Mammuthus Primigenius in the Cave and Portable Art

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Quaternary International 276-277 (2012) 61e76

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Mammuthus primigenius in the cave and portable art: An overview with a shortaccount on the elephant fossil record in Southern Europe during the last glacial

Ingmar M. Braun a,*, Maria Rita Palombo b,c

aWyhlenweg 4, CH e 4126 Bettingen, SwitzerlandbDipartimento di Scienze della Terra, Università degli Studi di Roma “La Sapienza”, Roma, ItalycCNR, “Istituto di Geologia Ambientale e Geoingegneria”, Montelibretti, Roma, Italy

a r t i c l e i n f o

Article history:Available online 20 July 2012

* Corresponding author.E-mail address: [email protected] (I.M. Braun

1040-6182/$ e see front matter � 2012 Elsevier Ltd ahttp://dx.doi.org/10.1016/j.quaint.2012.07.010

a b s t r a c t

A rich Upper Paleolithic iconography testifies to a long coexistence of humans and Mammuthus pri-migenius during the last glacial in most of Europe, including northern Spain, and supplies additionalinformation for a better understanding of the dispersion and last occurrence of woolly mammoths insouthernmost Europe (i.e. in the Iberian and Italian peninsulas) during this time. In Italy, where thescanty M. primigenius findings are likely not younger than 38 ka (except for the Gravettian remains fromthe Arene Candide cave, eastern Liguria), no representations of woolly mammoths have been reported todate. An exception is the carved mammoth objects (a few Gravettian ornaments and female figurines),recorded in Ligurian sites, but the hypothesis that they could have been imported from some distant areacannot be ruled out. Conversely, in Spain along the northern Atlantic coast, M. primigenius remains havebeen found in some sites yielding mammoth representations. In southern Spain, where M. primigeniuswas present in the Padul basin (Granada) during most of MIS 3 (between 40.4 and 30.6 cal ka BP), artisticrepresentations of woolly mammoths are unknown. As regard to Palaeoloxodon, some populations werepresent during the late MIS 3 in the Iberian Peninsula as in Western Europe, whereas no sound datasupport the persistence of straight-tusked elephants on mainland during MIS 2. Therefore, whether theintriguing elephant painting of the Spanish El Castillo cave could represent a straight-tusked elephant esuggesting a survival of the species in Northern Spain during the Last Glacial Maximum e or an unusualrepresentation of a woolly mammoth, still remains an unanswered question.

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1. Introduction

The iconic elephant of the last glacial, the woolly mammothMammuthus primigenius, is a species widely spread during the LatePleistocene 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 southern-most maximum extension during MIS 3a, when the woollymammoth is recorded at 36� 350 N in China (Ji’nan, ShandongProvince) (Takahashi et al., 2007), while the southernmost recordsof M. primigenius in Western Europe are those of Padul (37� 010N,Granada Basin, Spain) (Álvarez-Lao et al., 2009) and Cardamone(40� 210N, Apulia, Italy) (Rustioni et al., 2003). During the lateglacial, in particular during the Last Glacial Maximum (LGM), thedistribution of M. primigenius findings across Europe (Fig. 1) onlypartiallymatches that of vestiges of the Upper Paleolithic art, left by

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our direct ancestor Homo sapiens, from about 35 ka to 11 ka BP(Fig. 2).

This evidence is cave art products, known especially in WesternEurope, and portable art objects which are documented all overEurope and as far as Siberia. The cave art includes figurativerepresentations on walls and ceilings of caves and rock shelters aswell as those on cave floors (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 figurines of animals andhumans. Although the figures of woolly mammoths do not domi-nate among the zoomorphic Paleolithic representations, M. pri-migenius was depicted by Paleolithic artists using a variety ofdifferent materials and techniques.

The paper aims to present a short overview of the main depic-tions of M. primigenius thus far known in the Eurasian Paleolithicart, and to discuss some intriguing issues such as the perplexingrepresentation of the El Castillo elephant, and the lack of anyrepresentation in Italy, where the woolly mammoth is recordeddefinitively 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 (modified and updated from Markova et al., 2010).

I.M. Braun, M.R. Palombo / Quaternary International 276-277 (2012) 61e7662

2. M. primigenius in the portable and cave art: an overview

The first discovery of aM. primigenius representation dates backto the second half of the 19th century, when, in 1864 Edouard Lartetand Henry Christy, during the archaeological excavation of the Abride La Madeleine (Dordogne, France), found a piece of mammothivory with the engraving of a woolly mammoth (Fig. 3.1 and 3.2). Atthat time, this finding was regarded as proof that prehistoric menand woolly mammoths lived contemporaneously and interactedwith each other: “This new fact will not, indeed, add anything toalready acquired convictions as to the coexistence of Man with thefossil Elephant (Elephas primigenius) and other great Herbivores and

Fig. 2. Map showing the distribution of selected Upper Paleolithic sites with the main reprfrom Bosinski and Fischer, 1980). 1: Las Caldas; 2: Pindal; 3: El Castillo; 4: Arco B cave; 5:Latrone; 11: Bayol; 12e14: Oulen, Le Figuier, Chabot; 15e16: Chauvet, Ebbou; 17: BruniPigeonnier, Le Mammouth, Cavaille; 25e33: Croze-à-Gontrand, La Mouthe, Cournazac, La GRaymonden, Rouffignac, Labattut; 37e39: Les Vachons, Les Bernous, Jovelle; 40: Pair-non-Cheval, Grande Grotte d’Arcy-sur-Cure; 46: Solutré; 47: La Colombière; 48: Trou de ChaleKlause; 54e55: Dolní V�estonice, Pavlov; 56: P�redmostí; 57: Avdeevo; 58: Sungir’; 59e61: K

Carnivores which geologists regard as having lived together in theearly phases of the Quaternary Period” (Lartet and Christy, 1875, p.207).

Today, the Paleolithic art of Eurasia counts at least 561 repre-sentations of M. primigenius known from about 76 sites (Berdin,1970; Bosinski and Fischer, 1980; Bellier et al., 1999; Gély andAzéma, 2005) (Fig. 2). This number likely underestimates theactual number of mammoth paintings, figurines and engravings asin some site investigations are still in progress (e.g. in France thecaves of Chauvet, Ardèche, and Cussac, Dordogne), old publisheddata might be incomplete, and unknown mammoth representa-tions might exist in either private or public collections.

esentations (cave and portable art) of Mammuthus primigenius (modified and updatedLos Casares; 6: Isturitz; 7: Gargas; 8: Les Trois-Frères; 9: Canecaude I; 10: La Baume-quel, Abri Montastruc; 18e20: Pech-Merle, Cougnac, Roucadour; 21e24: Cussac, Lerèze, Bernifal, Les Combarelles, Font-de-Gaume, Laugerie-Haute, La Madeleine; 34e36:Pair; 41: Chanlat; 42e43: La Marche-Réseau Guy Martin, La Marche cave; 44e45: Leux; 49e50: Gönnersdorf, Andemach; 51e52: Vogelherd, Geissenklösterle; 53: Obereostienki 1, Kostienki 4, Kostienki 11; 62: Kapova.

