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Metapodial Bones of Cave CRANIUM 22, 2 - 2005 First morphometric data from the metapodial bones of Ursus from Cerè Cave (Venetia Region, North Italy) Mario Rossi & Giuseppe Santi Summary North Numerous metapodial remains of bears from the Cerè Cave (Verona Province, Venetia Region, Italy), are morphologically and morphometrically analysed here. Three species of Ursus are represented (U. deningeri Von Reichenau, 1904; U. spelaeus Rosenmüller-Heinroth, 1794 and U. arctos Linnaeus, 1758), but in different proportions, than the arctoid forms. Two of U. of with the deningerian and speloid being more numerous populations spelaeus different shown different In the sizes are and this could reflect their (geological) ages. particular, significant presence when data from other in the of deningerian fossils in the Cerè Cave, compared to caves Lombardy and Piedmont these for the Venetia with limited numbers in regions, could indicate a strong presence of species solely region very until reach Central in the Cerè Cave from than surrounding areas we Italy. Lastly, the fossil findings derive more one level of the Pleistocene and the of U. the extension of the lower to the presence deningeri justifies boundary Early Pleistocene. Samenvatting aantal middenhands- middenvoetsbeenderen beren uit de Cerè Een groot overblijfselen van en van grot (provincie hier morfometrisch Drie Verona, regio Venetië, Noord-Italië) worden morfologisch en geanalyseerd. soorten van Ursus zijn vertegenwoordigd (U. deningeri VonReichenau, 1904; U. spelaeus Rosenmüller-Heinroth, 1794 en U. arctos de dan de laatste. Linnaeus, 1758), maar in verschillende verhoudingen, waarbij twee eerste soorten meer talrijk zijn U. verschillende worden verschillende Twee populaties van spelaeus van grootte aangetoond, hetgeen op een geologische ouderdomkan wijzen. Ursus deningeri komt in de Cère-grot veelvuldig voor, in vergelijking met andere de Dit sterke deze in de ditin grotten in Lombardije en Piedmont. wijstop een aanwezigheid van soort regio Venetië; U. veel zeldzamere tegenstelling met het gebied tot aan Midden-Italië, waar deningeri een verschijning was. Tenslotte, het Pleistoceen de de fossiele vondsten in de Cerè grot zijn afkomstig van meer danéén niveau van en aanwezigheid de het Pleistoceen. van U. deningeri rechtvaardigt de uitbreiding van ondergrens tot Vroeg low Introduction a statistically significant morphodynamic index. Bear fossils from a number of caves in North rich of information Italy are a source to explain the behavioural and the evolutionary paths of Previous works this vertebrate. Caves in the Venetia Region, to those found in the compared nearby fossils the richest Studies on U. spelaeus are most Lombardy and Piedmontregions, have parti- numerous but they frequently concern the cular value because contain remains both they cranial and mandibular remains. We refer to the of archaic and recent forms of Ursus. probable studies by Santi and Rossi (2001a) and Rossi The Cerè Cave (Verona Province) (fig. 1) is the and Santi (2001) for a more extensive review. most significant from this point of view. Altoge- We here list the main works that have empha- of ther these caves contain a great quantity sised on Ursus limb and metapodial bones. mammal fossils which "...sembrano rappresen- Dubois and Stehlin (1933), and Fritz tare in realta piü momenti del Pleistocene" (Bon Koby in metatarsial and et al, 1991:189). (1950) analysed particular phalange remains. Subsequently, Koby (1950), This work will confirm that different species of after studying different fossils, was able to bears lived inside the Cere Cave and indicate a re-assemble an anterior, almost complete paw possible evolutionary step through morphome- (with only one missing uncinate) and compared tric and of morphological analyses metapodial it with that from the modern brown bear. bones. The morphodynamic analyses of the studies the bones of the loco- teeth following the Rabeder's (1999) method is Many concerning of these focus sexual in motor bears, on in progress, but the former data at least this apparatus (i.e. Kurten, 1955). In this preliminary phase of the work, seem to present dimorphism regard 13 First morphometric data from the metapodial bones of Ursus from Cere Cave (Venetia Region, North Italy) Fig 1 Geographic position of the Cerè Cave (Venetia Region, Northern Italy) Geografische positie van de Cerè grot (regio Venetië, noord Italië) Cuggiani's (1981) analysis of the long bones had dimensions correlating with juveniles), from Equi Cave (Tuscany Region) is worth noting sexual differentiation. Using a mathema- noting. Recently, other interesting hypotheses tical model, Viranta (1994) highlighted some concerning the sexual dimorphism were correlations between limb proportions and advanced by: Kunst (1992) through bivariate mass of cave bears and compared them to those distal from modern bears. The shafts wider analysis using greatest metapodal length were in and width as bears' limbs and their variables; Grandal D'Anglade cave hence, body mass who (1993, 2001), indicated that this was greater. dimorphism does not equally affect the various In the morpho-dimensional analysis of the bones of the skeleton and who noted a different remains from Arcy-sur-Cure (Yonne, France) of degree sexual dimorphism in separate popu- Baryshnicov and David (2000) confronted the lations; Rustichelli (1993) and, more in parti- problem of classifying fossils of small dimen- cular Reisinger & Hohenegger (1998), who sions with certainty. Argant (1991) and Bonifay proposed a sexual dimorphism hypothesis (1975) described the different osteological based on biometrical characters of the limb remains of Ursus from Bourgogne and from bones. Correze (France). We again refer to research carried out by Torres Di Canzio & Petronio (2001) studied the carpals, (1988) and by Kunst (1996) which involved the metacarpals, tarsal and metatarsal remains, biometrical analysis of femurs from Pleistocene most likely from a female, coming from Cola bears in Europe. These authors discovered that Cave (Aquila, Abruzzo Region, Central Italy). this bone structure originally sought to provide Lastly, Rossi and Santi (2001) preliminarily static stability. Weinstock (1999) described the proposed a biomedical analysis of the long bones of locomotor apparatus bears from the bones of the Ursus from the Buco dell'Orso material gathered in the Grosse Grotte (Blau- Cave (Laglio, Como Province, Lombardy, Italy). beuren, SW Germany). He compared their Gerhard undertook (2001) a most detailed study morpho-dimensional characteristics with fossils Ursus bones from on metapodial Austrian caves from other German caves (most of the remains 14 CRANIUM 22, 2 - 2005 - - Lateral wall of the Cerè B of the Cerè Cave from within Fig 2 A cave showing bone breccias. Entry seen - bottenbrecccie. - Cerè A Zijwand van de Cerè grot met B ingang van de grot gezien van binnenuit advancing an evolutionary hypothesis. Later, n. ssp. and Ursus spelaeus eremus n. ssp. Germonpré and Sablin (2001) studied fossils in the Totes Gebirge (Rabeder et al. 2004). from Goyet (Belgium). Santi et al. (2003) analysed metapodial remains from the Buco dell'Orso Cave. Jambresiae and Paunoviae (2002) Geographic-stratigraphical setting performed detailed osteometrical, biomecha- Cere Cave, also known as "Tana dell'orso" or nical and locomotion analyses on the Ursus limb "La is about 750mabove from Withalm Tanasela", positioned remains Croatian caves. (2001) level. It is 12m and 150 east of sea deep opens m undertook a thorough study of the metapodial Ceredo Anna inside the while Athen village (S. d'Alfaedo) evolutionof the cave bears group, Rosso Ammonitico Limestones. Its opening is et al. (2004), with biostatistical methods, circular with small dimensions, and in the analysed the different features among the cave proximity of an evident slope breach that bear species, especially Ursus spelaeus and Ursus characterises the right side of the Vajo dei deningeri. Lastly, Perego et al. (2001) extensively Falconi. This leads to wide door with studied number of bones entry a a a great U. spelaeus sub-quadrangular floor about 6,5m x 5m and is from Campo dei Fiori Massif (Varese, 6m in Walls often covered over height. are by Lombardy Region, Italy) highlighting new and older and deteriorated concretions. On the interesting biometrical correlations between southern side of the hollow, a chimney lm in Ursus limbs of different geological ages. More diameter is present that was originally recently, the evolution of the metapodial bones connected to the exterior. The Cere Cave is together with the morphodynamic indices of considered a karst hole filled to its apex by the teeth and the fossil mtDNA analyses, has bone-breccias It was allowed the of the (fig. 2). partially dug to support hypothesis further a second karst a of within the bear during cycle forming presence new taxa cave smaller cave. Today it opens across a short group of the Alps (Rabeder & Nagel 2001; tunnel which had evolved from a Rabeder Ursus originally et al, 2004). They are: ingressus n. cone From bottom to which occurred in the eastern of the pyroclastic (Pasa, 1954). sp., parts of: top the stratigraphic succession consists Alpine region and in the Dinarids of Slovenia and Croatia and the Ursus subspecies ladinicus spelaeus 15 First morphometric data from the metapodial bones of Ursus from Cere Cave (Venetia Region, North Italy) O 1. Concretionated ferrous-manganesiferous clay and the transversal width of the proximal (to the karst bed rock contact) epiphysis
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