphysis and
percussion marks (white arrow); E) tibia of Bos/Bison
with cut-marks (3).

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Fig. 6. Bovinae bone retouchers from Fumane Cave (A), San Bernardino Cave (B) and De Nadale Cave (C–D). A) femur of Bison priscus with pits produced during
knapping (1); B) tibia of Bos/Bison with close-up of area used like retoucher (2); C) scapula of Bos/Bison with close-up on the areas used for retouching stone artifacts
(3); D) left humerus of Bos/Bison with cut-marks (4) and pits produced during knapping on diaphysis (5).

first steps of butchering would have occurred at the kill site. The pre- considerations about age at death.
sence of elements of relatively poor nutritional value like the few skull Our data and analyses thus indicate that large bovids were hunted in
and mandible fragments at Fumane and De Nadale might correlate to the Middle Palaeolithic of northern Italy but were not specially targeted
the transport of the complete carcass from the kill site. The latter should by Neanderthals as their main prey. A similar situation has been ob-
be located a short distance from the settlement, as the skull has the served in this region where variable evidence of large bovid exploita-
greatest decrease in probability of transport as a function of increasing tion by humans has been recorded, even if it is in some cases ephemeral,
distance (Schoville and Otárola-Castillo, 2014). However, the almost like at Riparo Tagliente (Thun-Hohenstein and Peretto, 2005). Large
complete absence of low survival elements (vertebrae and ribs) with bovids are also found at Grotta della Ghiacciaia in the Lessini Moun-
carpals and tarsals may be due in part to multiple attrition processes tains, despite the limited investigation carried out at the site (Bertola
generated by Neanderthals, the action of carnivores, and post-deposi- et al., 1999), Grotta del Broion (Peresani and Porraz, 2004) and Grotta
tional agents. A further reduction could result from the intensive di Paina (Gurioli et al., 2006) in the Berici Hills (however, these two
treatment of these anatomical parts by boiling (Morin and Soulier, caves preserve only ephemeral traces of human occupation). In the
2017b) or from their use as fuel. On the other hand, the absence of axial Fumane, San Bernardino, and De Nadale bone assemblages the ratios of
bones does not occur with the same incidence in these assemblages, so large bovid remains to other ungulates are −1:0.02, 1:0.05, 1:0.3 re-
their presence, although rare, indicates that at some point carcasses spectively. One reason for this is that other resources were available
were introduced into the caves intact. during the year (e.g. cervids), and may have been more abundant in the
Based on these observations, it seems that the anatomical re- surrounding areas or may have been easier to hunt (Delagnes and
presentation may be related to the sum of different carcass transport Rendu, 2011). In general, hunting and related activities are largely
patterns, where the size of the animal was one of a few determining reflected among ungulates relative to their taxonomic abundance and
factors. Fetal bones at De Nadale are indicative of at least one pregnant the ecological conditions found in the proximity of each specific site.
female that was brought to the cave in its entirety. The age at death of This shifted with climatic oscillations during the Late Pleistocene (Fiore
both species suggests a lack of individual selection. The presence of two et al., 2004). Red and roe deer were mainly targeted, with the occa-
infant remains from Fumane and De Nadale indicates that hunting oc- sional hunting of chamois and ibex (Fiore et al., 2004; Romandini et al.,
curred between spring and summer, the period when bovids disperse 2014) and the limited exploitation of giant deer, elk and, very rarely,
into small, highly mobile groups (counting only a few individuals in wild boar. Nevertheless, De Nadale partly diverges from this pattern in
some cases) and give birth (Rendu and Armand, 2009). The rarity of that it demonstrates that predation was mostly focused on large her-
available data and the low number of individuals do not support further bivores like giant deer, red-deer and bovids. As the faunal spectrum

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Fig. 7. NISP % of ungulates from Fumane Cave, San Bernardino Cave and De Nadale Cave.

suggests (Fig. 7), even if other medium sized prey – roe deer, chamois a wide territory including the lowest mountain ranges, and the karst
and ibex – were encountered near the surroundings, hunting seems to plateau and alluvial plains of the Po, Adige, and the rivers of the
have focused on large sized ungulates. This aspect is highlighted by the Friulian-Venetian plain (Fig. 8). Bovid remains (Bison priscus, Bos pri-
NISP and the MNI indices, which are higher for bovids and giant deer migenius and Bos/Bison) are reported in Late Pleistocene deposits
when compared to other taxa. Moreover, over half of the total amount (Fig. 8: Bon et al., 1991; Breda and Gallini, 2001; Cassoli and
of large sized ungulates bears human traces produced from butchering Tagliacozzo, 1994; Sala, 1990; Sami and Ghezzo, 2015) and both spe-
and the exploitation of the shafts as retouchers for shaping stone tools. cies are often found in association within the same site, for instance at
While the broad game spectrum possibly reflects the ecological varia- the Quinzano quarries and Grotta Tilde (Bon et al., 1991; Cassoli and
bility around the cave, with the co-existence of open landscapes on the Tagliacozzo, 1994). This suggests the existence of different ecological
Berici plateau and the presence of springs at the bottom of the Calto conditions, from dense forests with wetlands and small streams, eco-
Valley, the predominance of large sized taxa might be linked to a spe- logically more attractive to Bos primigenius, to hilly grasslands and
cific hunting behaviour, related to the mobility of Neanderthal groups. plains, populated by bison. According to this distribution, could topo-
In addition to this, cultural and technological differences might have graphy have influenced the hunting of both animals in relation to the
played a major role in the intense exploitation of these animals. structure of the settlement system in the area? What is certain is that
During MIS 4–3 in northeast Italy, large bovids were present across the Fumane, San Bernardino, and De Nadale caves are located in a

Fig. 8. Localization of Bovinae remains (Bison priscus, Bos

primigenius and Bos/Bison) in northern Italy during MIS
4–3, including the sites of this work (yellow star). Sea
level 70 m below the present-day coastline (courtesy by S.
Ricci, University of Siena. Based on the global sea-level
curve by Waelbroeck et al. (2002), but lacking estimation
of post-MIS 3 sedimentary thickness and eustatic magni-
tude). (For interpretation of the references to colour in
this figure legend, the reader is referred to the web ver-
sion of this article.)

