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Quaternary International 276-277 (2012) 61e76

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Quaternary International
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Mammuthus primigenius in the cave and portable art: An overview with a short account on the elephant fossil record in Southern Europe during the last glacial
Ingmar M. Braun a, *, Maria Rita Palombo b, c
a

Wyhlenweg 4, CH e 4126 Bettingen, Switzerland Dipartimento di Scienze della Terra, Universit degli Studi di Roma La Sapienza, Roma, Italy c CNR, Istituto di Geologia Ambientale e Geoingegneria, Montelibretti, Roma, Italy
b

a r t i c l e i n f o
Article history: Available online 20 July 2012

a b s t r a c t
A rich Upper Paleolithic iconography testies to a long coexistence of humans and Mammuthus primigenius during the last glacial in most of Europe, including northern Spain, and supplies additional information for a better understanding of the dispersion and last occurrence of woolly mammoths in southernmost Europe (i.e. in the Iberian and Italian peninsulas) during this time. In Italy, where the scanty M. primigenius ndings are likely not younger than 38 ka (except for the Gravettian remains from the Arene Candide cave, eastern Liguria), no representations of woolly mammoths have been reported to date. An exception is the carved mammoth objects (a few Gravettian ornaments and female gurines), recorded in Ligurian sites, but the hypothesis that they could have been imported from some distant area cannot be ruled out. Conversely, in Spain along the northern Atlantic coast, M. primigenius remains have been found in some sites yielding mammoth representations. In southern Spain, where M. primigenius was present in the Padul basin (Granada) during most of MIS 3 (between 40.4 and 30.6 cal ka BP), artistic representations of woolly mammoths are unknown. As regard to Palaeoloxodon, some populations were present during the late MIS 3 in the Iberian Peninsula as in Western Europe, whereas no sound data support the persistence of straight-tusked elephants on mainland during MIS 2. Therefore, whether the intriguing elephant painting of the Spanish El Castillo cave could represent a straight-tusked elephant e suggesting a survival of the species in Northern Spain during the Last Glacial Maximum e or an unusual representation of a woolly mammoth, still remains an unanswered question. 2012 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction The iconic elephant of the last glacial, the woolly mammoth Mammuthus primigenius, is a species widely spread during the Late Pleistocene at the middle and northern latitudes of Eurasia (Fig. 1) as well as in North America, where it may have originated (Debruyne et al., 2008). In Eurasia its range reached its southernmost maximum extension during MIS 3a, when the woolly mammoth is recorded at 36 350 N in China (Jinan, Shandong Province) (Takahashi et al., 2007), while the southernmost records of M. primigenius in Western Europe are those of Padul (37 010 N, Granada Basin, Spain) (lvarez-Lao et al., 2009) and Cardamone (40 210 N, Apulia, Italy) (Rustioni et al., 2003). During the late glacial, in particular during the Last Glacial Maximum (LGM), the distribution of M. primigenius ndings across Europe (Fig. 1) only partially matches that of vestiges of the Upper Paleolithic art, left by

* Corresponding author. E-mail address: IngmarBraun@gmx.ch (I.M. Braun). 1040-6182/$ e see front matter 2012 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2012.07.010

our direct ancestor Homo sapiens, from about 35 ka to 11 ka BP (Fig. 2). This evidence is cave art products, known especially in Western Europe, and portable art objects which are documented all over Europe and as far as Siberia. The cave art includes gurative representations on walls and ceilings of caves and rock shelters as well as those on cave oors (e.g. engravings on clay ground). Objects of portable art are engravings of animals, rarely of humans, and various other representations (e.g. dots, lines) carved on stones, ivory, bones, reindeer antlers, as well gurines of animals and humans. Although the gures of woolly mammoths do not dominate among the zoomorphic Paleolithic representations, M. primigenius was depicted by Paleolithic artists using a variety of different materials and techniques. The paper aims to present a short overview of the main depictions of M. primigenius thus far known in the Eurasian Paleolithic art, and to discuss some intriguing issues such as the perplexing representation of the El Castillo elephant, and the lack of any representation in Italy, where the woolly mammoth is recorded denitively not later than MIS 3.

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Fig. 1. Map showing the distribution of the main Late Pleistocene sites with woolly mammoth remains (modied and updated from Markova et al., 2010).

2. M. primigenius in the portable and cave art: an overview The rst discovery of a M. primigenius representation dates back to the second half of the 19th century, when, in 1864 Edouard Lartet and Henry Christy, during the archaeological excavation of the Abri de La Madeleine (Dordogne, France), found a piece of mammoth ivory with the engraving of a woolly mammoth (Fig. 3.1 and 3.2). At that time, this nding was regarded as proof that prehistoric men and woolly mammoths lived contemporaneously and interacted with each other: This new fact will not, indeed, add anything to already acquired convictions as to the coexistence of Man with the fossil Elephant (Elephas primigenius) and other great Herbivores and

Carnivores which geologists regard as having lived together in the early phases of the Quaternary Period (Lartet and Christy, 1875, p. 207). Today, the Paleolithic art of Eurasia counts at least 561 representations of M. primigenius known from about 76 sites (Berdin, 1970; Bosinski and Fischer, 1980; Bellier et al., 1999; Gly and Azma, 2005) (Fig. 2). This number likely underestimates the actual number of mammoth paintings, gurines and engravings as in some site investigations are still in progress (e.g. in France the caves of Chauvet, Ardche, and Cussac, Dordogne), old published data might be incomplete, and unknown mammoth representations might exist in either private or public collections.

Fig. 2. Map showing the distribution of selected Upper Paleolithic sites with the main representations (cave and portable art) of Mammuthus primigenius (modied and updated from Bosinski and Fischer, 1980). 1: Las Caldas; 2: Pindal; 3: El Castillo; 4: Arco B cave; 5: Los Casares; 6: Isturitz; 7: Gargas; 8: Les Trois-Frres; 9: Canecaude I; 10: La BaumeLatrone; 11: Bayol; 12e14: Oulen, Le Figuier, Chabot; 15e16: Chauvet, Ebbou; 17: Bruniquel, Abri Montastruc; 18e20: Pech-Merle, Cougnac, Roucadour; 21e24: Cussac, Le Pigeonnier, Le Mammouth, Cavaille; 25e33: Croze--Gontrand, La Mouthe, Cournazac, La Grze, Bernifal, Les Combarelles, Font-de-Gaume, Laugerie-Haute, La Madeleine; 34e36: Raymonden, Roufgnac, Labattut; 37e39: Les Vachons, Les Bernous, Jovelle; 40: Pair-non-Pair; 41: Chanlat; 42e43: La Marche-Rseau Guy Martin, La Marche cave; 44e45: Le Cheval, Grande Grotte dArcy-sur-Cure; 46: Solutr; 47: La Colombire; 48: Trou de Chaleux; 49e50: Gnnersdorf, Andemach; 51e52: Vogelherd, Geissenklsterle; 53: Obere stonice, Pavlov; 56: P Klause; 54e55: Doln Ve redmost; 57: Avdeevo; 58: Sungir; 59e61: Kostienki 1, Kostienki 4, Kostienki 11; 62: Kapova.

