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The Saalian Ursus thibetanus from France and Italy
i
THE SAALIAN URSUS THIBETANUS F R O M F R A N C E A N D ITAIN
EVELYNECRI~GUT-BONNOURE CRt~GUT-BONNOURE E. 1997. The Saalian Ursus thibetanus from France and Italy. [UUrsus thibetanus saalien de France et d'Italie]. GEOBIOS, 30, 2 : 285-294. Villeurbanne, le 30.04.1997. Manuscrit d6pos6 le 01.08.1994; accept6 d6finitivement le 21.12.1995. ABSTRACT - Fragmentary small ursid remains were found in the Saalian site of the C6dres Cave (Le Plan d'Aups, Var). Their comparison with Middle Pleistocene Ursidae from Italy (Reale Cave), assigned to Ursus rnediterraneus F. MAJOR, and from France (Bruges), related to U stehlini (KRETZOi),shows that all these bears should be referred to U thibetanus CUVIER.The differences in the morphometry support the hypothesis of the existence of three subspecies at the end of the Middle Pleistocene in Southern Europe: Ursus thibetanus kurteni nov. subsp, from the C6dres Cave, Ursus thibetanus mediterraneus (F. MAJOR) from Reale Cave and Ursus thibetanus vireti nov. subsp, from Bruges loam pit. Such differentiation is assumed to indicate a geographic isolation of the populations. KEYWORDS: URSIDAE, TAXINOMY, MIDDLE PLEISTOCENE, FRANCE, ITALY, GEOGRAPHIC ISOLATION. RI~SUM]~ - Des restes fragmentaires d'un petit Ursidae ont 6t6 d6couverts dans le site saalien de la Grotte des C6dres (Le Plan d'Aups, Var). Leur comparaison avec Ursus mediterraneus F. MAJOR d'Italie (Grotte de Reale) et Ursus stehlini (KRETZODde France (Argiles de Bruges) montre que ces Ursidae correspondent fi Ursus thibetanus CUVIER. Les diff6rences morphom6triques relev6es sont en faveur de l'hypoth6se de l'existence de trois sous-esp6ces d'ours du Tibet fi la fin du P16istoc6ne moyen dans le Sud de l'Europe: Ursus thibetanus kurteni nov. subsp, de la grotte des C6dres, Ursus thibetanus rnediterraneus (F. MAJOR) de la grotte de Reale et Ursus thibetanus vireti nov. subsp, des argiles de Bruges. Ces faits sugg6rent l'isolement g6ographique de ces populations. MOTS-CL]~S: URSIDAE, TAXINOMIE, PLI~ISTOC]~NE MOYEN, FRANCE, ITALIE, ISOLAT Gt~OGRAPHIQUE.
INTRODUCTION Recently, t h e C6dres Cave, in t h e S a i n t e - B a u m e M o u n t a i n r a n g e (Le P l a n d'Aups, Var), h a s yielded a M i d d l e P l e i s t o c e n e f a u n a in association with a Mousterian industry. Our preliminary s t u d y allows us to i n d i c a t e t h e p r e s e n c e of b r o w n bear, U r s u s arctos LINNE, a n d cave bear, U. spel a e u s ROSENM~LLER • HEINROTH ( D e f l e u r et al. 1988). S o m e u r s i d pieces c o r r e s p o n d to a n o t h e r U r s i d a e . T h e m o r p h o l o g i c a l f e a t u r e s allowed us to i n c l u d e t h e C 6 d r e s b e a r in t h e U. t h i b e t a n u s species ( C r 6 g u t - B o n n o u r e in D e f l e u r & Cr6gutB o n n o u r e 1995). A c o m p a r i s o n w i t h t h e R e c e n t Ursus t h i b e t a n u s a n d w i t h t h e little b e a r s assig n e d to U. m e d i t e r r a n e u s F. MAJOR a n d U. stehlini (KRETZOI)allows a n e w i n t e r p r e t a t i o n of t h e s e little bears.
THE BEAR FROM THE CEDRES CAVE (FRANCE) T h e m a t e r i a l belongs to t h r e e individuals. It is composed by eleven pieces ( u p p e r t e e t h : left I3 (C6 SII), left P4 (C6 SI), r i g h t P4 (CIII 239), i n c o m p l e t e m a x i l l a r y w i t h P4, M1 a n d M2 (right, C6), r i g h t M2 (C6 SII), distal p a r t of a left M2 (C6 SII) - lower teeth: left P4 (C3-4-5 SI), left M3 (C56 SI) - post-cranial: I ° left m e t a t a r s a l , II ° & I I I ° left m e t a t a r s a l (CII), r i g h t V ° m e t a t a r s a l (C3-4
sI). LOWER TEETH T h e m o r p h o l o g y is t h e s a m e as in t h e R e c e n t U. thibetanus. T h e protoconid of P4 is s i t u a t e d anteriorly. T h e m e s i a l r i m is concave a n d t h e distal one p r e s e n t s a little slot. T h e lingual face of t h e
286
1 cnl
la
3a
1 cm
2a
3c
3b
1 cm
1 cm
FIGURE 1-3 - 1. Ursus thibetanus kurteni nov. subsp., C~dres Cave. 1. Left lower M3, C5-6 SI, occlusal view. Scale in cm. 2. Ursus thibetanus kurteni nov. subsp., C~dres Cave. Holotype, " upper right row, C6. a: vestibular view, b: occlusal view. Scale in cm. 3. Ursus thibetanus kurteni nov. subsp., C~dres Cave. a: upper right P4, C6 SI, lingual view, b: upper left P4 CIII 239, lingual view, e: upper right M2, C6 SII, occlusal view. Scale in cm. Photographies A. Guerrand (Calvet Museum). 1. Grotte des 1 cm
2b
t h i r d m o l a r , o v a l in o u t l i n e , p r e s e n t s l i t t l e w r i n k l e s , well d e v e l o p e d on t h e a n t e r o - v e s t i b u l a r face (Fig. 1). A cuspid, p r o b a b l y t h e protoconid, is p r e s e n t . A d i a g o n a l r i d g e r u n s f r o m it to t h e m i d d le of t h e crown. T h e v e s t i b u l a r conids set internally: consequently, t h e v e s t i b u l a r face is oblique. (Tab71) UPPER TEETH AND MAXILLARY T h e I3 is s l e n d e r a n d h i g h e r t h a n in U. a r c t o s . T h e p a l a t a l face is convex w i t h a ridge r u n n i n g off. An i m p o r t a n t c i n g u l u m exists a l o n g t h e l i n g u a l base. On t h e m a x i l l a r y , t h e i n f r a - o r b i t a l f o r a m e n is l o c a l i z a t e d quite p e r p e n d i c u l a r l y to t h e m i d d l e of
C~dres. M3 inf~rieure droite, C5-6 SI, rue occlusale. 2. Grotte des C~dres. Holotype, rangge dentaire supgrieure droite, C6. a: vue vestibulaire, b: vue occlusaIe. 3. Grotte des C~dres. a: P4 supgrieure droite, C6 SI, rue linguale, b: P4 sup~rieure gauche, CIII 239, vue linguale, e" M2 supgrieure gauche, C6 SII, rue
t h e M1 p a r a c o n e as in t h e R e c e n t b l a c k b e a r w h o s e m u z z l e is s h o r t (Fig. 2a). T h i s f o r a m e n in U. a r c t o s is s i t u a t e d p e r p e n d i c u l a r l y to t h e m i d d le of t h e M1 m e t a c o n e or to t h e b o u n d a r y of t h e two molars. T h e M2 v e s t i b u l a r cones a r e l o w e r t h a n t h o s e of t h e first molar. T h i s f e a t u r e is char a c t e r i s t i c of U. t h i b e t a n u s ( F i s t a n i & Cr~gutB o n n o u r e 1993). T h e P4 consists of t h r e e cusps, g e n t l y d i l a t e d a n d w i t h o u t a n y a c c e s s o r y denticles. T h e c o n e - s h a p e d d e u t e r o c o n e is p l a c e d f o r w a r d t h e n o t c h s e p a r a t i n g p a r a - m e t a c o n e (Fig. 2b,3a,b). C o n s e q u e n t l y t h e outline of t h e c r o w n is m o r e or less t r i a n g u l a r . T h e p a r a c o n e , slightly h i g h e r t h a n t h e m e t a c o n e ,
287 ! ,Lower teeth L IP4 ,Les C ~ d r e s !C3-4-5 SI Recent. n Variation Mean
10,61 9 8,88-10,96 9,82
5,91 9 5,24-6,16 5,85
6,2 9 5,22-6,54 5,78
7 8 5,12-9,43 7,16
iM2 Reale IGF 4808V IGF4BO7V Recent - n Variatien [Mean
20,62 21,45 9 17,38-21,37 18,7
14 14,61 7 11L14,12 12
7,29 7,41 8 5,49-7,11 6,34
7,33 7,3 8 4,33-6,99 i 5,61
17,4 9 12,71-17,06 14,52
12,8 9 9,96-12,19 11,11
'LMe3sC~dres CS-6 SI Recent - n Variation Mean I
1
2
3
4
5
6
7
6,64 5,81 9 4,28-5,62 5,11
6,85 6,9 9 5,1-6,57 5,94
8
12,96 12,28 9 10,26-11,91 10,92
9
12,24 11,81 9 10,11-11,66 10,93
7,32 11,62 8 9,89411,43 10,75
TABLE 1 - Dimensions of the lower teeth in Ursus thibetanus kurteni nov. subsp, from C~dres Cave, Ursus thibetanus mediterraneus from Reale Cave and Recent asiatic black bear (in ram). For P4 - total length (1), total width (2), protoconid height (3), distance from the protoconid cusp to the distal point (4). For M2 - Length: total (1), trigonid (2); height: protoconid (3), hypoconid (4), metaeonid (5),entoconid (6); width: trigonid (7), talonid (8), minimal (9). For M3 - total length (1), total width (2). Dimensions des dents jugales infdrieures d ~ r s u s t h i b e t a n u s kurteni nov. subsp, de la grotte des C~dres (c), d'Ursus t h i b e t a n u s m e d i t e r r a n e u s de la grotte de Reale et de Fours du Tibet actuel (en mm). Rgfdrence des mesures: P4 - Longueur totale (1), largeur totale (2), hauteur du protoconide (3), distance du sommet du protoconide au bord distal (4). M2 - Longueur: totale (1), trigonide (2); hauteur: protoconide (3), hypoconide (4), m4taconide (5), entoconide (6); largeur: trigonide (7), talonide (8), minimale (9). M3 - Longueur totaIe(1), largeur totale (2).
is characterized by a concave mesial rim. The edge r u n n i n g from the top to the base is curved outward (Fig. 2b). The metacone and the deuterocone are higher t h a n in the Recent species and t h a n in the two other Middle Pleistocene little bears examined (Fig. 4, parameters 8,9, Tab. 2) and the paracone is comparatively lower (parameter 7). A basal cingulum is present on each side of the deuterocone. It runs along the vestibular face.
