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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 et Gloria Cuenca‑Bescós
p. 13-24

Résumés

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.

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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, 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 Gómez-Alba 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 her 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.

1 - Introduction

1At 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 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

2The 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 clast-supported 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.

3The site was discovered by members of CEA (Centro Aragonés de Espeleología) in 2005 while they were mapping the cave. The cave of Los Rincones is described by the CEA members Gisbert and 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.

Fig. 1: Geographical location of the Los Rincones cave.

Fig. 1: Geographical location of the Los Rincones cave.


Fig. 2: Elevation and plan views of the Los Rincones cave.

Fig. 2: Elevation and plan views of the Los Rincones cave.

3 - Material and methods

4Studied 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.

5The 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 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).

6The 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.

7For 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.

8Abbreviations

9Coll: 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.

4 - Systematic paleontology

  • 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)

4.1 - Material and measurements

10Description: The right mandible from Los Rincones (Ri10/C1/2010), attributed to P. pardus on the basis of its morphology and measurements (tab. 1 & 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 well-preserved coronoid, articular and angular apophyses.

11In 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.

12In 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.

13The 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 semi-cylindrical, anteroposteriorly narrow and lateromedially elongated.

14In 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.

15The canine belonging to (Ri10/C1/2010) has a relatively rounded transverse section, and presents one carina in the lingual distal edge.

16The 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.

17The 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.

18

Tab. 1: Measurements of mandible and teeth of Panthera pardus from Los Rincones (Ri10/C1/2010).

Tab. 1: Measurements of mandible and teeth of Panthera pardus from Los Rincones (Ri10/C1/2010).

19

Fig. 3: Fossil remains of Panthera pardus from Los Rincones (Ri10/C1/2010).

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

4.2 - Discussion

20The mandible from Los Rincones (Ri10/C1/2010) shows morphological similarities to present-day snow leopard, Panthera uncia. As in this species, moreover, 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.

21The 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 & 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).

22The 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).

23In dorsal view, the p4 from Los Rincones (Ri10/C1/2010) presents an oval outline, with a slight double-pinched 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).

24The 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).

25In 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).

26The 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 univariate representation of this character, it can be seen that Ri10/C1/2010 falls within the range of variation of P. uncia (tab. 2 & 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).

27The 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 & fig. 5). Low robustness indices are a consequence of the relative elongation of m1 in relation to the mandible (Testu, 2006).

28The 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.

29The 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).

Tab. 2: Measurements (in mm) of mandibles and teeth of Panthera pardus and Panthera uncia.

Tab. 2: Measurements (in mm) of mandibles and teeth of Panthera pardus and Panthera uncia.

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: 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. 5: Box-plot representing the robustness index (height of mandibular body posterior to m1 versus DMD m1 x 100).

Fig. 5: Box-plot representing the robustness index (height of mandibular body posterior to m1 versus DMD m1 x 100).

Fig. 6: Graph representing length p3-m1 versus the total length of the mandible.

Fig. 6: Graph representing length p3-m1 versus the total length of the mandible.

5 - The palaeontological record of p. pardus-p. uncia in Europe

30The 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).

31Other authors have described P. pardus in the south of Europe with characters similar to those of P. uncia, 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 Pleistocene, 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.

32By 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).

33On the other hand, the microfauna 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 characterized 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

34Most 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).

35The 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 & fig. 7). 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).

36The 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 & fig. 8). At the end of the last glaciation (MIS 2), its presence in Europe decreased, it being found sporadically at the sites of Olivier2017-10-19T12:02:00OVraona 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 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).

37These 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 & 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 & fig. 9). The leopard is present in the Aurignacian levels of the sites of Hornos de la Olivier2017-10-19T12:02:00OPeña, 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 (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).

38

Fig. 7: Distribution area of Panthera pardus in Europe during the Lower and Middle Pleistocene.

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/ Aze 1-5, 8/ Lazaret, 9/ Vallonet; ITALY: 10/ Valdemino Cave, 11/ Romagnano, 12/ Grotta di Cere, 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/ Repolusthohle, 21/ Hundsheim, HUNGARY: 22/ Tarko, CZECH REPUBLIC: 23/ Stranska Skala, 25/ Koneprusy C718; POLAND: 24/ Biśnik Cave; GERMANY: 26/ Mauer 5, 27/ Mosbach 2.

Tab. 3: Localities with Panthera pardus in Europe.

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.

Fig. 8: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Middle Palaeolithic).

