Quaternaire
Revue de l'Association française pour l'étude du
Quaternaire
vol. 24/1 | 2013
Volume 24 Numéro 1
The Iberian Peninsula, the last european refugium
of Panthera pardus Linnaeus 1758 during the
Upper Pleistocene
La péninsule Ibérique, le dernier refuge européen de Panthera pardus Linnaeus
1758 durant le Pléistocène supérieur
Víctor Sauqué and Gloria Cuenca-Bescós
Publisher
Association française pour l’étude du
quaternaire
Electronic version
URL: http://quaternaire.revues.org/6468
DOI: 10.4000/quaternaire.6468
ISSN: 1965-0795
Printed version
Date of publication: 1 mars 2013
Number of pages: 35-48
ISSN: 1142-2904
Electronic reference
Víctor Sauqué and Gloria Cuenca-Bescós, « The Iberian Peninsula, the last european refugium of
Panthera pardus Linnaeus 1758 during the Upper Pleistocene », Quaternaire [Online], vol. 24/1 | 2013,
Online since 01 March 2016, connection on 30 September 2016. URL : http://
quaternaire.revues.org/6468 ; DOI : 10.4000/quaternaire.6468
The text is a facsimile of the print edition.
© Tous droits réservés
Quaternaire, 24, (1), 2013, p. 35-48
THE IBERIAN PENINSULA, THE LAST EUROPEAN REFUGIUM
OF PANTHERA PARDUS LINNAEUS 1758
DURING THE UPPER PLEISTOCENE
n
Víctor SAUQUÉ1 & Gloria CUENCA-BESCÓS1
ABSTRACT
The new Quaternary site of Los Rincones in the region of Moncayo (Zaragoza, north-east Spain) has provided a well-preserved mandible of Panthera pardus. This mandible shows morphological similarities with that of snow leopard, Panthera uncia. The
resemblance between specimens described as P. pardus in the European Pleistocene and P. uncia raises the question of whether the
morphological variability of P. uncia includes the specimens from the European Pleistocene or whether it is rather a case of convergence in which the European leopard acquired particular characters of the snow leopard, such as the flattened snout, the short dental
diastema and the elongation of the carnassial, through an adaptation process to a mountain environment. In addition, the Moncayo
mandible led to the revision of the P. pardus material from the Iberian Peninsula. Though this region is one of the most densively
inhabited by P. pardus in Europe, it has been poorly studied in the past. This shows that the Cantabrian region in the north of the
Iberian Peninsula might have been the last refuge for this species prior to its complete disappearance from Europe.
Keywords: Panthera pardus, Panthera uncia, European Pleistocene, Iberian Peninsula, last refuge
RÉSUMÉ
LA PÉNINSULE IBÉRIQUE, LE DERNIER REFUGE EUROPÉEN DE PANTHERA PARDUS LINNAEUS 1758 DURANT LE
PLÉISTOCÈNE SUPÉRIEUR
Le nouveau gisement quaternaire de Los Rincones dans la région de Moncayo (Saragosse, nord-est de l’Espagne) a livré une
hémi-mandibule bien préservée de Panthera pardus. Cette mandibule présente des similitudes morphologiques avec celle du léopard
des neiges, Panthera uncia. La similitude entre les spécimens décrits comme P. pardus dans le Pléistocène européen et P. uncia
soulève une question importante : cette mandibule appartient-elle à P. uncia, ou bien est-ce un cas de convergence morphologique
dans lequel le léopard européen acquiert des caractères spécifiques du léopard des neiges, comme l’aplatissement du museau, un
court diastème, et l’allongement de la carnassière en réponse à un processus d’adaptation aux environnements de montagne ? En
outre, la mandibule de Moncayo permet un réexamen du matériel de P. pardus de la péninsule Ibérique. Il est à noter que bien
qu’étant l’une des régions les plus densément peuplées par P. pardus en Europe, celle-ci a été peu étudiée dans les travaux antérieurs.
Enfin, l’analyse du registre fossile indique que la zone cantabrique, au nord de la péninsule Ibérique, pourrait représenter le dernier
refuge de cette espèce avant sa disparition complète en Europe.
Mots-clés : Panthera pardus, Pantehra uncia, Pléistocène européen, péninsule Ibérique, dernier refuge
1 - INTRODUCTION
At present the genus Panthera is made up of five
species: P. leo (lion), P. tigris (tiger), P. onca (jaguar),
P. pardus (leopard) and P. uncia (snow leopard). The
fossil record is known to contain another five species that
are now extinct: P. gombaszoegensis, P. palaeosinensis,
P. spelaea, P. fossilis and P. atrox. Of these, it is the
leopard P. pardus that has the greatest distribution area,
covering almost all of Africa and Asia (Kingdon, 1977;
Nowell & Jackson, 1996; Turner & Antón, 1997). It is
found in all habitats except very dry ones, and exhibits
a great diversity of behaviour associated with the habitat
type (Pocock, 1932; Kingdon, 1977; Turner & Antón,
1997). Leopards are solitary and territorial (Bertram,
1999; Hayward et al., 2006). They protect captured prey
from hyenas in ways that vary with the habitat: in open
environments they drag their prey up trees, whereas in
areas with caves they haul the carcasses inside (see references in Darryl & Berger, 2000). During the Pleistocene,
P. pardus occupied southern Europe and various areas
in central Europe corresponding to Germany, Austria,
Hungary, the Czech Republic and Switzerland (Hemmer,
1971; Jánossy, 1986; Spassov & Raychev, 1997). It is
difficult to reconstruct the last occurrence and extinction
of P. pardus in Europe. The imprecisely dated Mesolithic record from northem Spain (Altuna, 1972) and the
Holocene bone remains from Greece (Nagel, 1999) are
1
Grupo Aragosaurus-IUCA. Paleontología. Facultad de Ciencias. Universidad de Zaragoza. C/ Pedro Cerbuna 12. E-50009 Zaragoza.
Emails: [email protected], [email protected]
Manuscrit reçu le 17/09/2012, accepté le 08/01/2013
1301-066 - Mep 1-2013.indd 35
20/02/13 12:32
36
two indicators that the P. pardus could have survived into
the Holocene (Sommer & Benecke, 2006). The discovery
of a fossil of P. pardus at the site of Los Rincones, a new
Upper Pleistocene locality in the region of Moncayo in
Zaragoza (Spain), extends the palaeontological record
of the European leopard. The unique morphology of
the fossil has led us to revise the published material as
well as present-day collections of P. pardus fossils in
the Iberian Peninsula. The aim of the present paper is to
study the fossil from Los Rincones and discuss the affinities between the leopards of the European Pleistocene
and the present-day snow leopard.
2 - GEOGRAPHICAL AND GEOLOGICAL
SITUATION AND HISTORY
OF THE DISCOVERY OF THE SITE
The cave of Los Rincones is located in the Sierra del
Moncayo, in the central part of the Iberian Range in the
north of the Iberian Peninsula (fig. 1), between the Duero
and Ebro river basins. The mouth of the cave is situated
at the head of the ravine of Los Rincones, at an altitude
of 1010 m, in the Lower Jurassic material of the Cortes
de Tajuña Formation. The cave is oriented SW-NE and
is divided into three chambers. The large main chamber
descends 28 m and presents large blocks that have fallen
from the ceiling. A large sedimentary cone deposited
directly at the cave mouth. This cone is composed mainly
of autochthonous clastic sediments, consisting of clastsupported limestone gravels, clay, and occasional pieces
of speleothem and bones. They are heterometric and
light in colour. The cone closes off the main entrance to
the cave, probably at the end of the Pleistocene (fig. 2).
Among the fallen blocks, near to the top of the cone of
the large main chamber, is the “Ursus Gallery”, a passage
that is small yet of great interest, since there are numerous fossil remains at the surface inside. To the northeast
of the main chamber is the “East Gallery”, which is
characterized by the presence of speleothems.
The site was discovered by members of CEA (Centro
Aragonés de Espeleología) in 2005 while they were
Fig. 2: Elevation and plan views of the Los Rincones cave.