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G. Terlato, et al. Journal of Archaeological Science: Reports 25 (2019) 129–143

region that included large bovid habitats; however, the exploitation of scenario in relation to faunal and human movements. The mobility
these resources could effectively have been influenced by the physical patterns of large bovids in this wide and flat land are still unknown, due
landscape of the site's surroundings. Fumane Cave was situated on a to the lack of MIS 4 and MIS 3 faunal assemblages between the northern
steep slope in proximity to one of the flat ridges of the Monti Lessini, a and the southern edge of the Po plain, and to the still incomplete data
fan-shaped high plateau gently raising from the Adige alluvial plain, on seasonality, isotope compositions, and other sources for inferring
dissected by large valleys that converge at higher elevation on the dietary patterns. Although the northern Adriatic Plain was more ex-
plateau. Herds of animals may have migrated seasonally along these tended than it is today, it still did not support seasonal movements of
ridges to and from the alluvial plain. Similarly, the location of De Na- bovid herds at a large scale, unlike less constrained landscapes in
dale must have played a fundamental role in the acquisition of certain western Eurasia. The possibility that large herbivores did not show
preys. The excellent view and the strategic location on the Calto Valley extensive mobility patterns is furtherly enhanced by the ecological
would have allowed hunters to easily monitor herd movements. This conditions of MIS 4, which never reached the level of the arctic open-
situation may explain the presence of entire carcasses at the site. At the landscapes of Western and Northern Europe, thus providing a sustain-
same time, the location of the sites in an ecologically varied landscape able biomass to maintain local subsistence patterns and the adaptation
should have favoured access to a spectrum of resources. These con- of hominins bearers of the Quina techno-typological system.
siderations lead us to question if the selection and hunting of certain The aim of this study was to discuss the contribution of large bovids
kinds of prey could have been influenced by other factors, such as cli- to Neanderthal subsistence in northern Italy and the conditions that
mate or seasonality. Climatic conditions fail to explain this phenom- might have affected the presence or absence of these animals during
enon for San Bernardino and Fumane, which both date to the first part MIS 4 and MIS 3. It has been shown here that Bison priscus and Bos
of MIS 3, demonstrated by the presence of the same resources in the primigenius were an important resource for different groups of Middle
region. De Nadale Cave provides a different pattern. A study of its Palaeolithic hunters that lived across a wide territory of northern Italy,
micromammal assemblage (López-García et al., 2018) shows that it was although they were not the most commonly exploited animals.
likely surrounded by open woodland and open-dry habitats, probably at Furthermore, they were present and, in some cases, hunted in a variety
the onset of MIS 4, which may explain the higher incidence of bovids of landscapes, influenced by the behaviour of the species themselves. A
when compared to the MIS 3 evidence from San Bernardino and Fu- significant set of evidence shows that Neanderthals were able to face
mane caves. An increase in bison populations during MIS 4 in Western environmental and topographical constraints and change over time by
Europe was previously suggested by Delpech (1999) on the base of adapting their hunting behaviour, technological lithic system, and
zooarchaeological data. mobility.
It has also been considered that seasonality may have been one of
the factors that made Neanderthals hunt specific animals at certain time Acknowledgements
of the year, thus exploiting the less migratory and more predictable
animals such as bovines in some periods. However, studies on season- Research at Grotta Fumane, Grotta San Bernardino and Grotta De
ality are still incomplete; therefore, a more detailed analysis based on Nadale is designed by the University of Ferrara (M.P.) and supported by
cementochronology or isotopes, similar to those undertaken for some the Ministry of Culture e Veneto Archaeological Superintendency,
European sites (Bocherens et al., 2015; Julien et al., 2012; Lieberman, public institutions (Lessinia Mountain Community and Regional
1994; Naji et al., 2015; Niven et al., 2012; Rendu, 2010; Rendu and Natural Park, Fumane Municipality, Zovencedo Municipality, the
Armand, 2009; Rendu et al., 2012) is necessary to provide further in- Province of Vicenza, Veneto Region - Department for Cultural Heritage,
formation. the H. Obermaier Society) and local private companies (R.A.A.S.M. and
Culture is often considered an important factor in explaining the Saf). Author contributions: M.P. designed the research; A.L., M.R. and
variability of Neanderthal subsistence patterns. Of the most common G.T. analysed data; G.T., A.L., M.R. and M.P. wrote the paper. We are
techno-typological systems adopted to fulfill mobility patterns, the grateful to Claudine Gravel and Britt M. Starkovich for proofreading of
Quina Mousterian is traditionally considered as the expression of a the manuscript.
highly mobile technological system related to hunting strategies fo-
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