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Fig. 3. The rst discovered representation of a woolly mammoth found in the Abri de La Madeleine (Dordogne, France) by E. Lartet and H. Christy in 1864 (modied from Lartet and Christy, 1875) (above). A drawing of the engraving of the mammoth of the Abri de La Madeleine modied from Roussot, 2002) (below).

Although in the cave and portable art animals were often depicted in a detailed, realistic way, some representations consist of schematic outlines and the species can only be recognised because the silhouette highlights some specic, diagnostic characteristics. Concerning woolly mammoths, Bellier et al. (1999, p.108) wrote: [.] le mammouth est essentiellement voqu par sa silhouette caractristique, marque par un massif frontal lourd et puissant, se prolongeant vers le bas par la masse de la trompe, par la ligne cervicodorsale soulignant les bosses de la tte et du dos, spares par la dpression nuchale, et par laffaissement de larrire-train. Les dtails tels que les pattes, la toison laineuse ou les dfenses ne sont souvent pas indiqus. 2.1. The mammoth in portable art Figurines representing woolly mammoths are especially frequent in during the early and middle Upper Paleolithic (Aurignacian and Gravettian). Most engravings and tools decorated with mammoths date back to the late Upper Paleolithic (Magdalenian). 2.1.1. Figurines of woolly mammoths The oldest portable art objects thus far known in Eurasia are the ivory gurines, dating back to the Aurignacian (ca. 35 to 30 ka BP), found in the Lone and Ach valleys in the Swabian Alps (BadenWrtemberg, South West Germany). In the Lone valley, the Vogelherd cave, completely excavated in 1931 by Riek (1934) yielded some important ndings, including quite small ivory gurines of various animals (Hahn, 1986). Among those representing woolly mammoths there is a nearly complete gurine (5 cm long, 2.2 cm wide, 3.1 cm high) (Fig. 4.1), found in horizon V, that was carved on the core of a mammoth tusk, which shows X signs, and alignments of dots and grooves, decorating the mammoth body (Bosinski, 1982). A large fragment of ivory, representing the backside and the feet of a mammoth found at the same site, was also decorated with different signs (Bosinski, 1982; Hahn, 1986). Of particular interest is a perforated oval bone, regarded as a pendant, showing a demi-relief of a mammoth (Bosinski, 1982).

Since 2005, a team of the University of Tbingen has been reexploring the Vogelherd cave. The most important nding of these new investigations is represented by a complete gurine discovered in 2006 (Fig. 4.2). With a total length of 3.7 cm, it is the smallest gurine of a woolly mammoth thus far recorded from the Swabian Alps and the only complete among the 20 Aurignacian gurines of animals recorded in the Lone and Ach valleys. In this gurine of Vogelherd cave, the soles of the feet are decorated with ne crossing lines, and six ne lines are also engraved on the top of the head (Conard, 2009). In the Ach valley, archaeological investigations were made by the University of Tbingen from 1973 to 2002 in the Geissenklsterle (surroundings of Blaubeuren). The objects of portable art are of great interest, albeit less numerous than those found in the Vogelherd cave. More than 40 pieces of ivory were found, belonging to a nearly complete gurine of a mammoth, about 7 cm long and carved in the inner part of a tusk. Although the gurine is fragmentary and not perfectly preserved, the shape clearly indicates a woolly mammoth. As are the other gurines of the Swabian Alps, it is decorated with different signs as well as with points of red ochre, as shown by microscopic investigations (Hahn, 1986, 1988). In the middle Upper Paleolithic (Gravettian, from about 28 to 22 ka BP), the gurines of woolly mammoths became more numerous. Most are reported from open air sites in Moravia (Czech Republic) and in Eastern Europe (Russia and Ukraine). Conversely to the Aurignacian gurines from the Swabian Alps, these gurines do not show any geometric sign and were carved not only in ivory but also in bones, clay, and stone. In the Pavlovian of Moravia, female and animal gurines (some representing the woolly mammoth), formed in clay then burned, are known from the sites stonice and Pavlov (Valoch and Lzni of Doln Ve ckov-Galetov, 2009) (Fig. 5). The use of clay for the fabrication of portable art objects is only known from the Pavlovian sites. At Pavlov, as well at P redmost (Moravia), an ivory gurine of a woolly mammoth was also found (Valoch and Lzni ckov-Galetov, 2009) (Fig. 6). More eastward, about 40 gurines of M. primigenius were discovered in some sites situated in the Russian plain, such as those of the Kostienki area, and Avdeevo, Eliseevitchi, and Sungir (Abramova, 1995). The animal and female gurines recorded from this area date to the so-called Kostenkian, corresponding to the European late Gravettian (from about 24 to 22 ka BP) (Djindjian et al., 1999). In the Kostienki area, some open air sites are located along the Don River north of the Black Sea. The gurines of mammoth from these sites are all marly representations (Abramova, 1995), which clearly show the characteristic shape of M. primigenius (Fig. 7), as shown by the only woolly mammoth gurine found in the open air site of Eliseevitchi (Abramova, 1995). Interesting objects were also found in the open air site at Avdeevo. The sample includes two mammoth gurines carved on sandstone (Fig. 8.2), and one in a spongeous bone, probably a vertebra of mammoth (Fig. 8.1). The shape of the later is similar to that of gurines from P redmost (Moravia) (Gvozdover, 1995). An ivory gurine regarded as a woolly mammoth was discovered in a completely different context, a richly equipped double tomb of two children, at Sungir (Bogoljubova, outskirts of Vladimir city, Russia) (Abramova, 1995). The most eastward site yielding a mammoth representation is the Siberian site Ust Kova, where a gurine carved in ivory was found (Abramova, 1995). 2.1.2. Engravings of mammoths Engravings of mammoths in portable art are especially carved on stone, but also on other materials. In some engravings, it is difcult to decode the mammoth representations because of lines or other animal gures overlying them. Except for four sites dating

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Fig. 4. 1) Figurine of a woolly mammoth found by G. Riek in the Vogelherd cave (Baden-Wrttemberg, Germany) in 1931 (modied from Floss, 2009). 2) Figurine of mammoth found in the Vogelherd cave (Baden-Wrttemberg, Germany) in 2006 (modied from Conard, 2008).