Fig. 2 : Vues en élévation et en plan de la grotte de Los Rincones.
mapping the cave. The cave of Los Rincones is described
by the CEA members Gisbert & Pastor (2009). The
presence of bones in the cave was reported to one of us
(G. C.-B.), who visited the cave in 2006 in company of
Juan Luis Arsuaga and Milagros Algaba and members
of CEA. During this visit we observed that there were
indeed many bone remains scattered in the surface of
several galleries. Subsequently we conducted a several
geological surveys during 2009 and 2010 to collect the
stratigraphic, taphonomic, and cartographic-photographic data. During one of these surveys, in June 2010,
we found the fossil remains of the Felidae studied in the
present work.
3 - MATERIAL AND METHODS
Fig. 1: Geographical location of the Los Rincones cave.
Fig. 1 : Situation géographique de la grotte de Los Rincones.
1301-066 - Mep 1-2013.indd 36
Studied material includes an almost complete right
mandible of a large Felidae, with the canine, the fourth
premolar and the first molar. The material is provisionally housed at the University of Zaragoza and has been
given the field specimen number Ri10/C1/2010.
The mandible was found in the main gallery, at the base
of the sedimentary cone at the cave mouth. The upper part
of the cone is dated by means of the faunal assemblages
found in the sediments of the “Ursus Gallery” that are
probably the lateral end of the upper layers of the cone. It
consists of the rodents Microtus, Iberomys and Pliomys
lenki, which means that the mandible would be at least
20/02/13 12:32
37
as old as Late Pleistocene. The species Pliomys lenki
disappears between 50-40 ka in Central Iberia where it
is found only in Mousterian localities (Cuenca-Bescós et
al., 2010a).
The description of the carnivore mandible follows the
terminology proposed by Schmid (1940), Clot (1980) and
Testu (2006), using the terms “anterior” and “posterior”
for proximal or mesial and distal positions, respectively.
The terms “labial” and “lingual” are used to describe the
elements of the dentition, and the terms “lateral” and
“medial” to describe the dentary bone.
For the biometric section, the measurements were
taken using a digital calliper, following the methodology
proposed by Schmid (1940) and Clot (1980) for teeth and
Testu (2006) for mandible measurements.
H md behind m1 (mm)
12,81 28,28 25,18 24,98 12,98 9,76 12,68 15,49 17,25 8,85 18,81 19,65 8,67 14,79 27,46 26,89
H condilo (mm)
H md in p4-m1 (mm)
DT condilo(mm)
DT md m1 (mm)
DVL m1 (mm)
L alv p3 (mm)
DVL c1 (mm)
32
DMD c1 (mm)
long p3-p4 (mm)
49
H br in p4 (mm)
L alv p3-m1 (mm)
52,54
H br md in front p4 (mm)
H. proc ang Co (mm)
122
H md in c1-p3 (mm)
L Ap Co (mm)
132
L c1-p3 (mm)
L total (mm)
Description: The right mandible from Los Rincones
(Ri10/C1/2010), attributed to P. pardus on the basis of
its morphology and measurements (tab. 1 and fig. 3),
is well preserved and practically complete; the mandibular ramus presents the canine, p4 and m1. These two
elements show wear caused by attrition. Part of the
mandibular symphysis and the incisors are missing. The
vertical ramus of the mandible has complete and wellpreserved coronoid, articular and angular apophyses.
DMD m1 (mm)
4.1 - MATERIAL AND MEASUREMENTS
L alv m1 (mm)
Order Carnivora Bowdich, 1821
Family Felidae Fischer von Waldheim, 1817
Subfamily Felinae Fischer von Waldheim, 1817
Genus Panthera Oken, 1816
Species Panthera pardus (Linnaeus 1758)
DVL p4 (mm)
4 - SYSTEMATIC PALEONTOLOGY
DMD p4 (mm)
Abbreviations
Coll: Collection; MPZ: Museo Paleontologico, University of Zaragoza, M.N.H.N.: National Museum of Natural
History, Paris; MusMZB: Museu de Ciències Naturals de
Barcelona; URV: Universitat Rovira i Virgili de Tarragona; md: mandible, DMD: meso-distal diameter; DVL:
vestibulo-lingual diameter; DT: transverse diameter; H:
height; L: length; Ap: apophysis; Co: coronoid; proc:
process; ang: angular; al: alveolus; LC-p3: length of
diastema; c1: canine; p3: third lower premolar; p4: fourth
lower premolar; m1: first lower molar.
L alv p4 (mm)
In occlusal view, the upper edge of the horizontal
ramus only runs from the alveolus of the canine to the
start of the vertical ramus. After the canine is the diastema, which is the area included between the posterior
edge of the canine and the anterior edge of the third
premolar, which is not preserved. In vertical section,
after the diastema the mandible widens slightly, presenting three alveoli, the first two of which correspond to
the premolars and the third to m1. The vertical ramus of
the mandible extends from the posterior edge of m1. The
base of this area forms the anterior and upper limits of the
vertical ramus, where the start of the coronoid apophysis
is situated. This presents a subtriangular morphology, and
in our specimen the upper part of it is fractured. In lateral
view, the lower edge of the horizontal ramus is practically rectilinear, with a slight convexity that changes
to the slightest of concavities at the height of the coronoid apophysis, extending caudally as far as the angular
process, which is fairly robust.
In labial view, the horizontal ramus in its posterior
part presents a deep masseteric fossa with a triangular
morphology that extends from the posterior edge of m1
to the angular process and the condyle. The lower insertion of the external masseter muscle is well developed
and displays a bony support called the torus. Mandible
(Ri10/C1/2010) is lacking part of this torus, which has
been gnawed away, as can be inferred from the tiny
parallel marks characteristic of rodents (Cáceres, 2002).
In the anterior part of the horizontal ramus in labial view,
three mentonian foramina can be seen, two of which
are situated at the height of the diastema and the third
caudally at the height of p3.
The articular condyle is situated posteriorly and above
the angular process, on a line that would be an extension
of the edge of the mandibular body. The condyle is semicylindrical, anteroposteriorly narrow and lateromedially
elongated.
In lateral and medial view, the horizontal ramus in its
anterior part presents a rugose area with protuberances
and depressions known as the symphyseal face. This
divides the mandible into two hemi-mandibles that do not
fuse completely, in such a way that there is a permanent
symphysis. In the specimen only a small fragment of the
symphyseal face can be seen. The rest of the horizontal
ramus is flat except for a slight convexity situated in the
posterior area in the part opposite the masseteric fossa
and at the height of the condyle. The masseteric fossa is a
shallow depression with a subcircular outline.
11
27,25
Tab. 1: Measurements of mandible and teeth of Panthera pardus from Los Rincones (Ri10/C1/2010).
Measurements are expressed in millimetres, to the nearest 0.1 mm.
Tab. 1 : Dimensions de la mandibule et des dents de Panthera pardus de Los Rincones (Ri10/C1/2010). Les mesures sont exprimées en millimètres, au
10e de millimètre près.
1301-066 - Mep 1-2013.indd 37
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38
Fig. 3: Fossil remains of Panthera pardus from Los Rincones (Ri10/C1/2010).
Buccal (A), lingual (B) and occlusal (C) views of the right hemi-mandible.
Fig. 3 : Restes fossiles de Panthera pardus de Los Rincones (Ri10/C1/2010). Vues buccale (A), linguale (B) et occlusale (C) de l’hémi-mandibule droite.
The canine belonging to (Ri10/C1/2010) has a relatively rounded transverse section, and presents one carina
in the lingual distal edge.
The p4 is relatively long in comparison with m1 (90%
of its length). In labial view, it is fairly symmetrical,
presenting a paraconid and a metaconid that are similar
in length and height. The protoconid is at the height of
the paraconid of m1 and presents a slanting anterior
edge that is displaced slightly backwards, and an almost
vertical posterior edge. In the caudal part it has a marked
cingulum that is developed above all on the lingual face.
