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Journal of Archaeological Science 168 (2024) 106007
Available online 29 May 2024
0305-4403/© 2024 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC license (http://creativecommons.org/licenses/by-
nc/4.0/).
Neanderthal hunting grounds: The case of Teixoneres Cave (Spain) and Pi´
e
Lombard rockshelter (France)
Antigone Uzunidis
a
,
b
,
*
, Ruth Blasco
b
,
c
, Jean-Philip Brugal
d
, Tiffanie Fourcade
e
,
f
,
Juan Ochando
g
,
h
, Jordi Rosell
b
,
c
, Audrey Roussel
i
, Anna Ruf`
a
j
,
k
, Maria Fernanda S´
anchez
Go˜
ni
f
,
l
, Pierre-Jean Texier
d
, Florent Rivals
b
,
c
,
m
,
**
a
Universit´
e de Paris 1 Panth´
eon-Sorbonne & UMR 7041 ArScAn-Arch´
eologiesEnvironnementales, Paris, 75006, France
b
Institut Catal`
a de Paleoecologia Humana i Evoluci´
o Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edici W3), 43007, Tarragona, Spain
c
Departament d’Hist`
oria i Hist`
oria de l’Art, Universitat Rovira i Virgili, Av. Catalunya 35, 43002, Tarragona, Spain
d
CNRS, Aix Marseille University, Minist. Culture, UMR 7269 LAMPEA, 13097, Aix-en-Provence, France
e
Arch´
eosciences Bordeaux, UMR 6034, Universit´
e Bordeaux Montaigne – CNRS – EPHE – Universit´
e de Bordeaux, 33607, Pessac Cedex, France
f
Environnements et Pal´
eoenvironnements Oc´
eaniques et Continentaux (EPOC), UMR 5805, Universit´
e de Bordeaux, CNRS, Bordeaux INP, EPHE, Universit´
e PSL,
33600, Pessac, France
g
Department of Plant Biology (Botany Area), Faculty of Biology, University of Murcia, Campus de Espinardo, 30100, Murcia, Spain
h
Department of Environmental Biology, Sapienza University of Rome, Piazzale Aldo Moro 5, 00185, Rome, Italy
i
Universit´
e Paul Val´
ery Montpellier 3, UMR 5140 ASM, Montpellier, France
j
ICArEHB – Interdisciplinary Center for Archaeology and the Evolution of Human Behaviour, Universidade do Algarve, Campus de Gambelas 8005-139 Faro, Portugal
k
PACEA UMR 5199, Univ. Bordeaux, CNRS, MCC, F-33600, Pessac, France
l
Ecole Pratiques des Hautes Etudes, EPHE, PSL University, 75000, Paris, France
m
ICREA, Pg. Lluís Companys 23, 08010, Barcelona, Spain
ARTICLE INFO
Keywords:
Subsistence strategies
Dental microwear
Habitats
Herbivores
Prey
Ungulates
ABSTRACT
The study of Neanderthal-Environment interactions very often lacks precise data that match the chrono-
geographical frame of human activities. Here, we reconstruct Neanderthals’ hunting grounds within three
distinct habitats using dental microwear analysis combined with zooarchaeological data. The predation patterns
toward ungulates are discussed in term of frequency (NISP/MNI) and potential meat intake (MAM). Unit IIIa of
Teixoneres Cave (MIS 3, NE Spain) corresponds to a mosaic landscape, Unit IIIb was more forested, and, in the
“Ensemble” II of Pi´
e Lombard (MIS 4, SE France), forest cover dominated. At Pi´
e Lombard, Neanderthals rely on a
high diversity of taxa from closed and semi-open hunting grounds, mostly two ungulate species as well as rabbits
and several bird taxa. At Teixoneres Cave, mainly open areas are exploited in summer with a predation mostly
focused on large gregarious ungulates. The larger size of ungulate herds in open spaces may have allowed Ne-
anderthals to restrict their subsistence behaviour only to very few species, in specic hunting strategies. In Unit
IIIa, they do not appear to have made any selection within the most abundant species, while in Unit IIIb, they
focused on aurochs and also opportunistically and heavily on newborn red deer. Neanderthal subsistence stra-
tegies seem, therefore, only partially linked to the hunting grounds they had access to. While it impacted the
diversity of the prey they selected, Neanderthal groups were able to develop distinct hunting strategies within
similar environments.
1. Introduction
The subsistence behaviours of human populations during the Middle
Palaeolithic are commonly attributed to the intricate connection
between cultural and biotic factors (Discamps and Royer, 2017; Gaud-
zinski-Windheuser and Roebroeks, 2011; Hardy et al., 2012; Rolland
and Dibble, 1990; Tzedakis et al., 2007; Vettese et al., 2022). The latter
factor is investigated through the reconstruction of environmental
* Corresponding author. Universit´
e de Paris 1 Panth´
eon-Sorbonne & UMR 7041 ArScAn-Arch´
eologiesEnvironnementales, Paris 75006, France.
** Corresponding author. Institut Catal`
a de Paleoecologia Humana i Evoluci´
o Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edici W3),
43007 Tarragona, Spain.
E-mail addresses: antigone.uzunidis@wanadoo.fr (A. Uzunidis), orent.rivals@icrea.cat (F. Rivals).
Contents lists available at ScienceDirect
Journal of Archaeological Science
journal homepage: www.elsevier.com/locate/jas
https://doi.org/10.1016/j.jas.2024.106007
Received 12 March 2024; Received in revised form 22 May 2024; Accepted 23 May 2024
Journal of Archaeological Science 168 (2024) 106007
2
parameters using a wide range of proxies such as pollen from long
terrestrial and deep-sea sedimentary sequences (e.g., Pons and Reille,
1988; S´
anchez Go˜
ni et al., 2008), pollen from archaeological sequences
(Carri´
on et al., 2018, 2022a, 2022b; Ochando et al., 2020a, 2020b,
2022), or small mammals (e.g., Discamps and Royer, 2017; Fern´
an-
dez-García et al., 2022; L´
opez-García et al., 2021). The accuracy of the
correlations suggested by these proxies between the reconstructed en-
vironments and those experienced by humans can greatly vary
depending on their resolution. Indeed, the chronological framework
they address (millennial to yearly) never corresponds fully to the period
of the site occupation by human groups. Likewise, the size of the terri-
tory they cover varies signicantly, ranging from continental, regional
to local scales. Proxies that reconstruct environmental contexts on large
scales may overlook the diversity of local landscapes that
hunter-gatherer groups had access to (Uzunidis and Rivals, 2023).
Neanderthals have exploited a wide range of resources present in
various habitats. Populations that inhabited woodland consumed a
higher proportion of plant food (El Zaatari et al., 2011; Hlusko et al.,
2013), while the ones from cold steppe environments had a higher meat
intake (El Zaatari et al., 2011, 2013; Lalueza-Fox and P´
erez-P´
erez, 1993;
Williams et al., 2018). In terms of predation, the most common prey
found in Neandertal camp, in general, were large and medium-sized
ungulates (Gamble, 1999; Mellars, 1996; Rivals et al., 2022). In South-
ern Europe, these hunts were completed with a signicant acquisition of
small game, while it appears less common in the Northern territories
(Blasco et al., 2022). In the Western Mediterranean region notably, birds
and leporids were commonly preyed upon to access both meat and
pelt/feather (Blasco et al., 2013; Blasco and Fern´
andez Peris, 2009;
Fiore et al., 2020; Martínez Valle et al., 2016; Morin et al., 2019). Thus,
Neandertal dietary and predation habits are strongly dependent on cli-
matic and environmental contexts. Accurate environmental re-
constructions at the local scale are then essential to comprehensively
grasp the impact of biocoenosis on Neanderthal subsistence strategies.