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

I.M. Braun, M.R. Palombo / Quaternary International 276-277 (2012) 61e76 63

Although in the cave and portable art animals were oftendepicted in a detailed, realistic way, some representations consist ofschematic outlines and the species can only be recognised becausethe silhouette highlights some specific, diagnostic characteristics.Concerning woolly mammoths, Bellier et al. (1999, p.108) wrote:“[.] le mammouth est essentiellement évoqué par sa silhouette car-actéristique, marquée par un massif frontal lourd et puissant, se pro-longeant vers le bas par la masse de la trompe, par la ligne cervico-dorsale soulignant les bosses de la tête et du dos, séparées par ladépression nuchale, et par l’affaissement de l’arrière-train. Les détailstels que les pattes, la toison laineuse ou les défenses ne sont souventpas indiqués”.

2.1. The mammoth in portable art

Figurines representing woolly mammoths are especiallyfrequent in during the early and middle Upper Paleolithic (Auri-gnacian and Gravettian). Most engravings and tools decorated withmammoths date back to the late Upper Paleolithic (Magdalenian).

2.1.1. Figurines of woolly mammothsThe oldest portable art objects thus far known in Eurasia are the

ivory figurines, dating back to the Aurignacian (ca. 35 to 30 ka BP),found in the Lone and Ach valleys in the Swabian Alps (Baden-Würtemberg, South West Germany).

In the Lone valley, the Vogelherd cave, completely excavated in1931 by Riek (1934) yielded some important findings, includingquite small ivory figurines of various animals (Hahn, 1986). Amongthose representing woolly mammoths there is a nearly completefigurine (5 cm long, 2.2 cm wide, 3.1 cm high) (Fig. 4.1), found inhorizon V, that was carved on the core of a mammoth tusk, whichshows X signs, and alignments of dots and grooves, decorating themammoth body (Bosinski, 1982). A large fragment of ivory, repre-senting the backside and the feet of a mammoth found at the samesite, was also decorated with different signs (Bosinski, 1982; Hahn,1986). Of particular interest is a perforated oval bone, regarded asa pendant, showing a demi-relief of a mammoth (Bosinski, 1982).

Since 2005, a team of the University of Tübingen has been re-exploring the Vogelherd cave. The most important finding ofthese new investigations is represented by a complete figurinediscovered in 2006 (Fig. 4.2). With a total length of 3.7 cm, it is thesmallest figurine of a woolly mammoth thus far recorded from theSwabian Alps and the only complete among the 20 Aurignacianfigurines of animals recorded in the Lone and Ach valleys. In thisfigurine of Vogelherd cave, the soles of the feet are decorated withfine crossing lines, and six fine lines are also engraved on the top ofthe head (Conard, 2009).

In the Ach valley, archaeological investigations were made bythe University of Tübingen from 1973 to 2002 in the Geissenklös-terle (surroundings of Blaubeuren). The objects of portable art areof great interest, albeit less numerous than those found in theVogelherd cave.More than 40 pieces of ivorywere found, belongingto a nearly complete figurine of a mammoth, about 7 cm long andcarved in the inner part of a tusk. Although the figurine is frag-mentary and not perfectly preserved, the shape clearly indicatesa woolly mammoth. As are the other figurines of the Swabian Alps,it is decorated with different signs as well as with points of redochre, as shown by microscopic investigations (Hahn, 1986, 1988).

In the middle Upper Paleolithic (Gravettian, from about 28 to22 ka BP), the figurines of woolly mammoths became morenumerous. Most are reported from open air sites in Moravia (CzechRepublic) and in Eastern Europe (Russia and Ukraine). Converselyto the Aurignacian figurines from the Swabian Alps, these figurinesdo not show any geometric sign and were carved not only in ivorybut also in bones, clay, and stone. In the Pavlovian of Moravia,female and animal figurines (some representing the woollymammoth), formed in clay then burned, are known from the sitesof Dolní V�estonice and Pavlov (Valoch and Lázni�ckovà-Galetová,2009) (Fig. 5). The use of clay for the fabrication of portable artobjects is only known from the Pavlovian sites. At Pavlov, as well atP�redmost (Moravia), an ivory figurine of a woolly mammoth wasalso found (Valoch and Lázni�ckovà-Galetová, 2009) (Fig. 6).

More eastward, about 40 figurines of M. primigenius werediscovered in some sites situated in the Russian plain, such as thoseof the Kostienki area, and Avdeevo, Eliseevitchi, and Sungir’(Abramova, 1995). The animal and female figurines recorded fromthis area date to the so-called Kostenkian, corresponding to theEuropean late Gravettian (from about 24 to 22 ka BP) (Djindjianet al., 1999).

In the Kostienki area, some open air sites are located along theDon River north of the Black Sea. The figurines of mammoth fromthese sites are all marly representations (Abramova, 1995), whichclearly show the characteristic shape of M. primigenius (Fig. 7), asshown by the only woolly mammoth figurine found in the open airsite of Eliseevitchi (Abramova, 1995). Interesting objects were alsofound in the open air site at Avdeevo. The sample includes twomammoth figurines carved on sandstone (Fig. 8.2), and one ina spongeous bone, probably a vertebra of mammoth (Fig. 8.1). Theshape of the later is similar to that of figurines from P�redmost(Moravia) (Gvozdover, 1995).

An ivory figurine regarded as a woolly mammoth was discov-ered in a completely different context, a richly equipped doubletomb of two children, at Sungir’ (Bogoljubova, outskirts of Vladimircity, Russia) (Abramova, 1995). The most eastward site yieldinga mammoth representation is the Siberian site Ust’ Kova, wherea figurine carved in ivory was found (Abramova, 1995).