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The cusps of the molars set internally relative to the base of the crown and are weakly dilated:
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FIGURE 4 - Ratio diagram of the proportions of the upper P4 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c), Ursus thibetanus mediterraneus from Reale Cave (m) and Ursus. thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in Table 2. Length: vestibular (1), paracone (2), metacone (3), deuterocone (4); width: paracone (5), in deuterocone axis (6); height: paracone (7), metacone (8), deuterocone (9); distance from the deuterocone cusp to the mesial point (10). Diagramme des rapports des proportions de la P4 supdrieure d ~ r s u s thibetanus kurteni nov. subsp, de la grotte des C~dres (c), d~Ursus thibetanus mediterraneus de la grotte de Reale (m) et d ~ r s u s thibetanus vireti nov. subsp, des argiles de Bruges (b). Les diffgrences sont exprimdes en logarithmes ddcimaux. Base de rgf4rence O: Ursus thibetanus actuel. Dimensions dans le tableau 2. Longueur: externe (1), parac6ne (2), mdtac6ne (3), deutdrocSne (4); largeur: paracSne (5), au niveau du deutdroc6ne (6); hauteur: parac6ne (7), mgtacSne (8), deutdrocOne (9); distance milieu deut4roc6ne-bord mgsial (10).
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0,0
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288 Upper teeth
66•
1
P4 Les C6dres CII1239 14,37 9,65 5,7 10,11 S1 C6 15,4 C6 14,11 9,29 ' 5,66 Reale 9,45 6,8 5,94 IGF-4806V 15,43 Bruges 9,16 5,8 6,05 JRight 14,38 8,9 5,67 6,17 Left 14,76 I0 I0 10 Recent - n 10 Variation 111,41-13,25 6,71-8,47 4,08-5,65 4,48-7,32 Mean 12,21 5,84
/
L
M1 ,Les C~dres C6ruges Right Recent ,n /Variation .Mean
iLes C~dres C~SII
6
10
8,41 7,92 7,93
10,97 10,47 10,21
8,17 8,56 8
7,04 7,2 7,28
5,9 5,99 5,78
10,27 10,17
7,14
11,25
9,12
7,82
6,49
10,82
8,11 7,46 10 6,58-7,43 7,03
9,82 10,04 10 8,21-9,17 8,51
8,9 9,1 10 6,61-8,09 7,26
6,93 7,179
5,58 5~74
11,64 I 10,48 I
5,03-6,21 5,59
] 3,36
19,49 18.75 10 16,67-19,07 17,99
6,375-79 5,%1y2
28,51 28,37
2,4 I0 2,12-2.52 2,30
14,14
15,52
14,32
8.9
14.3 14.79 14,27 9 10 10 9 10 11,61-13,41 12,13-14,33 11,99-13,22 7,54-9,01 12,58 13,39 12,69 8,28
9,5
3,87-5,33 7,6910,64 4,49 8,46 I i
8.75
8,26
8,46
8;42
8~15
7,516
7,54-8,69 8,09
6,571,73 %7,;7?2
4,95 3,71
16,99 16,59 16,38
10,32 11,15 10,91
16,36 15,97
15,66 15,18 15,48
11,84 11,71 11,46
7,65 7,48
7,56 7.46 ' 7,56
3,75
14,87
10,14
15,42
15,68
10,55
7,34
7,66
17,68 19~18
3,4 14,699
15,29 15~33
7,05 7~2
8
C6 SI1 IReale IGF 4806V . 26,02 7,05 7,44 Bruges Right 28,62 8,24 10,21 7~9 Left 29,29 9,03 Recent - n 10 9 Variation 23,09-29,04 7,72-11,05 5,61-9,69 !Mean 25,61 i 9,26 7,4
11
2,85 2,85
16,01 16,14 9 13,07-17,84 9,93-13,27 10,19-14,8 15,24 11,18 13,63
12,9 6,97 13,98 9 12,93-14,52 9,06-13,4 13,78 10,91
! 7£
5,73-6,8] 5,68-6,84 6,23 ] 6,13
TABLE 2 - Dimensions of the upper teeth in Ursus thibetanus kurteni nov. subsp, from C6dres Cave, Ursus thibetanus mediterraneus from Reale Cave, Ursus thibetanus vireti nov. subsp, from Bruges and Recent asiatic black bear (in mm). For P4 - Length: yestibular (1), paracone (2), metacone (3), deuterocone (4); width: paracone (5), in deuterocone axis (6); height: paracone (7), metacone (8), deuterocone (9); distance from the deuterocone cusp to the mesial point (10) For M1 - Length: vestibular (1), paracone (2), metacone (3), parastyle (4); width: first lobe (5), second lobe (6), minimal (7); height: paracone (8), metacone (9), protoeone (10), hypocone (11).- For M2 - Length: vestibular (1), paracone (2), metacone (3), parastyle (4), from the metacone top to the distal point (5), from the metacone distal base to the distal point (6); width: first lobe (7), second lobe (8), talon (9); height: paraeone (10), metacone (11). D i m e n s i o n s des dents j u g a l e s supgrieures d'Ursus thibetanus kurteni nov. subsp, de la grotte des C~dres (c), d~Ursus thibetanus mediterraneus de la grotte de Reale, d'Ursus thibetanus vireti nov. subsp, de Bruges et de Fours d u Tibet actuel (en ram). Rdf~rence des mesures: P 4 - longueur externe (1), du parac6ne (2), du mgtacdne (3), d u deutdrocSne (4), largeur d u parac(me (5), au niveau d u deutdrocSne (6), h a u t e u r du parac6ne (7), d u m~tacSne (8), d u deutdroc6ne (9), distance milieu deutdrocSne-bord mdsial (10) - M 1 - longueur externe (1), d u parac6ne (2), du mgtac6ne (3), du parastyle (4), largeur d u p r e m i e r lobe (5), d u deuxi~me lobe (6), m i n i m a l e (7% h a u t e u r d u parac6ne (8), du m~tacSne (9), du protoc6ne (10), de l'hypocSne (11) - M 2 - longueur: externe (1), paracSne (2), m~tacSne (3), parastyle (4); distance: de la pointe d u mdtac~ne au bord distal de la couronne (5), de la base distale d u mgtacdne au bord distal de la couronne (6); largeur: p r e m i e r lobe (7), second lobe (8), talon (9); h a u t e u r parac~ne (10), mgtac6ne (11).
consequently, the medial channel is larger t h a n in U. arctos (Fig. 2b). The lingual cones constitute a thin and sharp edge. A well developed parastyle is present and the vestibular cingulum is weak. The paracone and metacone of the upper M1 are equal in size. A little slot exists on the distal rim of the metacone suggesting a differenciation of the metastyle. Two accessory cusps exist between the protocone and hypocone. The second lobe is broader t h a n in the Recent black bear (Fig. 5, parameter 6, Tab. 2); the parastyle is stronger (parameter 4) and in consequence the paracone is shorter (parameter 2). Finally all the cones are higher,
!
especially the hypocone (parameter l l ) . T h e length of the M2 is shorter t h a n the combined length of M1 and P4. A transversal ridge runs from the paracone to the hypocone (Fig. 2b, Fig. 3c). It corresponds to the mesial limit of the lower M3 which is placed under the talon of the second upper molar. This ridge is not very developed or missing in the Recent species. On the right isolated M2 (Fig. 3c), an edge exists on the internal side of the paracone. The parastyle is powerful: it is absent on nine of the ten recent specimens taken in comparison. Erdbrink indicates t h a t it is "very often present" but without a n y precision on
289 The length of the dental row is stronger than in the Recent U. thibetanus. Effectively, the P4 and the M2 are well developed. On the other hand, the first molar is shorter. This is a characteristic of all the Pleistocene bears belonging to the Tibetan group (Fistani & Cr6gut-Bonnoure 1993).
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POST-CRANIAL
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The post cranial material is scarce. The diaphysis of the metapodes is slight and short. The articular facets of the upper articulation are small and the distal condyle is globular. These features are present on the recent specimens seen. The third metatarsal shows an oblique proximal articular side: the medial side is inner than the lateral one. A same morphology is to be observed on the Gajtan specimens (Fistani & Cr6gut-Bonnoure 1993), but the metapodes of this site are stronger. CONCLUSION
I 0
11
The dental features indicate that the C~dres bear clusters with U. thibetanus b u t differences occur between the Pleistocene and the Recent forms. I propose for this bear the following diagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAEViret, 1955 Genus Ursus LINNE, 1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus kurteni nov. subsp.
FIGURE 5 - Ratio diagram of the proportions of the upper M1 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c) and Ursus thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in table 2. Length: vestibular (1), paracone (2), metacone (3), parastyle (4); width: first lobe (5), second lobe (6), minimal (7); height: paracone (8), metacone (9), protocone (10), hypocone (11). Diagramme des rapports des proportions de la M1 supdrieure d~Ursus thibetanus kurteni nov. subsp, de la grotte des C~dres (c) d~Ursus thibetanus vireti nov. subsp, des argiles de Bruges (b). Les diffd-
rences sont exprimdes en logarithmes dgcimaux. Base de r~fdrence O: Ursus thibetanus actuel. Dimensions dans le tableau 2. Longueur: externe (1), du paracSne (2), du mdtacSne (3), du parastyle (4); largeur: premier lobe (5), deuxi~me lobe (6), minimale (7); hauteur: paracSne (8), mdtacSne (9), protocSne (10), hypoe6ne (11).
this frequency in the sub-species (1953: 150). An important cingulum is present at the lingual base of the crown.The distal rim of the metacone is longer t h a n the mesial compared to the case in the Recent black bear. Consequently the length from the top of the cusp to the distal side (Fig. 6, parameter 5, Tabl. 2) is stronger while the length from the base of the cusp to the distal side (parameter 6), which corresponds to the talon length, is shorter. The first lobe is broader than the second which is broader than the talon: consequently the width decreases from the first to the third lobe (parameters 7,8,9).
D e r i v a t i o n o m i n i s - In respect to late Professor BjSrn Kurt6n. L o c u s t y p i c u s - C~dres Cave (=Baume du Couloir Ovale), situated in the Mesozoic Sainte Baume Mountain, Provence, Le Plan d'Aups, Var, France. S t r a t u m t y p i c u m - Archeological levels II and III of Alban Defleur excavations, and archeological levels 5 and 6 of Max Escalon de Fonton excavations, sequence of the end of the Saale. H o l o t y p e - Upper right tooth row with the carnassial and the
two molars, level 6. Conserved in the Museum Requien (M.R.) Avignon, France. Fig. 2 a & b and fig. 52 in Defleur & Cr6gutBonnoure 1995. D i a g n o s i s - Ursus thibetanus with strong dentition compared with the Recent black bear. Denticles of the upper teeth with moderate dilatation. Metacone of the upper P4 high, and paracone relatively low. Paracone characterized by a concave mesial rim and an edge running from the top to the base curved outwards. Cone-shaped deuterocone high and strait, situated anteriorly to the vestibular notch. Metacone and deuterocone higher than in the Recent species and deuterocone lower. Upper M1 with powerful parastyle. Paracone shorter and hypocone higher than in the Recent black bear. Second lobe thicker than the first and thicker than in the Recent species. Upper M2 with a powerful parastyle and a transversal ridge running from the
290 paracone to the hypocone. Distal rim of the metacone longer than the mesial one, compared to Recent U thibetanus and consequently a short talon. Decreasing anteroposterior width of the lobes. Upper I3, lower P4 and metapodes identical as in the Recent species. +0,20
Synonymy 1954 Ursus arctos, Charles, p. 294 1971 Ursus arctos L. fossilis GOLDFUSS, Bonifay, p. 243 1995 Ursus thibetanus ssp., Cr~gut-Bonnoure in Defleur & Cr~gut-Bonnoure, p. 58-60
+0,15
THE BEAR FROM REALE CAVE (ITALY)
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2~ .:11 ...., +0,05
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LOWER TEETH
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Two M2, belonging to two different individuals, and one M3 were studied.