to the middle Upper Paleolithic (see Bosinski and Fischer, 1980), most of engravings known thus far in portable art date to the late Upper Paleolithic. During the Magdalenian (from about 18 to 12.5 ka), their geographical distribution is mainly restricted to Western and Central Europe. The best known representation is that on a fragment of a tusk found in the Abri de La Madeleine (southwestern France). The greatest number of mammoth engravings (76 representations) was found in the Magdalenian open air site of Gnnersdorf near Koblenz (Rheinland-Pfalz, Germany) (Fig. 9), where mammoths, various animals and schematic female gurines were engraved on slate plates (see Bosinski, 2008; Bosinski et al., 2001). Adult mammoths and also young individuals were represented both alone and in groups. Although the representations are very

detailed, all mammoths of Gnnersdorf are tuskless. Following Guthrie (in Bosinski, 1984) the tusklessness would affected the Gnnersdorf mammoth population as a result of starvation caused by low vegetation productivity at the end of the late glacial period. Bellier et al. (1999) believed that the representations of Gnnersdorf document the latest population of mammoths in Western Europe. The southernmost site yielding engravings of woolly mammoths likely is Los Casares (Guadalajara, northern Spain) (Angulo and Moreno, 2011). The westernmost site is Las Caldas (Asturias, northern Spain), where an engraving on a stone plate was found in a Middle Magdalenian context (Corchn, 1991e1992). Molariform teeth of M. primigenius have been dated to 21.8 cal ka BP (lvarezLao and Garca, 2012). Eastwards, mammoth engravings are recorded in Siberia, where an ivory perforated plaquette is recorded from the open air site of Malta near Irkutsk (Fig. 10), and a representation craved on a tusk of a young mammoth was found at Berelkh A (Sakha Republic, Yakutia) (Abramova, 1995). The later is the most northern nding of a woolly mammoth engraving known. With its overlong legs, the very long trunk and the proportionally small body, this

stonice and Fig. 5. eFigurines of Mammuthus primigenius in burned clay from Doln Ve Pavlov in the Moravian region, Czech Republic (modied from Valoch and Lzni ckovGaletov, 2009).

Fig. 6. Ivory gurine of Mammuthus primigenius from P redmost in the Moravian region, Czech Republic (modied from Absolon and Klima, 1977).

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Fig. 7. Two marly gurines of Mammuthus primigenius from Kostienki 1, Russia (modied from Bosinski, 1990).

Fig. 9. Engraving of Mammuthus primigenius on a slate plaquette from Gnnersdorf (Rheinland-Pfalz, Germany) (modied from Bosinski and Fischer, 1980).

representation differs clearly from the other engravings of woolly mammoth known in Eurasia (Fig. 11). 2.1.3. Tools decorated with woolly mammoths In contrast to the gurines and engravings, only a few tools decorated with mammoths are known to date, mostly from France. A fragment of a bton perc, found in the Protomagdalenian layer of the Abri Laugerie-Haute (Dordogne, France), shows two deeply

engraved mammoths, facing each other and touching their heads (Peyrony and Peyrony, 1938), which were regarded by Bosinski and Fischer (1980) as two ghting animals (Fig. 12). Two Magdalenian spear-throwers in the shape of a mammoth were found in France at Canecaude I (Aude) and at the Abri Montastruc (Tarn-et-Garonne). The rst, perfectly reproducing the silhouette of a woolly mammoth (Fig. 13), was carved on an antler of the reindeer (Sacchi, 1986). The second, more schematic, was discovered, with other Magdalenian objects, in the second half of the 19th century (Sieveking, 1987) (Fig. 14). Two Magdalenian perforated bone disks, supposed to be ornaments (Bellier et al., 1991) but whose actual use is still unclear, have been found at Raymonden (Dordogne, France) and Trou de Chaleux in Belgium. The rst disk shows engravings of a mammoth on both sides (Fig.15), whereas on the second only a gure of a mammoth is present.

2.2. The mammoth in cave art In the cave art, the representations of animals dominate, especially those of herbivores. About 6.3% of the gurative representations depict woolly mammoths (Tosello and Fritz, 2006), and have been recorded from 46 caves, 15% of all sites with cave art vestiges. Horses are the most frequently represented animals, followed by bison and mammoths (Gly and Azma, 2005). In northern Spain, deer representations are much more frequent than those of M. primigenius. Caves showing the greatest number of mammoth representations are the Roufgnac cave (Dordogne, France) with 158 gures (Plassard, 1999) followed by the Chauvet cave (Ardche, France) with 76 specimens (Gly and Azma, 2005). Only a few representations of mammoth are known from the Iberian Peninsula, while none has been discovered in Italy so far.

Fig. 8. eBone (1) and sandstone (2a,b,c) gurine of Mammuthus primigenius from Avdeevo, Russia (modied from Gvozdover, 1995).

Fig. 10. Engraving of Mammuthus primigenius on a perforated plaquette of ivory from Malta near Irkutsk, Russia (modied from Abramova, 1995).

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2.2.1. Discovery of engravings in the Chabot cave In 1878, Lopold Chiron discovered some engravings on the walls of the Chabot cave (Gard, France) that he published in 1889, providing, for the rst time, a graphic and photographic documentation of paleolithic cave art and identifying some of the Chabot engravings as birds and humans. Chiron did not realize the old age of the representations of Chabot cave (Aujoulat, 1987), some of which nowadays are known as depicting woolly mammoths (Combier, 1984a) (Fig. 16.1 and 16.2). 2.2.2. The cave art In Europe, vestiges of Paleolithic cave art are recorded in about 300 caves, whose geographical distribution is much more limited than that of the portable art. Most are located in Western Europe, in France and in Spain. A dozen caves with cave art are known in Italy, especially from Sicily (Roussot, 2002). In 2003, caves with Paleolithic cave art were discovered for the rst time in Great Britain (Bahn and Pettitt, 2009) and in 2009 in Romania (Besesek et al., 2010). The caves of Kapova and Ignatievka, both in the Southern Ural Mountains, are the most  eastward caves with cave art known to date (S celinskij and  Sirokov, 1999). 2.2.2.1. Techniques in cave art: painting, engraving, and sculpture. Three main techniques were used in cave art: painting, engraving and sculpture. In most caves, there are gures made by means of different techniques. Painting, engraving and sculpture can be present in each cave in combination or singularly. In many cases the Paleolithic artists either modied or integrated the natural relief of the walls in their art works. The Paleolithic painting is based on three fundamental colours: yellow, red and black. Other nuances of colour could be achieved by mixing these colours. Colours were often obtained from mineral components. The yellow colour was obtained from either limonite or goethite, the red from ferric oxides, especially hematite, while the black colour used manganese oxide and charcoal. Pigments

Fig. 11. The most northern depiction of Mammuthus primigenius from Berelkh A in the Sakha Republic, Russia (modied from Bosinski and Fischer, 1980).

Fig. 12. Bton perc with two depicted wolly mammoths from Laugerie-Haute (Dordogne, France) (modied from Bosinski, 1990).

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Fig. 15. Bone disk with engravings of Mammuthus primigenius on each side from Raymonden (Dordogne, France) (modied from Bosinski and Fischer, 1980).