2.1.2. Engravings of mammothsEngravings of mammoths in portable art are especially carved

on stone, but also on other materials. In some engravings, it isdifficult to decode the mammoth representations because of linesor other animal figures overlying them. Except for four sites dating

Fig. 4. 1) Figurine of a woolly mammoth found by G. Riek in the Vogelherd cave (Baden-Württemberg, Germany) in 1931 (modified from Floss, 2009). 2) Figurine of mammothfound in the Vogelherd cave (Baden-Württemberg, Germany) in 2006 (modified 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 lateUpper Paleolithic. During the Magdalenian (from about 18 to12.5 ka), their geographical distribution is mainly restricted toWestern and Central Europe.

The best known representation is that on a fragment of a tuskfound in the Abri de La Madeleine (southwestern France). Thegreatest number of mammoth engravings (76 representations) wasfound in the Magdalenian open air site of Gönnersdorf nearKoblenz (Rheinland-Pfalz, Germany) (Fig. 9), where mammoths,various animals and schematic female figurines were engraved onslate plates (see Bosinski, 2008; Bosinski et al., 2001). Adultmammoths and also young individuals were represented bothalone and in groups. Although the representations are very

Fig. 5. eFigurines of Mammuthus primigenius in burned clay from Dolní V�estonice andPavlov in the Moravian region, Czech Republic (modified from Valoch and Lázni�ckovà-Galetová, 2009).

detailed, all mammoths of Gönnersdorf are tuskless. FollowingGuthrie (in Bosinski, 1984) the tusklessness would affected theGönnersdorf mammoth population as a result of starvation causedby low vegetation productivity at the end of the late glacial period.Bellier et al. (1999) believed that the representations of Gönners-dorf document the latest population of mammoths in WesternEurope.

The southernmost site yielding engravings of woollymammothslikely is Los Casares (Guadalajara, northern Spain) (Angulo andMoreno, 2011). The westernmost site is Las Caldas (Asturias,northern Spain), where an engraving on a stone plate was found ina Middle Magdalenian context (Corchón, 1991e1992). Molariformteeth of M. primigenius have been dated to 21.8 cal ka BP (Álvarez-Lao and García, 2012).

Eastwards, mammoth engravings are recorded in Siberia, wherean ivory perforated plaquette is recorded from the open air site ofMal’ta near Irkutsk (Fig. 10), and a representation craved on a tuskof a young mammoth was found at Berelëkh A (Sakha Republic,Yakutia) (Abramova,1995). The later is the most northern finding ofa woolly mammoth engraving known. With its overlong legs, thevery long trunk and the proportionally small body, this

Fig. 6. Ivory figurine of Mammuthus primigenius from P�redmost in the Moravianregion, Czech Republic (modified from Absolon and Klima, 1977).

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

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


representation differs clearly from the other engravings of woollymammoth known in Eurasia (Fig. 11).

2.1.3. Tools decorated with woolly mammothsIn contrast to the figurines and engravings, only a few tools

decorated with mammoths are known to date, mostly from France.A fragment of a bâton percé, found in the Protomagdalenian layer ofthe Abri Laugerie-Haute (Dordogne, France), shows two deeply

Fig. 8. eBone (1) and sandstone (2a,b,c) figurine of Mammuthus primigenius fromAvdeevo, Russia (modified from Gvozdover, 1995).

engraved mammoths, facing each other and touching their heads(Peyrony and Peyrony, 1938), which were regarded by Bosinski andFischer (1980) as two fighting animals (Fig. 12).

Two Magdalenian spear-throwers in the shape of a mammothwere found in France at Canecaude I (Aude) and at the Abri Mon-tastruc (Tarn-et-Garonne). The first, perfectly reproducing thesilhouette of a woolly mammoth (Fig. 13), was carved on an antlerof the reindeer (Sacchi, 1986). The second, more schematic, wasdiscovered, with other Magdalenian objects, in the second half ofthe 19th century (Sieveking, 1987) (Fig. 14).

Two Magdalenian perforated bone disks, supposed to be orna-ments (Bellier et al., 1991) but whose actual use is still unclear, havebeen found atRaymonden (Dordogne, France) andTroudeChaleux inBelgium. Thefirst disk shows engravings of amammothonboth sides(Fig.15),whereason thesecondonlyafigureof amammoth ispresent.

2.2. The mammoth in cave art

In the cave art, the representations of animals dominate, espe-cially those of herbivores. About 6.3% of the figurative representa-tions depict woolly mammoths (Tosello and Fritz, 2006), and havebeen recorded from 46 caves, 15% of all sites with cave art vestiges.Horses are the most frequently represented animals, followed bybison and mammoths (Gély and Azéma, 2005). In northern Spain,deer representations are much more frequent than those ofM. primigenius. Caves showing the greatest number of mammothrepresentations are the Rouffignac cave (Dordogne, France) with158 figures (Plassard, 1999) followed by the Chauvet cave (Ardèche,France) with 76 specimens (Gély and Azéma, 2005).

Only a few representations of mammoth are known from theIberian Peninsula, while none has been discovered in Italy so far.

Fig. 10. Engraving of Mammuthus primigenius on a perforated plaquette of ivory fromMal’ta near Irkutsk, Russia (modified from Abramova, 1995).

Fig. 11. The most northern depiction of Mammuthus primigenius from Berelëkh A inthe Sakha Republic, Russia (modified from Bosinski and Fischer, 1980).

Fig. 12. Bâton percé with two depicted wolly mammoths from Laug


2.2.1. Discovery of engravings in the Chabot caveIn 1878, Léopold Chiron discovered some engravings on the

walls of the Chabot cave (Gard, France) that he published in 1889,providing, for the first time, a graphic and photographic docu-mentation of paleolithic cave art and identifying some of the Cha-bot engravings as birds and humans. Chiron did not realize the oldage of the representations of Chabot cave (Aujoulat, 1987), some ofwhich nowadays are known as depicting woolly mammoths(Combier, 1984a) (Fig. 16.1 and 16.2).

2.2.2. The cave artIn Europe, vestiges of Paleolithic cave art are recorded in

about 300 caves, whose geographical distribution is much morelimited than that of the portable art. Most are located inWestern Europe, in France and in Spain. A dozen caves with caveart are known in Italy, especially from Sicily (Roussot, 2002). In2003, caves with Paleolithic cave art were discovered for thefirst time in Great Britain (Bahn and Pettitt, 2009) and in 2009in Romania (Besesek et al., 2010). The caves of Kapova andIgnatievka, both in the Southern Ural Mountains, are the mosteastward 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, engravingand sculpture. In most caves, there are figures made by means ofdifferent techniques. Painting, engraving and sculpture can bepresent in each cave in combination or singularly. In many cases thePaleolithic artists either modified or integrated the natural relief ofthe 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 bymixing these colours. Colours were often obtained from mineralcomponents. The yellow colour was obtained from either limoniteor goethite, the red from ferric oxides, especially hematite, whilethe black colour used manganese oxide and charcoal. Pigments

erie-Haute (Dordogne, France) (modified from Bosinski, 1990).