Approaches based on the study of ungulates benet from the strict
contemporaneity between the archaeological site occupation and the
accumulation of prey resulting from hunter-gatherer activities. While
environmental reconstructions based on large mammals can be
misleading due to the high ecological exibility of many of these taxa
(Rivals et al., 2010, 2015a; Rivals and Lister, 2016; Ungar et al., 2020;
Uzunidis, 2021; Uzunidis et al., 2022), approaches based on biogeo-
chemical and dental wear analyses have emerged. These methods enable
the examination of different life stages of the animals, allowing for the
inclusion of their inherent variability (Bocherens et al., 2005, 2016;
Britton, 2018; Britton et al., 2023; S´
anchez-Hern´
andez et al., 2016).
Dental microwear stands out among these methods as a tool capable of
reconstructing an individual’s diet from a few days to a month prior to
its death (Solounias and Semprebon, 2002; Winkler et al., 2020).
Consequently, it provides a way to explore past ecologies, vegetal
structure and to infer paleoenvironments within the biome where un-
gulates were present when and where they were preyed upon.
In this study, our aim is to provide insights into the subsistence
strategies of late Middle Palaeolithic Neanderthals, focusing on three
distinct archaeological units. Specically, we examine Teixoneres Cave
Units IIIa and IIIb (MIS 3, Spain) and Pi´
e Lombard rockshelter (MIS 4,
France), where Neanderthals inhabited environments ranging from
mosaic to forested biome habitats (Fern´
andez-García et al., 2022;
Ochando et al., 2020a; Renault-Miskovsky and Texier, 1980; Texier
et al., 2011; Uzunidis and Rivals, 2023). Our approach involves recon-
structing the hunting grounds selected by Neanderthal groups within
their environment and comparing them to the species and individuals
they targeted as prey. The objective is to assess the diversity of Nean-
derthal subsistence strategies, considering the potential inuence of
environmental contexts and discerning the role of selective choices in
shaping these strategies.
2. Archaeological sites
2.1. Pi´
e Lombard rock shelter
Pi´
e Lombard rock shelter (Tourrettes-sur-Loup, Alpes-Maritimes,
France) is an archaeological site located near the city of Nice (Fig. 1),
discovered in 1962 (Lumley, 1969) and excavated between 1971 and
1986 under the supervision of P.-J. Texier (Texier, 1974; Texier et al.,
2011). It is located at the foot of a rather steep cliff at a relatively low
altitude: 250 m above sea level (a.s.l) in a very rugged area surrounded
by the Mediterranean Sea to the south and the Loup River, along with
mountainous areas exceeding 1000 m in elevation to the east and north.
Throughout the Pleistocene, as it is today, its topographical situation
gives rise to ecotone landscapes and conditions.
Pi´
e Lombard has yielded a 3-m thick archaeological sequence that
was separated into two main stratigraphic complexes, or ensembles
(Texier et al., 2011).
-“Ensemble” I: dated by accelerator mass spectrometry (AMS) to be-
tween 20,639 and 22,073 cal BP (Tomasso, 2014). This “ensemble”
has been attributed to the Epigravettian culture and yielded very few
faunal remains (for more details, see Pelletier et al., 2019).
-“Ensemble” II, identied as Mousterian complex, is dated to a tran-
sitional phase between Marine Isotope Stage (MIS) 5 and 4 based on
a thermoluminescence (TL) date of 70 ±8 kyr BP (Valladas et al.,
1987). Stalagmites underlying the Mousterian sediments yielded two
electron spin resonance (ESR) dates of 147 ±10 kyr and 130 ±20
kyr BP (Yokoyama et al., 1983). The present study focuses on this
ensemble.
Several ungulate species are present at Pi´
e Lombard. Capra ibex and
Cervus elaphus are the main represented large herbivore species while
Bos primigenius, Capreolus capreolus, Equus caballus, and Rupicapra rupi-
capra are less abundant (Texier et al., 2011; Uzunidis et al., 2022;
Uzunidis and Rivals, 2023). Few remains of carnivores were also iden-
tied belonging to Ursus arctos, Felis silvestris, Lynx sp., Panthera pardus,
Canis lupus, and some remains from Vulpes vulpes and Cuon sp. (Texier
et al., 2011). Small game is very well represented, including 225 rabbits
(Oryctolagus cuniculus) and at least 86 individuals from various bird
species (Pelletier et al., 2019; Romero et al., 2017). Zooarchaeological
and taphonomical studies indicate that both ungulates and small game
were exploited for meat, skin, and feathers, while carnivores were not
exploited (Pelletier et al., 2019; Romero et al., 2017; Roussel, 2023).
While there is no evidence of carnivore denning at Pi´
e Lombard, some
bones exhibit marks of carnivore activities suggesting their partial
involvement in the accumulation of the caprid remains (Roussel, 2023).
Additionally, two Neanderthal deciduous incisors were discovered
(Lumley de, 1976; Texier et al., 2011). The Mousterian lithic industry,
produced from various raw materials like int, chert and quartzite,
originates from both local and semi-local origin. Most raw materials
come from the littoral area, with already nished tools abandoned at the
site (Porraz, 2005, 2009). The “ensemble” II of Pi´
e Lombard is inter-
preted as a seasonal camp, with limited carnivore incursions, primarily
dedicated to hunting activities of both small game and ungulates by
Neanderthals (Porraz, 2005, 2009; Roussel et al., 2021; Texier et al.,
2011) that occurred mostly in autumn but also in spring (Roussel, 2023;
Roussel et al., 2021; Uzunidis and Rivals, 2023).
2.2. Teixoneres Cave
Teixoneres Cave, located in Moi`
a (Barcelona, Spain), is part of the
Toll caves karstic system (Fig. 1). It was discovered as an archaeological
site in the 1950s and some intermittent excavation seasons by different
researchers took place until the 1990s. Since 2003, annual excavations
have been conducted by a team from the Institut Catal´
a de Paleoecologia
Humana i Evoluci´
o Social (IPHES-CERCA; Rosell et al., 2008; 2010a;
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
3
2010b, 2014). The Toll caves karstic system is located 780 m a.s.l.
Nowadays, the landscape around Teixoneres Cave corresponds to small
hills and minor streams, with the exception of the Llobregat River to the
south and the Ter River to the north, which border the area (Rosell et al.,
2017), implying a more homogenous environment compared to the
partitioned landscape at Pi´
e Lombard.
Teixoneres cave has yielded a 6-m deep sedimentary sequence
divided into eight stratigraphic units, further categorized into sub-units
(Rosell et al., 2008, 2010b; Zilio et al., 2021). At the top of the sequence,
Unit I has an age of ca. 14–16 ka (Tissoux et al., 2006). Unit II has been
radiocarbon dated between 44,210 and 33,060 cal BP and Unit III was
radiocarbon dated from >51,000 BP to 44,210 cal BP (Talamo et al.,
2016). Unit III, the focus of this study, is the richest unit in Teixoneres.
