OJS 615 Perez et al def3 figsbarate.pub


   

 Available online http://amq.aiqua.it 
ISSN (print): 2279-7327, ISSN (online): 2279-7335 
 
 

Alpine and Mediterranean Quaternary, 35 (2), 2022, 135-155 

1. INTRODUCTION 
 

Marine Isotope Stage (MIS) 3 is recognized to be a 
chronological stage (ca. 60-30 ka) characterized by 
rapid climatic shift. This variability is associated with 
cyclical and sudden fluctuations in Greenland tempera-
tures called Dansgaard-Oeschger (D/O) events, which 
have led to episodes of iceberg detachment in the North 
Atlantic, called Heinrich (HEs) events, resulting in a 
cooling of temperatures and drying climate conditions. 
This rapid alternation of warming and subsequent cool-
ing of temperatures results in a strong climatic instabil-
ity, alternating temperate-humid with cold-arid phases at 
mid-low latitudes (Heinrich, 1988; Bond et al., 1993; 
Dansgaard et al., 1993; Fleitmann et al., 2009; Naugh-
ton et al., 2009; Fletcher et al., 2010). It is in this context 
that the Middle to Upper Palaeolithic Transition (MUPT; 
ca. 50-40ka) occurred and the last Neanderthals, proba-
bly facing a demographic decrease while occupying 
smaller and smaller areas, were replaced by the first 
Anatomically Modern Humans (AMHs) (Higham et al., 
2014; Benazzi et al., 2015; Hublin 2015; Been et al., 
2017; Douka & Higham 2017; but see also Slimak et al., 
2022) 

How this transition occurred is widely debated 

(Mellars, 2006; Hoffecker, 2009; Higham et al., 2014; 
Villa & Roebroeks, 2014; Benazzi et al., 2015; Hublin, 
2015; Rey-Rodríguez et al., 2016; Greenbaum et al., 
2019; Timmermann, 2020). Mediterranean Europe, par-
ticularly the Italian peninsula, is a key region for under-
standing the dynamics of AMHs peopling and the disap-
pearance of the Neanderthals (Marciani et al., 2020). 
Italy, while allowing the dispersion of AMHs along the so
-called 'Mediterranean route' (Douka et al., 2012), has 
been characterised by an environmental asymmetry 
between the eastern Adriatic side and the western Tyr-
rhenian side; this is – and was – mainly due to the pres-
ence of two geographical barriers, the Alps and the Ap-
ennines mountains (Badino et al., 2020). This ecological 
dichotomy could have played a dramatic evolutionary 
role by providing 'refuge' zones (Columbu et al., 2020; 
Bicho & Carvalho, 2022; Jones, 2022) to the last Nean-
derthal groups. Among these refuge zones, the Liguria 
region, located on the north-western Tyrrhenian coast of 
Italy, appears to be a key area for understanding the 
MUPT and the subsequent AMHs peopling of the Medi-
terranean Europe. Liguria has a very narrow territorial 
surface, closed from the south by the Tyrrhenian Sea 
and from the North by the Alps. Thanks to its orography, 
Liguria could have largely played the role of a climatic 

 

https://doi.org/10.26382/AMQ.2022.08 

THE FAUNAL ASSEMBLAGE FROM THE RIPARO MOCHI SITE (BALZI ROSSI): 
NEW INSIGHTS ON THE MOUSTERIAN-AURIGNACIAN 

HUMAN-ENVIRONMENT RELATIONSHIP. 
 

Andrea Perez1, Fabio Santaniello1,2, Ursula Thun Hohenstein3, Stefano Grimaldi1,2 
 

1 LaBAAF, Dipartimento di Lettere e Filosofia, Università di Trento, Italy. 
2 Istituto Italiano di Paleontologia Umana, Anagni, Italy. 

3 Dipartimento di Studi Umanistici, Università degli Studi di Ferrara, Italy. 
 

Corresponding author: A. Perez <andrea.perez@unitn.it> 

 
ABSTRACT: Due to its geography, the Liguria region represented an obligatory pathway for animals and human groups that moved along 
the northern Mediterranean route, connecting the central Italian peninsula to the South-eastern France. Among the several Ligurian sites 
yielding traces of palaeolithic human activities, Riparo Mochi is a key site to understand the human peopling dynamics occurred during the 
Middle to Upper Palaeolithic Transition (MUPT). Its archaeological deposit is in fact one of the most complete and well dated MUPT se-
quences in this region. This study will contribute to increase our knowledge about the behavioural differences between the last Neander-
thals and the first Anatomically Modern Humans (AMHs) who inhabited the sites as well as the palaeoenvironmental changes that occurred 
from Marine Isotope Stage (MIS) 5 to 3. To do so, our study is focused on the zooarchaeological analysis of faunal remains coming from 
the Mousterian (Unit I), Protoaurignacian (Units H and G), and Aurignacian (Unit F) units of the site. Neanderthals inhabited the site during 
the early phases (MIS 5-4), hunting mainly Cervus elaphus and other middle-large size ungulates. A great variability in the faunal spectrum 
is shown during the coldest phases of the Mousterian. A decrease in variability is observed in the upper layers of Unit H, corresponding to 
the first AMHs occupation of the site. Despite the warmer climatic conditions, a decrease in diversity of faunas is observed, maybe due to a 
different hunting strategy operated by the Protoaurignacian occupants. Red deer is still the most common prey, but hunting seems also 
oriented on alpine taxa, such as Capra ibex. Deer hunting in the Proto- and Aurignacian economy might be highlighted by the production of 
antler tools founded only in the Upper Palaeolithic (UP) layers. Zooarchaeological and palaeoenvironmental data from Riparo Mochi shows 
a region characterized by an overall climatic and biological stability, reflected in the almost constant presence of certain species of large 
mammals during the MUPT. Nevertheless, the variations observed since the beginning of the UP appear to be related to an economic 
behavioural change attributable to the disappearance of Neanderthals and the arrival of AMHs at the site. 
 
Keywords: Balzi Rossi; Protoaurignacian; Middle to Upper Palaeolithic transition; Thyrrenian region; Zooarchaeology. 



   

 

 
 

136 

 
 

Perez A. et al. 

refugium even during the coldest phases of MIS 3 
(Negrino & Tozzi, 2008; Berto, 2019). Likewise, geogra-
phy probably made Liguria an obligatory pathway for 
both MUPT animals and human groups resulting in a 
bidirectional route along the East-West axis that linked 
Italy and southern France (as also strongly suggested 
by the distribution of lithic raw materials; see Negrino & 
Starnini, 2003; Porraz et al., 2010; Negrino et al., 2016).  

In this paper, we will focus on the study of faunal 
remains recovered from the Mousterian, Protoaurigna-
cian, and Aurignacian layers of Riparo Mochi, a Ligurian 
site which has yielded one of the most complete and 
well dated MUPT sequences in this region (Douka et al., 
2012; Frouin et al., 2022). This units had been investi-
gated with a zooarchaeological approach while combin-
ing identified and unidentified bone remains together 
with data from previous studies (Alhaique, 2000; Arella-
no, 2004, 2009). The zooarchaeological analysis carried 
out on such a long sequence will contribute to the inter-
pretation of the site by providing data about the sur-
rounding palaeoenvironment and by defining both the 
behaviour and the economic choices of the hunter-
gatherers groups during the MUPT: How did the fauna 
and the ecosystem around Riparo Mochi evolved? 
Which species were mostly hunted by Neanderthals and 
which by AMHs? Which hunting adaptations were imple-
mented by humans between the Mousterian and 
throughout the Aurignacian? Answering these questions 
will provide a better understanding of how the continu-
ous and rapid climate changes that occurred in Liguria 
during MIS 3 affected the region's palaeoenvironment 
and how human-environment interactions changed be-
tween the final MP and the beginning of the UP. 

2. THE SITE 
 

Riparo Mochi is a rockshelter located at the base of 
a dolomitic limestone cliff called Balzi Rossi (also known 
as Grimaldi Caves), an archaeological complex with 
over 15 sites located in Liguria (Ventimiglia, Italy), close 
to the Italian-French border (Fig. 1). 