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2
3
5
6
Some elements belonging to a little ursid were discovered in the Reale Cave (Elbe Island, Porto Longone or Porto Azzuro) and attribuated by Forsyth Major (1873) to a new species: U. mediterraneus. This material, conserved in the Geological and Paleontological M u s e u m of Firenze, was recently revised by Rustioni & Mazza (1993) and referred to Ursus cf. thibetanus.
7
8
9
10
11
4
Fmu~E 6 - Ratio diagram of the proportions of the upper M2 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c), Ursus thibetanus mediterraneus from Reale Cave (m) and Ursus thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in Table 2. Length: vestibular (1), paracone (2), metacone (3), parastyle (4), from the metacone top to the distal point (5), from the metacone distal base to the distal point (6); width: first lobe (7), second lobe (8), talon (9); height: paracone (10), metacone (11).
The talonid of the left M2 is partially broken. All the conids set internally and are relatively high. Their lateral longitudinal profiles constitute a broken line. The protoconid is connected to the metaconid by a rugged transverse '~" shaped ridge outline.The mesial '~g" top is localized on the mesio-distal axis of the crown. An other ridge runs from this point forward and ends with a little accessory cusp. In mesial view, the sides of the transverse ridge are oblique from the external cusps toward the center, forming a large valley. The anterior part of the trigonid is conform with U thibetanus because "In front of the protoconid and the metaconid a semicircular ridge along the edge of the molar encloses a hollowed part of the mastication surface" (Erdbrink 1953: 153). An oblique ridge runs from the base of the protoconid distal rim to the middle of the crown, at the level of the lateral notches corresponding to the trigonid-talonid separation.The hypoconid is well developed and a ridge runs from it to the preDiagramme des rapports des proportions de la M 2 supgrieure d'Ursus t h i b e t a n u s kurteni nov. subsp, de la grotte des C~dres (c), d~tJrsus t h i b e t a n u s m e d i t e r r a n e u s de la grotte de Reale (m) et d'Ursus t h i b e t a n u s vireti nov. subsp, des argiles de Bruges (b). Les diffdrences sont exprimges en logarithmes ddcimaux. Base de rgfdrence O: Ursus t h i b e t a n u s actuel. Dimensions dans le tableau 2. Longueur: externe (1), paracSne (2), mdtacdne (3), parastyle (4); distance: de la pointe du mdtac6ne au bord distal de la couronne (5), de la base distale du mdtacSne au bord distal de la couronne (6); largeur: premier lobe (8), second lobe (8), talon (9); hauteur: paracSne (10), mgtacOne (11).
291 cedent crest. Two accessory cusps exist in front of the metaconid, which is the highest denticle of the tooth, and two other at the base of the distal rim.The endoconid, higher than the hypoconid, is enclosed by two little cusps. The morphology of the right M2 is the same (Tab. 2). The left M3 described originally as U mediterraneus (Del Campana 1910:18 and pl. III, fig. 6) belongs in fact to U spelaeus. Bonifay (1971: 218) pointed its great dimensions. The crown is bilobated and the occlusa] surface presents numerous denticles as in the cave bear (Clot 1980; Ballesio 1983). UPPER TEETH Forsyth Major pointed out a left M2. It is necessary to add to the material a left P4. They belong to a same individual which is young and the same that the lower left M2. The P4 was assigned to a cave bear milk tooth Cdi latte" on the label). According to the morphometry, this specimen belongs to U. thibetanus. This premolar, constituted by three cusps weakly dilated, has a triangular shape. The paracone has a straight mesial rim which shows a thin edge. At the base a slight ridge, coming from the top of the denticle, runs along the middle of the lingual face. The paracone is higher than the metacone. These two cusps are respectively lower and higher than in the Recent black bear (Fig. 4, parameters 7,8, Tab. 2) and the deuterocone is higher (parameter 9). Compared to the U. thibetanus from the C~dres Cave, the proportions are the same but the paracone is shorter (parameter 2) and the metacone is longer (parameter 3). A basal cingulum is present on each side of the deuterocone and runs along the vestibular base. A small accessory cusp exists at the mesial base of this cusp which is quite in the axis of the vestibular notch. The cones of the M2 is more internally offset relative to the base of the crown than in the Recent U thibetanus and in the C~dres bear, whose vestibular face is less oblique. The parastyle (Fig. 6, parameter 4, Tab. 2) is smaller and lower than in U thibetanus kurteni and consequently the paracone is longer (parameter 2). The paracone and the metacone are weakly dilated. A transversal ridge runs from the top of each cone to the lingual base, reaching about the middle of the crown. An accessory cusp is present at the base of the distal rim of the paracone. The lingual cusps, lower than the vestibular one, constitute a thin sinuous ridge fringed by an important cingulum. The protocone is the lowest cone. Two accessory cusps exist between it and the hypocone. The mesial rim of this last one is short (1,97 mm) compared with its distal counterpart (6,20 ram). Little wringles occupies all the occlusal face. The second lobe is slightly broader than the first
one (Fig. 6, parameters 7,8). The talon is very strait as in U. thibetanus kurteni (parameter 9) but shorter (parameter 5). CONCLUSION The Reale bear can be assigned to U. thibetanus. The material shows analogies with the C~dres ursid but morphological and metrical differences exist between them. In consequence, I propose the following neodiagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAEViret, 1955 Genus Ursus LINN]~,1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus mediterraneus (FoRSYTH Derivatio nominis
MAJOR,
1873).
- Ursid from Mediterranean
zone.
Locus typicus - From the Reale Cave situated in Porto Longone (=Porto Azzuro), Elb Island (Italy). Stratum typicum - End of the Saalian, or early beginning of the Upper Pleistocene. Holotype - Left upper P4 and M2 (IGF 4806 V), and left lower M2 (IGF 46), Museum of Geology and Paleontology, University of Firenze, Italy.
Neodiagnosis U thibetanus with strong dentition compared to the Recent black bear. Denticles of the lower and the upper teeth few dilated. Paracone of the upper P4 higher than the metacone and with straight mesial rim. Deuterocone localized in the axis of the vestibular notch. Paracone and metacone respectively lower and higher and deuterocone higher than in the Recent black bear. Paracone shorter than in U thibetanus kurteni and metacone longer. Upper M2 with a moderate parastyle and a short distal metacone rim; paracone longer than in U. thibetanus kurteni; talon short and strait. Lower M2 with an angulous transversal mesial ridge extended forward by an antero-posterior edge. Simplified synonymy 1873 Ursus mediterraneus, Forsyth Major, p. 8. 1910 Ursus sp., Del Campana, p. 17-19, pl. III, fig. 1-7. 1958 Ursus (Euarctos) mediterraneus, Thenius, p. 638. 1971 Ursus mediterraneus, Bonifay, p. 218-219. 1977 Ursus thibetanus mediterraneus, Kurten, p. 64-65. 1993 Ursus cf. thibetanus, Rustioni & Mazza, p. 35-38, fig. 1.
THE BEAR FROM THE BRUGES LOAM PIT (FRANCE) Some teeth discovered in the loam pit at Bruges (Gironde, France) were a t t r i b u a t e d by Viret (1947) to U. stehlini (KRETZOD. The material is
292 composed by two upper canines (left and right), a right maxillary with P4, M1 and M2, two isolated upper left teeth (one P4 and one M2) corresponding to a same animal, two lower canines (left and right) belonging to an other individual. The isolated upper M1 described by Viret is apparently lost. The two lower and the right upper canines were not described by this author. LOWER TEETH The two canines, laterally compressed, are more curved t h a n the upper ones. A thin sharp lingual ridge runs from the top to the base of the crown as in the Recent black bear.
UPPER
TEETH
AND
MAXILLARY
The two upper canines show a sharp ridge on their lingual face like in the Recent U. thibetanus. According to Viret, the paracone of the P4, sharppointed, is higher t h a n the metacone. But, compared with the C~dres and Reale bears, the proportions of these cusps and of the deuterocone are quite the same as in the Recent black bear (Fig. 4, parameter 7,8,9, Tab. 2). A transversal ridge is present on the lingual face of the paracone whose mesia] rim is rectilinear. The deuterocone, situated in the axis of the vestibular notch, is slightly dilated and elongated. On the maxillary, it is prolongated forward by a thin edge. On the isolated P4 an accessory cusp is present. A cingnlum runs on both sides of the lingual cone and exists on the vestibular base. The morphology of the M1 is the same as in the Recent U. thibetanus and in the C~dres bear. In comparison with the last one, the parastyle is smaller (Fig. 5, p a r a m e t e r 4, Tab. 2), the second lobe straiter (parameter 6), the paracone higher and the metacone lower (parameters 8,9). The protocone and the hypocone are worned out. An important cingulum occupies the lingual base of the crown. The M2 is without parastyle (Fig. 6, parameter 4, Tab. 2). However the mesial rim of the paracone is slighty in zigzag in occlusal view. A thin edge occupies the middle of the paracone lingual face. It is absent on the metacone. The paracone is longer t h a n in the C~dres and Reale bears but stays shorter t h a n in the Recent U thibetanus (parameter 2). The hypocone is well developed. The second lobe is broader t h a n the first (parameters 7,8). The talon, which is long (parameters 5,6), is broader than in the Recent U. thibetanus (parameter 9). CONCLUSION Viret thinks t h a t 'Tours de Bruges montre une similitude non seulement de taille mais encore de
structure" with U. thibetanus (Viret 1947). The morphology is effectively the same as in the Recent species. Erdbrink has the same opinion (1953). The differences pointed out by Viret (great development of the upper P4, presence of an important cingulum on the upper M1 and M2) are not specific but concern individual variation. The last author referred the Bruges material to U. stehlini (KRETZOD,discovered in the Lower Middle Pleistocene sites of M a u e r and Mosbach in Germany. The general features of this ursid, never revised since the yon Reichenau description, seem to be conform with U. thibetanus (Kurt~n 1957; Erdbrink 1953; Ficarrelli 1979). In m y opinion, it is not possible to identify the Bruges bear with the Mauer and Mosbach species: this last ursid is older, the material is composed by lower jaws and the morphologic variation existing in the upper teeth of the C~dres, Reale and Bruges ursids suggests an important plasticity of the Pleistocene black bear. During the Saalian period, U. thibetanus is present in France in Achenheim (Wernert 1957) where it is assigned to U. schertzi by Dehm (1943). Unfortunately, the material, a lower jaw, is lost: it is not possible to correlate the Bruges upper teeth with the Achenheim lower ones. Nevertheless, in U. schertzi the carnassial is very short (Fistani & Cr6gnt-Bonnoure 1993) which m u s t have a repercussion on the upper row proportions. Now the dental proportions of the upper tooth row of the Bruges bear are the same as in the other Pleistocene small bears examined: consequently it is not possible to refer the materials of these two sites to a same form. If we take into account the particularities of the Bruges bear and its differences with the C~dres and Reale U thibetanus, it is possible to propose the following diagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAE Viret, 1955 Genus Ursus LINNE, 1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus vireti nov. subsp. Derivatio nominis
- In respect to late Professor Jean Viret.