Two sub-units are identied in Unit III: Unit IIIa, dated between 44,210
and 45,000 cal BP, and Unit IIIb dated to 45,000 cal BP at the top and its
base has an age of >52,000 years BP (Ochando et al., 2020a; Talamo
et al., 2016; Tissoux et al., 2006).
Both sub-units, IIIa and IIIb, exhibit a similar representation of
ungulate species, including Bos primigenius, Capra pyrenaica, Cervus
elaphus, Capreolus capreolus, Equus caballus, Equus hydruntinus, and
Coelodonta antiquitatis. The species diversity is higher in Unit IIIa where
few remains of Rupicapra pyrenaica and Mammuthus primigenius were
identied (´
Alvarez-Lao et al., 2017; Uzunidis et al., 2022; Uzunidis and
Rivals, 2023). Carnivores are present in both sub-units: Ursus spelaeus,
Crocuta crocuta, Canis lupus, Vulpes vulpes, Lynx spelaea, and Meles meles
(Rosell et al., 2017), along with remains of leporids and birds (Ruf`
a
et al., 2014, 2016). Several Neanderthal remains of both juveniles and
adults were found in sub-unit IIIb. Carnivores seem to have occupied the
inner parts of the cave in alternation with Neanderthals who had fav-
oured the porch area (Rosell et al., 2010a, 2017; Ruf`
a et al., 2014, 2016,
2014; S´
anchez-Hern´
andez et al., 2014, 2016; Zilio et al., 2021). Despite
occasional human action, non-human predators are mainly responsible
for the accumulation of small game (Ruf`
a et al., 2014, 2016), while
ungulate bones display evidence of anthropogenic activities (Rosell
et al., 2017). The lithic raw materials are mostly of local origin (Mu˜
noz
del Pozo et al., 2023). The Mousterian industry consists of hunting and
Fig. 1. A: Geographical position of the site of Pi´
e Lombard rockshelter and Teixoneres Cave, B: General view of Pi´
e Lombard rockshelter within its current envi-
ronment (Photo: Pierre-Jean Texier); C: General view of Teixoneres cave within its current environment (Photo: Gabriel Cifuentes-Alcobendas).
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
4
cutting tools used in the early phases of butchery (Picin et al., 2020).
Porch areas of units IIIa and IIIb were interpreted as Neanderthal
hunting camps (Rosell et al., 2017) predominantly occupied during the
summer (S´
anchez-Hern´
andez et al., 2014, 2016, 2020; Uzunidis and
Rivals, 2023).
3. Material and methods
In both sites, specic-level identication of ungulates was success-
fully achieved for some cranial and post-cranial elements. However, the
identication of post-cranial elements is notably limited due to a high
fragmentation rate, particularly in Teixoneres Cave. Consequently, the
analysis heavily relies on dental elements.
In the case of Teixoneres Cave, the studied material exclusively
originates from the outer area of the cave, following the denition by
Zilio et al. (2021). Thus, the analysis was restricted to the anthropic part
of the site.
The material from Pi´
e Lombard material is curated at the Maison
M´
editerran´
eenne des Sciences de l’Homme (Aix-en-Provence, France)
and the Teixoneres Cave collection is located at the Institut Catal`
a de
Paleoecologia Humana i Evoluci´
o Social (Tarragona, Spain).
3.1. Quantication of ungulate representation
The representation of the ungulates in each accumulation was
evaluated using two indices for comparative purposes: Minimum
Number of Individuals (MNI) (Badgley, 1986) and the total Number of
Identied Specimens (NISP) (Lyman, 2008). The NISP was calculated by
considering all elements that were complete enough to secure anatom-
ical and specic identication, including both post-cranial and cranial
remains. The MNI is derived only from dental elements, which were the
best-preserved and most represented. The calculation is based on the
most abundant tooth from either the left or right side and includes age
class attributions. MNI for Teixoneres cave were previously published by
Uzunidis and Rivals (2023) and for Pi´
e Lombard it was re-evaluated
since the studies of Uzunidis and Rivals (2023), Roussel (2023) and
Pelletier et al. (2019).
Both NISP and MNI are indicative of Neanderthal predation toward
precise individuals. It can blur, however, the potential contribution of
each of them to meat intake. Thus, the representation of each species
regarding their meat mass was estimated following several steps.
1) MNI was calculated for each cohort and individuals were categorized
into two age groups (for the methodology associated to age estima-
tion, see below): ‘newborn’ and ‘grown’. The ‘newborn’ group
comprises individual aged between 0 and 4 months, while the
‘grown’ group includes all other individuals. The distinction between
these age groups aims to account for potential differences in meat
quantity related to age-biased ungulate accumulation. Juveniles
older than 4 months and sub-adult individuals are classied with the
adults in the ‘grown’ group. Indeed, those age classes were very
scarce in our samples and growing processes can vary signicantly
between populations according to several factors that are difcult to
assess in Pleistocene studies (see for example Gidney, 2013; Poppi
et al., 2018). As a consequence, the weight of growing individuals
(juveniles older than 4 months and subadults) was over-estimated.
2) The body mass of the ‘grown’ age-group was estimated using Janis
(1990) equations based on measurements from the second and third
lower molars taken at the collar. In one case, for B. primigenius of Pi´
e
Lombard, since no permanent teeth were available to estimate the
body mass, we used the transversal diameter of the metatarsal taken
in the middle of the diaphysis and used the equations from Scott
(1990). For each cohort, a mean mass was calculated by averaging
the results.
3) The body mass of the ‘newborn’ age group was calculated using the
mean mass of the ‘grown’ age group from each cohort and the
equation published by Traill et al. (2021).
4) For the large ungulates, the proportion of meat available on carcasses
was considered to be 60% of the live mass (Brugal, 2005; Lyman,
1979). This proportion was calculated both for the ‘newborn’ and
‘grown’ age groups.
5) The meat mass obtained for both the ‘newborn’ and the ‘grown’ age
groups was multiplied by the MNI obtained for the 0–4 months age
class and for the others, respectively.
This method presented provides a means to estimate the meat
available from each archaeological cohort of ungulates. However, it
operates under the assumption that all ungulates from Pi´
e Lombard and
the outer part of Teixoneres cave were preyed upon by Neanderthals.
While most evidence supports the idea that these assemblages were
primarily accumulated by Neanderthals (Pelletier et al., 2019; Rosell
et al., 2017; Roussel, 2023; Ruf`
a et al., 2014, 2016), other occasional
contributions from other predators cannot be entirely ruled out. Also,
the sedimentary layers likely encompass several distinct occupation
periods over time, but are concatenated here within each assemblage:
‘Ensemble’ II of Pi´
e Lombard and Units IIIa and IIIb of Teixoneres.
Additionally, the method does not account for potential selection of
specic body parts that may have occurred at the kill site. Indeed, it
assumes that the entire carcass contributed to the Neanderthal diet.