In occlusal view, the tooth presents an oval outline, with a
slight double-pinched constriction in the contact between
the paraconid and the protoconid.
The carnassial molar, or m1, has a typically feline
morphology, with two aligned conules, the paraconid
and the protoconid, which are separated by a notch of
4.71 mm. The paraconid is short in relation to the protoconid and is situated at the height of the cusp of p4,
presenting an anterior edge that is somewhat backwardly
inclined and a slanting posterior edge. The protoconid
is longer and moderately higher than the paraconid; its
posterior edge is slightly serrated and ends caudally in a
small, scarcely developed talonid. In occlusal view, m1
displays an oval morphology, with the development of a
small bulb in the lingual face at the height of the notch.
4.2 - DISCUSSION
The mandible from Los Rincones (Ri10/C1/2010)
shows morphological similarities to present-day snow
leopard, Panthera uncia. As in this species, moreover,
1301-066 - Mep 1-2013.indd 38
the mandibular body is robust, a character that both
species share with Puma concolor and P. pardus
(Madurell-Malapeira et al., 2010). The short diastema
from Los Rincones (Ri10/C1/2010) bear a resemblance
to P. uncia as well as to P. onca (Spassov & Raychev,
1997), characters also present in the P. pardus of the
Upper Pleistocene, as evidenced by the Iberian sites of
Zafarraya (Testu, 2006; Testu et al., 2011) and Algar da
Manga Larga (Cardoso & Regala, 2006) and the Bulgarian site of Triagalnata (Spassov & Raychev, 1997). In
lateral view, it has three mentonian foramina, two of
which are beneath the distal edge of c1 and the third
beneath the root of p3. This arrangement is seen in the
material from Saint-Vallier, as well as in P. concolor
and P. uncia, whereas in general P. pardus only presents
two foramina (Testu et al., 2011; Madurell-Malapeira
et al., 2010). The mandible from Aragó attributed to
P. uncia by Hemmer (2003) also has three mentonian
foramina. In the mandible from Los Rincones (Ri10/
C1/2010), the masseteric fossa reaches the level of
the protoconid of m1, as in P. concolor and P. uncia
(Madurell-Malapeira et al., 2010). The mandible from
Los Rincones (Ri10/C1/2010) shares characters with
P. pardus, such as the lesser height of the horizontal
ramus in front of p3 than behind m1 (Christiansen,
2008), and it also lacks the rectilinear basal outline, as
in P. uncia (Hemmer, 1972), its basal outline showing a
slight convexity that becomes a slight concavity at the
level of the coronoid apophysis.
The relatively rounded transverse section and the
diameter of the canine from Los Rincones (Ri10/C1/2010)
fall within the range for P. uncia (Hemmer, 1972) (tab 1
20/02/13 12:32
39
Ri 10,C1,1
Triagalnata cave 3859
Geinsheim
Torrejones
Observatoire 1858
Observatoire 1837
Observatoire 1861
Abric Romani
Zafarraya 93-P14-47
Zafarraya 90-R8/9-2356
Manga Larga
Cueva Allekoaitze
Observatoire 1857
Observatoire 1860
Coll M.N.H.N1883-1738
Coll M.N.H.N1928-3
Coll M.N.H.N A 1860
Coll M.N.H.N A 621
Coll M.N.H.N 1873-311
Coll M.N.H.N 1944-165
Coll M.N.H.N 1931-67
Coll M.N.H.N 001-1997
Coll M.N.H.N 1941-47
Coll M.N.H.N 1960-88
Coll M.N.H.N 1905-49
Coll M.N.H.N 1898-100
Coll M.N.H.N 1934-12
Coll M.N.H.N 1910-72
BCN 82-7004
BCN 94-0437
BCN 91-0064
BCN 94-0895
BCN 91-307
BCN 94-0461
BCN 82-376
BCN 82-376
BCN 88- 0027
BCN 82-7103
BCN 82-7122
Wien Museum Nr 11920
Choukoutien
Coll M.N.H.N A 14527
Coll M.N.H.N 1970-102
M.N.H.N Zoothèque 1998-1248
Coll lab Tarragona
BCN 2000-0482
BCN 2006-0317
BCN 2003-0096
URV
Animal diversity web 157589
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Upper Pleistocene
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Recent
Spassov & Raychev, 1997
Koenigswald et al ., 2006
Arribas et al ., 1997
Testu, 2006
Testu, 2006
Testu, 2006
Cáceres et al ., 1993
Barroso et al ., 2006
Barroso et al ., 2006
Cardoso & Regala, 2006
Corral, 2012
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Thenius, 1969
Thenius, 1969
Testu, 2006
Testu, 2006
Testu, 2006
Testu, 2006
Hemmer, 1972
P.pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus fossil
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. pardus
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
P. uncia
19,65
18,1
18,8
17,7
21,7
21,6
19,2
19,5
20,3
17,9
18,6
20,5
21,9
19
14,7
16,1
15,8
16,7
17,9
15,6
15,9
19,6
20,2
19,3
17,7
15,5
20,8
18,3
15,68
16,69
16,75
17,83
15,4
18,13
20,96
18,65
20,91
17,42
18,75
16
19,1
18,5
19,6
19
18,9
17,28
19,12
18,27
18,9
18,25
27,25
27,98
32,39
28,00
31,80
31,10
29,60
30,50
34,10
28,80
27,77
30,41
23,00
24,90
23,60
29,10
32,10
25,6
25,81
31,7
33,2
35,1
30,5
24,2
32,8
30,8
23,66
28,79
26,23
30,49
25,76
29,29
36,68
33,12
36,26
29,94
33,46
23,00
26,20
24,60
24,80
26,73
28,10
26,00
26,62
23,63
28,00
24,99
132
134
161
134
155
154,2
140,6
145
147,6
126
141
148
132
111
117,7
121,5
150,9
143
118
119,5
144
165,4
176,2
144,8
123,3
157,5
148,3
122,35
133,97
129,05
137,61
139,4
137,59
163,01
162,91
168,51
138,99
152,56
114
127
122,3
129,4
134
134,6
126,71
129,85
130,28
136,29
129
49
49,03
50,8
44,7
53,4
57,2
50,3
49,3
53,7
44,2
50,68
56,5
48,3
38,1
42,6
40,2
51,5
46,4
42,9
41,6
50,6
52,5
50,6
46,9
41,3
53,3
46,2
42,09
44,3
43,5
45,35
44,29
45,64
52,77
52,57
49,97
44,53
49,33
42
44
46,3
48,5
49,1
48,7
44,63
48,83
46,3
48,63
47
138,68
154,60
172,30
158,19
146,54
143,98
154,17
156,41
167,98
160,89
149,30
148,34
156,46
154,66
149,37
174,26
179,33
164,10
162,35
161,73
164,36
181,87
172,32
156,13
157,69
168,31
150,89
172,50
156,60
171,00
167,27
161,55
175,01
177,57
173,43
171,87
178,43
143,75
137,17
132,97
126,53
140,72
148,68
150,61
139,23
129,34
148,60
136,99
DMD Pro p4 / DMD p4 %
DMD Pa m1/ DMD m1 %
index DMD m1 / L total
md
(H md body after m1 / DMD m1 x
100)
Robustness index
L p3-m1 (mm)
L total (mm)
H behind m1 (mm)
The m1 of the mandible Ri10, C1, 1 has the following
characteristics in common with P. uncia: the presence
of a short paraconid, i.e. the length of the paraconid
in relation to the total length of m1 in P. uncia ranges
from 46-52 % (Hemmer, 1972), while in the specimen
from Los Rincones (Ri10/C1/2010) the ratio totals
9.94/19.65 x 100 = 50.54 %; the m1 of Ri10, C1, 1 has a
notch located in a low position characteristic of P. uncia
(Hemmer, 1972).
In P. uncia the talonid of m1 is divided by a transverse groove at the posterior edge of the protoconid, and
the edge of the protoconid is serrated (Schmid, 1940).
The protoconid of the m1 of Ri10, C1, 1 also presents a
slightly serrated edge. The talonid of the m1 of Ri10, C1,
1 is less marked than in P. uncia and does not display the
typical triangular morphology (Schmid, 1940).