The site was discovered by A.C. Blanc and L. Car-
dini of the Istituto Italiano di Paleontologia Umana (IIPU) 
in 1938 and was initially excavated in the same year 
(Blanc, 1938; see a brief history of research at the site in 
Douka et al., 2012). The archaeological deposit is ap-
proximately 10m deep and the sequence consists of 
nine cultural macrounits named from I to A from bottom 
to top by G. Laplace (1977) (Fig. 2). Unit I, associated 
with Mousterian lithic industries, is the thickest unit 
(about 5 m) (Kuhn & Stiner, 1998; Grimaldi & 
Santaniello, 2014). Following Cardini’s fieldnotes, Unit H 
(about 50-60 cm) is a semi-sterile layer where a mix of 
Mousterian and Protoaurignacian artifacts was found; 
recently, field work directed by one of us (S.G.) 
(Grimaldi et al., 2017) shows that the upper part of Unit 
H may represent the earliest UP occupation of the site 
above a thin sterile layer. Following these new data, it is 
reasonable to associate to the base of Unit G the faunal 
remains found during the 1959 L. Cardini excavations 
originally ascribed to Unit H; even if we will maintain the 
distinction in tables and graphs, this hypothesis is con-
firmed by the striking resemblance of the features evi-
denced in the lithic assemblages (see details in Grimaldi 
et al., 2014). Unit G (about 50 cm) provided Protoaurig-
nacian (Laplace, 1977) or Early Aurignacian with Dufour 
bladelets (Bietti et al., 2004) lithic assemblages. The 

Fig. 1 - Geographical location of Riparo Mochi and other archaeological sites cited in the paper (in red). 



   

 

 
 

137 

 
 
The faunal assemblage from the Riparo Mochi site.  

sequence continues with Unit F (about 1 m) defined as 
Middle Aurignacian (Laplace, 1977) or Classic Aurigna-
cian (Bietti & Negrino, 2007). Unit E is a semi-sterile and 
very thin unit, with rare artefacts and faunal remains. 
Unit D (about 1,65 m) is defined as Gravettian with 
Noailles burins (Laplace, 1977) while Unit C (about 55 
cm) provides a Final Gravettian assemblage without 
Noailles burins (Palma di Cesnola, 1993; Santaniello & 
Grimaldi, 2019). Unit B (about 60 cm) is semi-sterile with 
few poor characterizable artefacts, and few bone re-
mains. Finally, the upper Unit A (about 60 cm), previous-
ly defined as Proto-Mesolithic (Laplace, 1977), is now 
defined as Epigravettian (Palma di Cesnola, 1993; To-
masso, 2014). 
 
2.1. Chronology 

Previous radiocarbon dates were obtained using 
the Acid-Base-Oxidation/Stepped Combustion (ABOx-
SC) (Higham et al., 2009, 2011; Douka et al., 2012). 
Using a Bayesian statistical model built with OxCal 4.1 
(Bronk Ramsey, 2009), it has been possible to calculate 
that the Mousterian Unit I ended between 44 and 41.8 
ka cal BP (68.2%), the earliest Aurignacian from the 

bottom of Unit G was dated at 42.7 - 41.6 ka cal BP 
(68.2%), while the transition from G to F occurs between 
37.3 and 36.4 ka cal BP (68.2%). Gravettian Unit D 
started at 30.5 – 30.2 ka cal BP (68.2%) (Douka et al., 
2012). Recently, a new chronological assessment of 
Riparo Mochi’s sequence has been published on the 
base of both radiocarbon ABOx-SC and luminescence 
technique (Frouin et al., 2022). These dates provide new 
information about the timing of the MUPT at this site: 
while the bottom of the Mousterian sequence starts at 
the beginning of MIS 4 - probably earlier (MIS 5) - the 
end of the Mousterian is dated between 45.8-41.9 ka. 

 
2.2. Palaeoenvironmental studies 

Palaeoenvironmental data concerning small mam-
mals (Berto et al., 2019), large mammals (Alhaique, 
2000; 2004; Arellano, 2004; 2009; Zeppieri, 2009; Ta-
gliacozzo et al., 2012), and palynological studies 
(Renault-Miskovsky, 1972), allow a first reconstruction of 
the sequence from the bottom to the top: 

Unit I: small mammal data coming from the upper-
most spits of this Unit suggest the presence of cold wet 
conditions with a landscape characterized by forest 

Fig. 2 - Stratigraphic profile of the Riparo Mochi (original unpublished drawing by A. Segre 1949, modified from Douka et al., 2012). 



   

 

 
 

138 

patches with shrubs surrounded by open and rocky 
areas. Large fauna remains from this Unit were studied 
only from a palaeontological perspective (Arellano, 
2004; 2009) depicting a wide and diversified animal 
spectrum, mainly composed of ungulates. According to 
Berto et al. (2019), by grouping the artificial spits into 
macro groups, it is possible to obtain useful data from 
the variations of the fauna for palaeoenvironmental ob-
servations. At the bottom part of Unit I (spits 52 to 64, A. 
C. Blanc excavations) the abundance of cervids, wild 
boar, and roe deer shows a relatively humid climate. In 
the middle sequence (spits 37 to 46, A. C. Blanc exca-
vations), the higher presence of horse, chamois and 
ibex, as well as the decrease of wild boar suggests a 
shift towards a colder and arid climate. Finally, in the 
upper part of the unit (spits 32 to 36, A. C. Blanc exca-
vations), the presence of Dama dama suggests a milder 
climate.  

Unit H-G: Above Unit I, a change through warmer 
conditions is observed in Units H and G (Berto et al., 
2019), leading to a reduction of the forest environment 
and an increase of the open areas. Faunal remains from 
the semisterile Unit H are missing in the A. C. Blanc 
excavations, while data regarding the UP layers comes 
from Alhaique (2000). To sum up, Cervus elaphus is the 
most common taxon in Unit G, followed by Capra ibex. 
Marmota marmota is also attested but, due to its differ-
ent degree of fragmentation and fossilisation, which 
differs from other remains of the same layer, its pres-
ence is defined as intrusive (Alhaique, 2000). Due to the 
low frequency of remains recovered in the Protoaurigna-
cian level, it is difficult to integrate large faunal data into 
the palaeoenvironmental reconstruction.  

Unit F: A shift to colder temperatures and a gen-
eral reduction of precipitations are observed during the 
transition between Unit G and F; small faunal and paly-
nological data suggest the presence of sparse wooded 
areas without shrubs and a decrease of open meadows 
(Renault-Miskovsky, 1972; Berto et al., 2019). Subse-
quently, milder climatic conditions were observed during 
the evolution of Unit F. Large faunal remains are few but 
more abundant in this unit than in Unit G (Alhaique, 
2000) Cervus elaphus is still the common species in the 
faunal assemblage and an increase of Capreolus capre-
olus is observed.  

Unit E: Here, while decreasing temperatures and 
humidity followed by an improvement of the climatic 
conditions has been suggested (Renault-Miskovsky, 
1972), small mammals remains and large faunal assem-
blage (mainly composed of red deer remains, Alhaique, 
2000) are very scarce.  

Unit D: following (Berto et al, 2019), cold and dry 
conditions are still present in the lower stage of Unit D, 
followed by a decrease of open environments at the 
final phases. Large faunal remains were extensively 
studied by Zeppieri (2009): forest fauna like deer, roe 
deer, and wild boar coexist together with mountain fau-
na like ibex, chamois, and marmot. 

Unit C: Small mammal species are comparable to 
the previous Unit D, but a decrease of the open environ-
ments has been suggested (Berto et al, 2019). The unit 
C is poor in large mammals remains, but cold species 
such Capra ibex and Marmota marmota are attested 

(Alhaique, 2000).  
Unit B-A: while no changes in the small fauna com-

position from unit C to unit B are reported, while an in-
crease of temperature and a decrease of the open envi-
ronment is observed in unit A, leading to a more forest-
ed environment (Berto et al., 2019). Here, the only avail-
able information about large faunal remains comes from 
the original 1938 excavation notes; the presence in unit 
A of remains identified as Marmota marmota, Capreolus 
capreolus, Cervus elaphus and Capra ibex is reported. 