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

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


were used to draw outlines or to paint the whole figure. Therepresentations could be either monochrome, bichrome, orpolychrome.

Interesting silhouettes of woolly mammoths, drawn morefrequently in black than in red (Bosinski in �S�celinskij and �Sirokov,1999), are present in the Rouffignac (Fig. 17) (see Barrière, 1982;Plassard, 1999) and in Pech-Merle caves (Lot, France) (Lorblanchet,2010). Red painted representations are present in the GrandeGrotte of Arcy-sur-Cure (Yonne), Oulen (Gard), and Cougnac caves(Lot), in Spain in the Pindal cave (Asturias) and in Russia in thecaves of Kapova and Ignatievka (Combier, 1984b; Baffier and Girad,1998; �S�celinskij and �Sirokov, 1999; Saura and Múzquiz, 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) (modified from Sieveking, 1987).

(Ardèche, France) are represented as both black silhouettes, andpaintings (Gély and Azéma, 2005).

Of particular interest are the strange representations ofmammoths (Fig. 18) and other animals found at the Baume-Latronecave (Gard, France), which were drawn using clayey material (Louisand Drouot, 1953).

Engraving is a technique of abrasion, performed with the aid ofa pointed object, in general a flint tool, used to carve on the rock

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

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

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


wall one or more, fine or very deep lines (Fig. 19). The lines drawnon soft rocks with fingers are called finger flutings. The scrapingtechnique (raclage in French) is another type of abrasion performedto obtain a kind of chromaticity by scraping the cave wall.

Engravings of mammoths are fairly frequent in the cave art.Several of such configurations are present in the French caves ofRouffignac, Les Combarelles I (Dordogne), Font-de-Gaume (Dor-dogne), Jovelle (Dordogne), Chauvet (Ardèche), Cussac (Dordogne),Gargas (Haute-Garonne) and Les Trois-Frères (Ariège) (Capitanet al., 1910; Bégouën and Breuil, 1958; Barrière, 1976, 1982, 1993,1997; Delluc and Delluc, 1991; Plassard, 1999; Gély and Azéma,2005). In the Chauvet cave there are the only representations ofwoolly mammoths made using the scraping technique (Gély andAzéma, 2005), while some others were engraved with the fingersinto the soft cave walls.

A very deep technique of abrasion is the sculpture. In this casethe figures are strongly raised from the plane of the cave wall, ina plastic way (demi-relief). The sculpture technique is known froma few sites, especially from rock shelters in southwestern France.The only sculpted representation of mammoth was discovered inthe 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 indi-vidual animal or groups. The latter are present, for instance, in theRouffignac cave where numerous animals were depicted close

Fig. 18. Archaic representation of Mammuthus primigenius from the Baume-Latronecave (Gard, France) (modified from Bosinski, 1999).

together as if they were interacting (see Barrière, 1982; Plassard,1999).

Delluc and Delluc (2004a, 2004b) described two styles ofrepresentations: a realistic and an archaic style. There are also thedessins cursifs where the trunk, the head and the back of themammoth were drawn in a continuous, single line.

In a realistic representation, woolly mammoths are depicted ina very detailed way, a number of anatomical details were repre-sented such as eyes, ears, tusks, the fur, tail and even the “finger” atthe end of the trunk (Fig. 17). In most of these realistic represen-tations, usually dating to the middle or late Magdalenian, a numberof individuals are represented, apparently interacting. Realisticrepresentations of mammoth are known from Rouffignac, Font-de-Gaume, Les Combarelles and Bernifal (Dordogne, France).

Conversely to the realistic representations, the archaic ones aredepicted in a schematic way, without indication of anatomicaldetails (Figs. 21, 1e9 and 22, 1e2), and characterized by a silhouetteshowing a not bloated abdomen and extreme long legs in propor-tion to the rest of the body (Delluc and Delluc, 2004a, 2004b). Based

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

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

Fig. 22. Archaic representations of Mammuthus primigenius in the Jovelle cave (Dor-dogne, France) (modified from Delluc and Delluc, 1991).


on the shape of the abdomen, four main different types of repre-sentations can be recognized, showing an arched, nearly symmetricabdomen, or an ogival abdomen, or a straight abdomen, or evena weakly arched abdomen, with some exceptions. The archaicrepresentations date to the Aurignacian, Gravettian or Solutrean(Delluc and Delluc, 2004a, 2004b).

The richest sample of archaic representations of woollymammoths is known from the Chauvet cave (Gély and Azéma,2005). In other caves, such as La Grèze (Delluc and Delluc, 1991),and Arco B caves (González Sainz and San Miguel Llamosas, 2001),a mammoth representation is present among other figures ofanimals depicted in the archaic way. Following Delluc and Delluc(2004b), the archaic mammoth representations would representindividuals 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 representationson 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 hallof the El Castillo cave exposed a detailed, 18 m long stratigraphicsuccession, with archaeological layers ranging in age from theLower Paleolithic to the Bronze Age (Pérez and Smith, 2002).

A graphic documentation of the El Castillo Paleolithic repre-sentations was provided by the Abbé Henri Breuil (see Alcade delRio et al., 1911). Among others, the red-brown linear drawing ofa proboscidean has been matter of a still open debate (Fig. 23).According to Breuil (1952), the outline of the animal was drawn byjoining a number of short lines. Only one foreleg and hind leg aredepicted, the tusks are short and not much curved, the apex of theskull is only slightly higher than the back, and the outline of the

back is gently curved, differing in this from the outlines frequentlydepicting woolly mammoths with a roundish head, higher than theshoulders and with a back profile more or less backwards inclined.Breuil (1952) suggested the figure could represent a straight-tuskedelephant (Palaeoloxodon antiquus) because of the general shape ofthe body. Other researchers thought that the figure representseither a young mammoth or an individual with reduced fur (Leo-nardi, in: Ripoll-Perelló, 1964; Ripoll-Perelló, 1984; Pérez andSmith, 2002). Some others, as González (2001), claimed that it isnot clear whether the figure represents a mammoth, albeitconsidering this hypothesis as the most reasonable. Actually theprofile does not clearly depict either a woolly mammoth ora straight-tusked elephant. Although recent datings demonstratethat at El Castillo the tradition of decorating caves extends back toabout 40.8 ka (early Aurignacian period) (Pike et al., 2012), teeth ofP. antiquus found in the cave dated to about 42.7 � 3.5 ka BP, asconsistently with the presence in the same layer of Mousterianlithic implements (Liberda et al., 2010). Moreover, there is no firmevidence of the actual persistence of P. antiquus in Northern Spainduring the Upper Paleolithic (see below). Therefore, whether thepainting of the El Castillo cave represents a straight-tuskedelephant still remains an unanswered question.