The ratio between the length of m1 and the total length
of the mandible is a character used to separate P. uncia
from all other pantherines (Christiansen, 2008). In the
DMD m1 (mm)
Taxon
References
Chronology
Label
& 2). The mandible from Los Rincones (Ri10/C1/2010)
differs from the fossil species P. gombaszoegensis in the
canine, which, as in P. uncia, is smaller (García, 2003).
The short protoconid of the p4 from Los Rincones
(Ri10/C1/2010), measured as the length of the protoconid in relation to the total length of p4, falls within
the range of variability of P. uncia (see measurements
compared in the tabs.) (Hemmer, 1972). Moreover, the
p4 from Los Rincones (Ri10/C1/2010) shows a wide
anterior area also within the range shown by P. uncia
(fig. 3).
In dorsal view, the p4 from Los Rincones (Ri10/
C1/2010) presents an oval outline, with a slight doublepinched constriction at the height of the contact between
the protoconid and the paraconid. This characteristic is
not seen in P. uncia, which has an oval outline without
any constrictions (Spassov & Raychev, 1997). By
contrast, the presence of this double-pinched constriction
is typical of P. pardus (Spassov & Raychev, 1997).
0,149
0,135
0,117
0,132
0,140
0,140
0,137
0,134
0,138
0,148
0,145
0,148
0,144
0,132
0,137
0,130
0,111
0,125
0,132
0,133
0,136
0,122
0,110
0,122
0,126
0,132
0,123
0,128
0,125
0,130
0,130
0,110
0,132
0,129
0,114
0,124
0,125
0,123
0,140
0,150
0,151
0,151
0,142
0,140
0,136
0,147
0,140
0,139
0,141
45-52
43-50
Tab. 2: Measurements (in mm) of mandibles and teeth of Panthera pardus and Panthera uncia.
Tab. 2 : Mesures (en mm) des mandibules et des dents de Panthera pardus et Panthera uncia.
1301-066 - Mep 1-2013.indd 39
20/02/13 12:32
40
univariate representation of this character, it can be
seen that Ri10/C1/2010 falls within the range of variation of P. uncia (tab. 2 and fig. 4), although it is close to
the maximum values for P. pardus, such as those from
Manga Larga (Cardoso & Regala, 2006) and Observatoire (Testu, 2006).
The robustness index of the mandible from Los Rincones
(Ri10/C1/2010) falls within the range of P. uncia, very
close to its mean value (tab. 2 and fig. 5). Low robustness
indices are a consequence of the relative elongation of m1
in relation to the mandible (Testu, 2006).
The mandible Ri10/C1/2010 lies within the 95 % confidence ellipses of the populations of fossil P. pardus and
P. uncia, though it is closer to the population of P. uncia,
since it lies within the convex hull for this (fig. 6). This is
due to the fact that the jugal series occupies a great part of
the characteristic mandible typical of P. uncia.
The specimen from Los Rincones (Ri10/C1/2010) presents mosaic characteristics. On the one hand, it presents
characteristics typical of P. uncia, such as: a short diastema; a similar robustness index; relative elongation of the
molar series in relation to the total length of the mandible;
a DMD m1 / total length of mandible index that is very
similar to what is shown by P. uncia, these uncioid features
might be related with the specialization of the masticatory apparatus that is developed in a mountain environment where preys are scarce and the consumption of the
carcasses is common (Spassov & Raychev, 1997; Testu,
2006; Testu et al., 2011), these features are observed in
other P. pardus that developed in southern Europe in the
course of the Pleistocene. On the other hand, however, it
displays characteristics similar to P. pardus such as: basal
outline of the horizontal ramus concavo-convex; p4 with
double-pinched constriction; m1 with a scarcely marked
talonid and without metaconid; a mandibular ramus that is
higher behind m1 than in front of p3. The specimen Ri10,
C1, 1 would fall within the P. pardus despite the mandibular body resemble P. uncia the m1 without metaconid
cuspid permit to exclude P. uncia because the presence of
this cuspid is a diagnostic character of P. uncia (Schmid,
1940; Testu et al., 2011).
Fig. 4: Box-plot representing the ratio between the length of m1
and the total length of the mandible of Felidae (Ri10/C1/2010 and
Christiansen, 2008).
Fig. 4 : Boîte à moustaches représentant le rapport entre la longueur de
la m1 et la longueur totale de la mandibule de Felidae (Ri10/C1/2010
et Christiansen, 2008).
Fig. 5: Box-plot representing the robustness index (height of mandibular body posterior to m1 versus DMD m1 x 100).
Fig. 5 : Boîte à moustaches représentant l’indice de robustesse (hauteur
du corps mandibulaire postérieur à la m1 par rapport au DMD de la
m1 x 100).
5 - THE PALAEONTOLOGICAL RECORD
OF P. PARDUS-P. UNCIA IN EUROPE
The possible presence of P. uncia in the European
Pleistocene is a question that has periodically been raised
ever since Woldrich (1893) assigned the fossils from the
site of Turská Maštal (Czech Republic) to this taxon,
though these were subsequently re-ascribed to Lynx
(Thenius, 1957). Years later, Thenius (1969) ascribed to
P. uncia two mandibles from Stránska Skála, which were
subsequently re-assigned to P. pardus by Hemmer (1971).
Recently, Hemmer (2003) assigned the mandible from La
Caune de l’Arago to P. uncia; this was then re-assigned to
Panthera pardus by Testu et al., (2011).
Other authors have described P. pardus in the south
of Europe with characters similar to those of P. uncia,
1301-066 - Mep 1-2013.indd 40
Fig. 6: Graph representing length p3-m1 versus the total length of
the mandible.
Fig. 6 : Graphe représentant la longueur p3-m1 par rapport à la
longueur totale de la mandibule.
examples including the specimen from Manga Larga
in Portugal (Cardoso & Regala, 2006), that from Triagalnata Cave in Bulgaria (Spassov & Raychev, 1997),
the specimens of P. pardus vraonensis from Vraona in
Greece (Olive, 2006), as well as the skeleton found in
Bosnia Herzegovina (Malez & Pepeonik, 1970; von
Koenigswald et al., 2006). There are two possible
hypotheses that might explain the presence of these specimens with uncioid characteristics during the Upper Pleis-
20/02/13 12:32
41
tocene, the first assuming that P. uncia reached Europe
and the second assuming morphological convergence by
P. pardus during the Upper Pleistocene. Hemmer (2003)
has argued that P. uncia reached the south of Europe by
migrating at a time of maximum glacial expansion during
the Middle Pleistocene, in the course of which it crossed
the Persian Mountains, the Balkans and the Alps to reach
as far as the Pyrenees.
By contrast, the hypothesis positing the acquisition of
characters typical of P. uncia is based on the fact that certain
characteristics of the snow leopard, such as the robustness
of the mandible, the verticality of the symphysis, the short
diastema, and the elongation of the jugal teeth in relation
to the mandible, can be considered adaptations to mountainous regions, where prey are scarce and difficult to
capture, (probably Caprinae) leading to the consumption
of the whole carcass (Spassov & Raychev, 1997; Testu,
2006; Testu et al., 2011). Such characteristics are to be
expected in felines that have evolved in environments of
this sort like the P. pardus from Caune de l’Arago (Testu
et al., 2011). Most authors agree that P. pardus shows
great versatility in adapting itself to a variety of habitats,
displaying broad variations in morphology and size that
would have facilitated its adaptation to all types of environments except the most arid and desert-like (Pocock,
1930; Nowell & Jackson, 1996; Turner & Antón, 1997).
On the other hand, the microvertebrate discovered in other
levels of the cave at Los Rincones contains, among other
insectivores and rodents, the arvicoline Pliomys lenki, a
fossil species that was probably an inhabitant of open lands
on mountains (Cuenca-Bescós et al., 2010a). Large herbivores have also been found in other levels at Los Rincones
such as those in the “Ursus Gallery”, where Capra pyrenaica is dominant. This would suggest that during the
Upper Pleistocene the cave at Los Rincones was charac-
terized by mountainous conditions, possibly representing a
habitat similar to that of the present-day snow leopard.