(Ardche, France) are represented as both black silhouettes, and paintings (Gly and Azma, 2005). Of particular interest are the strange representations of mammoths (Fig. 18) and other animals found at the Baume-Latrone cave (Gard, France), which were drawn using clayey material (Louis and Drouot, 1953). Engraving is a technique of abrasion, performed with the aid of a pointed object, in general a int tool, used to carve on the rock

Fig. 13. Spear thrower in the shape of Mammuthus primigenius from Canecaude I (Aude, France) (modied from Sacchi, 1986).

were used to draw outlines or to paint the whole gure. The representations could be either monochrome, bichrome, or polychrome. Interesting silhouettes of woolly mammoths, drawn more   frequently in black than in red (Bosinski in S celinskij and Sirokov, 1999), are present in the Roufgnac (Fig. 17) (see Barrire, 1982; Plassard, 1999) and in Pech-Merle caves (Lot, France) (Lorblanchet, 2010). Red painted representations are present in the Grande Grotte of Arcy-sur-Cure (Yonne), Oulen (Gard), and Cougnac caves (Lot), in Spain in the Pindal cave (Asturias) and in Russia in the caves of Kapova and Ignatievka (Combier, 1984b; Bafer and Girad,   1998; S celinskij and Sirokov, 1999; Saura and Mzquiz, 2007; Lorblanchet, 2010). The woolly mammoths in the Chauvet cave

Fig. 14. Spear thrower in the shape of Mammuthus primigenius from Abri Montastruc (Tarn-et-Garonne, France) (modied from Sieveking, 1987).

Fig. 16. Engraving (1) of Mammuthus primigenius and other lines in the Chabot Cave (Gard, France) (modied from Bosinski, 1999), and its scientic drawing (2) (modied from Roussot, 2002).

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Fig. 17. Outline of a realistic representation of Mammuthus primigenius from the Roufgnac cave (Dordogne, France) (Photo and Jean Plassard).

wall one or more, ne or very deep lines (Fig. 19). The lines drawn on soft rocks with ngers are called nger utings. The scraping technique (raclage in French) is another type of abrasion performed to obtain a kind of chromaticity by scraping the cave wall. Engravings of mammoths are fairly frequent in the cave art. Several of such congurations are present in the French caves of Roufgnac, Les Combarelles I (Dordogne), Font-de-Gaume (Dordogne), Jovelle (Dordogne), Chauvet (Ardche), Cussac (Dordogne), Gargas (Haute-Garonne) and Les Trois-Frres (Arige) (Capitan et al., 1910; Bgoun and Breuil, 1958; Barrire, 1976, 1982, 1993, 1997; Delluc and Delluc, 1991; Plassard, 1999; Gly and Azma, 2005). In the Chauvet cave there are the only representations of woolly mammoths made using the scraping technique (Gly and Azma, 2005), while some others were engraved with the ngers into the soft cave walls. A very deep technique of abrasion is the sculpture. In this case the gures are strongly raised from the plane of the cave wall, in a plastic way (demi-relief). The sculpture technique is known from a few sites, especially from rock shelters in southwestern France. The only sculpted representation of mammoth was discovered in the Grotte du Mammouth (Dordogne) (Delluc and Delluc, 1983) (Fig. 20). 2.3. Archaic and realistic representations of mammoth in cave art The mammoth was represented in the caves either as an individual animal or groups. The latter are present, for instance, in the Roufgnac cave where numerous animals were depicted close

Fig. 19. Engraving of Mammuthus primigenius from the Roufgnac cave (Dordogne, France) (Photo and Jean Plassard).

together as if they were interacting (see Barrire, 1982; Plassard, 1999). Delluc and Delluc (2004a, 2004b) described two styles of representations: a realistic and an archaic style. There are also the dessins cursifs where the trunk, the head and the back of the mammoth were drawn in a continuous, single line. In a realistic representation, woolly mammoths are depicted in a very detailed way, a number of anatomical details were represented such as eyes, ears, tusks, the fur, tail and even the nger at the end of the trunk (Fig. 17). In most of these realistic representations, usually dating to the middle or late Magdalenian, a number of individuals are represented, apparently interacting. Realistic representations of mammoth are known from Roufgnac, Font-deGaume, Les Combarelles and Bernifal (Dordogne, France). Conversely to the realistic representations, the archaic ones are depicted in a schematic way, without indication of anatomical details (Figs. 21, 1e9 and 22, 1e2), and characterized by a silhouette showing a not bloated abdomen and extreme long legs in proportion to the rest of the body (Delluc and Delluc, 2004a, 2004b). Based

Fig. 18. Archaic representation of Mammuthus primigenius from the Baume-Latrone cave (Gard, France) (modied from Bosinski, 1999).

Fig. 20. Sculptered mammoth in the demi-relief technic from the Grotte du Mammouth (Dordogne, France) (modied from White, 1993).

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Fig. 21. Examples of archaic representations of Mammuthus primigenius. 1e2: Chabot cave; 3. Chauvet cave; 4: Pech-Merle Cave; 5: Cougnac cave; 6e7: Roucadour cave; 8: La Grze cave; 9: Jovelle cave (modied from Lorblanchet, 1995).

on the shape of the abdomen, four main different types of representations can be recognized, showing an arched, nearly symmetric abdomen, or an ogival abdomen, or a straight abdomen, or even a weakly arched abdomen, with some exceptions. The archaic representations date to the Aurignacian, Gravettian or Solutrean (Delluc and Delluc, 2004a, 2004b). The richest sample of archaic representations of woolly mammoths is known from the Chauvet cave (Gly and Azma, 2005). In other caves, such as La Grze (Delluc and Delluc, 1991), and Arco B caves (Gonzlez Sainz and San Miguel Llamosas, 2001), a mammoth representation is present among other gures of animals depicted in the archaic way. Following Delluc and Delluc (2004b), the archaic mammoth representations would represent individuals without or reduced fur because of seasonal moulting, which lost weight because of a long winter starvation. 2.4. The intriguing elephant painting of the El Castillo cave In 1903, H. Alcade del Rio discovered Paleolithic representations on the walls of the El Castillo cave, located on the Mount Castillo (Puento Viesgo, Cantabria, northern Spain) as in the Las Chimineas, La Pasiega and Las Monedas caves. Excavations in the entrance hall of the El Castillo cave exposed a detailed, 18 m long stratigraphic succession, with archaeological layers ranging in age from the Lower Paleolithic to the Bronze Age (Prez and Smith, 2002). A graphic documentation of the El Castillo Paleolithic representations was provided by the Abb Henri Breuil (see Alcade del Rio et al., 1911). Among others, the red-brown linear drawing of a proboscidean has been matter of a still open debate (Fig. 23). According to Breuil (1952), the outline of the animal was drawn by joining a number of short lines. Only one foreleg and hind leg are depicted, the tusks are short and not much curved, the apex of the skull is only slightly higher than the back, and the outline of the

Fig. 22. Archaic representations of Mammuthus primigenius in the Jovelle cave (Dordogne, France) (modied from Delluc and Delluc, 1991).