Also, as explained in the methodology above, the weight of the juvenile
individual above 4 months and of the subadult was purposely over
estimated. Therefore, this approach should be interpreted as an
approximation of the Maximum Available Meat (MAM) from the un-
gulate cohorts, acknowledging its limitations and assumptions.
3.2. Estimation of the age at death
The estimation of the age at death of the ungulates was conducted
based on the identication of dental remains, including isolated teeth for
equids and cervids, as well as teeth on maxilla and mandibles for the
other taxa. The age estimation for red deer and equids involved the use
of specic equations. We followed the work of Steele (2004) to establish
the age of red deer based on the fourth lower decidual tooth and the rst
lower molar. For equids, we employed the equations published by Fer-
nandez (2009). Both methods provide an estimation of the individual
age associated with a certainty rate. We include in this study only the
teeth with a certainty rate above 85%. For both taxa, unworn or very
slightly worn decidual teeth were included into the 0–4 month age
group. For the other taxa, we used methods based on tooth eruption and
wear patterns: B. primigenius (Brugal and David, 1993), Capra sp.
(Pailhaugue, 2003; P´
erez Ripoll, 1988), Rupicapra sp. (Pailhaugue,
1998) and Capreolus capreolus (Paulus, 1973).
3.3. Dental microwear
Dental microwear involves studying the microscopic marks on tooth
occlusal surfaces, which are produced by food and masticatory pro-
cesses. In herbivores, these dietary micro-traces, such as scratches and
pits, result from the presence of phytoliths (Rivals et al., 2015b; Walker
et al., 1978). The quantication of these traces provides insights into
three dietary types: grazers (consuming grass), browsers (consuming
ligneous and herbaceous plants), and mixed-feeders (consuming both
categories seasonally or not). When applied to a diverse range of her-
bivorous taxa from the same site, dental wear analysis can indicate the
contribution of dicots and monocots to the environment (Mainland,
1998, 2000; Semprebon et al., 2004b; Solounias and Semprebon, 2002;
Uzunidis and Rivals, 2023). Dental microwear helps reconstruct the diet
of an individual a few days before its death (Hoffman et al., 2015;
Teaford and Oyen, 1989; Winkler et al., 2020). Indeed, it serves as a
valuable proxy for discussing the environmental setting of Neanderthal
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
5
groups at the moment of their prey acquisition (Uzunidis and Rivals,
2023).
The study followed the protocol established by Solounias and Sem-
prebon (2002) and Semprebon et al. (2004a). The occlusal surface of
each tooth was cleaned using acetone, then 96% ethanol, followed by
moulding using high-resolution silicone (vinylpolysiloxane). Trans-
parent casts were created using epoxy resin and observed with a ste-
reomicroscope at x35 magnication. The microwear features were
identied and quantied on a standard surface of 0.16 mm
2
using an
ocular reticule, preferably on the upper tooth paracone and the lower
tooth protoconid, excluding the second and third premolars (Xas et al.,
2017). Individuals deemed too old or too young individuals and teeth
bearing taphonomic damages were discarded (Fortelius and Solounias,
2000; King et al., 1999; Rivals et al., 2007). Microscopic taphonomical
defects were also discarded following the morphological descriptions of
Mic´
o et al. (2023); Uzunidis et al. (2021). For the identication of the
microwear features, the classication of Solounias and Semprebon
(2002) and Semprebon et al. (2004a) was followed, quantifying pits
(small and large), scratches (ne, coarse and hypercoarse), and gouges.
The scratch width score (SWS) was also estimated, giving a score of ‘0’ to
teeth with predominantly ne scratches, ‘1’ to those with mixed ne and
coarse scratches, and ‘2’ to those with predominantly coarse scratches.
Microwear features are compared to a referential of present-day herbi-
vores with known diet (Solounias and Semprebon, 2002).
4. Results
4.1. Representation of the species in Teixoneres Cave Units IIIa/IIIb and
Pi´
e Lombard
In Teixoneres Cave Unit IIIb, C. elaphus represents 55.88% of the
NISP and 52.94% of the MNI (Table 1; Fig. 2). E. caballus and
C. capreolus follow as the second most represented species, accounting
for 14.57% and 12.17% of the NISP and 13.23% and 11.76% of the MNI,
respectively. B. primigenius and C. pyrenaica (NISP =6.79%; MNI =
10.29% and 5.88%, respectively) are slightly more represented than
E. hydruntinus (NISP =3.79%; MNI =5.88%). A signicant part of the
C. elaphus cohort comprises very young individuals below 4 months
(Table 2; Fig. 3), strongly inuencing the MAM. B. primigenius domi-
nates the MAM with 50.72%, followed by C. elaphus at 27.6%. The other
species have a more limited contribution: E. caballus, 10.36%,
E. hydruntinus, 6.73%, C. pyrenaica, 2.96% and C. capreolus, 1.60%
(Table 1; Fig. 2).
In Teixoneres Cave Unit IIIa, E. caballus and C. elaphus represent
42.35% and 35.29% of the NISP and 39.28% and 28.57% of the MNI,
respectively (Table 1; Fig. 2). They are the most abundant species in this
unit followed by Bos primigenius and E. hydruntinus, representing 12.94%
and 2.35% of the NISP, respectively, and both accounting for 10.71% of
the MNI. Finally, C. pyrenaica, C. capreolus and R. pyrenaica are relatively
scarce, each comprising 2.35% of the NISP and 3.57% of the NMI. In
terms of available meat, E. caballus, B. primigenius and C. elaphus
contributed the most to the MAM with 36.8%, 28.37% and 24.56%,
respectively (Table 1; Fig. 2). They are followed by E. hydruntinus
(7.56%), C. pyrenaica (1.04%), R. pyrenaica (1.02%) and C. capreolus
(0.6%).
In Pi´
e Lombard, C. ibex is the main represented species (NISP =
60.84%; MNI =49%), followed by C. elaphus (NISP =32.49%; MNI =
28.3%). The other species are less common with R. rupicapra (NISP =
4.6%; MNI =11.32%), B. primigenius (NISP =1.13%; MNI =3.77%),
C. capreolus (NISP =0.47%; MNI =3.77%) and E. caballus (NISP =
0.37%; MNI =3.77%) (Table. 1; Fig. 2). C. elaphus and C. ibex contribute
most to the MAM representing 52.69% and 31.16%, respectively, fol-
lowed by B. primigenius (8%), E. caballus and R. rupicapra (3.74%) and
C. capreolus (0.63%).
4.2. Diet of the ungulates at Teixoneres Cave Units IIIa and IIIb and Pi´
e
Lombard at the moment of death
In Teixoneres Cave Unit IIIb, B. primigenius, C. elaphus and E. caballus
exhibit a relatively high number of small pits and scratches, the presence
of large pits, a mixed scratch texture and an absence of gouges (Fig. 4;
Table 3). Equus hydruntinus displays the same characteristics but lacks
large pits. Their diet is therefore considered abrasive and falls within the
range of the extant mixed-feeders. Capra pyrenaica and C. capreolus are
characterized by a low number of scratches and a ne scratch texture,
indicative of a soft diet. The number of pits for C. capreolus is lower
compared to C. pyrenaica, which could suggest the ingestion of dust for
the latter, while the former may have focused on leaves (Semprebon
et al., 2011; Uzunidis et al., 2022). Both species fall within the con-
dence ellipse of extant browsers.