6 - PALEOGEOGRAPHIC DISTRIBUTIONS AND
ORIGIN OF PANTHERA PARDUS IN EUROPE
Most of the works on P. pardus in Europe during the
Pleistocene (Schmid, 1940; Fischer, 2000; Sommer &
Benecke, 2006; von Koenigswald et al., 2006; tab. 3) lack
data on P. pardus in the Iberian Peninsula, even though
this area has one of the largest records of archaeological
and palaeontological remains in Europe for the Upper
Pleistocene. Above all in the Upper Palaeolithic, a greater
density of sites is recorded than in other parts of Europe.
The Iberian localities are mainly in the Basque-Cantabrian region, where the number of citations of P. pardus
is greater than all the rest of Europe at this time (tab. 3).
The oldest reference to P. pardus in Europe is from
the Lower Pleistocene of Le Vallonnet (France), dated to
980-910 ka (Moullé et al., 2006; Turner, 2009). This is
the only citation of this species in the European Early
Pleistocene (Palombo & Valli, 2003). From the beginning of the Middle Pleistocene P. pardus started the
progressive replacement of P. pardoides (Hemmer, 2004;
Palombo et al ., 2008), P. pardus is found at various
Middle Pleistocene sites in Italy (Kotsakis & Palombo,
1979) and France (Palombo & Valli, 2003; Palombo et
al., 2008), and it is also sporadically recorded in England
(Currant & Jacobi, 2001), Germany (Hemmer & Schütt,
1969), the Czech Republic (Thenius, 1972), Hungary
(Kahlke et al., 2011) Austria (Schmid, 1940; Döppes &
Rabeder, 1997), Georgia (Baryshnikov, 2011), Greece
(Kurten & Poulianos, 1980; Kurten 1983) and Spain
(Altuna, 1972; Falguères et al., 2005) (tab. 3 and fig. 7).
Fig. 7: Distribution area of Panthera pardus in Europe during the Lower and Middle Pleistocene.
UK: 1/ Pontnewydd Cave, 2/ Bleadon Cavern; SPAIN: 3/ Lezetxiki; FRANCE: 4/ Caune de l’Arago, 5/ Lunel-Viel, 6/ Orgnac 3, 7/ Azé 1-5, 8/ Lazaret,
9/ Vallonet; ITALY: 10/ Valdemino Cave, 11/ Romagnano, 12/ Grotta di Cerè, 13/ Soave-Sentiero, 14/ Soave Monte Tenda, 15/ Monte Sacro (Prati Fiscali),
16/ Isernia; GREECE: 17/ Petralona Cave; GEORGIA: 18/ Treugolnaya, 19/ Kudaro; AUSTRIA: 20/ Repolusthöhle, 21/ Hundsheim, HUNGARY:
22/ Tarkö, CZECH REPUBLIC: 23/ Stránska Skála, 25/ Koneprusy C718; POLAND: 24/ Biśnik Cave; GERMANY: 26/ Mauer 5, 27/ Mosbach 2.
Fig. 7 : Aire de répartition de Panthera pardus en Europe au cours du Pléistocène moyen et inférieur.
1301-066 - Mep 1-2013.indd 41
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42
Chronology
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
Early Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
M.Pl
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Country
United Kingdon
United Kingdon
Spain
France
France
France
France
France
France
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Greece
Georgia
Georgia
Austria
Austria
Hungary
Czech Republic
Poland
Czech Republic
Germany
Germany
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Gibraltar (UK)
Gibraltar (UK)
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Monaco
Monaco
Italy
Italy
Italy
Italy
Germany
Italy
Switzerland
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Switzerland
Site
Pontnewydd Cave
Bleadon Cavern
Lezetxiki
Caune de l`Arago
Lunel-Viel
Orgnac 3
Grotte d'Aze I-5
Grotte du Lazaret
Grotte du Vallonet
Valdemino cave
Romagnano
Grotta di Cerè
Soave-Sentiero
Soave-Monte Tenda
Prati Fiscali
Isernia
Petralona Cave
Treugolnaya Cave
Kudaro 1, 3
Repolusthöhle
Hundsheim
Tarkö
Stránska Skála
Biśnik Cave
Koneprusy C718
Mauer 5
Mosbach 2
Escoural
Figueira Brava
Prado das Salemas
Casa da Moura
Furinha
Caldeirão
Algar da Manga Larga
Lorga da Dine
Gorham Cave
Vanguard Cave
Zafarraya
Carigüela
Cova Negra
Cova Foradada
Pinarillo I
Pinilla del Valle
Cueva de la Zarzamora
Cueva del Búho
Torrejones
Los Casares
Aguilón P-7
Los Rincones
Ermita
Valdegoba
Cueva de "El Castillo"
Prado Vargas
Abrigo de Olha
Coscobilo
Ekain
Axlor
Arrillor
Amalda
Cueva Allekoaitze
Lezetxiki
Gabasa
Les Muricers
Cova del Gegant
Abric Romani
L`Arbreda
Ermitons
Mollet I
Grotte de la Carrière
Coupe-Gorge
La Crouzade
Tournal
L' Hortus
L'Arago
Cave of Jaurens
d' Artenac
Moula-Guercy
Chauvet
Grotte de l' Adaouste
Verzé
Blanot 2
Etrigny
La Grotte des Enfants
L’Observatoire
Grotte du Prince
Madonna dell' Arma
Santa Lucia Superiore
Cave of Fate
Cave of Equi
Rittersaal Höhle
Groticelle di Sambughetto
La Grotte de Cotencher
Grotta de Fossellone
S. Agostino
Grotta di Castelcivita
Ingarano
Grotta di Fumane
Riparo Tagliente
Grotta di San Bernardino
Zandobbio
Wildkirchli
Nº in the map
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
Level
Archaeo
Chronology
L.VI
Ms.
ESR: 234 ± 32 ka
MIS 14
MN23a
MN23b (late Galerian)
MN24 (early Aurelian)
OIS 7; MN24 (early Aurelian)
EPR: 980-910 ka
L.5
Riss
Biostratigraphy: 400-160 ka
K/Ar: 736 ± 40 ka
L.5c
Lo. Pa.
Ms.
TL: 360 ± 90 ka
230
Th/U : 230 +13-12 ka
OIS 5-9
Elsterian (550-450 ka)
Biostratigraphy: MIS 15 (600 ka)
U/Th : 48.9 + 5.8 / -5.5 and 26.4 +11 / -10 ka
14
Ms.
14
C: 30.93 ± 0.70 C ka BP
C: 24.85 ± 0.55 14C ka BP
14
C: 25 ± 0.22 C ka BP
14
14
L.1 b E
Ms.
Ms.
14
14
C: 27 ± 0.6 C ka BP
14
C: 35-20 C ka BP
14
L.G; L.K, M, Q
L.UB; L.UE
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
14
14
14
14
C: 51-45 C ka BP
OSL: 93.3-70.3 ka
C: 50-42 C ka BP
14
L VII
14
C: 33.9 ± 0.3 C ka BP
Ms.
Ms.
14
L.5_4
L.18
L.D
L.Lmc
L.VII
L.VI
L.e-g
Unit A.
L.B-I
L.Eb-Gbc.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
Ms.
14
14
C : > 34.76 C ka BP
Biostratigraphy: 50-40 ka
Würm II-III
U/Th : < 73.2 ± 5 ka
14C: 40.7 ± 1.5 and 39.5 ± 2 ka BP
Amino acid dating: 46.4 ka
14
14
14
14
C: > 30.6 C ka BP
C: > 45.9 C ka BP
OIS 5e - OIS 3 (128-40 ka)
14
14
C: 47.1 ± 2.1 and 35.4 ± 0.8 C ka BP
14
L.IV-VI
Ms.
Ms.
Lo. Au.
Ms.
Ms.
Ms.
Ms.