back is gently curved, differing in this from the outlines frequently depicting woolly mammoths with a roundish head, higher than the shoulders and with a back prole more or less backwards inclined. Breuil (1952) suggested the gure could represent a straight-tusked elephant (Palaeoloxodon antiquus) because of the general shape of the body. Other researchers thought that the gure represents either a young mammoth or an individual with reduced fur (Leonardi, in: Ripoll-Perell, 1964; Ripoll-Perell, 1984; Prez and Smith, 2002). Some others, as Gonzlez (2001), claimed that it is not clear whether the gure represents a mammoth, albeit considering this hypothesis as the most reasonable. Actually the prole does not clearly depict either a woolly mammoth or a straight-tusked elephant. Although recent datings demonstrate that at El Castillo the tradition of decorating caves extends back to about 40.8 ka (early Aurignacian period) (Pike et al., 2012), teeth of P. antiquus found in the cave dated to about 42.7 3.5 ka BP, as consistently with the presence in the same layer of Mousterian lithic implements (Liberda et al., 2010). Moreover, there is no rm evidence of the actual persistence of P. antiquus in Northern Spain during the Upper Paleolithic (see below). Therefore, whether the painting of the El Castillo cave represents a straight-tusked elephant still remains an unanswered question. 3. Mammoths and straight-tusked elephants in Southern Europe during the last glacial: a short account In Southern Europe, only woolly mammoths are represented in Paleolithic art, except for the problematic silhouette of the El

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deciduous temperate forest or Mediterranean evergreen woodland spread during the interglacial phases. Middle to late Pleistocene populations of mammoths (Mammuthus trogontherii and M. primigenius) mainly inhabited more open steppe-tundra environments. Therefore, the marked climate changes that characterised the transition from the Middle to the Upper Paleolithic differently affected the expansion/contraction, the dispersal, and the extinction of the representatives of these elephant lineages. Since the late last interglacial (MIS 5a), in response to climatic cooling and in keeping with vegetation changes over most of Europe, populations of Palaeoloxodon antiquus progressively reduced their range to core refugial areas, mainly, but not exclusively, located in Southern Europe. Straight-tusked elephants lasted until about 37e?32 ka on mainland, but the extinction of Mediterranean insular species was much later (see below). Therefore, some residual straight-tusked elephant populations might be present at the time of the rst dispersal of modern humans towards and across Europe possibly shortly before the Aurignacian (see e.g. Stringer, 2008).
Fig. 23. The intriguing elephant painting of the El Castillo cave (Cantabria, Spain). Drawing by H. Breuil (modied from Alcade del Rio et al., 1911).

3.1. The last straight-tusked elephants In the Iberian Peninsula (Fig. 24), the most recent Palaeoloxodon remains have mainly been found in cave sedimentary successions, whose layers frequently yielded late Mousterian or Aurignacian artefacts, but not paintings, engravings or objects representing elephants, except for the El Castillo cave mentioned above (Fig. 24). Although the silhouette painted on the wall of this Spanish cave somehow evokes a straight-tusked elephant (e.g. small, not-curved tusks, head nearly in line with the gently arched back, position of the tie), a weak support to this hypothesis can be found in the fossil record. In the El Castillo cave, three molariform teeth of P. antiquus, two belonging to a young individual, were retrieved along with Aurignacian artefacts from layer 18 (Altuna, 1972; Bernaldo de Quiros, 1982). A number of numerical dates performed with radiocarbon and ESR on some remains found in levels 18 and 20, as well as in the underlining Mousterian levels 22 and 23 (Cabrera-

Castillo cave. This is consistent with the Late Pleistocene fossil record of Elephantini in each region, particularly with the differences characterizing the Iberian and Italian artistic and fossil records. Since their appearance, mammoths and straight-tusked elephants have been an important component of the Eurasian fauna, but their evolutionary history differs in terms of dispersal, dispersion, last stands in different regions, and timing of nal extinction. During the long period of their coexistence, from the latest Early Pleistocene to about MIS 3, representatives of Mammuthus and Palaeoloxodon lineages were infrequently found together in the same local faunal assemblages (LFAs), although their ranges largely overlap. In the LFAs where both were present, their relative abundance was very different because of their different ecological behaviour. Straight-tusked elephants prevailed in regions where

Fig. 24. Map showing the distribution of Iberian and Italian sites yielding the latest remains of Palaeoloxodon antiquus (as source of data see references in text).

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Valds et al., 1996; Rink et al., 1997; Stuart, 2005; Liberda et al., 2010), indicate that the Palaeoloxodon teeth are slightly older than 40 ka. Accordingly, taking into account that the youngest known stratigraphical record of Palaeoloxodon in El Castillo cave shortly predates the rst appearance of Homo sapiens in the area (see Pike et al., 2012), and that the doubtful identication of the elephant painted on the wall of the cave cannot condently support the hypothesis of a longer survival of the species in northern Spain, whether H. sapiens and P. antiquus coexisted in the region still remains to be conrmed. It is worth noting, indeed, that the radiocarbon date of 23.57 ka obtained for a partial skeleton found in marine deposits of Cueva de la Silluca Buelna (Asturias) could be rejected in the absence of further conrmatory evidence, because the marine sediments were likely deposited during an interglacial substage (either MIS 5e or 5c) predating the last glacial (Pinto Llona and Aguirre, 1999; Stuart, 2005). The other most recent Iberian remains of P. antiquus are not signicantly younger. At Olha (Laburdi, Pas Vasco), for instance, the species is reported from levels yielding Mousterian artifacts (Saenz de Buruaga, 2000; Made van der and Mazo, 2001 and references therein). A large fragment of tusk and a molariform tooth were found in layer V of Cova Negra (Valencia, southeastern Spain), already correlated with MIS 4 (Aguire, 1968/69; Perez Ripoll, 1977), but likely much older (see Stuart, 2005 for a discussion). In the eastern Iberian Peninsula, the presence of P. antiquus during late MIS 3 is documented by the ndings of Foz do Enxarrique, where an unworn upper molar plate was found in layers dated at about 33e34 ka. Conversely, the not signicantly younger molar plate from Gruta da Figueira Brava (ca. 30e31 ka) cannot condently be ascribed to the species (Cardoso, 1996; Brugal and Raposo, 1999; Sousa and Figueiredo, 2001). In the Italian peninsula, there is no compelling evidence for a survival of straight-tusked elephants later than MIS 5a to MIS 4 transition (Fig. 24). The molariform tooth from the Neanderthal site of Grotta Guattari (Circeo, southern Latium), was assigned to the latest MIS 5a (Caloi and Palombo, 1995) or to the beginning of MIS 4, as supported by the radiometric (U-series) and ESR dating available for this site (average age ca. 57 ka (6 ka), Schwarcz et al., 1991), while an age older than 54 ka has been suggested for the specimens found in the uppermost portion of the sedimentary succession cropping out at Canale delle Acque Alte (Pontina plain, southern Latium) (Blanc et al., 1957; Caloi and Palombo, 1995 and references therein; Farina, 2011). P. antiquus has also been reported from Layer b of Ingarano cave (Apulia, southern Italy), claimed to have deposited during MIS 4 (Petronio and Sardella, 1998). The specimen described by these authors as a molar plate actually is a portion, transversally cut, of a fallow deer antler. Another small fragment of an unworn plate is too incomplete to allow any rm identication. The age of this remain is uncertain because the stratigraphic relationship of Layer b with the sedimentary succession assigned to MIS 2 and 3 is unclear, and any radiometric dating failed because the molar fragment contains no collagen. In Greece, P. antiquus is among the dominant taxa during the last interglacial, but it is not recorded during the last glacial phase (Doukas and Athanassiou, 2003 and references therein, Evangelia Tsoukala et al., 2011). Therefore it seem that the latest record of P. antiquus in Southern Europe was the molariform tooth found at Foz do Enxarrique (Portugal), whose age would be slightly younger than the youngest radiocarbon dates available for specimens from the Netherland (37,440 350, 310 BP; Mol et al., 2007). As a result, the available data indicate that straight-tusked elephants had disappeared from Southern Europe long before the onset of the LGM. Conversely, their dwarf insular descendants lasted on some Mediterranean islands much more than their ancestors on the mainland. Palaeoloxodon mnaidriensis disappeared on Sicily