 
3. MATERIALS AND METHODS  
 

Previous zooarchaeological data from the Riparo 
Mochi allow a general assessment of the sequence but 
the lack of data about anthropic activities and their im-
pact on the bone assemblages prevents more accurate 
interpretations. In fact, while large mammals from Unit D 
were extensively studied by a zooarchaeological ap-
proach (Zeppieri, 2009, summarized in Tagliacozzo et 
al., 2012), the same cannot be said for the Units below. 
Nowadays, Unit I was sampled and studied only from a 
paleontological perspective (Arellano, 2004, 2009) while 
data from Units G, F, and E (Alhaique, 2000) come only 
from limited archaeological samples. Finally, data from 
unit H (L. Cardini 1959 excavations) are still un-
published. The aim of this paper is to fill these gaps by 
analyzing the whole faunal assemblages coming from 
the site. 

 
3.1. The analysed faunal assemblage 

The analysed faunal assemblage counts a total of 
41.162 remains from Units E, F, G, H, and I. The re-
mains were found during the A. C. Blanc excavation 
campaigns (1938, 1941-2, and 1948-9), L. Cardini exca-
vation (1959), A. Bietti excavations (1995-2006), and 
current excavations led by one of us (SG) since the 
2007. A re-examination of previously studied materials 
(Alhaique, 2000; Arellano, 2004, 2009) was followed by 
the analysis of all other unstudied remains. As already 
said above, recent field work, allows to consider faunal 
remains from Unit H likely to be in association with the 
Protoaurignacian level found at the base of Unit G; fau-
nal remains from Unit E may be seen as a homogenous 
sample without any internal subdivision. On the contrary, 

Tab. 1 - Dating estimate made by luminescent determinations 
on feldspar from Mousterian (Unit I) and semi sterile (Unit H) of 
Riparo Mochi (see detailed discussion about bayesian models 
in Frouin et al., 2022). The subdivision in phases takes in ac-
count the variation of sedimentology and the techno-typological 
features of the lithic assemblages. 

 
 

Perez A. et al. 



   

 

 
 

139 

the remains from the Mousterian Unit I have been divid-
ed into three groups named: Unit I – base (Ib), Unit I – 
corpus (Ic), and Unit I – top (It) (Tab. 1). The reason for 
that is to be found in a clear sedimentological variability 
found during the earliest excavation activities as well as 
in techno-typological differences found in recent studies 
of the lithic assemblages (see details in Grimaldi et al., 
2017). In sum, the middle part of the Mousterian se-
quence in Unit I (that is to say, I-corpus) suggests an 
environmental change, possibly corresponding to a cold-
er/dryer landscape (Blanc, 1938; Laplace, 1977), pre-
sumably associated with a behavioural adaptation evi-
denced by the lithic production (Grimaldi & Santaniello, 
2014).  
 
3.2. The analytical approach 

Taxonomic and anatomical identifications were 
carried out using the reference collections of the Labora-
tory of Zooarchaeology of MUSE (Trento, Italy) and of 
the Laboratory of Large Vertebrate of the University of 
Ferrara (Ferrara, Italy); skeletal and anatomical atlases 
were also used (Pales & Lambert, 1971, 1981; Schmid, 
1972; Barone, 1976). Undetermined remains were sort-
ed into four weight size classes, following the criteria 
proposed by Bunn (1986) with some adaptations related 
to the studied materials: (1) small-size mammals weigh-
ing >20 kg (small carnivores, Lagomorpha, Rodentia); 
(2) medium-size mammals weighing between 20 and 
100 kg (Capridae, small cervids, Sus scrofa); (3) medi-
um-large size mammals weighing between 100 and 300 
kg (Cervus elaphus, Ursidae); (4) large size mammals 
weighing >300 kg (Equidae, Rhinocerontidae, Alces 
alces, Megaloceros giganteus). The remains were also 
arranged according to their maximum length: (1) 0 to 2,9 
cm; (2) 3 to 3,9 cm; (3) 4 to 4,9 cm; (4) 5 to 5,9 cm; (5) 6 
to 7,9 cm; (6) 8 to 9,9 cm; (7) 10≤ cm. Quantification of 
the identified remains were operated using: Number of 
remains (NR) and Number of identified specimens 
(NISP) Grayson (1984). Minimal number of individuals 
(MNI) was estimated from the observation of skeletal 
elements and teeth of the same laterality and osteologi-
cal development (Grayson, 1984; Klein & Cruz-Uribe, 
1984; Lyman, 1994), Minimal number of elements 
(MNE) was counted considering the side, size and on-
togeny of the elements (Binford, 1984; Klein & Cruz-
Uribe, 1984),  Minimal anatomical units (MAU) (Binford, 
1984). %MAU was calculated dividing the MAU value of 
each element by the highest MAU value of the assem-
blage. A percentage equal to 100% for all the body parts 
would imply the entire presence of a complete animal 
skeleton. Age at death estimation was made based on 
the degree of dental wear and the presence of decidu-
ous teeth, also the fusion stage of the epiphysis was 
considered (Habermehl, 1961, 1992; Silver, 1969; Nod-
dle, 1974; Mariezkurrena & Altuna, 1983; Stiner, 1998; 
Gipson et al., 2000; Azorit et al., 2002; Tomé & Vigne, 
2003; Weinstock, 2009; Geiger et al., 2016). Five age 
groups were defined according to the degree of dental 
wearing: Infant (animals with unworn deciduous teeth), 
Juvenile (animals with slight to moderate worn decidu-
ous teeth), Subadult (animals with highly worn decidu-
ous teeth or unworn permanent teeth according to the 
timing of dental eruption for each species), Adult 

(animals with permanent teeth with low or moderate 
wear) and Senile (animals with highly worn teeth). Un-
derstanding variations of evenness and diversity of the 
faunal assemblage throughout different archaeological 
units, correlated with the presence of anthropic modifica-
tions and other natural events (e.g. carnivores activity, 
post-depositional processes, etc.), allows us to better 
comprehend possible human-environment interactions 
that occurred between the MP and UP (e.g. the data 
might show if there was hunting activity was focused on 
specific animals).  To evaluate the diversity of the faunal 
assemblage Inverse Simpson’s index of diversity (1/D) 
was calculated; this index allows to measure the diversi-
ty of the animal population (Simpson, 1949) found in an 
archaeological site. We decided to use this index be-
cause is the most reliable when applied to a faunal as-
semblage as diverse as the one observed at Riparo 
Mochi (Faith & Du, 2018). It is also a very reliable index 
even for relatively small assemblages.  The resulting 
values permit to understand the taxonomical evenness 
of the faunal assemblage: the lower the value, the more 
the faunal assemblage is dominated by a single species; 
on the contrary, the higher the value, the more the diver-
sity of species in the faunal assemblage. To investigate 
pre- and post-depositional processes that occurred at 
the site, all specimens were examined with a 10x hand 
lens and with a binocular microscope. Modifications of 
major interest were captured with a Leica SD6 micro-
scope. Non-human biotic modifications made by ro-
dents, carnivores and roots were recorded as well as a-
biotic modifications such as concretions, weathering, 
manganese oxide and trampling (Behrensmeyer, 1978; 
Lyman, 1994; Fisher, 1995; Blumenschine et al., 1996; 
Domínguez-Rodrigo & Barba, 2006; Fernández-Jalvo & 
Andrews, 2018). Carnivores’ marks were classified as 
pits, punctures, gnawing, furrowing and digestion 
(Fisher, 1995; Domínguez-Rodrigo & Piqueras, 2003; 
Domínguez-Rodrigo & Barba, 2006; Coil et al., 2020). 
Anthropogenic modifications observed in the assem-
blage includes cut-marks, percussion marks and burned 
remains. These traces have been documented and ana-
lysed, however their morphological characterisation, 
location on the skeleton and their correlation to specific 
carcass exploitation activities will not be treated in this 
study (Binford, 1981; Potts & Shipman, 1981; Shipman, 
1981; Shipman & Rose, 1984; Blumenschine & Selvag-
gio, 1988; Pickering & Egeland, 2006; Galán et al., 
2009; Vettese, 2014; Vettese et al., 2017, 2020; Fernán-
dez-Jalvo & Andrews, 2018;Coil et al., 2020). Finally, 
burnt specimens were sorted according to the colour: 
brown (burnt), black (carbonized) and grey-white 
(calcinated) (Stiner et al., 1995). We also carried out a 
preliminary investigation to distinguish between remains 
truly affected by thermal alterations and remains dark-
ened by post-depositional processes (Perez et al., 
2020). 
 