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/ Caldeirao, 34/ Algar da Manga Larga,35/ Lorga da Dine, UK: 36/ Gorham Cave (Gibraltar), 37/ Vanguard Cave (Gibraltar), SPAIN: 38/ Zafarraya, 39/ Cariguela, 40/ Cova Negra, 41/ Cova Foradada, 42-45/  Pinarillo I, Pinilla del Valle, Cueva de la Zarzamora and Cueva del Buho respectively, 46/°Torrejones, 47/ Los Casares, 48/ Aguilon 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 Romani, 66/ L’Arbreda, 67/ Ermitons, 68/ Mollet I; FRANCE: 69/ Grotte de la Carriere, 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/ Verze, 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 Hohle, 102/ Hohlenstein-Stadel, 103/ Geinsheim, 104/ Biedermann cave, 106/ Petershohle, 107/ Teufelsbrucke, 108/ Taubach, 109/ Cave of Baumann, 110/ Niederlehme; AUSTRIA: 101/  Wildkirchli, 111/ Holzinger Hohle, 112/ Luegloch, 113/ Merkenstein, 116/  Grose Peggauerwandhohle, 117/ Kugelsteinhohle, 118/ Grose Badhohle, 119/ Funffenstergrotte, 120/ Ochsenhalt Hohle; 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/ Karain; GEORGIA: 130/ Mezmaiskaya, 132/ Bronzovaya Cave, 133/ Azokh I; RUSSIA: 131/ Akhstyrskaya Cave.

Fig. 9: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Upper Palaeolithic).

Fig. 9: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Upper Palaeolithic).

PORTUGAL: 134/ Fontainhas,135/ Casa da Moura, 136/ Caldeirao; UK: 137/ Gorham Cave (Gibraltar); SPAIN: 138/ La Riera, 139-143/ Hornos de la Pena, Ceuva de “El Castillo”, El Juyo, Cueva Morin and El Miron respectively, 144/ Las Pajucas, 145/ La Blanca, 146-150/ Oyalkoba, Atxuri, Bolinkoba, Lezetxiki and Amalda respectively, 151/ Urtiaga, 153/ L’Arbreda, 154/ Cau de Coces; FRANCE: 152/ Isturitz, 155/ L’aven du Charnier, 156/ La Balauziere; ITALY: 157/ Arene Candide; SWITZERLAND: 158/ Ettingen, GERMANY: 159/ Teufelsbrucke; CZECH REPUBLIC: 160/ Predmosti; BULGARIA: 161/ Triagalnata Cave; GREECE: 162/ Vraona.

7 - Conclusions

39The 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.

40During 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.