around 32 ka (Bonglio et al., 2008), on Crete a population slightly reduced in size has been claimed to be present during the LGM, ca. 18 ka or later (Palaeoloxodon chaniensis in Symeonides et al., 2001), dwarf elephants have been supposed to have survived on Cyprus to the end of the Pleistocene ca.11e10 ka (Reese, 1999), and on Tilos Palaeoloxon tiliensis persisted during the Holocene, maybe to about 3.5 ka (Theodorou and Symeonides, 2001; Theodorou et al., 2007), roughly the same time during which the latest M. primigenius inhabited Wrangel Island (Vartanyan et al., 2008). All in all, as continental Europe, the virtual lack of P. antiquus in the Paleolithic art is roughly consistent with the timing of its extinction on the mainland, and with the very reduced range of the few long- surviving populations of European straight-tusked elephants. 3.2. The woolly mammoth from the Iberian and Italian peninsulas During the last glacial woolly mammoth populations, mostly inhabiting treeless steppe-tundra environments, were widely spread across Northern Eurasia, while the species was rare in southernmost Europe especially during the LGM, when, at about 18 ka, woolly mammoth populations dramatically contracted their range or even disappeared in Western Europe (see e.g. Kahlke, 1999; Doukas and Athanassiou, 2003; Palombo and Ferretti, 2005; Markova et al., 2010; lvarez-Lao and Garca, 2012). In Greece, the woolly mammoth fossil record is extremely poor and limited to the northernmost Greek regions (Fig. 25). Scanty remains are reported in Eastern Macedonia, where a fragment of tusk and an incomplete molar were found at Aggtis (Drama) in a possibly late glacial fauna, including among others Coelodonta antiquitatis (Koufos, 1981), and in the Penios valley (Eastern Thessalia) near Stomio (Elephas (Mammontheus) cf. primigenius in Paraskevaidis, 1977). M. primigenius has been claimed as also present in Southern Greece at Megalopolis (Peloponnes), a site located at about 37 N (see Doukas and Athanassiou, 2003). A recent geochemical analysis performed on the remains stored in the Museum of Paleontology and Geology of Athens University demonstrated that these woolly mammoth specimens were not found in the Megalopolis region, but are part of the collection of Ukrainian fossils, coming from the Kiev-Telichka locality, given as a present to the Greek Universitys Museum (Iliopoulos et al., 2010). In the Iberian Peninsula, remains of M. primigenius are much more abundant and have been reported from at least 25 sites (see lvarez-Lao and Garca, 2012 and references therein for the most recent critical inventory) (Fig. 25). Woolly mammoth remains were mainly retrieved from karst deposits in a number of caves located in the northernmost Spanish regions (Cantabrian area and Catalonia), while a few nds, mainly coming from alluvial deposits, are recorded in the central Iberia peninsula (few sites in Portugal and in the Madrid area). The southernmost ndings, at about 37 N, are those of Padul (Granada) (lvarez-Lao et al., 2009) (see below). In most localities, only scanty remains were found, generally consisting of few molariform teeth, mandibles, and more or less fragmentary tusks, with complete tusks only recovered at Pmanes, Arriaga, Casa Eulogio and Padul sites (Fig. 25). Skulls and postcranial bones are less frequent, except for the rich samples of Pmanes (Lirganes, Cantabria) (Carballo, 1912, 1920; Harl, 1912) and Padul (Granada) (lvarez-Lao et al., 2009). At Pmanes, the woolly mammoth remains, dating to the late MIS 3 (25.4; 27.6 cal ka BP, lvarez-Lao and Garca, 2012) include, along with a nearly complete skull seriously damaged during the archaeological excavation, two tusks, one well preserved mandible with the last molariform teeth (M3), one pelvis, one tibia and one femur likely belonging to a single, fully adult individual (Carballo, 1912, 1920).

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Fig. 25. Map showing the distribution of selected Iberian, Italian Greek sites yielding Mammuthus primigenius remains (as source of data see references in text). Spain: 1 Bujn, 2 Las Caldas, 3e6 La Gelga, El Cierro, La Lloseta, Cueto de la Mina, 7e11 Udas, Mina Angel, Morim, Pmanes, Minas de Heras, 12 Labeco Coba, 13 Urtiagako Lleizea, 14 Clot de Llop, 15 LAlbreda, 16 cau de lesGoyes, 17 Butarque, 18 Casa Eulogio, 19 Aldehuela, 20 Arriaga, 21 Edar Culebro, 22 Padul, 23 Algar de Jo ao Ramos, 24 Figueria Brava, 25 Cruz Quebrada. Italy: 26 Arene Candide, 27 Po valley, various sites, 28 Adda Valley (Cremona), 29 Riparo Taglinte, 30 Asolo, 31 Vidor, 32 Settepolesini, 33 Buca della Iena, 35 Arezzo, various sites, 36 Torrente Conca, 37 Tarquinia, 38 Canale delle Acque Alte, 39 Veroli, 40 Cardamone; Greece: 41 Penios valley (Eastern Thessalia), 42 Aggtis (Drama) (see references in text).