4. RESULTS 
 
4.1. The faunal assemblage 

The taxonomically identified specimens are 792 
(NISP), belonging to 111 (MNI) individuals (Tab. 2). 
There were identified 22 taxa, mainly large mammals, 

 
 
The faunal assemblage from the Riparo Mochi site.  



   

 

 
 

140 

 

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Perez A. et al. 



   

 

 
 

141 

and few unidentified bird bones. Ungulates dominate the 
faunal assemblage of each unit, while carnivores are 
rare. Lagomorpha, Insectivora, and Rodentia were also 
observed.  

The three phases of Unit I are dominated by differ-
ent species: Unit Ib (100 NISP, 18 MNI, 11 taxa) by Cer-
vus elaphus (30% NISP, 3 MNI), Unit Ic (71 NISP, 15 
MNI, 11 taxa) by Equus sp. (39% NISP, 3 MNI), Unit It 
(127 NISP, 25 MNI, 12 taxa) by Sus scrofa (20% NISP, 
3 MNI) (Tab. 2). A single fragmented tooth of Rhinoc-
erontidae (1% NISP, 1 MNI) is only observed in Ib lev-
els, while large mammals are common in the entire Unit 
I. Bos/Bison is more common in Ib (20%NISP, 1 MNI) 
and It (11%NISP, 3 MNI) than in Ic (6%NISP, 1 MNI). 
On the contrary, Equus sp. NISP has a peak in Ic layers 
(39%NISP, 3 MNI). Medium-large size ungulates like 
Cervus elaphus are much more abundant in Ib and Ic 
layers (Tab. 2). As red deer belongs to the medium-
large size class, its presence in the archaeological rec-
ord may be underestimated. It is interesting to note that, 
within the Unit I, antler (Ib: 4 NR; Ic: 1 NR; It: 8 NR) and 
metapodial fragments (Ib: 14 NR; Ic: 4 NR; It: 12 NR) of 
undetermined cervidae show a similar distribution to that 
of Cervus elaphus (minor NISP in Ic than Ib and It). Cap-
reolus capreolus (Ib: 2% NISP, 1 MNI; It: 1% NISP, 1 
MNI) and Capra ibex (Ib: 6% NISP, 1 MNI; Ic: 11% 
NISP, 1 MNI; It: 4%, 2 MNI) are scarce, while Dama 
dama is observed only in It (1% NISP, 1 MNI). As far as 
weight size classes are concerned, small and medium 
size mammal remains are scarce; medium-large size 
remains are more common in Ib and It and this coin-
cides with the high presence of red deer and wild boar in 
the same layers; large size mammal remains are com-
mon in each phase of the unit. Carnivores are rare: few 
remains of Ursus spelaeus (Ib: 3% NISP, 1 MNI; It: 1% 
NISP, 1 MNI), Ursus arctos (1% NISP, 1 MNI),, Canis 
lupus (Ib: 1% NISP, 1 MNI; Ic: 1% NISP, 1 MNI; It: 2% 
NISP, 1 MNI), and Panthera sp. (Ic: 1% NISP, 1 MNI) 
are observed. Lepus sp. (12%NISP, 3 MNI) is present 
only in It, but a different preservation as well as the pres-
ence of three individuals coming from the same spits 
and from the same sector of the excavation, could sug-
gest their appearance in the stratigraphy after the depo-
sition of Unit I. The fossorial nature of this animal might 
support this hypothesis.  

From the semisterile Unit H (63 NISP, 12 MNI, 10 
taxa) it is possible to observe a variation in the faunal 
assemblage which will persists into the subsequently 
Unit G (199 NISP, 18 MNI, 12 taxa) and Unit F (178 
NISP, 15 MNI, 8 taxa). In these Units, large ungulates 
show a decrease in NISP and, in the case of Equus sp. 
(H: 1.6% NISP, 1 MNI; G: 1% NISP, 1 MNI; F: 1.1% 
NISP, 1 MNI), the reduction is remarkable (Tab. 2). Me-
dium size ungulates are highly represented as in the 
previous Unit I: Cervus elaphus is the most common 
species in each unit (H: 36.5% NISP, 1 MNI; G: 50.3% 
NISP, 4 MNI;  F: 53.4% NISP, 4 MNI). Also, cervids 
remains result noticeably well represented in Unit G (78 
NR) coinciding with the highest NISP observed for red 
deer in the entire deposit (100 NISP). It is also observed 
an increase of NISP of Capra ibex, which is the second 
most abundant taxon (H: 19% NISP, 1 MNI; G:  13.6% 
NISP, 3 MNI; F: 22.5% NISP, 3 MNI), while NISP of Sus 

scrofa decreases (H: 11.1% NISP, 1 MNI; G: 10.6% 
NISP, 2 MNI; F: 9.6% NISP, 2 MNI). Medium size ungu-
lates are more abundant than those observed in the 
lower unit: Capreolus capreolus is well represented in 
Units G and F (G: 11.1% NISP, 1 MNI; F: 6.2% NISP, 2 
MNI) while few remains of Rupicapra rupicapra (G: 0.5% 
NISP, 1 MNI; F: 0.6% NISP, 1 MNI) have been found. 
The abundance of red deer and roe deer in Protoaurig-
nacian /Aurignacian layers could be related to the high 

Fig. 3 - Inverse Simpson’s index (1/D) calculated on MNI for 
each archaeological unit of Riparo Mochi.  

Tab. 3 - Comparison between the inverse Simpson’s index (1/
D) calculated in Riparo Mochi and other contemporary sites of 
the Ligurian region. Both NISP and MNI were used to calculate 
the 1/D. (a) data from Holt et al. (2019). (b) data from Valensi & 
Psathi, (2004). (*) data not available. 

 
 
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142 

presence in unit G and F, of numerous antlers (G: 45 
NR; F: 6 NR) and metapodials fragments (G: 30 NR; F: 
17 NR) identified as Cervidae Specimen sorted by 
weight size classes might reflect the distribution of NISP 
from Units H, G, and F. Medium-large size ungulate 
remains are the most numerous (H: 30 NR; G: 82 NR; 
F: 56 NR), followed by medium (H: 15 NR; G: 35 NR; F: 
18 NR) and small size (H: 7 NR; G: 19 NR; F: 10 NR), 
while large size ungulates (H: 2 NR; G: 13 NR; F: 9 NR) 
decrease in number. Carnivores are rare and absent in 
Unit F. Ursus spelaeus is present only in Unit H (4.8% 
NISP, 1 MNI), while Ursus sp. is present in Unit H (3.2% 
NISP, 1 MNI) and Unit G (0.5% NISP, 1 MNI). Crocuta 
sp. (0.5% NISP, 1 MNI), Canis lupus (0.5% NISP, 1 
MNI), and Mustela sp. (0.5% NISP, 1 MNI) are only 
present in Unit G. Lepus sp. from Unit H (9.5% NISP, 2 
MNI) could be linked to the analogous remains recov-
ered in the top of Unit I as both seem to share different 
taphonomic characteristics from those of the fossil rec-
ord; moreover, they come from the same excavation 
sector. Two remains of Marmota marmota (H: 1.6% 
NISP, 1 MNI; G: 0.5% NISP, 1 MNI) and a mandible of 
Erinaceus sp (1% NISP, 1 MNI). from Unit G and F are 
observed. At last, few specimens (157 NR) come from 
the semisterile Unit E. Among the ungulates, only Cer-
vus elaphus (58.6% NISP, 3 MNI), Capra ibex (12.1% 
NISP, 1 MNI), Rupicapra rupicapra (1.7% NISP, 1 MNI), 
and Capreolus capreolus (24.1% NISP, 1 MNI) are pre-
sent (Tab. 2). Cervids remain the most common ungu-
late taxa, while carnivores are scarce with the presence 
of Ursus sp. (1.7% NISP, 1 MNI) and Canis lupus (1.7% 
NISP, 1 MNI). Weight size classes remains are differ-
ently distributed: medium size mammals (33 NR) are the 
most abundant followed by small size (17 NR), medium-
large (4 NR) and large size (1 NR) are scarce.  