- Loam pit at Bruges, Gironde,France. S t r a t u m t y p i c u m - Holsteinian or early beginning of the Saalian period. I-Iolotype - Upper right row with the carnassial and the two molars, National Museum of Natural History, Paris (France). Mold of this specimen in the Department of Earth Sciences, University Claude Bernard-LyonI, Villeurbanne, France. Locus typicus
U. thibetanus with strong dentition compared to the Recent black bear. Paracone of the upper P4 higher t h a n the metacone and with straight mesial rim. Deuterocone elongated and
Diagnosis
-
293 s i t uated in th e axis of the vestibular notch. Paracone, m e t a c o n e and de ut e r ocone hei ght s n e a r l y the same as in t he Recent U. thibetanus. U p per M1 with a m o d e r a t e parastyle and a weak metastyle. Second lobe broader t h a n the first one and s t r a i t e r t h a n in t he Recent black bear. Upper M2 with o u t parastyle. Talon elongated and broader compared to Recent U. thibetanus. Simplified s y n o n y m y
1947 Ursusstehlini, Viret, p.356-360. 1948 Ursusstehlini, Viret & Balland, p. 238-239. 1971 Plionarctosstehlini, Bonifay,p. 216-217.
GENERAL CONCLUSION T h r e e dental morphotypes exist in U. thibetanus l i n eag e d u r i n g t h e Middle Pl ei s t oc e ne in S o u t h e r n Europe. G reat analogies could be pointed out between the C~dres and Reale bears. However, the morphometrical characters are sufficiently different to refute the hypothesis of a continuous variation in a same population, such differences not existing in t h e r e c e n t m a t e r i a l exam i ne d. C o n s e q u e n t l y these two ursids can be r e f er r ed to different subspecies: U thibetanus kurteni nov. subsp, from the C~dres Cave and U thibetanus mediterraneus (F. MAJOR) from th e Reale Cave. According to the morphometry, the Bruges bear is clearly different. Therefore, it is possible to assign it to another sub-species: U. thibetanus vireti nov. subsp. The t h r e e sites are considered Middle Pleistocene in age, b u t exact chronological correlations are unknown. The C~dres Cave has yielded a well preserved fauna with Canis lupus, Vulpes vulpes, Ursus arctos, Ursus spelaeus, Lynx spelaea, Fells silvestris, Equus sp.,
Sus scrofa, Cervus elaphus, Capreolus capreolus, Rangifer tarandus, Bos primigenius, Bison sp., Hemitragus cedrensis, Rupicapra rupicapra, Oryctolagus cuniculus baumensis, Marmota marmota mesostyla, Apodemus sylvaticus, Pliomys gr. episcopalls, Terricola duodecimcostatus, Microtus agrestis cf. jansoni, Microtus brecciensis defleuri, Arvicola sapidus aupsensis, cf. Alectoris graeca, Columba cf. livia, cf. Turdus torquatus, Coccothraustes coccothraustes, Garrulus glandarius, Pyrrhocorax graculus cf. vetus. This assemblage date to the end of the
The Bruges loam pit has yielded Lynx sp., U arctos, Palaeoloxodon antiquus, D. hemitoechus, S. scrofa, C. eIaphus, C. capreolus, Castor fiber, Trogontherium sp. (Balland 1963; GuSrin 1980). According to the flora, its age is Holsteinian (Elhai 1966). A comparative study of the f a u n a is lacking and no more informations are available. A p p a r e n t l y the Bruges black bear is the oldest and t he Reale bear could be the younger. Hence if t hese t h r e e black bears are not exactly contamporaneous, th e differences between t h e m m a y correspond to a rapid evolution of the species and could have a chronological connotation. With regard to t he morphological analogies, it is clear t h a t the origin of t he C~dres and Reale bears is common. If t hey date to t he end of t he Saalian, the differences bet w een t h e m could reflect a geographical differenciation in consequence of th e Alpine Arch which can act as a geographical barrier. The origin of the Provence-Tuscany black bear group is unknown. T h e y could evolve from the Bruges group. May be the Rh6ne and th e Durance rivers, which during t he Pleistocene flowed separately into the M e d i t e r r a n e a n sea, could be at the origin of this chronological isolation. Such an isolation is suggested for t he t a h r from the C~dres Cave (Crggut-Bonneure 1989). In conclusion, two groups of black b e a r do exist d u r i n g t h e Middle P l e i s t o c e n e in S o u t h e r n Europe with at least t h r e e different subspecies clearly individualized. Thus it appears t h a t th e Pleistocene black bear shows various morphological and biometrical p a t t e r n s but more m a t e r i a l is required for a bet t er u n d e r s t a n d i n g of t he evolution of the species. Acknowledgments - I would like to thank Prof. A. Azzaroli, Prof. D. Torre and Dr. P. Mazza for the assistance and remarks they allow me during my study of the Italian collections. I am most indebted to Prof. B. Sala for his help in literature data, and to Prof. R. Ballesio and Dr. A. Argant for the critical reading of the manuscript. Also, I thank Prof. L. Ginsburg and Dr. A. Prieur who permit me to study the Bruges material. I express my gratitude to Dr. A. Defleur and Dr. M. Escalon de Fonton for having permit me to study the C~dres fauna which is preserved in the Museum Requien, thanks to Dir. A. Jacob from the D.R.A.C. My sincere gratitude to A. Guerrand for the photographies and to M. BuissenCatif for the illustration.
Saalian period (Defleur & Cr6gut-Bonnoure 1995). The Reale f a u n a is composed with U. spelaeus, Lynx sp., Panthera spelaea, S. scrofa, Hippopotamus sp., Dicerorhinus sp., C. elaphus, C. capreolus, Lepus sp. (Del C a m p a n a 1910) and could be of Saalian age, or could "come from two different stratigraphic levels" of the Upper Pleistocene (Rustioni & Mazza 1993: 37).
REFERENCES ARGANTA 1991 - Carnivores quaternaires de Bourgogne. Documents des Laboratoires de Ggologie de Lyon, 115: 1-301. BALLANDR. 1963 - Le gisement pleistocene des "Argiles de Bruges" (Gironde). Cahiers des Naturalistes, N.S., 19: 57-63.
294 BALLESIOR. 1983 - Le gisement pleistocene sup~rieur de la grotte de Jaurens ~ Nespouls, Corr~ze, France. Les Carnivores (Mammalia, Carnivora): Ursidae. Nouvelles Archives du Musge d'Histoire naturelle de Lyon, 21: 943. BONIFAY M.-F. 1971 - Carnivores q n a t e r n a i r e s du SudE s t de la France. Mdmoires du Musdum national d'Histoire naturelle, C, 21: 43-377. CHARLES R.P. 1952 - La faune Pleistocene de la grotte du Vallon des C~dres, Massif de la Sainte-Baume (Var). Bulletin de la Socidtg prghistorique fran~aise, 7: 294295. CLOT A. 1980 - La grotte de la Carri~re (Gerde, H a u t e s P y r e n e e s ) . S t r a t i g r a p h i e et pal~ontologie des Carnivores, 239 p., Th~se 3 '~° cycle, Toulouse (in~dit). CRI~Gu%BoNNOURE E. 1989 - Un n o u v e a u Caprinae, Hemitragus cedrensis nov. sp. ( M a m m a l i a , Bovidae) des n i v e a u x pl~istoc~nes moyen de la grotte des C~dres (Le P l a n d'Aups, Var). Int~r~t biog~ographique. Geobios, 22, 5: 653-663. DEFLEUR A., CRI~GUT-BONNOURE E., RADULESCU C. & V~ETTE Ph. 1988 - La grotte des C~dres, commune du P l a n d'Aups (Var). Bulletin de la Socidtd prdhistorique franfaise, 87, 9: 270-274. DEFLEUR A. & CRI~GUT-BONNOUREE. 1995 - Le g i s e m e n t Pal~olithique moyen de la grotte des C~dres (Var). Documents d'Archdologie fran~aise, 49: 1-182. DEHM R. 1943. Ein besonderes kleiner B a r (Ursus schertzi n. sp.) aus d e m 15ss von A c h e n h e i m bei S t r a s s b u r g in E l s a s s . Neuen Jahrbuch fi~r Mineralogie, Geologie und Pali~ontologie, B: 137-153. DEL CAMPANAD. 1 9 1 0 - M a m m i f e r i della g r o t t a di Reale presso Porto Longone (Isola d'Elba). Mondo sotteraneo, VI, 1-2, 23 p. ELHAI H. 1966 -Deux g i s e m e n t s du Q u a t e r n a i r e moyen: Bruges (S.-W. de la France), Bafiolas (Espagne).
Bulletin de l'Association fran~aise de l'Etude du Quaternaire, 6, 1: 69-78.