The Padul specimens, representing the southernmost remains of woolly mammoth thus far known in Iberia, as well as in Europe, were retrieved, along with a few remains of steppe bison (Bison priscus), red deer (Cervus elaphus) and a medium sized horse (Equus sp.), from swamp clayey deposits cropping out at different sites in the Padul Basin. The woolly mammoth remains, ranging in age from 30.6 to 40.4 cal ka BP, consist of a few molariform teeth, a complete and a fragmentary tusk, three mandibles and a number of limb bones, belonging to at least four adult males (lvarez-Lao et al., 2009). The Iberian ndings of M. primigenius mainly dated to the last glacial, although archaic representatives of the species have been recorded at the transition from the Middle to Late Pleistocene (Ses and Soto, 2002a, 2002b). In central Spain (Madrid province) woolly mammoth remains have been reported from sites whose age have been estimated by stratigraphic correlations to be either late Middle Pleistocene (e.g. Casa Eulogio and Arriaga localities), or early Late Pleistocene (e.g Aldehuela locality; Mammuthus cf. M. intermedius in Ses and Soto, 2002a), as well as at the sites of Butarque and Edar Culebro, dated at about 100 ka (lvarez-Lao and Garca, 2012 and references therein). During the following early Late Pleistocene, until the beginning of the last glacial, there is no rm evidence of the presence of M. primigenius in the Iberian Peninsula, although a molar plate ascribed to this species has been reported from a Mousterian level of the Arbreda cave (Gerona) (lvarez-Lao and Garca, 2010). During the last glacial, the Iberian fossil record of M. primigenius became much more abundant. Most of the Iberian woolly mammoth remains date to MIS 3 and 2, but whereas those ranging in age between about 30 and 43 ka cal BP spread from the northern to southern Iberian Peninsula, those dated between about 28 and 18 ka cal BP (mostly found in Late Gravettian, Solutrean and Lower Magdalenian archaeological contexts) are almost exclusively located in northern Spain, mainly in the extant Basque territories.

M. primigenius remains dated to the late LGM were found in the Asturias region (northern Spain) in the Lower Magdalenian levels of Las Caldas (about 20e17 ka cal BP), and La Lloseta (from about 20.2 to 17.6 cal ka BP). The molar fragment found at Cueto de la Mina is slightly older, dating at about 23.5 ka cal BP (lvarez-Lao and Garca, 2012 and references therein). In the westernmost Iberian Peninsula, M. primigenius was possibly present around 14 ka BP at o Romaos (Antunes and Cardoso, 1992). This nding, Algar de Joa a fragment of femur showing extensive gnawing, if not reworked, would represent the only occurrence of M. primigenius in Southern o Europe at that time. Whether the mammoth from Algar de Joa Ramos would belong to a population that dispersed from central Europe towards the Iberian Peninsula about 16 ka BP - when M. primigenius reoccupied some of the western European territories it had withdrawn during the LGM - or was the last representative of a former population that inhabited Portugal since MIS 3, is a still o unanswered question. Whatever the actual age of Algar de Joa Ramos M. primigenius, it is certain that woolly mammoth populations were spread in northern Iberia in a time span including the Heinrich event 2 and most of the LGM, when the climate registered cold and dry phases and steppe environments dominated the landscape (see lvarez-Lao and Garca, 2010, 2012). Woolly mammoths likely dispersed towards Iberia at that time following a northern dispersal route and extending their range to those territories where the most signicant Iberian representations of woolly mammoth are known. In Italy, M. primigenius is mostly recorded during MIS 4 and 3 by few, some isolated, ndings. Some other remains, older in age (MIS 6) (mainly teeth but also a skull with mandible), and showing some archaic features have been ascribed to this species (see Reggiani and Sala, 1992; Palombo and Ferretti, 2005; Mussi and Villa, 2009 and references therein) (Fig. 25). Woolly mammoth remains have been found associated with Mousterian tools, as at Riparo Tagliente (Veneto), Asolo (Treviso,

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Veneto), and Buca della Iena near Mommio (Lucca, Tuscany). At Riparo Tagliente (Grezzana, Valpantena, Verona) two fragments of plates of a young mammoth were found associated with Mousterian lithic implements within the lowermost layers (levels 52 to 31) of a long stratigraphic succession, ranging in age from the Mousterian to the Epigravettian (Capuzzi and Sala, 1980; Bartolomei et al., 1982). The abundance in these layers of Cervus elaphus, Capreolus capreolus and Alces alces, together with the presence of Marmota marmota, indicate a moderately humid, temperate-cold climate (Fiore et al., 2004). At Buca della Iena the fossiliferous bed, containing Mousterian lithic implements, is younger than 41 ka (Pitti and Tozzi, 1971). In the alluvial deposits dated to the Last Glacial (cf. Venzo, 1977) in the neighborhood of Pagano dAsolo (Asolo, Treviso), a skeleton of a female of M. primigenius about 32 years old (Reggiani and Sala, 1992), was found in 1878 together with a few Mousterian akes (Dal Piaz, 1922, 1931). The direct radiocarbon date of about 27.8 ka obtained for the mammoth remains has to be rejected because the bones were consolidated using sh glue, while the Levallois implements associated with the skeleton t well into either MIS 4 or 3 (see Mussi and Villa, 2009 for a discussion). Another partial skeleton of a 35 old male was retrieved in 1973 from lacustrine deposits, possibly correlatable with MIS 4 cropping out at S. Giovanni di Valdobbiadine (Vidor, Treviso, Veneto) (Venzo, 1977; Reggiani and Sala, 1992). The uncertainty regarding the precise stratigraphic position of the nding, the lack of numerical dates as well as of associated artifacts makes it difcult to ascertain the chronology of this woolly mammoth. Excluding Asolo and Vidor, most of the other MIS 4 and 3 localities yielded only isolated dental and skeletal remains. The material can be condently referred to M. primigenius on the basis of the derived character of the molars, such as the elevated number of plates, the high lamellar frequency and thin, highly creased enamel (Palombo and Ferretti, 2005). In north Italy, the presence of a mammoth steppe during MIS 3 is indicated by the fauna with M. primigenius (14C dates on mammoth bones range from ca. 35,800 to 33,830 BP), Coelodonta antiquitatis, Megaloceros giganteus, Bison priscus, and Alces alces found a deposit at Settepolesini di Bondeno (eastern Po Valley, Ferrara) (Gallini and Sala, 2001). The dispersal of cold adapted taxa throughout the Po Valley was likely permitted due to the lowering of the sea level during glacial stadials, leading to the emergence of an extensive part of the Adriatic platform. Together with other species (e.g. Sicista betulina, Microtus oeconomus, Coelodonta antiquitatis, B. priscus) M. primigenius spread southward along the Adriatic coast to about 40 220 N, as documented by the ndings of some bones of young mammoths at Cardamone (Lecce, Apulia). No numerical dates are available for the Cardamone fauna. Rustioni et al. (2003) hypothesized that the faunal assemblage could date to the LGM because of the absence of Equus hydruntinus and Dama dama, but the avifauna indicates temperate-cold, quite humid climate conditions. In central Italy, M. primigenius is known from the alluvial deposits cropping out at Torrente Conca (Cattolica); Montecatini Terme (Lucca), a few sites in the Marche region, and in the Arezzo basin (Tuscany) (De Giuli, 1983; Ferretti, 2000; Palombo and Ferretti, 2005 and references therein). The southernmost ndings on the Tyrrhenian side of the Italian peninsula are between 41 410 N (S, Anna, Veroli) and 41 280 N (Canale Acque alte, Latina, Pontina plane, southern Latium). Among the most interesting remains is an incomplete mandible with the penultimate molars found during a well excavation at S. Anna, near Veroli (Frosinone, Souther Latium) (Biddittu and Celletti, 2001; Palombo unpublished data). There is no stratigraphic control for the deposit, but the radiocarbon date of