In Teixoneres Cave Unit IIIa, the species exhibit similar dietary
patterns to those in unit IIIb, with B. primigenius, C. elaphus, E. caballus
and E. hydruntinus showing an abrasive diet and falling within the
mixed-feeder group, while C. pyrenaica and C. capreolus fall within the
condence ellipse of the extant browsers (Fig. 4; Table 3).
In Pi´
e Lombard, E. caballus and B. primigenius exhibit a high number
of scratches and a low number of pits, falling within the condence el-
lipse of the extant grazers (Fig. 4; Table. 3). Cervus elaphus is charac-
terized by moderate values for both pits and scratches, along with the
scarce presence of gouges and large pits, indicating a mixed-feeding diet.
Capreolus capreolus, R. rupicapra and C. ibex display few scratches and
pits, as well as a ne scratch texture, suggesting a soft browsing diet
Table 1
Representation of the species from Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard in term of NISP (number of identied specimens), MNI (minimum number of
individuals) and MAM (maximum of available meat, in kg).
Site/species Teixoneres Cave - IIIa Teixoneres Cave - IIIb Pi´
e Lombard
NISP MNI MAM NISP MNI MAM NISP MNI MAM
B. primigenius 11 3 954 34 7 2731.76 12 2 317.86
% 12.94 10.71 28.37 6.79 10.29 50.72 1.13 3.78 8
Capra sp. 2 1 35 34 4 159.61 649 26 1235.26
% 2.35 3.57 1 6.79 5.89 2.96 60.94 49 31.16
Rupicapra sp. 2 1 34.45 0 0 0 49 6 148.36
% 2.35 3.57 1 0 0 0 4.6 11.32 3.74
C. elaphus 30 8 826 280 36 1486.86 346 15 2088.42
% 35.29 28.57 24.56 55.89 52.94 27.6 32.49 28.3 52.69
C. capreolus 2 1 20.26 61 8 86.3 5 2 25.16
% 2.35 3.57 0.6 12.17 11.76 1.6 0.47 3.78 0.63
E. caballus 36 11 1238.4 73 9 558.34 4 2 148.62
% 42.35 39.28 36.83 14.57 13.23 10.37 0.37 2.78 3.75
E. hydruntinus 2 3 254.26 19 4 362.58 0 0 0
% 2.35 10.71 7.56 3.79 5.89 6.73 0 0 0
Total 85 28 3362.5 501 68 5385.46 1065 53 3963.67
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
6
focused on leaves (Semprebon et al., 2011).
5. Discussion
5.1. Habitats of the ungulates from Teixoneres Cave Units IIIa and IIIb
and Pi´
e Lombard
Teixoneres IIIb (~45–100 ka): studies on ungulate dental mesowear
and micromammals suggest a substantial amount of forest cover
(Fern´
andez-García et al., 2022; L´
opez-García et al., 2012a; Uzunidis and
Rivals, 2023). Dental mesowear also indicates the presence of dicot
meadows with few patches of grasslands (Uzunidis and Rivals, 2023).
According to pollen analysis from the cave, the forest cover is still
substantial but less developed compare to Unit IIIa, with a percentage of
arboreal pollen (AP) of 60% (Ochando et al., 2020a). Strong environ-
mental variations occurred during the accumulation of Unit IIIb at
Teixoneres Cave since it encompasses a wider period than Unit IIIa. For
this timeframe, the regional forest cover appears to generally decrease
from 60 ka to 48 ka, based on the decrease of the temperate forest pollen
percentage from 59 to 6% documented by the deep-sea pollen record
from the Gulf of Lion (Fourcade, 2022; S´
anchez Go˜
ni et al., 2020). At the
moment of their death, B. primigenius, C. elaphus, E. caballus, and
E. hydruntinus had a mixed-feeding diet (Fig. 4; Table 3). They fed on
both grass and ligneous plants, indicating the exploitation of an ecotone
at the transition between forest and grassland or of a dicot meadow. The
diet of C. pyrenaica and C. capreolus is focused on soft ligneous plants,
suggesting their presence in forested areas.
Teixoneres IIIa (45,000– 44.210 cal BP): the general environmental
context is characterized by the presence of both forested areas and
grassland in the region of the site, as indicated by dental mesowear
analysis (Uzunidis and Rivals, 2023). This description is further sup-
ported by studies on micromammals (Fern´
andez-García et al., 2022;
L´
opez-García et al., 2012a). Climatically, Unit IIIa experienced cooler
and more humid conditions compared to Unit IIIb (Fern´
andez-García
et al., 2022; L´
opez-García et al., 2012a, 2012b). The pollen analysis of
this unit, however, attests to a signicant forest cover with more than
87% of arboreal pollen (AP) (Ochando et al., 2020a). Regional pollen
data from the deep-sea sedimentary sequence MD99-2343 (40◦29
′
N,
4◦01
′
E, collected in the Gulf of Lion and partially fed by the terrestrial
sediments transported by the Rhˆ
one and Ebro rivers) indicates that the
pollen percentage of the temperate forest was 23% and the percentage of
the grassland was 55 % at ca. 45.1 ka, contemporaneous with the for-
mation of Unit IIIa. At 43.9 ka, the forest percentage was 7% and the
grassland one was 66 % (Fourcade et al., 2024; Fourcade, 2022; Four-
cade et al., 2024; S´
anchez Go˜
ni et al., 2020). These data indicate that
forested and grassland areas co-existed at a regional scale. Overall, the
Fig. 2. Comparison of the representation of the ungulates from Teixoneres Cave Units IIIa (A) and IIIb (B) and Pi´
e Lombard (C) based on the percentage of the NISP
(number of identied specimens), MNI (minimum number of individuals) and MAM (maximum of available meat, in kg).
Table 2
Minimum number of individuals per age group of the ungulates from Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard. The total number of individuals within the
juvenile age class is given and, among them, the number of individuals between 0 and 4 months that correspond to the “Newborn” age group. The mass of available
meat (in kg) per species for one carcass for the age group ‘newborn’ and ‘grown’ is also given.
Site Species Juvenile adult senile Meat available per carcass (kg)
Total 0–4 months ‘Newborn’ ‘Grown’
Teixoneres Cave - IIIa B. primigenius 1 1 2 27.16 463.42
C. pyrenaica 1 5 35.09
C. elaphus 4 3 4 10.70 158.78
C. capreolus 1 20.26
E. caballus 4 2 5 2 9.45 135.50
E. hydruntinus 1 2 6.71 84.76
R. pyrenaica 1 34.45
Teixoneres Cave - IIIb B. primigenius 3 1 3 1 26.48 450.88
C. pyrenaica 1 3 5.47 39.90
C. elaphus 26 24 9 1 7.96 107.98
C. capreolus 3 2 5 3 16.06
E. caballus 5 5 4 9.06 128.26
E. hydruntinus 3 1 7.03 90.64
Pi´
e Lombard B. primigenius 1 1 18.30 299.55
C. ibex 6 15 5 6.11 47.51
C. elaphus 6 2 8 1 10.72 159.00
C. capreolus 1 1 1 3.31 21.84
E. caballus 1 1 1 9.64 138.98
R. rupicapra 3 1 2 1 4.61 29.67
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
7
forest cover is regionally less important within the Unit IIIa time-span
compared to Unit IIIb. At the moment of their death, B. primigenius,
C. elaphus, E. caballus, and E. hydruntinus had a mixed-feeding diet. Bo-
vids and equids displayed more grazing trends, while cervids showed
more browsing tendencies (Fig. 4; Table 3). The inclusion of grass in
their diet indicates that they occupied grasslands and open areas in the
surrounding habitats of Teixoneres Cave. In contrast, C. capreolus and
C. pyrenaica had a diet focused on soft ligneous plants, suggesting their
presence in the forested areas of the habitat.