ERDBRINK D.P. 1953 - A review of fossil a n d r e c e n t b e a r s of the old world. 320 p., Deventer. FISTANI A. & CRI~GuT-BoNNOURE E. 1993 - D~couverte d'Ursus thibetanus ( M a m m a l i a , Carnivora, U r s i d a e ) dans le site Pleistocene moyen de G a j t a n (Shkoder, Albanie). Geobios, 26, 2: 241-263. FORSYTH MAJOR C.J. 1873 - R e m a r q u e s s u r quelques mammif'eres p o s t - t e r t i a i r e s de l'Italie, suivies de considerations g~n~rales s u r la f a u n e des m a m mif~res post-tertiaires. Atti de la Societa Italiana di Scienze Naturale, XV, 5, 27 p. GUI~RIN C. 1980 Les r h i n o c e r o s ( M a m m a l i a , Perissodactyla) du Miocene t e r m i n a l a u P15istoc~ne sup~rieur en E u r o p e Occidentale. Documents des Laboratoires de Ggologie de Lyon, 79, 3:785-1185. KURTEN B. & POULIANOSN. 1977 - New s t r a t i g r a p h i c a n d f a u n a l m a t e r i a l from P e t r a l o n a cave w i t h special reference to the Carnivora. Anthropos, 4, 1-2: 47-130. RUSTIONI M. • MAZZA P. 1993 - The T i b e t a n - l i k e b e a r from G r o t t a di Reale, Porto Azzuro (Isle of Elba, Italy). Il Quaternario, 6, 1: 35-38. THENIUS E. 1958 - ~ o e r eine Kleinb~iren aus dem Pleistoz~in von Slowenien. Razprave Dissertationes, IV: 333646. VIRET J. 1947 - S n r les U r s i d a e de Bruges (Gironde). Eclogae Geologicae Helvetiae, 40, 2: 356-360. VIRET J. & BALLAND R. 1948 - S u r quelques n o u v e a u x m a m m i f e r e s pl~istoc~nes des "Argiles de Bruges" (Gironde). Comptes-Rendus sommaires de la Socigtg ggologique de France, 12: 238-239. WERNERT P. 1957 - S t r a t i g r a p h i e pal~ontologique et pr6h i s t o r i q u e des s ~ d i m e n t s q u a t e r n a i r e s d'Alsace. Achenheim. Mdmoire du Service de la Carte ggologique d'Alsace Lorraine, 14, 262 p. E. C R I ~ G U T - B O N N O U R E
Museum Requien 67, rue Joseph-Vernet F-84000 Avignon
THE SAALIAN URSUS THIBETANUS F R O M F R A N C E A N D ITAIN
EVELYNECRI~GUT-BONNOURE CRt~GUT-BONNOURE E. 1997. The Saalian Ursus thibetanus from France and Italy. [UUrsus thibetanus saalien de France et d'Italie]. GEOBIOS, 30, 2 : 285-294. Villeurbanne, le 30.04.1997. Manuscrit d6pos6 le 01.08.1994; accept6 d6finitivement le 21.12.1995. ABSTRACT - Fragmentary small ursid remains were found in the Saalian site of the C6dres Cave (Le Plan d'Aups, Var). Their comparison with Middle Pleistocene Ursidae from Italy (Reale Cave), assigned to Ursus rnediterraneus F. MAJOR, and from France (Bruges), related to U stehlini (KRETZOi),shows that all these bears should be referred to U thibetanus CUVIER.The differences in the morphometry support the hypothesis of the existence of three subspecies at the end of the Middle Pleistocene in Southern Europe: Ursus thibetanus kurteni nov. subsp, from the C6dres Cave, Ursus thibetanus mediterraneus (F. MAJOR) from Reale Cave and Ursus thibetanus vireti nov. subsp, from Bruges loam pit. Such differentiation is assumed to indicate a geographic isolation of the populations. KEYWORDS: URSIDAE, TAXINOMY, MIDDLE PLEISTOCENE, FRANCE, ITALY, GEOGRAPHIC ISOLATION. RI~SUM]~ - Des restes fragmentaires d'un petit Ursidae ont 6t6 d6couverts dans le site saalien de la Grotte des C6dres (Le Plan d'Aups, Var). Leur comparaison avec Ursus mediterraneus F. MAJOR d'Italie (Grotte de Reale) et Ursus stehlini (KRETZODde France (Argiles de Bruges) montre que ces Ursidae correspondent fi Ursus thibetanus CUVIER. Les diff6rences morphom6triques relev6es sont en faveur de l'hypoth6se de l'existence de trois sous-esp6ces d'ours du Tibet fi la fin du P16istoc6ne moyen dans le Sud de l'Europe: Ursus thibetanus kurteni nov. subsp, de la grotte des C6dres, Ursus thibetanus rnediterraneus (F. MAJOR) de la grotte de Reale et Ursus thibetanus vireti nov. subsp, des argiles de Bruges. Ces faits sugg6rent l'isolement g6ographique de ces populations. MOTS-CL]~S: URSIDAE, TAXINOMIE, PLI~ISTOC]~NE MOYEN, FRANCE, ITALIE, ISOLAT Gt~OGRAPHIQUE.
INTRODUCTION Recently, t h e C6dres Cave, in t h e S a i n t e - B a u m e M o u n t a i n r a n g e (Le P l a n d'Aups, Var), h a s yielded a M i d d l e P l e i s t o c e n e f a u n a in association with a Mousterian industry. Our preliminary s t u d y allows us to i n d i c a t e t h e p r e s e n c e of b r o w n bear, U r s u s arctos LINNE, a n d cave bear, U. spel a e u s ROSENM~LLER • HEINROTH ( D e f l e u r et al. 1988). S o m e u r s i d pieces c o r r e s p o n d to a n o t h e r U r s i d a e . T h e m o r p h o l o g i c a l f e a t u r e s allowed us to i n c l u d e t h e C 6 d r e s b e a r in t h e U. t h i b e t a n u s species ( C r 6 g u t - B o n n o u r e in D e f l e u r & Cr6gutB o n n o u r e 1995). A c o m p a r i s o n w i t h t h e R e c e n t Ursus t h i b e t a n u s a n d w i t h t h e little b e a r s assig n e d to U. m e d i t e r r a n e u s F. MAJOR a n d U. stehlini (KRETZOI)allows a n e w i n t e r p r e t a t i o n of t h e s e little bears.
THE BEAR FROM THE CEDRES CAVE (FRANCE) T h e m a t e r i a l belongs to t h r e e individuals. It is composed by eleven pieces ( u p p e r t e e t h : left I3 (C6 SII), left P4 (C6 SI), r i g h t P4 (CIII 239), i n c o m p l e t e m a x i l l a r y w i t h P4, M1 a n d M2 (right, C6), r i g h t M2 (C6 SII), distal p a r t of a left M2 (C6 SII) - lower teeth: left P4 (C3-4-5 SI), left M3 (C56 SI) - post-cranial: I ° left m e t a t a r s a l , II ° & I I I ° left m e t a t a r s a l (CII), r i g h t V ° m e t a t a r s a l (C3-4
sI). LOWER TEETH T h e m o r p h o l o g y is t h e s a m e as in t h e R e c e n t U. thibetanus. T h e protoconid of P4 is s i t u a t e d anteriorly. T h e m e s i a l r i m is concave a n d t h e distal one p r e s e n t s a little slot. T h e lingual face of t h e
286
1 cnl
la
3a
1 cm
2a
3c
3b
1 cm
1 cm
FIGURE 1-3 - 1. Ursus thibetanus kurteni nov. subsp., C~dres Cave. 1. Left lower M3, C5-6 SI, occlusal view. Scale in cm. 2. Ursus thibetanus kurteni nov. subsp., C~dres Cave. Holotype, " upper right row, C6. a: vestibular view, b: occlusal view. Scale in cm. 3. Ursus thibetanus kurteni nov. subsp., C~dres Cave. a: upper right P4, C6 SI, lingual view, b: upper left P4 CIII 239, lingual view, e: upper right M2, C6 SII, occlusal view. Scale in cm. Photographies A. Guerrand (Calvet Museum). 1. Grotte des 1 cm
2b
t h i r d m o l a r , o v a l in o u t l i n e , p r e s e n t s l i t t l e w r i n k l e s , well d e v e l o p e d on t h e a n t e r o - v e s t i b u l a r face (Fig. 1). A cuspid, p r o b a b l y t h e protoconid, is p r e s e n t . A d i a g o n a l r i d g e r u n s f r o m it to t h e m i d d le of t h e crown. T h e v e s t i b u l a r conids set internally: consequently, t h e v e s t i b u l a r face is oblique. (Tab71) UPPER TEETH AND MAXILLARY T h e I3 is s l e n d e r a n d h i g h e r t h a n in U. a r c t o s . T h e p a l a t a l face is convex w i t h a ridge r u n n i n g off. An i m p o r t a n t c i n g u l u m exists a l o n g t h e l i n g u a l base. On t h e m a x i l l a r y , t h e i n f r a - o r b i t a l f o r a m e n is l o c a l i z a t e d quite p e r p e n d i c u l a r l y to t h e m i d d l e of
C~dres. M3 inf~rieure droite, C5-6 SI, rue occlusale. 2. Grotte des C~dres. Holotype, rangge dentaire supgrieure droite, C6. a: vue vestibulaire, b: vue occlusaIe. 3. Grotte des C~dres. a: P4 supgrieure droite, C6 SI, rue linguale, b: P4 sup~rieure gauche, CIII 239, vue linguale, e" M2 supgrieure gauche, C6 SII, rue
t h e M1 p a r a c o n e as in t h e R e c e n t b l a c k b e a r w h o s e m u z z l e is s h o r t (Fig. 2a). T h i s f o r a m e n in U. a r c t o s is s i t u a t e d p e r p e n d i c u l a r l y to t h e m i d d le of t h e M1 m e t a c o n e or to t h e b o u n d a r y of t h e two molars. T h e M2 v e s t i b u l a r cones a r e l o w e r t h a n t h o s e of t h e first molar. T h i s f e a t u r e is char a c t e r i s t i c of U. t h i b e t a n u s ( F i s t a n i & Cr~gutB o n n o u r e 1993). T h e P4 consists of t h r e e cusps, g e n t l y d i l a t e d a n d w i t h o u t a n y a c c e s s o r y denticles. T h e c o n e - s h a p e d d e u t e r o c o n e is p l a c e d f o r w a r d t h e n o t c h s e p a r a t i n g p a r a - m e t a c o n e (Fig. 2b,3a,b). C o n s e q u e n t l y t h e outline of t h e c r o w n is m o r e or less t r i a n g u l a r . T h e p a r a c o n e , slightly h i g h e r t h a n t h e m e t a c o n e ,
287 ! ,Lower teeth L IP4 ,Les C ~ d r e s !C3-4-5 SI Recent. n Variation Mean
10,61 9 8,88-10,96 9,82
5,91 9 5,24-6,16 5,85
6,2 9 5,22-6,54 5,78
7 8 5,12-9,43 7,16
iM2 Reale IGF 4808V IGF4BO7V Recent - n Variatien [Mean
20,62 21,45 9 17,38-21,37 18,7
14 14,61 7 11L14,12 12
7,29 7,41 8 5,49-7,11 6,34
7,33 7,3 8 4,33-6,99 i 5,61
17,4 9 12,71-17,06 14,52
12,8 9 9,96-12,19 11,11
'LMe3sC~dres CS-6 SI Recent - n Variation Mean I
1
2
3
4
5
6
7
6,64 5,81 9 4,28-5,62 5,11
6,85 6,9 9 5,1-6,57 5,94
8
12,96 12,28 9 10,26-11,91 10,92
9
12,24 11,81 9 10,11-11,66 10,93
7,32 11,62 8 9,89411,43 10,75
TABLE 1 - Dimensions of the lower teeth in Ursus thibetanus kurteni nov. subsp, from C~dres Cave, Ursus thibetanus mediterraneus from Reale Cave and Recent asiatic black bear (in ram). For P4 - total length (1), total width (2), protoconid height (3), distance from the protoconid cusp to the distal point (4). For M2 - Length: total (1), trigonid (2); height: protoconid (3), hypoconid (4), metaeonid (5),entoconid (6); width: trigonid (7), talonid (8), minimal (9). For M3 - total length (1), total width (2). Dimensions des dents jugales infdrieures d ~ r s u s t h i b e t a n u s kurteni nov. subsp, de la grotte des C~dres (c), d'Ursus t h i b e t a n u s m e d i t e r r a n e u s de la grotte de Reale et de Fours du Tibet actuel (en mm). Rgfdrence des mesures: P4 - Longueur totale (1), largeur totale (2), hauteur du protoconide (3), distance du sommet du protoconide au bord distal (4). M2 - Longueur: totale (1), trigonide (2); hauteur: protoconide (3), hypoconide (4), m4taconide (5), entoconide (6); largeur: trigonide (7), talonide (8), minimale (9). M3 - Longueur totaIe(1), largeur totale (2).
is characterized by a concave mesial rim. The edge r u n n i n g from the top to the base is curved outward (Fig. 2b). The metacone and the deuterocone are higher t h a n in the Recent species and t h a n in the two other Middle Pleistocene little bears examined (Fig. 4, parameters 8,9, Tab. 2) and the paracone is comparatively lower (parameter 7). A basal cingulum is present on each side of the deuterocone. It runs along the vestibular face.