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Table des illustrations

Titre Fig. 1: Geographical location of the Los Rincones cave.
URL http://quaternaire.revues.org/docannexe/image/6468/img-1.jpg
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Titre Fig. 2: Elevation and plan views of the Los Rincones cave.
URL http://quaternaire.revues.org/docannexe/image/6468/img-2.png
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Titre Tab. 1: Measurements of mandible and teeth of Panthera pardus from Los Rincones (Ri10/C1/2010).
URL http://quaternaire.revues.org/docannexe/image/6468/img-3.png
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Titre Fig. 3: Fossil remains of Panthera pardus from Los Rincones (Ri10/C1/2010).
Légende Buccal (A), lingual (B) and occlusal (C) views of the right hemi-mandible
URL http://quaternaire.revues.org/docannexe/image/6468/img-4.png
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Titre Tab. 2: Measurements (in mm) of mandibles and teeth of Panthera pardus and Panthera uncia.
URL http://quaternaire.revues.org/docannexe/image/6468/img-5.png
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Titre 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).
URL http://quaternaire.revues.org/docannexe/image/6468/img-6.png
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Titre Fig. 5: Box-plot representing the robustness index (height of mandibular body posterior to m1 versus DMD m1 x 100).
URL http://quaternaire.revues.org/docannexe/image/6468/img-7.png
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Titre Fig. 6: Graph representing length p3-m1 versus the total length of the mandible.
URL http://quaternaire.revues.org/docannexe/image/6468/img-8.png
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Titre Fig. 7: Distribution area of Panthera pardus in Europe during the Lower and Middle Pleistocene.
Légende UK: 1/ Pontnewydd Cave, 2/ Bleadon Cavern; SPAIN: 3/ Lezetxiki; FRANCE: 4/ Caune de l’Arago, 5/ Lunel-Viel, 6/ Orgnac 3, 7/ Aze 1-5, 8/ Lazaret, 9/ Vallonet; ITALY: 10/ Valdemino Cave, 11/ Romagnano, 12/ Grotta di Cere, 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/ Repolusthohle, 21/ Hundsheim, HUNGARY: 22/ Tarko, CZECH REPUBLIC: 23/ Stranska Skala, 25/ Koneprusy C718; POLAND: 24/ Biśnik Cave; GERMANY: 26/ Mauer 5, 27/ Mosbach 2.
URL http://quaternaire.revues.org/docannexe/image/6468/img-9.png
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Titre Tab. 3: Localities with Panthera pardus in Europe.
Légende 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.
URL http://quaternaire.revues.org/docannexe/image/6468/img-10.png
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Titre Fig. 8: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Middle Palaeolithic).
Légende PORTUGAL: 28/ Escoural, 29/ Figueira Brava, 30/ Prado des Salemas, 31/ Casa da Moura, 32/ Furinha, 33/ Caldeirao, 34/ Algar da Manga Larga,35/ Lorga da Dine, UK: 36/ Gorham Cave (Gibraltar), 37/ Vanguard Cave (Gibraltar), SPAIN: 38/ Zafarraya, 39/ Cariguela, 40/ Cova Negra, 41/ Cova Foradada, 42-45/  Pinarillo I, Pinilla del Valle, Cueva de la Zarzamora and Cueva del Buho respectively, 46/°Torrejones, 47/ Los Casares, 48/ Aguilon 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 Romani, 66/ L’Arbreda, 67/ Ermitons, 68/ Mollet I; FRANCE: 69/ Grotte de la Carriere, 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/ Verze, 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 Hohle, 102/ Hohlenstein-Stadel, 103/ Geinsheim, 104/ Biedermann cave, 106/ Petershohle, 107/ Teufelsbrucke, 108/ Taubach, 109/ Cave of Baumann, 110/ Niederlehme; AUSTRIA: 101/  Wildkirchli, 111/ Holzinger Hohle, 112/ Luegloch, 113/ Merkenstein, 116/  Grose Peggauerwandhohle, 117/ Kugelsteinhohle, 118/ Grose Badhohle, 119/ Funffenstergrotte, 120/ Ochsenhalt Hohle; 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/ Karain; GEORGIA: 130/ Mezmaiskaya, 132/ Bronzovaya Cave, 133/ Azokh I; RUSSIA: 131/ Akhstyrskaya Cave.
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Titre Fig. 9: Distribution area of Panthera pardus in Europe during the Upper Pleistocene (Upper Palaeolithic).
Légende PORTUGAL: 134/ Fontainhas,135/ Casa da Moura, 136/ Caldeirao; UK: 137/ Gorham Cave (Gibraltar); SPAIN: 138/ La Riera, 139-143/ Hornos de la Pena, Ceuva de “El Castillo”, El Juyo, Cueva Morin and El Miron respectively, 144/ Las Pajucas, 145/ La Blanca, 146-150/ Oyalkoba, Atxuri, Bolinkoba, Lezetxiki and Amalda respectively, 151/ Urtiaga, 153/ L’Arbreda, 154/ Cau de Coces; FRANCE: 152/ Isturitz, 155/ L’aven du Charnier, 156/ La Balauziere; ITALY: 157/ Arene Candide; SWITZERLAND: 158/ Ettingen, GERMANY: 159/ Teufelsbrucke; CZECH REPUBLIC: 160/ Predmosti; BULGARIA: 161/ Triagalnata Cave; GREECE: 162/ Vraona.
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Víctor Sauqué et Gloria Cuenca‑Bescós, « The Iberian Peninsula, the last European refugium of panthera pardus linnaeus 1758 during the Upper Pleistocene », Quaternaire, vol. 24/1 | 2013, 13-24.

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Víctor Sauqué et Gloria Cuenca‑Bescós, « The Iberian Peninsula, the last European refugium of panthera pardus linnaeus 1758 during the Upper Pleistocene », Quaternaire [En ligne], vol. 24/1 | 2013, mis en ligne le 01 mars 2016, consulté le 23 novembre 2017. URL : http://quaternaire.revues.org/6468 ; DOI : 10.4000/quaternaire.6468

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Víctor Sauqué

Grupo Aragosaurus-IUCA. Paleontología. Facultad de Ciencias. Universidad de Zaragoza. C/ Pedro Cerbuna 12. E-50009 Zaragoza. Email: vsauque@unizar.es;

Gloria Cuenca‑Bescós

Grupo Aragosaurus-IUCA. Paleontología. Facultad de Ciencias. Universidad de Zaragoza. C/ Pedro Cerbuna 12. E-50009 Zaragoza. Email: cuencag@unizar.es;

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