Pi´
e Lombard, ensemble II (~70 ka): dental mesowear analysis suggests
a forest-dominated environment with dicot meadows and very few
grasslands (Uzunidis and Rivals, 2023). Studies based on herpetofauna,
Fig. 3. Distribution of the age groups per species in Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard in MNI.
Fig. 4. Bivariate plot of the mean number of pits and scratches for the ungulates from Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard. The error bars correspond
to the standard deviation (±1 SD). The ellipses correspond to the Gaussian condence ellipse (p =0.95) on the centroids of extant grazers and browsers based on the
dataset published by (Solounias and Semprebon, 2002).
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
8
malacofauna, micromammals and avifauna indicate a correlation with a
Mediterranean-temperate forested habitat (Texier et al., 2011). How-
ever, the pollen analysis of the site suggests an open landscape domi-
nated by grassland and a cold and dry climate (Renault-Miskovsky and
Texier, 1980; Texier et al., 2011). In contrast with the pollen analysis of
Pi´
e Lombard, the pollen analysis of the deep-sea core from the Gulf of
Lion shows a high amount of forest cover in the adjacent northwestern
Mediterranean landmasses during MIS 4 (Fourcade, 2022; Fourcade
et al., 2024). At the moment of their death, E. caballus and B. primigenius
had an abrasive diet oriented toward grass, indicating that they occu-
pied the scarce open areas in the vicinity of the site (Fig. 4; Table 3). The
diet of C. elaphus falls within the variability of the mixed-feeders with
few scratches, allowing them to be characterized as browse-dominated
mixed-feeders. The topography around the site implies an ecotone
zone with inuences both from the valleys and the mountainous areas.
The habitat was mostly forested with some grassy areas or a meadow
rich in dicot plants. Capreolus capreolus, C. ibex, and R. rupicapra were
browsers that fed on ligneous plants in the forested aeras.
5.2. Neanderthal environmental exploitation for prey acquisition in
Teixoneres Cave Units IIIa-IIIb and Pi´
e Lombard
Teixoneres IIIb: the most represented species is C. elaphus in term of
NISP and MNI. The MAM is dominated by B. primigenius (Fig. 2). The
most important species for Neanderthal subsistence strategy, therefore,
occupied open or semi-open landscape areas (Fig. 5B, 5B
′
, 5B
″
).
Bos primigenius are mainly adults and also few juvenile and senile
individuals (Fig. 3). Cervus elaphus are mainly represented by juvenile
individuals, most of them under 4-month-old. This pattern is known in
other Mousterian sites from the Iberian Peninsula, such as Abric Romaní
level I (Marín et al., 2017), Cova Beneito levels XII-X (Tortosa et al.,
2002), and Gabasa (Steele, 2004). This type of accumulation has been
interpreted as a specialized hunt (Marín et al., 2017), where human
groups took advantage of predictive doe behaviour with newborn red
deer (Rendu, 2010). In the Unit IIIb, newborns seem to have been
trapped or ambushed while does were generally spared since adults are
not as well represented as the juveniles.
Teixoneres IIIa: C. elaphus and E. caballus were the most numerous
prey species preyed upon by Neanderthals, both in terms of NISP and
MNI. The MAM is distributed between E. caballus, B. primigenius, and
C. elaphus (Fig. 2). Thus, in both cases, the species that most contributed
to Neanderthal subsistence strategies inhabited open landscapes, prob-
ably grasslands (Fig. 5A, A
′
, 5A
″
).
Mainly adults B. primigenius and C. elaphus are represented, with few
juvenile and no senile individuals (Fig. 3). The Bos primigenius mortality
pattern is quite similar to the one in Unit IIIb while the one for C. elaphus
is very distinct. Equus caballus is represented mainly by adults although
very few senile individuals are present. This age distribution is similar to
that described for instance at level Ja of Abric Romaní (Marín et al.,
2017). Since adult individuals are the main represented age class in a
herbivore living population, hunter-gatherers will capture them more
frequently (Bunn and Gurtov, 2014). The age distribution of the main
ungulates from Unit IIIa is consistent with a non-selective hunt by
Neanderthal groups based on age criteria.
Pi´
e Lombard, ensemble II: C. ibex and C. elaphus are the most numerous
ungulates in NISP and MNI, and they are also the two species that
contributed the most to the MAM (Fig. 2). Here, the species that were
most frequently preyed upon occupied forested and semi-open
Table 3
Summary of dental microwear data of the ungulates from Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard. Abbreviations: n =Number of specimens; NP =Mean
number of pits; NS =Mean number of scratches; %LP =Percentage of specimens with large pits; %G =Percentage of specimens with gouges; SWS =Scratch width
score; %HC =Percentage of specimens with hypercoarse scratches; %XS =Percentage of specimens with cross scratches; n =number of specimens; m =mean; sd =
standard deviation.
Site/species n Microwear
NP NS %LP %G SWS %HC %XS
Teixoneres Cave Unit IIIa
B. primigenius 1 19.50 21.50 100.00 0.00 1.00 0.00 100.00
C. pyrenaica 1 34.00 6.00 100.00 0.00 0.00 0.00 0.00
C. elaphus m 14 24.96 17.64 78.57 0.00 0.71 7.14 64.28
sd 11.54 2.75
C. capreolus m 2 8.75 5.00 50.00 0.00 0.00 0.00 50.00
E. caballus m 31 25.18 22.48 74.19 3.22 0.74 6.45 77.19
sd 8.63 2.50
E. hydruntinus m 5 18.20 22.00 0.00 0.00 0.60 0.00 60.00
sd 9.02 7.20
Teixoneres Cave Unit IIIb
B. primigenius m 12 26.25 18.46 25.00 16.67 1.00 16.67 50.00
sd 9.00 6.84
C. pyrenaica m 6 27.58 8.75 33.33 0.00 0.67 16.67 16.67
sd 11.48 1.37
C. elaphus m 62 22.87 19.57 74.19 17.74 0.91 14.51 78.68
sd 5.70 4.00
C. capreolus m 12 20.66 10.25 16.67 0.00 0.83 0.00 8.33
sd 10.27 2.64
E. caballus m 33 23.71 20.8 42.42 0.00 0.84 6.00 54.54
sd 7.93 4.34
E. hydruntinus m 14 24.43 20.21 42.86 0.00 0.86 14.28 57.14
sd 5.62 5.94
Pi´
e Lombard
B. primigenius 1 5.00 21.00 0.00 0.00 0.00 0.00 100.00
C. ibex m 29 13.86 8.96 20.69 10.34 0.48 0.00 72.42
sd 3.41 2.92
R. rupicapra m 6 14.08 6.50 16.67 83.3 0.17 0.00 83.30
sd 4.36 1.41
C. elaphus m 30 15.43 15.70 16.67 23.33 0.84 3.33 70.00
sd 5.12 2.82
C. capreolus m 2 14.00 7.50 50.00 0.00 0.50 0.00 50.00
E. caballus m 2 10.75 20.75 50.00 50.00 1.00 50.00 0.00
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
9
landscapes (Fig. 5C, C
′
, 5C
″
). The age distribution follows the same
pattern as the main ungulates from Teixoneres Cave Unit IIIa, indicating
an opportunistic hunting strategy.