3. Mammoths and straight-tusked elephants in SouthernEurope during the last glacial: a short account

In Southern Europe, only woolly mammoths are represented inPaleolithic art, except for the problematic silhouette of the El

Fig. 23. The intriguing elephant painting of the El Castillo cave (Cantabria, Spain).Drawing by H. Breuil (modified from Alcade del Rio et al., 1911).


Castillo cave. This is consistent with the Late Pleistocene fossilrecord of Elephantini in each region, particularly with the differ-ences characterizing the Iberian and Italian artistic and fossilrecords. Since their appearance, mammoths and straight-tuskedelephants have been an important component of the Eurasianfauna, but their evolutionary history differs in terms of dispersal,dispersion, last stands in different regions, and timing of finalextinction.

During the long period of their coexistence, from the latest EarlyPleistocene to about MIS 3, representatives of Mammuthus andPalaeoloxodon lineages were infrequently found together in thesame local faunal assemblages (LFAs), although their ranges largelyoverlap. In the LFAs where both were present, their relative abun-dance was very different because of their different ecologicalbehaviour. Straight-tusked elephants prevailed in regions where

Fig. 24. Map showing the distribution of Iberian and Italian sites yielding the lates

deciduous temperate forest or Mediterranean evergreen woodlandspread during the interglacial phases. Middle to late Pleistocenepopulations of mammoths (Mammuthus trogontherii and M. pri-migenius) mainly inhabited more open steppe-tundra environ-ments. Therefore, the marked climate changes that characterisedthe transition from the Middle to the Upper Paleolithic differentlyaffected the expansion/contraction, the dispersal, and the extinc-tion of the representatives of these elephant lineages. Since the latelast interglacial (MIS 5a), in response to climatic cooling and inkeeping with vegetation changes over most of Europe, populationsof Palaeoloxodon antiquus progressively reduced their range to corerefugial areas, mainly, but not exclusively, located in SouthernEurope. Straight-tusked elephants lasted until about 37e?32 ka onmainland, but the extinction of Mediterranean insular species wasmuch later (see below). Therefore, some residual straight-tuskedelephant populations might be present at the time of the firstdispersal of modern humans towards and across Europe possiblyshortly before the Aurignacian (see e.g. Stringer, 2008).

3.1. The last straight-tusked elephants

In the Iberian Peninsula (Fig. 24), the most recent Palaeoloxodonremains have mainly been found in cave sedimentary successions,whose layers frequently yielded late Mousterian or Aurignacianartefacts, but not paintings, engravings or objects representingelephants, except for the El Castillo cave mentioned above (Fig. 24).Although the silhouette painted on the wall of this Spanish cavesomehow evokes a straight-tusked elephant (e.g. small, not-curvedtusks, head nearly in line with the gently arched back, position ofthe tie), a weak support to this hypothesis can be found in the fossilrecord. In the El Castillo cave, three molariform teeth of P. antiquus,two belonging to a young individual, were retrieved along withAurignacian artefacts from layer 18 (Altuna, 1972; Bernaldo deQuiros, 1982). A number of numerical dates performed withradiocarbon and ESR on some remains found in levels 18 and 20, aswell as in the underlining Mousterian levels 22 and 23 (Cabrera-

t remains of Palaeoloxodon antiquus (as source of data see references in text).


Valdés et al., 1996; Rink et al., 1997; Stuart, 2005; Liberda et al.,2010), indicate that the Palaeoloxodon teeth are slightly olderthan 40 ka. Accordingly, taking into account that the youngestknown stratigraphical record of Palaeoloxodon in El Castillo caveshortly predates the first appearance of Homo sapiens in the area(see Pike et al., 2012), and that the doubtful identification of theelephant painted on thewall of the cave cannot confidently supportthe hypothesis of a longer survival of the species in northern Spain,whether H. sapiens and P. antiquus coexisted in the region stillremains to be confirmed. It is worth noting, indeed, that theradiocarbon date of 23.57 ka obtained for a partial skeleton found inmarine deposits of Cueva de la Silluca Buelna (Asturias) could berejected in the absence of further confirmatory evidence, becausethe marine sediments were likely deposited during an interglacialsubstage (either MIS 5e or 5c) predating the last glacial (Pinto Llonaand Aguirre, 1999; Stuart, 2005). The other most recent Iberianremains of P. antiquus are not significantly younger. At Olha (Lab-urdi, País Vasco), for instance, the species is reported from levelsyielding Mousterian artifacts (Saenz de Buruaga, 2000; Made vander and Mazo, 2001 and references therein). A large fragment oftusk 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 (seeStuart, 2005 for a discussion).

In the eastern Iberian Peninsula, the presence of P. antiquusduring late MIS 3 is documented by the findings of Foz do Enxar-rique, where an unworn upper molar plate was found in layersdated at about 33e34 ka. Conversely, the not significantly youngermolar plate from Gruta da Figueira Brava (ca. 30e31 ka) cannotconfidently be ascribed to the species (Cardoso, 1996; Brugal andRaposo, 1999; Sousa and Figueiredo, 2001).

In the Italian peninsula, there is no compelling evidence fora survival of straight-tusked elephants later than MIS 5a to MIS 4transition (Fig. 24). The molariform tooth from the Neanderthal siteof Grotta Guattari (Circeo, southern Latium), was assigned to thelatestMIS 5a (Caloi and Palombo,1995) or to the beginning ofMIS 4,as supported by the radiometric (U-series) and ESR dating availablefor 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 specimensfound in the uppermost portion of the sedimentary successioncropping out at “Canale delle Acque Alte” (Pontina plain, southernLatium) (Blanc et al., 1957; Caloi and Palombo, 1995 and referencestherein; Farina, 2011). P. antiquus has also been reported from“Layer b” of Ingarano cave (Apulia, southern Italy), claimed to havedeposited duringMIS 4 (Petronio and Sardella, 1998). The specimendescribed by these authors as a molar plate actually is a portion,transversally cut, of a fallow deer antler. Another small fragment ofan unworn plate is too incomplete to allow any firm identification.The age of this remain is uncertain because the stratigraphic rela-tionship of Layer b with the sedimentary succession assigned toMIS 2 and 3 is unclear, and any radiometric dating failed becausethe molar fragment contains no collagen.