I
0,1
/(
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.."
The cusps of the molars set internally relative to the base of the crown and are weakly dilated:
< t, ' \
FIGURE 4 - Ratio diagram of the proportions of the upper P4 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c), Ursus thibetanus mediterraneus from Reale Cave (m) and Ursus. thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in Table 2. Length: vestibular (1), paracone (2), metacone (3), deuterocone (4); width: paracone (5), in deuterocone axis (6); height: paracone (7), metacone (8), deuterocone (9); distance from the deuterocone cusp to the mesial point (10). Diagramme des rapports des proportions de la P4 supdrieure d ~ r s u s thibetanus kurteni nov. subsp, de la grotte des C~dres (c), d~Ursus thibetanus mediterraneus de la grotte de Reale (m) et d ~ r s u s thibetanus vireti nov. subsp, des argiles de Bruges (b). Les diffgrences sont exprimdes en logarithmes ddcimaux. Base de rgf4rence O: Ursus thibetanus actuel. Dimensions dans le tableau 2. Longueur: externe (1), parac6ne (2), mdtac6ne (3), deutdrocSne (4); largeur: paracSne (5), au niveau du deutdroc6ne (6); hauteur: parac6ne (7), mgtacSne (8), deutdrocOne (9); distance milieu deut4roc6ne-bord mgsial (10).
/
0,0
]0
288 Upper teeth
66•
1
P4 Les C6dres CII1239 14,37 9,65 5,7 10,11 S1 C6 15,4 C6 14,11 9,29 ' 5,66 Reale 9,45 6,8 5,94 IGF-4806V 15,43 Bruges 9,16 5,8 6,05 JRight 14,38 8,9 5,67 6,17 Left 14,76 I0 I0 10 Recent - n 10 Variation 111,41-13,25 6,71-8,47 4,08-5,65 4,48-7,32 Mean 12,21 5,84
/
L
M1 ,Les C~dres C6ruges Right Recent ,n /Variation .Mean
iLes C~dres C~SII
6
10
8,41 7,92 7,93
10,97 10,47 10,21
8,17 8,56 8
7,04 7,2 7,28
5,9 5,99 5,78
10,27 10,17
7,14
11,25
9,12
7,82
6,49
10,82
8,11 7,46 10 6,58-7,43 7,03
9,82 10,04 10 8,21-9,17 8,51
8,9 9,1 10 6,61-8,09 7,26
6,93 7,179
5,58 5~74
11,64 I 10,48 I
5,03-6,21 5,59
] 3,36
19,49 18.75 10 16,67-19,07 17,99
6,375-79 5,%1y2
28,51 28,37
2,4 I0 2,12-2.52 2,30
14,14
15,52
14,32
8.9
14.3 14.79 14,27 9 10 10 9 10 11,61-13,41 12,13-14,33 11,99-13,22 7,54-9,01 12,58 13,39 12,69 8,28
9,5
3,87-5,33 7,6910,64 4,49 8,46 I i
8.75
8,26
8,46
8;42
8~15
7,516
7,54-8,69 8,09
6,571,73 %7,;7?2
4,95 3,71
16,99 16,59 16,38
10,32 11,15 10,91
16,36 15,97
15,66 15,18 15,48
11,84 11,71 11,46
7,65 7,48
7,56 7.46 ' 7,56
3,75
14,87
10,14
15,42
15,68
10,55
7,34
7,66
17,68 19~18
3,4 14,699
15,29 15~33
7,05 7~2
8
C6 SI1 IReale IGF 4806V . 26,02 7,05 7,44 Bruges Right 28,62 8,24 10,21 7~9 Left 29,29 9,03 Recent - n 10 9 Variation 23,09-29,04 7,72-11,05 5,61-9,69 !Mean 25,61 i 9,26 7,4
11
2,85 2,85
16,01 16,14 9 13,07-17,84 9,93-13,27 10,19-14,8 15,24 11,18 13,63
12,9 6,97 13,98 9 12,93-14,52 9,06-13,4 13,78 10,91
! 7£
5,73-6,8] 5,68-6,84 6,23 ] 6,13
TABLE 2 - Dimensions of the upper teeth in Ursus thibetanus kurteni nov. subsp, from C6dres Cave, Ursus thibetanus mediterraneus from Reale Cave, Ursus thibetanus vireti nov. subsp, from Bruges and Recent asiatic black bear (in mm). For P4 - Length: yestibular (1), paracone (2), metacone (3), deuterocone (4); width: paracone (5), in deuterocone axis (6); height: paracone (7), metacone (8), deuterocone (9); distance from the deuterocone cusp to the mesial point (10) For M1 - Length: vestibular (1), paracone (2), metacone (3), parastyle (4); width: first lobe (5), second lobe (6), minimal (7); height: paracone (8), metacone (9), protoeone (10), hypocone (11).- For M2 - Length: vestibular (1), paracone (2), metacone (3), parastyle (4), from the metacone top to the distal point (5), from the metacone distal base to the distal point (6); width: first lobe (7), second lobe (8), talon (9); height: paraeone (10), metacone (11). D i m e n s i o n s des dents j u g a l e s supgrieures d'Ursus thibetanus kurteni nov. subsp, de la grotte des C~dres (c), d~Ursus thibetanus mediterraneus de la grotte de Reale, d'Ursus thibetanus vireti nov. subsp, de Bruges et de Fours d u Tibet actuel (en ram). Rdf~rence des mesures: P 4 - longueur externe (1), du parac6ne (2), du mgtacdne (3), d u deutdrocSne (4), largeur d u parac(me (5), au niveau d u deutdrocSne (6), h a u t e u r du parac6ne (7), d u m~tacSne (8), d u deutdroc6ne (9), distance milieu deutdrocSne-bord mdsial (10) - M 1 - longueur externe (1), d u parac6ne (2), du mgtac6ne (3), du parastyle (4), largeur d u p r e m i e r lobe (5), d u deuxi~me lobe (6), m i n i m a l e (7% h a u t e u r d u parac6ne (8), du m~tacSne (9), du protoc6ne (10), de l'hypocSne (11) - M 2 - longueur: externe (1), paracSne (2), m~tacSne (3), parastyle (4); distance: de la pointe d u mdtac~ne au bord distal de la couronne (5), de la base distale d u mgtacdne au bord distal de la couronne (6); largeur: p r e m i e r lobe (7), second lobe (8), talon (9); h a u t e u r parac~ne (10), mgtac6ne (11).
consequently, the medial channel is larger t h a n in U. arctos (Fig. 2b). The lingual cones constitute a thin and sharp edge. A well developed parastyle is present and the vestibular cingulum is weak. The paracone and metacone of the upper M1 are equal in size. A little slot exists on the distal rim of the metacone suggesting a differenciation of the metastyle. Two accessory cusps exist between the protocone and hypocone. The second lobe is broader t h a n in the Recent black bear (Fig. 5, parameter 6, Tab. 2); the parastyle is stronger (parameter 4) and in consequence the paracone is shorter (parameter 2). Finally all the cones are higher,
!
especially the hypocone (parameter l l ) . T h e length of the M2 is shorter t h a n the combined length of M1 and P4. A transversal ridge runs from the paracone to the hypocone (Fig. 2b, Fig. 3c). It corresponds to the mesial limit of the lower M3 which is placed under the talon of the second upper molar. This ridge is not very developed or missing in the Recent species. On the right isolated M2 (Fig. 3c), an edge exists on the internal side of the paracone. The parastyle is powerful: it is absent on nine of the ten recent specimens taken in comparison. Erdbrink indicates t h a t it is "very often present" but without a n y precision on
289 The length of the dental row is stronger than in the Recent U. thibetanus. Effectively, the P4 and the M2 are well developed. On the other hand, the first molar is shorter. This is a characteristic of all the Pleistocene bears belonging to the Tibetan group (Fistani & Cr6gut-Bonnoure 1993).
0,1~ f
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0,1£
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~
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POST-CRANIAL
t
1 t
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I
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The post cranial material is scarce. The diaphysis of the metapodes is slight and short. The articular facets of the upper articulation are small and the distal condyle is globular. These features are present on the recent specimens seen. The third metatarsal shows an oblique proximal articular side: the medial side is inner than the lateral one. A same morphology is to be observed on the Gajtan specimens (Fistani & Cr6gut-Bonnoure 1993), but the metapodes of this site are stronger. CONCLUSION
I 0
11
The dental features indicate that the C~dres bear clusters with U. thibetanus b u t differences occur between the Pleistocene and the Recent forms. I propose for this bear the following diagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAEViret, 1955 Genus Ursus LINNE, 1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus kurteni nov. subsp.
FIGURE 5 - Ratio diagram of the proportions of the upper M1 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c) and Ursus thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in table 2. Length: vestibular (1), paracone (2), metacone (3), parastyle (4); width: first lobe (5), second lobe (6), minimal (7); height: paracone (8), metacone (9), protocone (10), hypocone (11). Diagramme des rapports des proportions de la M1 supdrieure d~Ursus thibetanus kurteni nov. subsp, de la grotte des C~dres (c) d~Ursus thibetanus vireti nov. subsp, des argiles de Bruges (b). Les diffd-
rences sont exprimdes en logarithmes dgcimaux. Base de r~fdrence O: Ursus thibetanus actuel. Dimensions dans le tableau 2. Longueur: externe (1), du paracSne (2), du mdtacSne (3), du parastyle (4); largeur: premier lobe (5), deuxi~me lobe (6), minimale (7); hauteur: paracSne (8), mdtacSne (9), protocSne (10), hypoe6ne (11).
this frequency in the sub-species (1953: 150). An important cingulum is present at the lingual base of the crown.The distal rim of the metacone is longer t h a n the mesial compared to the case in the Recent black bear. Consequently the length from the top of the cusp to the distal side (Fig. 6, parameter 5, Tabl. 2) is stronger while the length from the base of the cusp to the distal side (parameter 6), which corresponds to the talon length, is shorter. The first lobe is broader than the second which is broader than the talon: consequently the width decreases from the first to the third lobe (parameters 7,8,9).