MNI value was used to calculate 1/D, Lepus sp. 

and Marmota marmota are not included as they both 
appear to be intrusive. The graph in Figure 3 shows how 
Mousterian faunal spectrum is characterized by a high 
diversity of taxa. Unit H and G, have a similar lower 1/D 
value (Fig. 3, Tab. 3). In the Aurignacian layers (Unit F) 
a decrease in diversity is observed, while in Unit E the 
diversity drops drastically. On the light of this, a de-
crease of faunal variety seems to happen during the UP 
units, especially in Unit F. 
 
4.2. Mortality profile 

A total of 111 individuals were identified in the site 
(Tab. 4). Young individuals are observed only in Unit I. 
Four juvenile individuals of Sus scrofa were identified: in 
Ib, two juveniles by two left dp4 (age estimation 5-7 
weeks according to Silver 1969; Habermelh, 1961); in Ic, 
one individual by an unfused metatarsal distal epiphysis, 
and in It, one individual from a left DP3. A new-born 
individual of Equus sp. were identified from a left DP3 in 
Ic. Senile individuals are observed in Unit H and Unit G, 
respectively an old Capreolus capreolus, identified from 
a high worn molar, and a Cervus elaphus old individual 
identified from a marked worn M1. The other taxa show 
only adult individuals. Cervus elaphus appears to be the 
species with the most number of individuals (19) across 
the entire stratigraphy (Tab. 4). In units G and F the MNI 
of the red deer increases slightly (Tab. 4)  and antler 
fragments are present. The only observable variation is 
the presence of a senile individual in Unit G (Tab. 4). 
Despite the presence of undetermined cervids antler 
fragments in unit I, G and F, it was not possible to define 
whether they were antlers obtained by hunting the ani-
mal or simply collected. For this reason, it is currently 
not possible to hypothesise a hunting seasonality for 
Cervus elaphus. However, it must be taken into account 
that the undetermined antler fragments may have be-

Tab. 4 - Age classes for each taxon of Riparo Mochi. Infant (I), Juvenile (J), Sub-adult (Sad), Adult (Ad), Senile (S). 

 
 

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143 

longed to Capreolus capreolus (from unit I to E), Dama 
dama and Alces/Megaloceros (only in unit I). Sus scrofa 
is the second ungulate with the highest number of total 
individuals (17 MNI). It is noted that throughout the 
Mousterian sequence there is a large presence of indi-
viduals (12 MNI), four of which are juvenile. We ob-
served that, starting from unit H, its MNI value decreas-
es (Tab. 4).  Considering that the wild boar can give 
birth twice a year when conditions are optimal (Mauget, 
1981), we could theorise that during the deposition of 
Unit I the regional environmental conditions have been 
milder compared to the upper levels. To strenghten this 
hypothesis, we observed similar variations in MNIs of 
Equus sp., Bos primigenius, Bison priscus and the ge-
nus Bos/Bison sp.: these appear to have higher values 
in the Mousterian levels than in the Aurignacian levels 
(Tab. 4). In particular Equus sp., which in Unit I has a 
total MNI equal to 8, one of which is an infant, reduces 
its MNI to a single individual in units H, G and F. This 
decrease is also observable in the NISP (Tab. 2). Con-
versely, an increase in the MNI of Capra ibex is ob-
served in units G and F. This decrease in the MNI of 
species wich inhabitate forests and/or open environ-
ments, such as Equus sp. and Bos/Bison and the in-
crease in the MNI of mountain species such as the ibex, 
might suggests a change in the environmental condi-
tions and/or in human hunting strategies that occurred 
between the MP (Unit I) and the UP (Unit G and F), re-
flected in the composition of the faunal spectrum. 
 
4.3. General taphonomy 

Except for Unit E, Ib, and Ic which have few re-
mains, the whole assemblage is highly fragmented (Fig. 
4). Bone fragments smaller than 2.9 cm represent the 
87% of the total remains. Whole specimens are the 

0.3% (140 NR) of the total, which are primarly represent-
ed by small fat bearing bones, such as sesamoids, car-
pals, tarsals, phalanges, and metapodial of small fauna, 
in association with isolated teeth (Fig. 5); low density 
bones as ribs, scapulae, and vertebras are rare. We 
have divided them according to the weight size of the 
species to which they belong. The remains belonging to 
small animal (small carnivores, rodents, birds) are pri-
marily represented by short limb bones like tarsals and 
phalanges. Those concentrated in It and H (Fig. 5) are 
almost exclusively determined as Lepus sp. The scarcity 

Fig. 4 - Percentage of fragmented remains (NR%) for each unit 
of Riparo Mochi sorted in seven classes according to their 
length. 

Fig. 5 - Percentage of whole remains divided according to the animal size in each unit of Riparo Mochi. Small size (small carnivores, Lago-
morpha, Rodentia), Medium size (Capridae, small cervids, Sus scrofa), Medium-large size (Cervus elaphus, Ursidae), Large size (Equidae, 
Rhinocerontidae, Alces alces, Megaloceros giganteus). 

 
 
The faunal assemblage from the Riparo Mochi site.  



   

 

 
 

144 

or absence of whole remains belonging to small animals 
in the other units it might confirm the intrusive nature of 
Lepus sp. in these levels. Overall, the skeleton of small 
faunas is mainly represented by fore and hindlimb in 
Units It and H (Fig. 6), while the cranial portion is the 
only represented part in F. Isolated teeth are rare and 
only observed in Ic. The scarce representation of small 
faunas throught the stratigraphic sequence could be the 
result of a heavier fragmentation of these elements. 
Medium (caprids, canids, suids) and medium-large (red 
deer, ursids) size animal bones share a similar distribu-
tion (Fig. 5, Fig. 6). For both the size classes whole 
remains are mainly represented by isolated teeth, small 
fat bearing bones such as carpals, tarsals, sesamoid 
and falanges (Fig. 5). The major variations in the stratig-
raphy are observed in Ic, where theeth are the only 
whole specimens, and in It in which the representation 
of short limb bones increases (Fig. 5). This variation is 
observable also in the representation of the anatomical 
parts (Fig. 6). The cranial skeleton of both size classes 
starts to decrease in unit H, reaching its minimum in unit 

 

Fig. 6 - Percentage of remains belonging to a specific anatomical portion in each unit of Riparo Mochi. Data are divided according to the 
size of the animal: Small size (small carnivores, Lagomorpha, Rodentia), Medium size (Capridae, small cervids, Sus scrofa), Medium-large 
size (Cervus elaphus, Ursidae), Large size (Equidae, Rhinocerontidae, Alces alces, Megaloceros giganteus)  

Fig. 7 - Percentage of remains bearing concretions, manganese 
oxide and root-etching in each unit of Riparo Mochi. 

 
 

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145 

G (less than 50% of the total NR%). Also the axial skele-
ton, rarest than the other anatomical parts, increase is 
frequency in the UP levels. A different variation is ob-
served in Unit E where the representation of the cranial 
skeleton is highest for the medium size animals than for 
the medium-large (Fig. 6). Lastly, whole remains of large 
size animals are composed by isolated teeth concentrat-
ed in Ib and Ic and sesamoid in F (Fig. 5). The cranial 
skeleton, mainly represented by teeth, is the most abun-
dant anatomical part in every archaeological unit, with 
the exception of unit E (Fig. 6). Cranial skeleton of large 

faunas presents a decrease in unit H and elements of 
the forelimb and hindlimb increase in number in unit G-
F. Is also observed a higher presence of the axial skele-
ton, which becomes the only anatomical portion present 
in unit E. Medium, medium-large and large sized re-
mains seem to have been subject to a similar attrition 
due to intense fragmentation processes. It cannot be 
excluded that the skeletal representation are also due to 
anthropic activity, which may have concentrated on the 
intentional breakage of bones. Traces of percussion are 
in fact the most attested anthropic traces in the entire 

Fig. 8 - Remains showing anthropic and carnivore modifications. A: Capreolus capreolus metatarsal with scraping marks on its diaphysis 
side (Unit I). B: Cervus elaphus humerus with impact point on its diaphysis (Unit G). C: an unidentified long bone fragment corroded by 
carnivore digestion (Unit I). D: Unidentified long bone diaphysis of a medium-large size mammal with multiple cut-marks (Unit F). E: Cervus 
elaphus metacarpal fragment with a single cut-mark (Unit F). 