In Greece, P. antiquus is among the dominant taxa during the lastinterglacial, but it is not recorded during the last glacial phase(Doukas and Athanassiou, 2003 and references therein, EvangeliaTsoukala et al., 2011). Therefore it seem that the latest record ofP. antiquus in Southern Europe was the molariform tooth found atFoz do Enxarrique (Portugal), whose age would be slightly youngerthan the youngest radiocarbon dates available for specimens fromthe Netherland (37,440 þ 350, �310 BP; Mol et al., 2007). Asa result, the available data indicate that straight-tusked elephantshad disappeared from Southern Europe long before the onset of theLGM. Conversely, their dwarf insular descendants lasted on someMediterranean islands much more than their ancestors on themainland. Palaeoloxodon “mnaidriensis” disappeared on Sicily

around 32 ka (Bonfiglio et al., 2008), on Crete a population slightlyreduced in size has been claimed to be present during the LGM, ca.18 ka or later (¼“Palaeoloxodon chaniensis” in Symeonides et al.,2001), dwarf elephants have been supposed to have survived onCyprus to the end of the Pleistocene ca.11e10 ka (Reese, 1999), andon Tilos Palaeoloxon tiliensis persisted during the Holocene, maybeto about 3.5 ka (Theodorou and Symeonides, 2001; Theodorouet al., 2007), roughly the same time during which the latestM. primigenius inhabited Wrangel Island (Vartanyan et al., 2008).

All in all, as continental Europe, the virtual lack of P. antiquus inthe Paleolithic art is roughly consistent with the timing of itsextinction on the mainland, and with the very reduced range of thefew long- surviving populations of European straight-tuskedelephants.

3.2. The woolly mammoth from the Iberian and Italian peninsulas

During the last glacial woolly mammoth populations, mostlyinhabiting treeless steppe-tundra environments, were widelyspread across Northern Eurasia, while the species was rare insouthernmost Europe especially during the LGM, when, at about18 ka, woolly mammoth populations dramatically contracted theirrange or even disappeared in Western Europe (see e.g. Kahlke,1999; Doukas and Athanassiou, 2003; Palombo and Ferretti,2005; Markova et al., 2010; Álvarez-Lao and García, 2012).

In Greece, the woolly mammoth fossil record is extremelypoor and limited to the northernmost Greek regions (Fig. 25).Scanty remains are reported in Eastern Macedonia, wherea fragment of tusk and an incomplete molar were found atAggìtis (Drama) in a possibly late glacial fauna, including amongothers Coelodonta antiquitatis (Koufos, 1981), and in the Peniosvalley (Eastern Thessalia) near Stomio (¼Elephas (Mammontheus)cf. primigenius in Paraskevaidis, 1977). M. primigenius has beenclaimed as also present in Southern Greece at Megalopolis(Peloponnes), a site located at about 37� N (see Doukas andAthanassiou, 2003). A recent geochemical analysis performedon the remains stored in the Museum of Paleontology andGeology of Athens University demonstrated that these woollymammoth specimens were not found in the Megalopolis region,but are part of the collection of Ukrainian fossils, coming fromthe Kiev-Telichka locality, given as a present to the GreekUniversity’s Museum (Iliopoulos et al., 2010).

In the Iberian Peninsula, remains of M. primigenius are muchmore abundant and have been reported from at least 25 sites (seeÁlvarez-Lao and García, 2012 and references therein for the mostrecent critical inventory) (Fig. 25). Woolly mammoth remains weremainly retrieved from karst deposits in a number of caves located inthe northernmost Spanish regions (Cantabrian area and Catalonia),while a few finds, mainly coming from alluvial deposits, arerecorded in the central Iberia peninsula (few sites in Portugal and inthe Madrid area). The southernmost findings, at about 37� N, arethose of Padul (Granada) (Álvarez-Lao et al., 2009) (see below).

In most localities, only scanty remains were found, generallyconsisting of few molariform teeth, mandibles, and more or lessfragmentary tusks, with complete tusks only recovered at Pámanes,Arriaga, Casa Eulogio and Padul sites (Fig. 25). Skulls and post-cranial bones are less frequent, except for the rich samples ofPámanes (Liérganes, Cantabria) (Carballo, 1912, 1920; Harlé, 1912)and Padul (Granada) (Álvarez-Lao et al., 2009). At Pámanes, thewoollymammoth remains, dating to the lateMIS 3 (25.4; 27.6 cal kaBP, Álvarez-Lao and García, 2012) include, along with a nearlycomplete skull seriously damaged during the archaeological exca-vation, two tusks, one well preserved mandible with the lastmolariform teeth (M3), one pelvis, one tibia and one femur likelybelonging to a single, fully adult individual (Carballo, 1912, 1920).

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 ¼ Bujàn,2 ¼ Las Caldas, 3e6 ¼ La Güelga, El Cierro, La Lloseta, Cueto de la Mina, 7e11 ¼ Udìas, Mina Angel, Morim, Pámanes, Minas de Heras, 12 Labeco Coba, 13 ¼ Urtiagako Lleizea,14 ¼ Clot de Llop, 15 ¼ L’Albreda, 16 ¼ cau de lesGoyes, 17 ¼ Butarque, 18 ¼ Casa Eulogio, 19 ¼ Aldehuela, 20 ¼ Arriaga, 21 ¼ Edar Culebro, 22 ¼ Padul, 23 ¼ Algar de Jo�ao Ramos,24 ¼ Figueria Brava, 25 ¼ Cruz Quebrada. Italy: 26 ¼ Arene Candide, 27 ¼ Po valley, various sites, 28 ¼ Adda Valley (Cremona), 29 ¼ Riparo Taglinte, 30 ¼ Asolo, 31 ¼ Vidor,32 ¼ Settepolesini, 33 ¼ Buca della Iena, 35 ¼ Arezzo, various sites, 36 ¼ Torrente Conca, 37 ¼ Tarquinia, 38 ¼ Canale delle Acque Alte, 39 ¼ Veroli, 40 ¼ Cardamone; Greece:41 ¼ Penios valley (Eastern Thessalia), 42 ¼ Aggìtis (Drama) (see references in text).