L.8
L.XV
14
C: 32.51 ± 0.24 C ka BP
Würm I- II
OIS 3 ( 65-32 ka)
OIS5 - OIS 3 (125-35 ka)
Ms.
14
C: 33.19 ± 0.66 C ka BP
Würm I
Riss terminal
Würm MIS 4
14
14
C: 38-34 C ka BP
MIS 3
14
C: 32.6 + 2.9 / - 2.1 C ka BP
early Würm
MIS 5e Eemian
14
14
14
C: 37-30 C ka BP
Ms.
Würm I
Ms.
Ms.
Ms.
end of MIS 5
first part of Up. Pl.
end of MIS 5 - MIS 3
Riss-Würm
Middle Würm
Würm
Ms.
Ms.
Ms.
Würm III
14
L.8, L.S,L.A5+A6
L.VI
Ms.
Ms.
14
C: 40-38 C ka BP
14
C: 38.25 ± 0.7 C ka BP
MIS 3
ESR: 108 ± 16 ka
Eemian
Eemian interglacial
14
References
Currant & Jacobi, 2001
Currant & Jacobi, 2001
Altuna, 1972; Falgères et al ., 2005
Hemmer, 2003; Testu et al ., 2011 and references therein
Hemmer, 1972; Fosse, 1996
Aouraghe, 1999
Argant, 1991, 2000
Valensi & Abbassi, 1998
Turner & Antón, 1997; Moullé et al ., 2006
Nocchi & Sala, 1997
Schmid, 1940
Kotsakis & Palombo, 1979
Kotsakis & Palombo, 1979 and references therein
Kotsakis & Palombo, 1979 and references therein
Kotsakis & Palombo, 1979
Sala, 2006
Kurten & Poulianos, 1980; Kurten, 1983
Baryshnikov, 2011
Baryshnikov, 2011
Döppes & Rabeder, 1997
Schmid, 1940
Kahlke et al ., 2011
Thenius, 1972; García & Virgós, 2007
Marciszak et al ., 2011
García & Virgós, 2007
Schütt, 1969; Wagner et al ., 2011
Hemmer & Schütt, 1969
Cardoso, 1996; Yravedra, 2003; Cardoso, 2006
Cardoso, 1996; Yravedra, 2003
Cardoso, 1996; Yravedra, 2003
Valente, 2004
Altuna, 1972
Davis, 2002
Cardoso & Regala, 2006
Cardoso, 1996; Yravedra, 2003
Stringer et al ., 2008
Cáceres, 2002
Barroso et al ., 2003
Ruiz Bustos & García-Sánchez, 1977
Pérez Ripoll, 1977
Pantoja et al ., 2011
Arribas, 1997
Alférez et al ., 1982
Sala et al ., 2011
Molero et al ., 1989
Arribas, 1997; Arribas et al .,1997
Delibes, 1972
Cuenca-Bescós et al ., 2010b
This study
Alférez et al ., 1982
Arribas et al ., 1997
Altuna, 1990
Navazo et al ., 2005
Altuna, 1972
Altuna, 1972
Altuna & Mariezkurrena, 1984
Castaños, 1987
Castaños, 1987
Altuna, 1990
Corral, 2012
Cabrera, 1984
Blasco, 1997; Utrilla et al ., 2010
Estévez, 1979
Daura et al ., 2005
Cáceres et al ., 1993; Cáceres, 2002
Estévez, 1987
Maroto, 1993
Maroto et al ., 1987
Clot, 1980
Guadelli, 1990
Henry-Gambier & Sacchi, 2008
Patou-Mathis, 1994
Testu, 2006
Testu, 2006; Testu et al ., 2011
Guérin et al., 1979; Ballesio, 1980
Delagnes et al ., 1999
Valensi et al ., 2012
Fosse, 2003
Defleur et al ., 1998
Argant, 1991
Argant, 1991
Argant, 1991
Testu, 2006
Boule & Villeneuve, 1927
Boule, 1906-1919
Valensi & Pstabi, 2004
Valensi & Pstabi, 2004
Schmid, 1940
Kotsakis & Palombo, 1979 and references therein
Döppes & Rabeder, 1997; Koenigswald et al ., 2006
Kotsakis & Palombo, 1979 and references therein
Schmid, 1940
Mallegni, 1992
Kotsakis & Palombo, 1979 and references therein
Caloi et al ., 1986
Petronio et al ., 2006
Fiore et al ., 2004
Thun Hohenstein, 2006
Cassoli & Tagliacozzo,1994
Kotsakis & Palombo, 1979
Schmid, 1940
Tab. 3: Localities with Panthera pardus in Europe.
Abbreviations for Lower (Lo.), Middle (M.), Upper (Up.), Pleistocene (Pl.), Palaeolithic (Pa.), Mousterian (Ms.), Aurignacean (Au.), Gravettian (Gr.),
Solutrean (So.), Magdalenian (Mg.), Azilian (Az.); electron spin resonance (ESR), electron paramagnetic resonance (EPR), radiocarbon (14C) and
thermoluminescence (TL) datings.
Tab. 3 : Gisements avec Panthera pardus en Europe. Abréviations pour Inférieur (Lo.), Moyen (M.), Supérieur (Up.), Pléistocène (Pl.), Paléolithique
(Pa.), Moustérien (Ms.), Aurignacien (Au.), Gravettien (Gr.), Solutréen (So.), Magdalénien (Mg.), Azilien (Az.) ; datations par résonance de spin électronique (ESR), résonance paramagnétique électronique (EPR), radiocarbon (14C) et thermoluminescence (TL).
1301-066 - Mep 1-2013.indd 42
20/02/13 12:33
43
Chronology
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -M. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Up.Pl -Up. Pa
Country
Germany
Germany
Germany
Belgium
Germany
Germany
Germany
Germany
Germany
Austria
Austria
Austria
Czech Republic
Czech Republic
Austria
Austria
Austria
Austria
Austria
Croatia
Croatia
Italy
Bosnia & Herzegobina
Serbia
Bulgaria
Greece
Greece
Turkey
Georgia
Russia
Georgia
Georgia
Portugal
Portugal
Portugal
Gibraltar (UK)
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
France
Spain
Spain
France
France
Italy
Switzerland
Germany
Czech Republic
Bulgaria
Greece
Site
Höhlenstein-Stadel
Geinsheim
Biedermann cave
Scladina cave
Petershöhle
Teufelsbrücke
Taubach
Cave of Baumann
Niederlehme
Holzinger Höhle
Luegloch
Merkenstein
Cave of Schwedentisch
Sveduv Stul
Große Peggauerwandhöhle
Kugelsteinhöhle
Große Badhöhle
Fünffenstergrotte
Ochsenhalt Höhle
Krapina
Veternica Cave
Caverna degli Orsi
Vjetrenichia
Smolucka cave
Bacho Kiro
Asprochaliko
Apidima
Karaïn
Mezmaiskaya
Akhstyrskaya cave
Bronzovaya cave
Azokh I
Fontainhas
Casa da Moura
Caldeirão
Gorham Cave
La Riera
Hornos de la Peña
Cueva de "EL.Castillo"
El Juyo
Cueva Morín
El Mirón
Las Pajucas
La Blanca
Oyalkoba
Atxuri
Bolinkoba
Lezetxiki
Amalda
Urtiaga
Isturiz
L`Arbreda
Cau de Coçes
L' aven du Charnier
La Balauzière
Arene Candide
Ettingen
Teufelsbrücke
Predmostí
Triagalnata cave
Vraona
Nº in the map
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
131
132
133
134
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
Level
Archaeo
Chronology
14
C: 47-31 ka
Riss-Würm
TL: 117.2 ± 11.2 ka
Eem Interglacial
Unit 4-5
Eemian
early Würm
early Upper Weichselian
Middle Würm
Middle Würm
14
14
C: 47-31 C ka BP
Würm
Middle Würm
early Würm
Middle Würm
L.20-25
14
14
C: 42.29 ± 9.7 C ka BP
early Würm
L.7
MIS 5 - MIS 2
early Würm
L.3
14
14
C: 47-29 C ka BP
10_16
Layer 2-3
Bed II.