5.3. Predation pattern and selective choice
In the two units of Teixoneres Cave, Neanderthals primarily exploi-
ted open environments to acquire ungulates. While other species lived in
this habitat, only some of them are well represented at the site:
C. elaphus, B. primigenius and E. caballus in Unit IIIa; and only the rst
two in Unit IIIb. Equus hydruntinus is consistently rare both in terms of
frequency and meat available despite having also inhabited open land-
scapes. The scarcity of this species could be attributed to several reasons:
(1) It might be indicative of human choices that may have avoided this
species for cultural reasons; (2) E. hydruntinus may have not been
abundant in Western Europe (¨
Ozkan et al., 2023; Uzunidis et al., 2024),
and thus, the small amount of hunted individuals in Teixoneres Cave
may reect its weak representation in the environment; or (3)
E. hydruntinus may have exhibited an erratic behaviour compared to
gregarious animals, making it more difcult to hunt (Burke et al., 2003).
In Unit IIIb, E. caballus is not as well represented as in Unit IIIa despite
having inhabited open landscape, as observed for C. elaphus and
B. primigenius. This could reect a reduction in the horse population in
relation with the habitats that were more forested in Unit IIIb compared
to Unit IIIa as suggested by dental mesowear and micromammals ana-
lyses (Fern´
andez-García et al., 2022; L´
opez-García et al., 2012a; Uzu-
nidis and Rivals, 2023). Horses, however, are very exible and can adopt
a mixed feeding and even a browsing diet to adapt to seasonal changes
and various habitats (Rivals et al., 2009; Rivals and Lister, 2016; Strani
and DeMiguel, 2023; Uzunidis, 2021). It could, therefore, reect a se-
lective choice by human groups. Otherwise, since most of the deposits of
both units occurred in summer (S´
anchez-Hern´
andez et al., 2014, 2016,
2020; Uzunidis and Rivals, 2023), the scarcity of equids may reect
seasonal movements and their absence around the site during the
occupation of Unit IIIb.
In Pi´
e Lombard, Neanderthals exploited forested and semi-open bi-
omes for ungulate acquisition within a partitioned environment of cliff,
plateau and valley oor. They focused on C. elaphus from semi-open
habitats and C. ibex from forested ones in term of NISP, MNI and
MAM. While having very similar dietary characteristics to C. ibex,
Fig. 5. Bivariate plot of the mean number of pits and scratches of the ungulates from Teixoneres Cave Units IIIa (A) and IIIb (B) and Pi´
e Lombard (C). The error bars
correspond to the standard deviation (±1 SD). The ellipses correspond to the Gaussian condence ellipse (p =0.95) on the centroids of extant grazers and browsers
based on the dataset published by Solounias and Semprebon (2002). The size of the circle for each cohort represents its frequency in terms of number of identied
specimens (NISP), minimum number of individuals (MNI) and of maximum of available meat (MAM) as they are indicated in Table 1.
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
10
C. capreolus and R. rupicapra were not as frequently preyed upon. This
last species is often considered challenging to hunt and more commonly
found in sites corresponding to occupations by anatomically modern
humans (Freeman, 1973; Gamble, 1995; Straus, 1987). Recent studies,
however, have shown that Neanderthal groups were able to hunt this
species (Rodríguez-G´
omez et al., 2022; Yravedra and Cobo-S´
anchez,
2015). Capreolus capreolus is also regularly represented in Neanderthal
sites (Gaudzinski-Windheuser and Roebroeks, 2011; Livraghi et al.,
2022; Romandini et al., 2023; Terlato et al., 2021). Pi´
e Lombard un-
gulate accumulation occurred mainly in autumn (Roussel, 2023; Roussel
et al., 2021; Uzunidis and Rivals, 2023) which can lead to
seasonally-biased accumulation. The scarcity of species can, therefore,
reect either species selection by Neanderthal groups or periodical
absence and presence in the vicinity of the site due to seasonal
movements.
In the open-area hunting grounds corresponding to the two Teix-
oneres Units, Neanderthal subsistence strategies relied on a few species.
In this site, two to three taxa, including red deer, horse and aurochs,
constitute the main basis of the meat intake. The presence of large
mammal herbivores organized in herds tend to increase in open habitats
(Evans, 1979; Geist and Walther, 1974; Gerard et al., 2002; Jarman,
1974; Lagory, 1986; Leuthold, 1970), impacting hunter-gatherer choice
and prey selection. In open habitats, other Neanderthal groups have also
specialized their hunting practices toward large ungulates, such as large
bovine (Bos/Bison) or cervids (Cervus/Rangifer) (Berlioz et al., 2023;
Gaudzinski-Windheuser and Roebroeks, 2011). While occupying Units
IIIa and IIIb at the same season and targeting large ungulates within an
open environment, Neanderthal groups did not develop similar strate-
gies. In Unit IIIa, they focused on three species without targeting a
specic age group when in Unit IIIb they focused mostly on one species
(B. primigenius) and supplemented their meat intake by opportunistically
preying on newborn red deer. At this stage, more information con-
cerning occupation patterns and butchering activities is needed to sug-
gest hypotheses explaining such differences in subsistence strategies.
In the forested hunting ground around Pi´
e Lombard, Neanderthal
predation toward ungulates was primarily focused on C. elaphus and
C. ibex. Beside the forested environment not being suitable for large
herds, the scarcity of B. primigenius and E. caballus could be related to the
topographic situation of Pi´
e Lombard. Its position on a cliff in a moun-
tainous area may not be suitable for aurochs or horse herds either. The
main preyed upon taxa, however, in terms of NISP/MNI were the rabbits
(MNI =225) and several bird species (MNI =86) (Pelletier et al., 2019;
Texier et al., 2011). Both bear anthropic marks, and a study on the rabbit
remains revealed that they were targeted for both their skin and their
meat (Pelletier et al., 2019). If rabbit contribution to the meat intake
remains limited (7%), they were far more frequently eaten than the
ungulates. The higher diversity of the species exploited in Pi´
e Lombard is
related to the forest cover, preventing access to large herds. In other
forested landscape, the exploitation of other small game species (i.e.
turtles, rabbits or beavers) is documented, such as in Taubach, Weimar
Parktravertin (Gaudzinski-Windheuser and Roebroeks, 2011) or in
Arbreda level G (Kehl et al., 2014; Lloveras et al., 2016).