The Padul specimens, representing the southernmost remains ofwoolly mammoth thus far known in Iberia, as well as in Europe,were retrieved, along with a few remains of steppe bison (Bisonpriscus), red deer (Cervus elaphus) and a medium sized horse (Equussp.), from swamp clayey deposits cropping out at different sites inthe Padul Basin. Thewoollymammoth remains, ranging in age from30.6 to 40.4 cal ka BP, consist of a few molariform teeth, a completeand a fragmentary tusk, three mandibles and a number of limbbones, belonging to at least four adult males (Álvarez-Lao et al.,2009).

The Iberian findings of M. primigenius mainly dated to the lastglacial, although archaic representatives of the species have beenrecorded at the transition from the Middle to Late Pleistocene (Seséand Soto, 2002a, 2002b). In central Spain (Madrid province) woollymammoth remains have been reported from sites whose age havebeen estimated by stratigraphic correlations to be either lateMiddle Pleistocene (e.g. Casa Eulogio and Arriaga localities), orearly Late Pleistocene (e.g Aldehuela locality; ¼ Mammuthus cf.M. intermedius in Sesé and Soto, 2002a), as well as at the sites ofButarque and Edar Culebro, dated at about 100 ka (Álvarez-Lao andGarcía, 2012 and references therein). During the following earlyLate Pleistocene, until the beginning of the last glacial, there is nofirm evidence of the presence of M. primigenius in the IberianPeninsula, although a molar plate ascribed to this species has beenreported from a Mousterian level of the Arbreda cave (Gerona)(Álvarez-Lao and García, 2010).

During the last glacial, the Iberian fossil record ofM. primigeniusbecame much more abundant. Most of the Iberian woollymammoth remains date to MIS 3 and 2, but whereas those rangingin age between about 30 and 43 ka cal BP spread from the northernto southern Iberian Peninsula, those dated between about 28 and18 ka cal BP (mostly found in Late Gravettian, Solutrean and LowerMagdalenian archaeological contexts) are almost exclusivelylocated in northern Spain, mainly in the extant Basque territories.

M. primigenius remains dated to the late LGM were found in theAsturias region (northern Spain) in the LowerMagdalenian levels ofLas Caldas (about 20e17 ka cal BP), and La Lloseta (from about 20.2to 17.6 cal ka BP). The molar fragment found at Cueto de la Mina isslightly older, dating at about 23.5 ka cal BP (Álvarez-Lao andGarcía, 2012 and references therein). In the westernmost IberianPeninsula, M. primigenius was possibly present around 14 ka BP atAlgar de Jo�ao Romaos (Antunes and Cardoso, 1992). This finding,a fragment of femur showing extensive gnawing, if not reworked,would represent the only occurrence of M. primigenius in SouthernEurope at that time. Whether the mammoth from Algar de Jo�aoRamos would belong to a population that dispersed from centralEurope towards the Iberian Peninsula about 16 ka BP - whenM. primigenius reoccupied some of thewestern European territoriesit had withdrawn during the LGM - or was the last representative ofa former population that inhabited Portugal since MIS 3, is a stillunanswered question. Whatever the actual age of Algar de Jo�aoRamos M. primigenius, it is certain that woolly mammoth pop-ulations were spread in northern Iberia in a time span including theHeinrich event 2 and most of the LGM, when the climate registeredcold and dry phases and steppe environments dominated thelandscape (see Álvarez-Lao and García, 2010, 2012). Woollymammoths likely dispersed towards Iberia at that time followinga northern dispersal route and extending their range to thoseterritories where the most significant Iberian representations ofwoolly mammoth are known.

In Italy,M. primigenius is mostly recorded during MIS 4 and 3 byfew, some isolated, findings. Some other remains, older in age (MIS6) (mainly teeth but also a skull with mandible), and showing somearchaic features have been ascribed to this species (see Reggianiand Sala, 1992; Palombo and Ferretti, 2005; Mussi and Villa, 2009and references therein) (Fig. 25).

Woolly mammoth remains have been found associated withMousterian tools, as at Riparo Tagliente (Veneto), Asolo (Treviso,


Veneto), and Buca della Iena near Mommio (Lucca, Tuscany). AtRiparo Tagliente (Grezzana, Valpantena, Verona) two fragments ofplates of a young mammoth were found associated with Mouste-rian lithic implements within the lowermost layers (levels 52 to 31)of a long stratigraphic succession, ranging in age from the Mous-terian to the Epigravettian (Capuzzi and Sala, 1980; Bartolomeiet al., 1982). The abundance in these layers of Cervus elaphus,Capreolus capreolus and Alces alces, together with the presence ofMarmota marmota, indicate a moderately humid, temperate-coldclimate (Fiore et al., 2004). At Buca della Iena the fossiliferousbed, containing Mousterian lithic implements, is younger than41 ka (Pitti and Tozzi, 1971).

In the alluvial deposits dated to the Last Glacial (cf. Venzo, 1977)in the neighborhood of Pagano d’Asolo (Asolo, Treviso), a skeletonof a female of M. primigenius about 32 years old (Reggiani and Sala,1992), was found in 1878 together with a few Mousterian flakes(Dal Piaz, 1922, 1931). The direct radiocarbon date of about 27.8 kaobtained for the mammoth remains has to be rejected because thebones were consolidated using fish glue, while the Levalloisimplements associated with the skeleton fit well into either MIS 4or 3 (see Mussi and Villa, 2009 for a discussion).

Another partial skeleton of a 35 old male was retrieved in 1973from lacustrine deposits, possibly correlatable with MIS 4 croppingout at S. Giovanni di Valdobbiadine (Vidor, Treviso, Veneto) (Venzo,1977; Reggiani and Sala, 1992). The uncertainty regarding theprecise stratigraphic position of the finding, the lack of numericaldates as well as of associated artifacts makes it difficult to ascertainthe chronology of this woolly mammoth.

Excluding Asolo and Vidor, most of the other MIS 4 and 3localities yielded only isolated dental and skeletal remains. Thematerial can be confidently referred to M. primigenius on the basisof the derived character of the molars, such as the elevated numberof plates, the high lamellar frequency and thin, highly creasedenamel (Palombo and Ferretti, 2005).