L.1 b E
E+D
L.8
L.108
L.III, IV-V, VI
L.III-IV
L.V-VI
L.F
B-C
Ms.
Ms.
Ms.
Ms.
Ms.
M. Pa.
Up. Pa.
Up. Gr.
So., Mg.
So.
Az.
Au.
Au.
Mg.
Au.
Mg.
Me. ?
So.
Up. Pa.
Up. Pa.
Mg., So., Gr.
Au.
Gr.
Up. Mg.
Au.
Mg.
So.
14
14
C: > 45 C ka BP
14
14
C: 22.73 + 8.8 / -7.9 C ka BP
14
C: 25.09 ± 0.22 C ka BP
14
C: 27.6 ± 0.6 and 18.06 ± 0.14 C ka BP
14
14
14
14
C: 20.93 ± 0.37 C ka BP
14
C: 37.7 ± 1,8 and 41.1 ± 0.7 C ka BP
14
14
C : 11.4 ± 0.3 and 14.4 ± 0.18 C ka BP
14
14
C: 28.15 ± 0.73 C ka BP
14
14
C: 13.66 ± 0.07 and 14.85 ± 0.06 C ka BP
14
14
14
C: 27.4 ± 1 and 19.9 ± 0.34 C ka BP
C: 17.05 ± 0.14 14C ka BP
14
14
14
C: 17.32 ± 0.32 C ka BP
MIS 2
Würm II
14
C: 20.47 ± 0.23 14C ka BP; 22.5 ± 0.37 14C ka BC
L.p9.
Mg.
Mg.
14
C: 13.02 ± 0.08 14C ka BP
C: 26.1 ± 0.5 and 26.9 ± 1.6 14C ka BP
C: 15.57 ± 0.31 14C ka BP
14
C: 24.49 ± 1.35 and 7.07 ± 0.28 14C ka BP
14
14
References
Gamble, 1999
Koenigswald et al ., 2006
Adam & Berckhemer, 1983
Patou-Mathis, 1998; Döppes et al ., 2008
Schmid, 1940
Koenigswald et al ., 2006
Schmid, 1940; Hemmer, 1977
Schütt, 1969
Fischer, 2000
Döppes & Rabeder, 1997; Koenigswald et al ., 2006
Koenigswald et al ., 2006
Wettstein & Mühlhofer, 1938
Schmid, 1940
Musil, 1980
Döppes & Rabeder, 1997; Koenigswald et al ., 2006
Döppes & Rabeder, 1997
Döppes & Rabeder, 1997
Koenigswald et al ., 2006
Döppes & Rabeder, 1997; Koenigswald et al ., 2006
Malez & Pepeonik, 1970
Musil, 1980
Berto & Rubinato, 2011
Malez & Pepeonik, 1970
Dimitrijevic, 1991
Wiszniowska, 1982
Bailey et al ., 1983
Tsoukala, 1999
Testu, 2006
Baryshnikov et al ., 1996; Baryshnikov, 2011
Baryshnikov 2011
Baryshnikov, 2011
Baryshnikov, 2011
Cardoso, 1996
Valente, 2004
Davis, 2002
Carrión et al ., 2008
Altuna, 1972
Barandiarán & Sonneville, 1964
Altuna, 1990
Castaños, 1983
Castaños, 1983
Marín, 2009
Altuna, 1972
Yravedra, 2008
Castaños, 1987
Altuna, 1972; Castaños, 1987
Castaños, 1983; Yravedra, 2007
Cabrera, 1984
Klein & Cruz Uribe, 1985; Altuna, 1990
Altuna, 1970; Castaños, 2005
Altuna, 1972
Estévez, 1987
Yravedra, 2008
Boulestin et al ., 2006
Testu, 2006
Bietti & Molari, 1994; Cassoli & Tagliacozzo, 1994
Koenigswald et al ., 2006
Musil, 1980; Hedges et al ., 1998
Musil, 1994
Spassov & Raychez, 1997
Nagel, 1999
Tab. 3 (continuation)
Tab. 3 (suite)
The oldest presence recorded in the Iberian Peninsula is
found in the Middle Pleistocene, in level VI of the site of
Lezetxiki, dated to 234 ± 32 ka (Altuna, 1972; Falguères
et al., 2005), this being the only mention of this age in
the Iberian Peninsula. The latest record from the Late
Pleistocene is probably the one from Valdegoba (Burgos,
Spain), dated to < 73.2 ± 5 ka (Quam et al., 2001).
The distribution area of P. pardus increased notably
(Clot, 1980; Castaños, 1990) from the second third
of the Late Pleistocene onwards, with the species
becoming a common feature of the sites of this
time period (tab. 3 and fig. 8). At the end of the last
glaciation (MIS 2), its presence in Europe decreased,
it being found sporadically at the sites of Vraona in
Greece, dated to between 17.075 ± 0.285 14C ka BP
and 7.07 ± 0.28 14C ka BP (Nagel, 1999), Triagalnata
Cave in Bulgaria, dated to 15.57 ± 0.31 14C ka BP
(Spassov & Raychev, 1997), Arene Candide layer
p 9 in Italy, dated to between 20.47 ± 0.02 14C ka BP
and 22.52 ± 0.37 ka BC (Bietti & Molari, 1994;
Cassoli & Tagliacozzo, 1994), the Magdalenian site
of Ettingen in Switzerland (von Koenigswald et al.,
2006), and Teufelsbrücke in Germany, assigned to
the Dryas I / Bølling complex (Musil, 1980; Hedges
1301-066 - Mep 1-2013.indd 43
et al., 1998), as well as the site of Predmostí in the
Czech Republic, dated to 26.1 ± 0.5 14C ka BP and
26.9 ± 1.6 14C ka BP (Musil, 1994).
These data show that the Iberian Peninsula seems to have
constituted the last refuge for the European P. pardus, for
it is a relatively common species in the faunal associations
of the sites of the Upper Palaeolithic (tab. 3 and fig. 9)
whereas in the rest of Europe its presence is only sporadic.
In the Iberian Peninsula it is found in Aurignacian, Perigordian, Solutrean, Magdalenian, Azilian and even Mesolithic levels (tab. 3 and fig. 9). The leopard is present in
the Aurignacian levels of the sites of Hornos de la Peña,
level E+D, dated to 20.93 ± 0.37 14C ka BP (de Barandiarán & Sonneville-Bordes, 1964), Cueva Morín, level 8,
dated to 28.43 ± 0.54 14C ka BP (Castaños, 1983; Maíllo
Fernández et al., 2001 ), Lezetxiki, levels III-IV (Altuna,
1972; Falguères et al., 2005 ), the Cave of “El Castillo”
(Cabrera Valdés, 1984), Isturitz (Altuna, 1972) and
Amalda, levels V and VI, dated to 19.90 ± 0.34 14C ka BP
and 27.4 ± 1.0 14C ka BP respectively (Klein & Cruz
Uribe, 1985; Altuna, 1990). P. pardus has also been
recorded during the Gravettian at the sites of Bolinkoba,
level VI (Yravedra Sainz de los Terreros, 2008), and
Casa da Moura, level 1b, dated to 25.09 ± 0.22 14C ka BP
26/02/13 10:50
44
Fig. 8: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Middle Palaeolithic).