Moreover, dietary behaviours of Neanderthal populations living in
forested areas includes both meat as well as a high proportion of plant
food (El Zaatari et al., 2011; Hlusko et al., 2013). The diversity of the
species preyed upon, including large mammals, birds and rabbits, im-
plies subsistence strategies relying on multiple resources. In mixed
habitats, the proportion of plants in the Neanderthal diet is lower, and,
nally, in open habitats, the Neanderthal diet appears more restricted to
meat consumption (El Zaatari et al., 2011, 2013; Harvati et al., 2013;
Lalueza-Fox and P´
erez-P´
erez, 1993; Williams et al., 2018, 2019). Food
acquisition practices may have then been very oriented toward a narrow
range of resources. The toolkits from Teixoneres and Pi´
e Lombard
exhibit typical features of the Middle Paleolithic, characterized by
dominant discoid and Levallois reduction sequences. Both sites feature a
combination of local and semi-local raw materials. Semi-local raw
materials commonly include nished tools with some retouching ac-
tivities, primarily for tool maintenance. In contrast, local raw materials,
especially in Teixoneres, show a higher proportion of cores and knap-
ping activities. At Teixoneres, tools appear primarily dedicated to
hunting or cutting activities, whereas in Pi´
e Lombard, they were used for
cutting and processing meat and likely plant materials. Across all three
occupations, lithic tools suggest a consumption-based economy rather
than one focused on lithic tool production (Porraz, 2005; Texier et al.,
2011; Picin et al., 2020; Mu˜
noz del Pozo et al., 2023). Despite the
relatively similar habitats, prey, and seasons associated with similar
lithic toolkits and knapping activities, acquisition strategies differ be-
tween Units IIIa and IIIb. Unit IIIa shows a more opportunistic approach,
while Unit IIIb is more selective. Despite the relatively similar habitat
occupied by quite similar prey in the same season associated with
similar lithic toolkit and knapping activities, acquisition strategies differ
between Units IIIa and IIIb, with the rst being more opportunistic and
the latter more selective. Thus, Neanderthal subsistence strategies are
greatly diverse although only partially related to the biome they lived in.
While the paleoethology of the prey related to the environmental con-
ditions, notably the organization in herds, other factors, such as pro-
curement goals or intrinsic factors (cultural), are involved in the prey
acquisition patterns of Neanderthal groups.
6. Conclusions
In Teixoneres Cave Units IIIa and IIIb and Pi´
e Lombard, Neanderthals
exploited distinct hunting grounds. Here, prey acquisition mostly
occurred in an open landscape despite the presence of forested areas
around the cave, while it occurred in a more forested landscape around
Pi´
e Lombard. Distinct procurement strategies can be identied accord-
ing to the foraged biome. In open areas, Neanderthal groups focused on
two or three large ungulate species, while in closer habitats, they
exploited chiey two species of ungulates along with several species of
small game (rabbit and birds), surely reecting their abundance in the
environment. The lowest specic diversity in sites with open-area
hunting grounds can be the consequence of a modication in ungulate
ethology: gregarious species occurring commonly in large herds, while
their herds are reduced in closed landscapes. While focusing on large
gregarious ungulates, Neanderthal groups developed distinct strategies
in the two units of Teixoneres Cave. In Unit IIIa, they did not make any
selection regarding the age of prey. In Unit IIIb, their potential main
meat intakes are based on aurochs of all age groups, but they also
opportunistically and clearly preyed upon newborn red deer. Since the
two units were mainly occupied at the same season (summer) Nean-
derthal groups had access in both case to the same age groups. There-
fore, the differences in their prey acquisition patterns may reect
Neanderthal choices rather than their environmental constraints.
Data availability
Data are available to the link https://dataverse.csuc.cat/dataset.xht
ml?persistentId=doi:10.34810/data734.
Funding
This work was funded by an MSCA individual fellowship (Grant
101024230, EU Horizon, 2020) to AU to benet of the support of the
IRN Taphen (IRN 0871). R.Bl., F.R. and J.R. develop their work within
the project PID2022-138590NB-C41 funded by MCIN/AEI/10.13039/
501100011033/and FEDER, UE, the projects 2021-SGR-01237 and
CLT009/22/000045 funded by the Generalitat de Catalunya, and the
project ERCAdG SCAVENGERS, ref. 101097511 funded by the European
Union. R.Bl. is supported by a Ramon y Cajal research contract by the
Spanish Ministry of Science and Innovation (RYC2019-026386-I). This
work contributes to the “María de Maeztu” Program for Units of Excel-
lence of the Spanish Ministry of Science and Innovation awarded to the
A. Uzunidis et al.
Journal of Archaeological Science 168 (2024) 106007
11
Institut Catal`
a de Paleoecologia Humana i Evoluci´
o Social (CEX2019-
000945-M). A.Ru. is beneciary of a CEEC - 3rd Edition research con-
tract promoted by the Portuguese FCT (reference: 2020. 00877.CEE-
CIND), and participates in the Spanish MICIIN projects PID2020-
114462 GB-I00, and the Generalitat de Catalunya projects CLT009/22/
00044 and CLT009/22/00024. The work of JO was funded by the Eu-
ropean Union – NextGenerationEU (Margarita Salas postdoctoral grant,
Ministerio de Universidades of the Government of Spain). The work of
TF was supported by a doctoral contract (n◦12–18, Ecole Doctorale
Montaigne Humanit´
es), the French Research National Agency under the
Investissements d’Avenir Program (ANR-10-LABX-52) and the scientic
support framework of the University of Bordeaux’s IdEx "Investments for
the Future" program/GPR "Human Past".
Authors’ contributions
AU collected the data; designed and conducted the research; wrote
the main manuscript; and prepared all the gures, tables, and supple-
mentary information. FR, RB, ARu were responsible of the excavation in
Teixoneres caves, and PJT of the ones in Pi´
e Lombard. TF, MSG and JO
provided data about terrestrial and deep-core pollen. AR and JPB pro-
vided zooarchaeological information for Pi´
e Lombard fauna. All the
authors have discussed the integration of zooarchaeological, paleoeco-
logical, and palynological data throughout the study. All authors
reviewed, commented and participated in improving the manuscript.
CRediT authorship contribution statement
Antigone Uzunidis: Writing – original draft, Visualization, Valida-
tion, Methodology, Investigation, Funding acquisition, Formal analysis,
Data curation, Conceptualization. Ruth Blasco: Writing – review &
editing, Resources, Project administration, Investigation, Funding
acquisition. Jean-Philip Brugal: Writing – review & editing, Validation,
Project administration. Tiffanie Fourcade: Writing – review & editing,
Investigation, Formal analysis. Juan Ochando: Writing – review &
editing, Formal analysis. Jordi Rosell: Writing – review & editing, Re-
sources, Project administration, Funding acquisition. Audrey Roussel:
Writing – review & editing, Formal analysis. Anna Ruf`
a: Writing – re-
view & editing, Project administration. Maria Fernanda S´
anchez Go˜
ni:
Writing – review & editing, Supervision, Formal analysis. Pierre-Jean
Texier: Writing – review & editing, Project administration. Florent
Rivals: Writing – review & editing, Writing – original draft, Validation,
Supervision, Resources, Project administration, Investigation, Funding
acquisition, Conceptualization.
Declaration of competing interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
Acknowledgements
The authors would like to thank the two anonymous reviewers for
sharing their expertise and for their comments, which contributed to the
improvement of this manuscript.
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