D e r i v a t i o n o m i n i s - In respect to late Professor BjSrn Kurt6n. L o c u s t y p i c u s - C~dres Cave (=Baume du Couloir Ovale), situated in the Mesozoic Sainte Baume Mountain, Provence, Le Plan d'Aups, Var, France. S t r a t u m t y p i c u m - Archeological levels II and III of Alban Defleur excavations, and archeological levels 5 and 6 of Max Escalon de Fonton excavations, sequence of the end of the Saale. H o l o t y p e - Upper right tooth row with the carnassial and the
two molars, level 6. Conserved in the Museum Requien (M.R.) Avignon, France. Fig. 2 a & b and fig. 52 in Defleur & Cr6gutBonnoure 1995. D i a g n o s i s - Ursus thibetanus with strong dentition compared with the Recent black bear. Denticles of the upper teeth with moderate dilatation. Metacone of the upper P4 high, and paracone relatively low. Paracone characterized by a concave mesial rim and an edge running from the top to the base curved outwards. Cone-shaped deuterocone high and strait, situated anteriorly to the vestibular notch. Metacone and deuterocone higher than in the Recent species and deuterocone lower. Upper M1 with powerful parastyle. Paracone shorter and hypocone higher than in the Recent black bear. Second lobe thicker than the first and thicker than in the Recent species. Upper M2 with a powerful parastyle and a transversal ridge running from the
290 paracone to the hypocone. Distal rim of the metacone longer than the mesial one, compared to Recent U thibetanus and consequently a short talon. Decreasing anteroposterior width of the lobes. Upper I3, lower P4 and metapodes identical as in the Recent species. +0,20
Synonymy 1954 Ursus arctos, Charles, p. 294 1971 Ursus arctos L. fossilis GOLDFUSS, Bonifay, p. 243 1995 Ursus thibetanus ssp., Cr~gut-Bonnoure in Defleur & Cr~gut-Bonnoure, p. 58-60
+0,15
THE BEAR FROM REALE CAVE (ITALY)
J 2
/
+0,10
2~ .:11 ...., +0,05
if •I
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LOWER TEETH
',,,~
Two M2, belonging to two different individuals, and one M3 were studied.
,
-0,05
-0,10 1'
i! -0,15 1
2
3
5
6
Some elements belonging to a little ursid were discovered in the Reale Cave (Elbe Island, Porto Longone or Porto Azzuro) and attribuated by Forsyth Major (1873) to a new species: U. mediterraneus. This material, conserved in the Geological and Paleontological M u s e u m of Firenze, was recently revised by Rustioni & Mazza (1993) and referred to Ursus cf. thibetanus.
7
8
9
10
11
4
Fmu~E 6 - Ratio diagram of the proportions of the upper M2 in Ursus thibetanus kurteni nov. subsp, from C~dres Cave (c), Ursus thibetanus mediterraneus from Reale Cave (m) and Ursus thibetanus vireti nov. subsp, from Bruges loam pit (b). Differences in decimal logarithm with the mean values of Recent Ursus thibetanus. Based on data in Table 2. Length: vestibular (1), paracone (2), metacone (3), parastyle (4), from the metacone top to the distal point (5), from the metacone distal base to the distal point (6); width: first lobe (7), second lobe (8), talon (9); height: paracone (10), metacone (11).
The talonid of the left M2 is partially broken. All the conids set internally and are relatively high. Their lateral longitudinal profiles constitute a broken line. The protoconid is connected to the metaconid by a rugged transverse '~" shaped ridge outline.The mesial '~g" top is localized on the mesio-distal axis of the crown. An other ridge runs from this point forward and ends with a little accessory cusp. In mesial view, the sides of the transverse ridge are oblique from the external cusps toward the center, forming a large valley. The anterior part of the trigonid is conform with U thibetanus because "In front of the protoconid and the metaconid a semicircular ridge along the edge of the molar encloses a hollowed part of the mastication surface" (Erdbrink 1953: 153). An oblique ridge runs from the base of the protoconid distal rim to the middle of the crown, at the level of the lateral notches corresponding to the trigonid-talonid separation.The hypoconid is well developed and a ridge runs from it to the preDiagramme des rapports des proportions de la M 2 supgrieure d'Ursus t h i b e t a n u s kurteni nov. subsp, de la grotte des C~dres (c), d~tJrsus t h i b e t a n u s m e d i t e r r a n e u s de la grotte de Reale (m) et d'Ursus t h i b e t a n u s vireti nov. subsp, des argiles de Bruges (b). Les diffdrences sont exprimges en logarithmes ddcimaux. Base de rgfdrence O: Ursus t h i b e t a n u s actuel. Dimensions dans le tableau 2. Longueur: externe (1), paracSne (2), mdtacdne (3), parastyle (4); distance: de la pointe du mdtac6ne au bord distal de la couronne (5), de la base distale du mdtacSne au bord distal de la couronne (6); largeur: premier lobe (8), second lobe (8), talon (9); hauteur: paracSne (10), mgtacOne (11).
291 cedent crest. Two accessory cusps exist in front of the metaconid, which is the highest denticle of the tooth, and two other at the base of the distal rim.The endoconid, higher than the hypoconid, is enclosed by two little cusps. The morphology of the right M2 is the same (Tab. 2). The left M3 described originally as U mediterraneus (Del Campana 1910:18 and pl. III, fig. 6) belongs in fact to U spelaeus. Bonifay (1971: 218) pointed its great dimensions. The crown is bilobated and the occlusa] surface presents numerous denticles as in the cave bear (Clot 1980; Ballesio 1983). UPPER TEETH Forsyth Major pointed out a left M2. It is necessary to add to the material a left P4. They belong to a same individual which is young and the same that the lower left M2. The P4 was assigned to a cave bear milk tooth Cdi latte" on the label). According to the morphometry, this specimen belongs to U. thibetanus. This premolar, constituted by three cusps weakly dilated, has a triangular shape. The paracone has a straight mesial rim which shows a thin edge. At the base a slight ridge, coming from the top of the denticle, runs along the middle of the lingual face. The paracone is higher than the metacone. These two cusps are respectively lower and higher than in the Recent black bear (Fig. 4, parameters 7,8, Tab. 2) and the deuterocone is higher (parameter 9). Compared to the U. thibetanus from the C~dres Cave, the proportions are the same but the paracone is shorter (parameter 2) and the metacone is longer (parameter 3). A basal cingulum is present on each side of the deuterocone and runs along the vestibular base. A small accessory cusp exists at the mesial base of this cusp which is quite in the axis of the vestibular notch. The cones of the M2 is more internally offset relative to the base of the crown than in the Recent U thibetanus and in the C~dres bear, whose vestibular face is less oblique. The parastyle (Fig. 6, parameter 4, Tab. 2) is smaller and lower than in U thibetanus kurteni and consequently the paracone is longer (parameter 2). The paracone and the metacone are weakly dilated. A transversal ridge runs from the top of each cone to the lingual base, reaching about the middle of the crown. An accessory cusp is present at the base of the distal rim of the paracone. The lingual cusps, lower than the vestibular one, constitute a thin sinuous ridge fringed by an important cingulum. The protocone is the lowest cone. Two accessory cusps exist between it and the hypocone. The mesial rim of this last one is short (1,97 mm) compared with its distal counterpart (6,20 ram). Little wringles occupies all the occlusal face. The second lobe is slightly broader than the first
one (Fig. 6, parameters 7,8). The talon is very strait as in U. thibetanus kurteni (parameter 9) but shorter (parameter 5). CONCLUSION The Reale bear can be assigned to U. thibetanus. The material shows analogies with the C~dres ursid but morphological and metrical differences exist between them. In consequence, I propose the following neodiagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAEViret, 1955 Genus Ursus LINN]~,1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus mediterraneus (FoRSYTH Derivatio nominis
MAJOR,
1873).
- Ursid from Mediterranean
zone.
Locus typicus - From the Reale Cave situated in Porto Longone (=Porto Azzuro), Elb Island (Italy). Stratum typicum - End of the Saalian, or early beginning of the Upper Pleistocene. Holotype - Left upper P4 and M2 (IGF 4806 V), and left lower M2 (IGF 46), Museum of Geology and Paleontology, University of Firenze, Italy.
Neodiagnosis U thibetanus with strong dentition compared to the Recent black bear. Denticles of the lower and the upper teeth few dilated. Paracone of the upper P4 higher than the metacone and with straight mesial rim. Deuterocone localized in the axis of the vestibular notch. Paracone and metacone respectively lower and higher and deuterocone higher than in the Recent black bear. Paracone shorter than in U thibetanus kurteni and metacone longer. Upper M2 with a moderate parastyle and a short distal metacone rim; paracone longer than in U. thibetanus kurteni; talon short and strait. Lower M2 with an angulous transversal mesial ridge extended forward by an antero-posterior edge. Simplified synonymy 1873 Ursus mediterraneus, Forsyth Major, p. 8. 1910 Ursus sp., Del Campana, p. 17-19, pl. III, fig. 1-7. 1958 Ursus (Euarctos) mediterraneus, Thenius, p. 638. 1971 Ursus mediterraneus, Bonifay, p. 218-219. 1977 Ursus thibetanus mediterraneus, Kurten, p. 64-65. 1993 Ursus cf. thibetanus, Rustioni & Mazza, p. 35-38, fig. 1.
THE BEAR FROM THE BRUGES LOAM PIT (FRANCE) Some teeth discovered in the loam pit at Bruges (Gironde, France) were a t t r i b u a t e d by Viret (1947) to U. stehlini (KRETZOD. The material is
292 composed by two upper canines (left and right), a right maxillary with P4, M1 and M2, two isolated upper left teeth (one P4 and one M2) corresponding to a same animal, two lower canines (left and right) belonging to an other individual. The isolated upper M1 described by Viret is apparently lost. The two lower and the right upper canines were not described by this author. LOWER TEETH The two canines, laterally compressed, are more curved t h a n the upper ones. A thin sharp lingual ridge runs from the top to the base of the crown as in the Recent black bear.