PORTUGAL: 28/ Escoural, 29/ Figueira Brava, 30/ Prado des Salemas, 31/ Casa da Moura, 32/ Furinha, 33/ Caldeirão, 34/ Algar da Manga Larga,
35/ Lorga da Dine, UK: 36/ Gorham Cave (Gibraltar), 37/ Vanguard Cave (Gibraltar), SPAIN: 38/ Zafarraya, 39/ Carigüela, 40/ Cova Negra, 41/ Cova
Foradada, 42-45/ Pinarillo I, Pinilla del Valle, Cueva de la Zarzamora and Cueva del Búho respectively, 46/°Torrejones, 47/ Los Casares, 48/ Aguilón
P-7, 49/ Los Rincones, 50/ Ermita, 51/ Valdegoba, 52/ Cueva de “El Castillo”, 53-62/ Prado Vargas, Abrigo de Olha, Coscobilo, Ekain, Axlor, Arrillor,
Amalda, Cueva Allekoaitze, and Lezetxiki respectively, 62/ Gabasa, 63/ Les Muricers, 64/ Cova del Gegant, 65/ Abric Romaní, 66/ L’Arbreda, 67/ Ermitons, 68/ Mollet I; FRANCE: 69/ Grotte de la Carrière, 70/ Coupe-Gorge, 71/ La Crouzade, 72/ Tournal, 73/ L’Hortus, 74/ Caune de l’Arago, 75/ Jaurens,
76/ d’Artenac, 77/ Moula-Guercy, 78/ Chauvet, 79/ Grotte de l’Adaouste, 80-82/ Verzé, Blanot 2 and Etrigny respectively, 83/ La Grotte des Enfants;
MONACO: 84/ L’Observatoire, 85/ Grotte du Prince; ITALY: 86/ Madonna dell’Arma, 87/ Santa Lucia Superiore, 88/ Cave of Fate, 89/ Cave of Equi,
91/ Groticelle di Sambughetto Valstrona, 93/ Grotta de Fossellone, 94/ S. Agostino, 95/ Grotta di Castelcivita, 96/ Ingarano, 97/ Grotta di Fumane,
98/ Riparo Tagliente, 99/ Grotta di San Bernardino, 100/ Zandobbio, 123/ Caverna degli Orsi; SWITZERLAND: 92/ Cave of Cotencher; BELGIUM:
105/ Scladina cave; GERMANY: 90/ Rittersaal Höhle, 102/ Höhlenstein-Stadel, 103/ Geinsheim, 104/ Biedermann cave, 106/ Petershöhle, 107/ Teufelsbrücke, 108/ Taubach, 109/ Cave of Baumann, 110/ Niederlehme; AUSTRIA: 101/ Wildkirchli, 111/ Holzinger Höhle, 112/ Luegloch, 113/ Merkenstein,
116/ Große Peggauerwandhöhle, 117/ Kugelsteinhöhle, 118/ Große Badhöhle, 119/ Fünffenstergrotte, 120/ Ochsenhalt Höhle; CZECH REPUBLIC:
114/ Cave of Schwedentisch, 115/ Sveduv Stul; CROATIA: 121/ Kaprina, 122/ Veternica; BOSNIA AND HERZEGOVINA: 124/ Vjetrenichia;
SERBIA: 125/ Smolucka Cave; BULGARIA: 126/ Bacho Kiro; GREECE: 127/ Asprochaliko, 128/ Apidimia; TURKEY: 129/ Karaïn; GEORGIA:
130/ Mezmaiskaya, 132/ Bronzovaya Cave, 133/ Azokh I; RUSSIA: 131/ Akhstyrskaya Cave.
Fig. 8 : Aire de répartition de Panthera pardus en Europe au cours du Pléistocène supérieur (Paléolithique moyen).
Fig. 9: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Upper Palaeolithic).
PORTUGAL: 134/ Fontainhas,135/ Casa da Moura, 136/ Caldeirão; UK: 137/ Gorham Cave (Gibraltar); SPAIN: 138/ La Riera, 139-143/ Hornos de la
Peña, Ceuva de “El Castillo”, El Juyo, Cueva Morín and El Mirón respectively, 144/ Las Pajucas, 145/ La Blanca, 146-150/ Oyalkoba, Atxuri, Bolinkoba, Lezetxiki and Amalda respectively, 151/ Urtiaga, 153/ L’Arbreda, 154/ Cau de Coçes; FRANCE: 152/ Isturitz, 155/ L’aven du Charnier, 156/ La
Balauzière; ITALY: 157/ Arene Candide; SWITZERLAND: 158/ Ettingen, GERMANY: 159/ Teufelsbrücke; CZECH REPUBLIC: 160/ Predmostí;
BULGARIA: 161/ Triagalnata Cave; GREECE: 162/ Vraona.
Fig. 9 : Aire de répartition de Panthera pardus en Europe au cours du Pléistocène supérieur (Paléolithique supérieur).
1301-066 - Mep 1-2013.indd 44
20/02/13 12:33
45
(Valente, 2004). During the Solutrean it is present at
Caldeirao, with a radiocarbon age of 18-20 ka (Davis,
2002), at Bolinkoba, levels IV and V (Yravedra Sainz de
los Terreros, 2008), and at Gorham’s Cave (Stringer et al.,
2008). The leopard has also been identified in Magdalenian levels at the sites of Caldeirao, radiocarbon dated
to 10-16 ka (Davis, 2002), El Mirón 108 (Marín Arroyo,
2009), Urtiaga, level F, dated to 17.05 ± 0.14 14C ka BP
(Altuna, 1970; Castaños, 2005), L’Arbreda, levels B-C,
dated to 17.32 - 17.72 14C ka BP (Estévez Escalera, 1987),
Bolinkoba, level III (Castaños, 1983) and El Juyo, dated to
between 11.4 ± 0.3 14C ka BP and 13.29 ± 2.4 14C ka BP.
This latter is one of the most recent citations of P. pardus
in Europe. The leopard survived in Europe until the Holocene, although it became increasingly scarce (Sommer
& Benecke, 2006). In the Iberian Peninsula it is cited at
the Azilian site of La Riera (Vega del Sella, 1930, p. 38;
Altuna, 1972) and at the site of Las Pajucas in a possibly
Mesolithic level (Altuna, 1972).
7 - CONCLUSION
The mandible of Panthera pardus from Los Rincones
Ri10/C1/2010 shows morphometric similarities to
P. uncia, as it is the case with other specimens of leopard
from the Upper Pleistocene of southern Europe. The
leopards from southern Europe probably developed an
adaptation to mountainous environment, which distinguishes them from present-day leopards, yet without
having become a distinct species.
During the Pleistocene P. pardus distributed itself across
southern Europe. The Iberian Peninsula shows special
features in relation to the rest of the continent as regards
the distribution of P. pardus, for its first appearance
there was almost 700 ka later than its first appearance in
Europe. In Iberia we find it for the first time in level VI
of Lezetxiki, ESR-dated to 234 ± 32 ka, and there is no
further proof of its presence until the Upper Pleistocene,
when it is distributed throughout the whole of the Iberian
Peninsula and is found at more than forty sites, making
this one of the areas with the greatest density of sites with
P. pardus in Europe. Furthermore, the Iberian Peninsula
acted as a refuge for the leopard in the Upper Palaeolithic,
when it is found at nineteen sites, thirteen of which are
concentrated in the Basque-Cantabrian region. In the rest
of the continent, by contrast, its presence is sporadic, with
citations at only nine widely scattered sites. The climatic
conditions in Cantabria during the Upper Palaeolithic may
well have been favourable for the survival of this species.
It then went on to survive until the Mesolithic in the area,
this thus being the last citation of the species in Europe.
ACKNOWLEDGEMENTS
The Government of Aragon has partially subsidized
our geological activities in Los Rincones (Exp. 50/2006,
132/2010). This study forms part of the “H54, Grupos
Consolidados” project of the Government of Aragon and
1301-066 - Mep 1-2013.indd 45
European Social Fund, as well as of the Atapuerca project.
Víctor Sauqué is the beneficiary of a predoctoral grant
from the Government of Aragon. The CEA, in particular
Mario Gisbert, discovered and undertook the topography
of the cave Los Rincones. Thanks go to Julio GómezAlba of the Museu de Geología , to Eulàlia Garcia of
the Museo de Ciències Naturals de Barcelona, to Isabel
Cáceres of the archaeozoology laboratory of the Universitat Rovira i Virgili de Tarragona. We also thank to Jorge
Colmenar for his help taking the photographs. We would
also like to thank Olivier Moine, Pascale Ruffaldi, Alain
Argant and Patrick Auguste, reviewers, whose comments
improved the quality of this paper.
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