 
 
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146 

assemblage, especially on long bones of medium-large 
size animals (Tab. 5). Furthermore, the presence of all 
the anatomical portions might suggest a complete 
transport of the carcasses, which occurred continuously 
between MP and UP. 

The preservation of bone surfaces is low as a high 
presence of concretions (Fig. 7), which affects almost 
half of the total remains in Units I and H (Ib: 58%, Ic: 
48%, It: 49%, H: 49%), is observed; it highly increases 
from Unit G to Unit E (G: 87%, F: 93%, E: 98%). Manga-
nese dendrites are observed in each unit, with a major 
abundance in Units I, G, and E. Root-etching is scarce 
except in Unit E, but, like the manganese oxides, the 
scarce amount of remains in this unit could lead to an 
overrepresentation of the data.  

Carnivore modifications are present in Units I and 
G but they are extremely rare and observed only on the 
0.03% of remains (Tab. 5, Fig. 8). Pits and scores are 
the main alteration made by carnivores and only one 
remain shows digestion marks. Among the identified 
taxa, a Cervus elaphus metacarpal and a Cervidae met-
apodial show carnivore modifications, while other nine 
remains belong to medium-size mammals. No rodent 
marks are observed. Traces of anthropogenic activity on 
animal remains are rare but present throughout the 
stratigraphic sequence (Tab. 5). Among the identified 
species, the most exploited remains belong to the Cer-
vus elaphus, whose remains show few percussion 
marks and cut marks. The exploitation of red deer ap-
pears to be continuous, from the Mousterian to the Early 
Aurignacian. Only in Unit I, rare anthropogenic modifica-
tions are observed on Capreolus capreolus and Sus 
scrofa remains. Nevertheless, along the entire site se-
quence, anthropic modifications are mostly observed on 
diaphysis of medium-large unidentified mammals. Burnt 
remains are abundant in all units, predominantly indeter-
minate bone fragments. However, there are few remains 
of Bos/Bison, Cervus elaphus, Capreolus capreolus, 
Capra ibex, Sus scrofa and Lepus sp. showing traces of 
combustion. 
 
4.4. Skeletal representation 

Skeletal representation data covers only those 
species with abundant remains, throughout the whole 
stratigraphy, which are not uniquely teeth. Those well-
represented species are Bos/Bison sp., Cervus elaphus, 
Capreolus capreolus, Capra ibex and Sus scrofa. One 
can observe how cranial elements are frequent through-
out the sequence, especially mandibles, except for units 
Ic and H. Such abundance is coherent with the high 
number of cranial elements and isolated teeth in the 
entire faunal assemblage (Fig. 6). The axial skeleton is 
almost entirely absent (Fig. 9), with scarcely frequent 
ribs from Bos/Bison sp. (G: 8% MAU) and Cervus ela-
phus (G: 2% MAU; F: 2% MAU; E: 2% MAU) in the up-

 

__________________________________________      >>>>> 
 
Tab. 5 - Anthropic and carnivore modifications observed on the 
fossil assemblage. Cut-mark (Cm), percussion-mark (Pm), cut-
mark + percussion-mark (Cm+Pm), burned bones (B), calcinated 
bones (C), percentage of anthropic modifications (%Ant), carni-
vore modifications (Car), percentage of carnivore modifications 
(%Car). 

 
 

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147 

per levels, and totally absent for unit I and H. 
The limb skeleton is well represented: the 
hindlimb, specifically the tibia and metapodials, 
are quite frequent throughout the whole stratig-
raphy. The main variation of the skeletal repre-
sentation is in between and MP and UP. In 
fact, in Unit H, the number of limb elements 
represented in the skeletal profile increases, 
with a high frequency of hindlimb bones, such 
as tibias, tarsals, phalanges. The increase in 
the latter could be related to a general in-
crease of short limb bones observed already in 
Unit It (Fig. 5). In unit G and F there is a higher 
number of elements of the proximal forelimb, 
such as scapula, humerus and radio-ulna, as 
well as of the hindlimb, such as tibia and 
astragalus, and a high number of both core 
and hind metapodials. Large sized animals, as 
the Bos/Bison, are represented by radio-ulna, 
femur and tibia, while for the Medium-large 
and medium animals, like red deer, ibex, wild 
boar and roe deer, we have every limb ele-
ment, in particular the forelimbs. This data, 
together with the scarcity of elements of the 
axial skeleton, might suggest a differentiated 
transport and exploitation of preys defined by 
their weight size. Nevertheless, the high fre-
quency of mandibles (Fig. 9) and the presence 
of all anatomical parts in the whole archaeo-
logical assemblage (Fig. 6), suggests that the 
carcass was transported as a whole inside the 
site. In this case, the absence of fragile and 
low-density bones, such as vertebrae, ribs and 
cranial bones may be due to intense tapho-
nomical processes which have caused a high 
fragmentation of these elements (Fig. 4). 
 
5. DISCUSSION 
 
5.1. Faunal variability and palaeoenviron-
mental scenario 

The zooarchaeological data allow a better 
interpretation of a possible environmental con-
text during which the Riparo Mochi archaeo-
logical deposit was formed. Generally speak-
ing, the fauna spectrum all along the Riparo 
Mochi sequence - from the MP to the Initial UP 
- seems to reflect the presence of a landscape 
characterized by a high variety of ecosystems 
(Fig. 3; Tab. 3). 
 
5.1.1. Unit I 

Base: The earliest phase of the Mousteri-
an of Riparo Mochi is characterised by a great faunal 
diversity (Fig. 3). The copresence of species preferring 
open grassland environments such as Bison priscus and 
Equus sp. (McDonald, 1981; Berger, 1986), or ones 
more akin to temperate forest environments such as 
Capreolus capreolus (Danilkin, 1995), as well as alpine 
taxon such as Capra ibex, probably reflects great variety 
of habitats present in the region surrounding the site at 
the end of MIS 4. Carnivores are rare, but the presence 
of coprolites from Ib and Ic (spits from 39 to 64) and 

undetermined remains with traces of digestions in unit It 
(Tab. 5) could indirectly suggest a occupation of the site 
by large predators such as hyenas (Tab.2). The pres-
ence of young Sus scrofa individuals, which are largely 
present in Ib, as well as throughout the entire Mousteri-
an sequence (Tab. 4), could reflect a high environmental 
richness. Considering that this species can breed twice 
a year under favourable conditions (Mauget, 1981), we 
could suggest a high presence of wild boar at regional 
level, whose high reproductive rate could have been 
favoured by milder environmental conditions which 

 

Fig. 9 - Percentage of Minimum Anatomical Units of Bos/Bison, Cervus elaphus, 
Capreolus capreolus, Capra ibex and Sus scrofa in each unit of Riparo Mochi. 

 
 
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148 

seem to have been maintained in Liguria even in peri-
ods of general cooling and climatic dryness (Negrino 
and Tozzi, 2008; Berto et al., 2019). The faunal assem-
blage from Ib fits reasonably well into the Ligurian ar-
chaeozoological palimpsest straddling MIS 4 and the 
beginning of MIS 3. Sites such as Madonna dell’Arma, 
Arma delle Manie, Santa Lucia Superiore and San Fran-
cesco (Valensi & Psathi, 2004), where anthropogenic 
accumulation is widely attested, are also characterised 
by a similar level of faunal diversity (Tab. 3). 