38,950 600 BP (Stuart, personal communication to MRP in 2004) indicates a correlation with MIS 3. At Canale delle Acque Alte, a few M. primigenius remains has been retrieved in layers correlated with MIS 4 (the same layers with P. antiquus) and in the overlying level correlated with MIS 3 (Blanc et al., 1957; Caloi and Palombo, 1995; Farina, 2011). The available radiocarbon dates conrm that M. primigenius was still present in Italy during MIS 3, while, and except for eastern Liguria, no compelling evidence indicates its presence since the LGM. At the Gravettian Arene Candide site (Liguria, north-western Italy), a few proboscidean bones (a second phalanx, a possible fragment of a patella, an epiphysis fragment and two shaft splinters) have been found (Cassoli and Tagliacozzo, 1994). The bones are not diagnostic and have been ascribed to the woolly mammoth mainly because of the age of the deposit, believed too recent for yielding straight-tusked elephant remains. A few mammoth ivory ornaments and female gurines have been also recorded in easternmost Gravettian sites of the Ligurian coast, close to the modern boundary with southern France, but the carved objects have been supposed to be imported from elsewhere (Mussi et al., 2000). All things considered, whether some M. primigenius populations actually inhabited the Italian peninsula after MIS 3 is still an open question. Woolly mammoth was likely present in Liguria, and, although no radiocarbon dating is available, the hypothesis that a few individuals could have dispersed from the lower Rhone valley to reach the easternmost Ligurian territories is reasonable, especially considering, for instance, that Rangifer tarandus is recorded in Italy only in this region (Sala, 2005). The fact that some M. primigenius herds could reach Liguria from southern France does not necessarily imply that they further dispersed to other parts of the Italian peninsula. Woolly mammoths might be conned to eastern Liguria either because the low number of individuals inhabiting the region did not allowed a range extension, or because the Ligurian Apennines, which separates Liguria from the Po Valley, represented a severe barrier to be crossed by these elephants. Therefore, pending a conrmation about the actual age of the Cardamone woolly mammoth, the still unanswered question arises why any dispersion event from East Europe allowed the presence of M. primigenius at least in northwestern Italy during MIS 2. 4. Remarks Although a rich Upper Paleolithic iconography testies to a long coexistence of humans and M. primigenius during the last glacial in most of central Eurasia and even in southern Europe, the frequency of representations of M. primigenius left by H. sapiens during the Upper Paleolithic (ca. 35 ka to 11 ka) as portable and cave art varies in either number of representations and their geographical distribution. In general, objects of portable art are known from numerous European regions, while cave art is mostly known from South West Europe, especially in France and northern Spain. Moreover, the typology of portable art representations and the rough material used to produce the objects differed during time and across space. During the Aurignacian, for instance, depictions of mammoth are only known in the form of ivory gurines in the Swabian Alps region (South West Germany). During the Gravettian, ivory gurines are known from a number of sites in the Czech Republic and the Russian plain, while gurines of mammoth made by using clay are only known in the Moravian region and those of marl only more eastward in the Russian plain. Finally, most engravings and tools decorated with mammoths are reported from Western and Central European Magdalenian sites. As regards the three main techniques of cave art (painting, engraving and sculpture), the main documentation comes from numerous caves located in France and in Spain. The most eastward

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I.M. Braun, M.R. Palombo / Quaternary International 276-277 (2012) 61e76 Barrire, C., 1976. Lart parital de la Grotte de Gargas, vol. 2. BAR, International Series 14, Oxford. Barrire, C., 1982. Lart parital de Roufgnac. Picard, Paris. Barrire, C., 1993. Les proboscidiens. In: GRAPP (Groupe de Rexion sur lArt Parital Palolithique). Lart parital palolithique. Techniques et mthodes dtude. Comit des Travaux Historiques et Scientiques, Paris, pp. 151e156. Barrire, C., 1997. Lart parital des grottes des Combarelles. Palo, Hors srie, Angoulme. Bartolomei, G., Broglio, A., Cattani, L., Cremaschi, M., Guerreschi, A., Mantovani, E., Peretto, C., Sala, B., 1982. I depositi wrmiani del Riparo Tagliente, vol. 15. Annali dellUniversit di Ferrara (N.S.), pp. 61e105. Bgoun, H., Breuil, H., 1958. Les cavernes du Volp: Trois-Frres, Tuc dAudoubert Montesquieu-Avants (Arige). Arts & Mtiers Graphiques, Paris. Bellier, C., Bott, S., Cattelain, P., 1991. Cahier IV: Objets de parure. 5.1. Fiche Rondelles. 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caves with cave art are those of Kapova and Ignatievka in the Southern Ural Mountains. Comparing the distribution of woolly mammoth fossil record with its representation in the Upper Paleolithic art, no depictions have been reported not only from the regions where M. primigenius had already disappeared by the LGM, but also from regions where both woolly mammoth remains and Paleolithic art were documented, such as, for instance, from Switzerland and Austria. The raison behind this lack still remains an unsolved issue. In particular, as regards southernmost Europe, the fact that the Spanish woolly mammoth representations are mainly located in the northern Cantabrian and Asturias regions is consistent with the fossil record. During the LGM, M. primigenius remains have been found in the area northwest of the Pyrenees along the coast of the Bay of Biscay, which could also be regarded as among the most suitable dispersal routes of M. primigenius towards Iberia (lvarez-Lao et al., 2009). Apparently, during the LGM the range of woolly mammoth did not reach the easternmost Iberian Peninsula. The youngest dating of fossil remains is that of Figueira Brava, correlating the fauna with late MIS 3 (lvarez-Lao and Garca, 2012 and references therein), and woolly mammoths are not represented in the Portuguese Palaeolithic art. The hypothesis that the absence of tundra-like environments prevented the spread of M. primigenius herds towards low latitudes in Iberia during MIS 2 cannot be ruled out, although it needs to be substantiated. The absence of any representation of M. primigenius in Italy, except maybe for the easternmost Liguria, could be regarded as an indirect evidence of the disappearance of the species by the LGM. Ongoing research could answer the question whether the absence of M. primigenius in most of Italy during MIS 2 is due to difculties in reaching the Italian peninsula from eastern Europe, maybe because of a reduced availability of suitable environments in some territories along the potential dispersal routes. Acknowledgements We thank the two anonymous reviewers for their useful comments. References
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