In north Italy, the presence of a “mammoth steppe” duringMIS 3is indicated by the fauna with M. primigenius (14C dates onmammoth bones range from ca. 35,800 to 33,830 BP), Coelodontaantiquitatis, Megaloceros giganteus, Bison priscus, and Alces alcesfound a deposit at Settepolesini di Bondeno (eastern Po Valley,Ferrara) (Gallini and Sala, 2001). The dispersal of cold adapted taxathroughout the Po Valley was likely permitted due to the loweringof the sea level during glacial stadials, leading to the emergence ofan extensive part of the Adriatic platform. Together with otherspecies (e.g. Sicista betulina, Microtus oeconomus, Coelodonta anti-quitatis, B. priscus) M. primigenius spread southward along theAdriatic coast to about 40� 220N, as documented by the findings ofsome bones of young mammoths at Cardamone (Lecce, Apulia). Nonumerical dates are available for the Cardamone fauna. Rustioniet al. (2003) hypothesized that the faunal assemblage could dateto the LGM because of the absence of Equus hydruntinus and Damadama, but the avifauna indicates temperate-cold, quite humidclimate conditions.

In central Italy, M. primigenius is known from the alluvialdeposits cropping out at Torrente Conca (Cattolica); MontecatiniTerme (Lucca), a few sites in the Marche region, and in the Arezzobasin (Tuscany) (De Giuli, 1983; Ferretti, 2000; Palombo andFerretti, 2005 and references therein). The southernmost findingson the Tyrrhenian side of the Italian peninsula are between41� 410N (S, Anna, Veroli) and 41� 280N (Canale Acque alte, Latina,Pontina plane, southern Latium).

Among the most interesting remains is an incomplete mandiblewith the penultimate molars found during a well excavation at S.Anna, near Veroli (Frosinone, Souther Latium) (Biddittu andCelletti, 2001; Palombo unpublished data). There is no strati-graphic 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 fewM. primigenius remains has been retrieved in layers correlatedwith MIS 4 (the same layers with P. antiquus) and in the overlyinglevel correlated with MIS 3 (Blanc et al., 1957; Caloi and Palombo,1995; Farina, 2011).

The available radiocarbon dates confirm thatM. primigeniuswasstill present in Italy during MIS 3, while, and except for easternLiguria, no compelling evidence indicates its presence since theLGM. At the Gravettian Arene Candide site (Liguria, north-westernItaly), a few proboscidean bones (a second phalanx, a possiblefragment of a patella, an epiphysis fragment and two shaft splin-ters) have been found (Cassoli and Tagliacozzo,1994). The bones arenot diagnostic and have been ascribed to the woolly mammothmainly because of the age of the deposit, believed too recent foryielding straight-tusked elephant remains. A few mammoth ivoryornaments and female figurines have been also recorded in east-ernmost Gravettian sites of the Ligurian coast, close to the modernboundary with southern France, but the carved objects have beensupposed to be imported from elsewhere (Mussi et al., 2000).

All things considered, whether someM. primigenius populationsactually inhabited the Italian peninsula after MIS 3 is still an openquestion. Woolly mammoth was likely present in Liguria, and,although no radiocarbon dating is available, the hypothesis thata few individuals could have dispersed from the lower Rhone valleyto reach the easternmost Ligurian territories is reasonable, espe-cially considering, for instance, that Rangifer tarandus is recorded inItaly only in this region (Sala, 2005). The fact that someM. primigenius herds could reach Liguria from southern France doesnot necessarily imply that they further dispersed to other parts ofthe Italian peninsula. Woolly mammoths might be confined toeastern Liguria either because the low number of individualsinhabiting the region did not allowed a range extension, or becausethe Ligurian Apennines, which separates Liguria from the Po Valley,represented a severe barrier to be crossed by these elephants.Therefore, pending a confirmation about the actual age of theCardamone woolly mammoth, the still unanswered question ariseswhy any dispersion event from East Europe allowed the presence ofM. primigenius at least in northwestern Italy during MIS 2.

4. Remarks

Although a rich Upper Paleolithic iconography testifies to a longcoexistence of humans and M. primigenius during the last glacial inmost of central Eurasia and even in southern Europe, the frequencyof representations of M. primigenius left by H. sapiens during theUpper Paleolithic (ca. 35 ka to 11 ka) as portable and cave art variesin either number of representations and their geographical distri-bution. In general, objects of portable art are known fromnumerous European regions, while cave art is mostly known fromSouth West Europe, especially in France and northern Spain.

Moreover, the typology of portable art representations and therough material used to produce the objects differed during timeand across space. During the Aurignacian, for instance, depictionsof mammoth are only known in the form of ivory figurines in theSwabian Alps region (SouthWest Germany). During the Gravettian,ivory figurines are known from a number of sites in the CzechRepublic and the Russian plain, while figurines of mammoth madeby using clay are only known in the Moravian region and those ofmarl only more eastward in the Russian plain. Finally, mostengravings and tools decorated with mammoths are reported fromWestern and Central European Magdalenian sites.

As regards the three main techniques of cave art (painting,engraving and sculpture), the main documentation comes fromnumerous caves located in France and in Spain. The most eastward


caves with cave art are those of Kapova and Ignatievka in theSouthern Ural Mountains.

Comparing the distribution of woolly mammoth fossil recordwith its representation in the Upper Paleolithic art, no depictionshave been reported not only from the regions whereM. primigeniushad already disappeared by the LGM, but also from regions whereboth woolly mammoth remains and Paleolithic art were docu-mented, such as, for instance, from Switzerland and Austria. Theraison behind this lack still remains an unsolved issue.

In particular, as regards southernmost Europe, the fact that theSpanish woolly mammoth representations are mainly located inthe northern Cantabrian and Asturias regions is consistent withthe fossil record. During the LGM, M. primigenius remains havebeen found in the area northwest of the Pyrenees along the coastof the Bay of Biscay, which could also be regarded as among themost suitable dispersal routes of M. primigenius towards Iberia(Álvarez-Lao et al., 2009). Apparently, during the LGM the range ofwoolly mammoth did not reach the easternmost Iberian Penin-sula. The youngest dating of fossil remains is that of FigueiraBrava, correlating the fauna with late MIS 3 (Álvarez-Lao andGarcía, 2012 and references therein), and woolly mammoths arenot represented in the Portuguese Palaeolithic art. The hypothesisthat the absence of tundra-like environments prevented thespread of M. primigenius herds towards low latitudes in Iberiaduring MIS 2 cannot be ruled out, although it needs to besubstantiated.

The absence of any representation of M. primigenius in Italy,except maybe for the easternmost Liguria, could be regarded as anindirect evidence of the disappearance of the species by the LGM.Ongoing research could answer the question whether the absenceof M. primigenius in most of Italy during MIS 2 is due to difficultiesin reaching the Italian peninsula from eastern Europe, maybebecause of a reduced availability of suitable environments in someterritories along the potential dispersal routes.

Acknowledgements

We thank the two anonymous reviewers for their usefulcomments.

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