Corpus: The main variation observed in the Mous-
terian levels is in Ic. Here, several features allow to de-
tect an environmental change, towards dryer and colder 
conditions (Tab. 3). In fact, it is shown an abrupt in-
crease of Equus sp. and a general decrease in the pres-
ence of other species; the absence of roe deer, of which 
the primary habitat is forest clearings, hedges, and 
woodland edges (Mauget, 1981; Danilkin, 1995), may 
support the hypothesis of a colder and less tree-covered 
environment during this phase. Finally, the presence of 
Alces/Megaloceros could also reflect colder environ-
mental conditions. Due to its chronology, this part of the 
Mousterian sequence seems to fall into a the Greenland 
Stadial 15.1 (Rasmussen et al., 2014) which could have 
led to a decrease in temperature and a consequent 
regression of temperate taxa. Unfortunately, no palaeo-
ecological nor archaeobotanical data are available nei-
ther for Ic nor for Ib phases. As in the previous Mousteri-
an layers, hyena coprolites have been found and the 
presence of carnivores such as Panthera sp., a high 
presence of Sus scrofa, and a high MNI 1/D value, may 
be a proxy for a wide biodiversity (Tab. 3). 

Top: Similarly, to what happens in Ib as well as 
palaeoenvironmental data also suggested (Renault-
Miskovsky, 1972; Berto et al., 2019), the region sur-
rounding the site was characterized by a patchwork of 
habitats of open and forested areas with a cold-humid 
environment during the final phase of the Mousterian at 
Riparo Mochi. The inverse Simpsons’ index is the high-
est for the Mousterian Unit (Fig. 3), showing a wide di-
versity in the faunal composition. Moreover, this diversi-
ty has been also recognized in the Mousterian level “M” 
of the nearby site of Riparo Bombrini (Holt et al., 2019), 
where a NISP 1/D appears to be very similar to what 
has been found in It of Riparo Mochi (Tab. 3). This data 
confirms how the region's great environmental variety 
persists during the coldest climatic phases. 
 
5.1.2. Unit H 

As far as the assemblage coming from Cardini’s 
Unit H is concerned as a whole, it is worth to mention 
that few remains were clearly identified showing few 
anthropogenic modifications. Generally speaking, the 
data coming from this Unit are very close to the later 
Unit G (see below). A general decrease in the faunal 
spectrum diversity is observed (Fig. 3), particularly with 
species such as Equus sp. and Sus scrofa (Tab. 2). 
Similarly, this evidence is also present in the semi-
sterile 'MS' levels of Riparo Bombrini (Holt et al., 2019) 
(Tab. 4). Other studies show a rise in temperatures and 
a change in the rainfall regime which should have led to 
a reduction in forest cover in favour of an expansion of 
open meadows (Berto et al., 2019). As this change is 

also observed in the later Unit G, it is therefore difficult 
to determine whether the variation in the faunal spec-
trum observed at the beginning of the Protoaurignacian 
level was mainly caused by a change in environmental 
conditions or by a change in hunting strategies that oc-
curred with the arrival of the first AMHs at the site.  
 
5.1.3. Unit G 

As seen in Unit H, the large fauna assemblage of 
the Protoaurignacian Unit G has a minor diversity than 
the Unit I (Fig. 3). Variability between the two Units is 
mainly represented by the presence of Rupicapra rupi-
capra, by a greater presence of Cervus elaphus and 
Capra ibex as well as an increase in the number of an-
thropogenic modifications in Unit G. As shown by small 
fauna studies (Berto et al., 2019), the general warming 
and reduction of forestall cover seen in unit H is still in 
progress during the formation of Unit G. The lower diver-
sity of the faunal spectrum (Fig. 3, Tab. 3) as well as the 
decreasing presence of Equus sp. and Sus scrofa re-
mains (Tab. 2), may testify a mutation of the biodiversity 
of the region, maybe characterized by more presence of 
alpine faunas (as suggested by the higher frequence 
ibex and chamois remains). As seen in the oldest ar-
chaeological units, the remains of carnivores are still 
rare; moreover, the absence of coprolites from these 
levels could indicate a reduced frequentation of the site 
by hyenas.  
 
5.1.4. Unit F 

No major changes are observed in the species 
represented in Unit F. Although small mammals and 
palynological data indicate a colder and drier environ-
ment in Unit F than in Unit G (Renault-Miskovsky, 1972; 
Berto et al., 2019), the representation of species ap-
pears constant through the UP sequence. Nevertheless, 
the faunal assemblage shows an increasing loss of di-
versity, which reaches the lowest value in Unit F (Fig. 3). 
Another interesting value is the lack of carnivores (Tab. 
2) that could be speculatively explained by a more in-
tense human occupation, which is also supported by a 
higher number of anthropogenic modifications observed 
on bone remains (Tab. 5). Cervus elaphus, Capra ibex, 
and Capreolus capreolus are the main taxa found in the 
early Aurignacian unit. The coexistence of ibex and roe 
deer - species that frequent alpine grassland and wood-
ed areas, respectively - may indicate the persistence of 
different environments, relatively close to each other, in 
the Riparo Mochi region. However, the presence of 
chamois, already found in Unit G, may reflect a greater 
presence of mountain faunas in the UP units than seen 
in the MP unit. 

 
5.1.5. Unit E 

Given the ephemeral nature of this deposit and the 
scarce presence of bone remains, Unit E may have 
been formed by an intrusion of materials from overlying 
and underlying units. It should be noted that Gravettian 
unit D has similar faunal data to units E and F 
(Tagliacozzo et al., 2012). In fact, the faunal spectrum of 
the Gravettian levels continues to be dominated by Cer-
vus elaphus (37.69% NISP), and there is a similar pro-
portion of Capra ibex remains (19.59% NISP) to that 

 
 

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149 

observed for the ibex in Unit F (22.5% NISP). Unlike the 
older units, carnivores are present with a greater num-
ber of species: fox, wild cat and lynx but also Panthera 
pardus and Crocuta crocuta.  

The faunas of the Gravettian unit continue to reflect 
a mosaic of environments, in which there is a co-
presence of forest species such as deer and wild boar 
and 'cold' or alpine environment species such as cham-
ois, ibex and marmot (Tagliacozzo et al. 2012), confirm-
ing that the climatic conditions of the region remain mild 
even with the cold and dry conditions present in the 
lower stage of Unit D (Berto et al. 2019).  
 
5.2. Human-faunal interaction 
 
5.2.1 Middle Palaeolithic 

Despite the presence of hyena coprolites during 
the Ib and Ic phases, and the rare traces of digestion 
and fracturing by carnivores, human groups certainly 
acted as the main agent for the accumulation of bone 
remains of the site. Unfortunately, the concretions as 
well as the high rate of bone fragmentation hamper a 
clear recognition of the anthropogenic modifications, 
making archaeozoological interpretation difficult. Thus, 
traces of faunal exploitation are rare all along the Mous-
terian sequence of the Riparo Mochi. Without consider-
ing the abundant burnt bone fragments, anthropogenic 
traces are predominantly percussion marks; only one 
scraping mark, from the deeper spits, was recorded 
(Tab. 5). The skeletal representation observed in Unit I 
seems to indicate the arrival of the whole carcass at the 
site. Both medium, medium-large and large animals 
(Fig. 6) present anatomical elements from all anatomical 
portions, with a large presence of the cranial skeleton. 
Regarding the skeletal frequency of the most represent-
ed species (Fig. 9), we see a lower presence of anatom-
ical elements in the Ic level compared to Ib and It. Thus, 
there does not appear to be a selection of specific ana-
tomical elements in all three mousterian phases. 

The Neanderthals who frequented Riparo Mochi 
had at their disposal a wide variety of prey, from large 
game such as horse to smaller game such as wild boar, 
the latter being extremely abundant according to the 
MNI in Unit I (Tab. 4). It does not seem to be possible to 
extrapolate data on a seasonal occupation of the site. 
The mere presence of young boar individuals (Tab. 4) 
does not allow to define a seasonality of death due to 
the animal's biological cycle, which doesn't give birth in 
a single time of the year (Mauget, 1981). In addiction, 
the sole presence of the fragments of cervid antlers, 
although distributed throughout the sequence, did not 
allow a seasonality of occupation of the site to be de-
fined. Exploitation seems have a focus on Cervus ela-
phus, whose remains show the highest number of im-
pact marks observed in the Mousterian unit. Further 
impact marks are observed mainly on the long bones of 
medium-large size mammals. As red deer belong to this 
size class, its exploitation by Neanderthals may be un-
derestimated. The high presence of Cervus elaphus in 
the archaeological assemblage is a very common fea-
ture of Middle and Upper Pleistocene sites in the Liguri-
an region. In coeval sites such as Riparo Bombrini, Ar-
ma delle Manie, and San Francesco, red deer is the 

main species among ungulates (Isetti, 1961; Valensi, 
2000; Psathi, 2003; Valensi & Psathi, 2004; Holt et al., 
2019) (Fig.1). A possible reason for this could be its high 
environmental adaptability, with a mix-feeder diet, plastic 
in its choice of habitats (Straus, 1981; Sommer et al., 
2008), and the better “economical return” that this spe-
cies could offer in terms of quality and quantity of the 
materials obtained, ease of hunting and/or transport 
(Valensi & Psathi, 2004). The Neanderthal economy at 
Riparo Mochi therefore seems to have focused on big 
game, a factor that seems to be common to coeval sites 
of Ligurian region. We also note the paucity of exploita-
tion of small faunas such as birds or rodents; although it 
cannot be ruled out that such remains have not survived 
due to the intense taphonomic processes that character-
ise the site, this scanty evidence could be speculatively 
related to a possible distinctive cultural-economic differ-
ence between the Tyrrhenian and Po-Adriatic regions. 
Several sites of Adriatic region provide evidence of the 
exploitation of both avifauna (Fiore et al., 2004, 2016; 
Peresani et al., 2011; Romandini et al., 2014) and small 
fauna (Alhaique et al., 2004; Romandini et al., 2018). 
Currently, only at the French Pié-Lombard site (Pelletier 
et al., 2019), geographically close to Riparo Mochi and 
integrated into the Ligurian-Provençal region, provided 
Mousterian levels - dated to the MIS 5-4 transition - 
showing the exploitation of both avifauna and small 
mammals (specifically Oryctolagus cuniculus). 
 
5.2.2. Upper Palaeolithic 

The beginning of the UP is marked by an increase 
of anthropogenic modifications that are mainly concen-
trated on the long bones of medium-large sized mam-
mals; determined remains of Cervus elaphus bearing cut
-marks and percussion marks are also present and, 
similarly to Unit I, the exploitation of red deer could be 
underestimated, a feature which seems to persist during 
the UP. It was not possible to determine a seasonality of 
occupation of the site due to the absence of juvenile 
individuals (Tab. 4) or antler fragments still in connection 
with the skull. Compared to the earlier Mousterian unit, 
the UP units display a more diversified and abundant 
skeletal profile (Fig. 6, Fig. 9). As in the MP, the skeleton 
appears to be complete, suggesting that the carcasses 
entered the site as whole (Fig. 6). Unlike the MP, ele-
ments of the limbs are more abundant. The skeletal 
representation of ungulates such as Cervus elaphus, 
Capra ibex and Capreolus capreolus, shows a higher 
frequency of specific elements such as tibiae and meta-
tapodials (Fig. 9). In general, the posterior appendicular 
skeleton is highly represented, suggesting a selection of 
specific anatomical elements or a higher degree of 
preservation of these. The change in faunal composition 
from Unit I to Unit H-G may be linked to a shift of anthro-
pogenic activity. In fact, if we accept the presence of a 
sterile layer within the former Unit H, see Grimaldi et al. 
(2014), the variation in the faunal spectrum that oc-
curred in Unit G may be mainly caused by a change in 
hunting strategies linked to the arrival of the earliest 
AMHs, rather than by climatic and environmental varia-
tions. For this reason, it cannot be excluded that the 
general increase in the number of remains belonging to 
cervids (red deer, roe deer or other cervids) is evidence 

 
 
The faunal assemblage from the Riparo Mochi site.  



   

 

 
 

150 

of a selection of prey which becomes more evident in 
Unit F where the variety of the faunal spectrum is lower. 
To support this hypothesis, the finding of antler industry 
in Early Aurignacian Unit F (Tejero & Grimaldi, 2015), 
specifically related to the production of split-based 
points, should be remarked. The production of such 
tools, namely hunting-oriented specialised tools, may 
have largely influenced the selection of prey, also useful 
for obtaining the non-lithic raw-material. In addition, the 
increase of Capra ibex and Capreolus capreolus at the 
expense of species such as Equus sp. and Sus scrofa, 
which are widely present in the Mousterian levels, could 
testify the change in hunting strategies that occurred 
during the UP. Interestingly, from Unit G to Unit E, more 
distinctly alpine taxa, such as Capra ibex and Rupicapra 
rupicapra, are more abundant than in Unit I. The AMHs 
inhabitants of Riparo Mochi may have preferred to hunt 
prey at higher altitudes than the Neanderthals, probably 
facilitated by a milder climate that made more accessi-
ble the mountainous areas close to the site. Evidence of 
hunting focused on alpine species can also be observed 
in the upper Unit D. Although surface preservation is as 
'bad' as that observed in the Unit I to E sequence, the 
Gravettian levels have yielded numerous traces of ex-
ploitation of ungulates such as red deer and ibex that 
seem to suggest the persistence of exploitation of 
mountain faunas (Tagliacozzo et al., 2012) 

Cervus elaphus and Capra ibex are the most ex-
ploited species, together with other ungulates such as 
Capreolus capreolus, Rupicapra rupicapra and Bos 
primigenius. Unlike what was observed in the Unit I-E 
sequence, in Unit D anthropogenic traces are found on 
the remains of carnivores such as Vulpes vulpes, Felis 
silvestris and Lynx lynx. These traces are attributable to 
skinning activities and could indicate a further variation 
in the hunting spectrum also linked to the obtaining of 
specific resources. A more in-depth study of the an-
thropic exploitation of preys in the Aurignacian levels will 
shed light on possible similarities and divergences be-
tween Unit F and Unit D. 
 
6. CONCLUSIONS 
 

This research allows a more specific environmen-
tal contextualization of Riparo Mochi throughout the 
formation of the Mousterian, Proto-Aurignacian and 
Aurignacian archaeological deposits. Data from the 
Riparo Mochi fit well with the palaeoenvironmental 
framework today available for the Liguro-provençal arc 
during the MIS 4 and 3. Despite rapid climatic fluctua-
tions, the region is characterized by more temperate 
environmental conditions during the MUPT, favouring 
the persistence of a mosaic of environments and rele-
vant biodiversity of large fauna. This heterogeneity can 
be observed as early as the end of MIS 5 (Unit I-base) 
and persists until MIS 3 (Unit F). During the earliest 
phases of the Mousterian, the site was also occasionally 
occupied by carnivores, as evidenced by the finding of 
hyena coprolites. The Neanderthals that inhabited Ri-
paro Mochi, although the presence of a wide range of 
prey, hunted mainly medium-large ungulates, in particu-
lar red deer. With the arrival of AMHs, attested since the 
top of Unit H - base of Unit G, a change in the composi-

tion of the faunal spectrum is observed with a decrease 
in diversity. Nevertheless, Cervus elaphus remains the 
most hunted species; this could be tentatively related to 
an ecological adaptation of the hunters, probably linked 
also to the production of antler tools. Future studies 
such as a fully developed functional analysis linking 
lithics and butchering marks, palynological analysis, 
malacological and microwear dental studies, will be use-
ful for a wider palaeoenvironmental contextualization 
and will provide a better understanding of the different 
human-environment interactions that occurred in the 
archaeological units at the Riparo Mochi. 
 
Author Contributions: A.P. carried out the Riparo Mo-
chi zooarchaeological study as part of his “The exploita-
tion of animal resources in the Upper Pleistocene in 
northern Italy: reconstruction of hunting strategies in the 
Middle and Upper Palaeolithic sequence at Riparo Mo-
chi (Balzi Rossi) and Riparo Tagliente (Monti Lessini)” 
PhD project which was designed and supervised by 
S.G. and U.T.H.; all authors wrote the paper; Riparo 
Mochi research project is directed by S.G under the 
permission of the Italian Ministry of Culture. 
 
ACKNOWLEDGMENTS 

The research was funded by the PhD course 
“European Cultures: Environment, Contexts, Histories, 
Arts, Ideas” of the Department of Humanities, University 
of Trento (Italy). Authors acknowledge the Laboratory of 
Zooarchaeology of MUSE (Trento, Italy) and the Labora-
tory of Large Vertebrate of the University of Ferrara 
(Ferrara, Italy) for providing facilities and the access to 
their paleontological reference collections. 
 
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