UPPER
TEETH
AND
MAXILLARY
The two upper canines show a sharp ridge on their lingual face like in the Recent U. thibetanus. According to Viret, the paracone of the P4, sharppointed, is higher t h a n the metacone. But, compared with the C~dres and Reale bears, the proportions of these cusps and of the deuterocone are quite the same as in the Recent black bear (Fig. 4, parameter 7,8,9, Tab. 2). A transversal ridge is present on the lingual face of the paracone whose mesia] rim is rectilinear. The deuterocone, situated in the axis of the vestibular notch, is slightly dilated and elongated. On the maxillary, it is prolongated forward by a thin edge. On the isolated P4 an accessory cusp is present. A cingnlum runs on both sides of the lingual cone and exists on the vestibular base. The morphology of the M1 is the same as in the Recent U. thibetanus and in the C~dres bear. In comparison with the last one, the parastyle is smaller (Fig. 5, p a r a m e t e r 4, Tab. 2), the second lobe straiter (parameter 6), the paracone higher and the metacone lower (parameters 8,9). The protocone and the hypocone are worned out. An important cingulum occupies the lingual base of the crown. The M2 is without parastyle (Fig. 6, parameter 4, Tab. 2). However the mesial rim of the paracone is slighty in zigzag in occlusal view. A thin edge occupies the middle of the paracone lingual face. It is absent on the metacone. The paracone is longer t h a n in the C~dres and Reale bears but stays shorter t h a n in the Recent U thibetanus (parameter 2). The hypocone is well developed. The second lobe is broader t h a n the first (parameters 7,8). The talon, which is long (parameters 5,6), is broader than in the Recent U. thibetanus (parameter 9). CONCLUSION Viret thinks t h a t 'Tours de Bruges montre une similitude non seulement de taille mais encore de
structure" with U. thibetanus (Viret 1947). The morphology is effectively the same as in the Recent species. Erdbrink has the same opinion (1953). The differences pointed out by Viret (great development of the upper P4, presence of an important cingulum on the upper M1 and M2) are not specific but concern individual variation. The last author referred the Bruges material to U. stehlini (KRETZOD,discovered in the Lower Middle Pleistocene sites of M a u e r and Mosbach in Germany. The general features of this ursid, never revised since the yon Reichenau description, seem to be conform with U. thibetanus (Kurt~n 1957; Erdbrink 1953; Ficarrelli 1979). In m y opinion, it is not possible to identify the Bruges bear with the Mauer and Mosbach species: this last ursid is older, the material is composed by lower jaws and the morphologic variation existing in the upper teeth of the C~dres, Reale and Bruges ursids suggests an important plasticity of the Pleistocene black bear. During the Saalian period, U. thibetanus is present in France in Achenheim (Wernert 1957) where it is assigned to U. schertzi by Dehm (1943). Unfortunately, the material, a lower jaw, is lost: it is not possible to correlate the Bruges upper teeth with the Achenheim lower ones. Nevertheless, in U. schertzi the carnassial is very short (Fistani & Cr6gnt-Bonnoure 1993) which m u s t have a repercussion on the upper row proportions. Now the dental proportions of the upper tooth row of the Bruges bear are the same as in the other Pleistocene small bears examined: consequently it is not possible to refer the materials of these two sites to a same form. If we take into account the particularities of the Bruges bear and its differences with the C~dres and Reale U thibetanus, it is possible to propose the following diagnosis. Family URSIDAE Gray, 1825 Sub-family URSINAE Viret, 1955 Genus Ursus LINNE, 1758 Species Ursus thibetanus CUVIER, 1823 Sub-species Ursus thibetanus vireti nov. subsp. Derivatio nominis
- In respect to late Professor Jean Viret.
- Loam pit at Bruges, Gironde,France. S t r a t u m t y p i c u m - Holsteinian or early beginning of the Saalian period. I-Iolotype - Upper right row with the carnassial and the two molars, National Museum of Natural History, Paris (France). Mold of this specimen in the Department of Earth Sciences, University Claude Bernard-LyonI, Villeurbanne, France. Locus typicus
U. thibetanus with strong dentition compared to the Recent black bear. Paracone of the upper P4 higher t h a n the metacone and with straight mesial rim. Deuterocone elongated and
Diagnosis
-
293 s i t uated in th e axis of the vestibular notch. Paracone, m e t a c o n e and de ut e r ocone hei ght s n e a r l y the same as in t he Recent U. thibetanus. U p per M1 with a m o d e r a t e parastyle and a weak metastyle. Second lobe broader t h a n the first one and s t r a i t e r t h a n in t he Recent black bear. Upper M2 with o u t parastyle. Talon elongated and broader compared to Recent U. thibetanus. Simplified s y n o n y m y
1947 Ursusstehlini, Viret, p.356-360. 1948 Ursusstehlini, Viret & Balland, p. 238-239. 1971 Plionarctosstehlini, Bonifay,p. 216-217.
GENERAL CONCLUSION T h r e e dental morphotypes exist in U. thibetanus l i n eag e d u r i n g t h e Middle Pl ei s t oc e ne in S o u t h e r n Europe. G reat analogies could be pointed out between the C~dres and Reale bears. However, the morphometrical characters are sufficiently different to refute the hypothesis of a continuous variation in a same population, such differences not existing in t h e r e c e n t m a t e r i a l exam i ne d. C o n s e q u e n t l y these two ursids can be r e f er r ed to different subspecies: U thibetanus kurteni nov. subsp, from the C~dres Cave and U thibetanus mediterraneus (F. MAJOR) from th e Reale Cave. According to the morphometry, the Bruges bear is clearly different. Therefore, it is possible to assign it to another sub-species: U. thibetanus vireti nov. subsp. The t h r e e sites are considered Middle Pleistocene in age, b u t exact chronological correlations are unknown. The C~dres Cave has yielded a well preserved fauna with Canis lupus, Vulpes vulpes, Ursus arctos, Ursus spelaeus, Lynx spelaea, Fells silvestris, Equus sp.,
Sus scrofa, Cervus elaphus, Capreolus capreolus, Rangifer tarandus, Bos primigenius, Bison sp., Hemitragus cedrensis, Rupicapra rupicapra, Oryctolagus cuniculus baumensis, Marmota marmota mesostyla, Apodemus sylvaticus, Pliomys gr. episcopalls, Terricola duodecimcostatus, Microtus agrestis cf. jansoni, Microtus brecciensis defleuri, Arvicola sapidus aupsensis, cf. Alectoris graeca, Columba cf. livia, cf. Turdus torquatus, Coccothraustes coccothraustes, Garrulus glandarius, Pyrrhocorax graculus cf. vetus. This assemblage date to the end of the
The Bruges loam pit has yielded Lynx sp., U arctos, Palaeoloxodon antiquus, D. hemitoechus, S. scrofa, C. eIaphus, C. capreolus, Castor fiber, Trogontherium sp. (Balland 1963; GuSrin 1980). According to the flora, its age is Holsteinian (Elhai 1966). A comparative study of the f a u n a is lacking and no more informations are available. A p p a r e n t l y the Bruges black bear is the oldest and t he Reale bear could be the younger. Hence if t hese t h r e e black bears are not exactly contamporaneous, th e differences between t h e m m a y correspond to a rapid evolution of the species and could have a chronological connotation. With regard to t he morphological analogies, it is clear t h a t the origin of t he C~dres and Reale bears is common. If t hey date to t he end of t he Saalian, the differences bet w een t h e m could reflect a geographical differenciation in consequence of th e Alpine Arch which can act as a geographical barrier. The origin of the Provence-Tuscany black bear group is unknown. T h e y could evolve from the Bruges group. May be the Rh6ne and th e Durance rivers, which during t he Pleistocene flowed separately into the M e d i t e r r a n e a n sea, could be at the origin of this chronological isolation. Such an isolation is suggested for t he t a h r from the C~dres Cave (Crggut-Bonneure 1989). In conclusion, two groups of black b e a r do exist d u r i n g t h e Middle P l e i s t o c e n e in S o u t h e r n Europe with at least t h r e e different subspecies clearly individualized. Thus it appears t h a t th e Pleistocene black bear shows various morphological and biometrical p a t t e r n s but more m a t e r i a l is required for a bet t er u n d e r s t a n d i n g of t he evolution of the species. Acknowledgments - I would like to thank Prof. A. Azzaroli, Prof. D. Torre and Dr. P. Mazza for the assistance and remarks they allow me during my study of the Italian collections. I am most indebted to Prof. B. Sala for his help in literature data, and to Prof. R. Ballesio and Dr. A. Argant for the critical reading of the manuscript. Also, I thank Prof. L. Ginsburg and Dr. A. Prieur who permit me to study the Bruges material. I express my gratitude to Dr. A. Defleur and Dr. M. Escalon de Fonton for having permit me to study the C~dres fauna which is preserved in the Museum Requien, thanks to Dir. A. Jacob from the D.R.A.C. My sincere gratitude to A. Guerrand for the photographies and to M. BuissenCatif for the illustration.
Saalian period (Defleur & Cr6gut-Bonnoure 1995). The Reale f a u n a is composed with U. spelaeus, Lynx sp., Panthera spelaea, S. scrofa, Hippopotamus sp., Dicerorhinus sp., C. elaphus, C. capreolus, Lepus sp. (Del C a m p a n a 1910) and could be of Saalian age, or could "come from two different stratigraphic levels" of the Upper Pleistocene (Rustioni & Mazza 1993: 37).
REFERENCES ARGANTA 1991 - Carnivores quaternaires de Bourgogne. Documents des Laboratoires de Ggologie de Lyon, 115: 1-301. BALLANDR. 1963 - Le gisement pleistocene des "Argiles de Bruges" (Gironde). Cahiers des Naturalistes, N.S., 19: 57-63.
294 BALLESIOR. 1983 - Le gisement pleistocene sup~rieur de la grotte de Jaurens ~ Nespouls, Corr~ze, France. Les Carnivores (Mammalia, Carnivora): Ursidae. Nouvelles Archives du Musge d'Histoire naturelle de Lyon, 21: 943. BONIFAY M.-F. 1971 - Carnivores q n a t e r n a i r e s du SudE s t de la France. Mdmoires du Musdum national d'Histoire naturelle, C, 21: 43-377. CHARLES R.P. 1952 - La faune Pleistocene de la grotte du Vallon des C~dres, Massif de la Sainte-Baume (Var). Bulletin de la Socidtg prghistorique fran~aise, 7: 294295. CLOT A. 1980 - La grotte de la Carri~re (Gerde, H a u t e s P y r e n e e s ) . S t r a t i g r a p h i e et pal~ontologie des Carnivores, 239 p., Th~se 3 '~° cycle, Toulouse (in~dit). CRI~Gu%BoNNOURE E. 1989 - Un n o u v e a u Caprinae, Hemitragus cedrensis nov. sp. ( M a m m a l i a , Bovidae) des n i v e a u x pl~istoc~nes moyen de la grotte des C~dres (Le P l a n d'Aups, Var). Int~r~t biog~ographique. Geobios, 22, 5: 653-663. DEFLEUR A., CRI~GUT-BONNOURE E., RADULESCU C. & V~ETTE Ph. 1988 - La grotte des C~dres, commune du P l a n d'Aups (Var). Bulletin de la Socidtd prdhistorique franfaise, 87, 9: 270-274. DEFLEUR A. & CRI~GUT-BONNOUREE. 1995 - Le g i s e m e n t Pal~olithique moyen de la grotte des C~dres (Var). Documents d'Archdologie fran~aise, 49: 1-182. DEHM R. 1943. Ein besonderes kleiner B a r (Ursus schertzi n. sp.) aus d e m 15ss von A c h e n h e i m bei S t r a s s b u r g in E l s a s s . Neuen Jahrbuch fi~r Mineralogie, Geologie und Pali~ontologie, B: 137-153. DEL CAMPANAD. 1 9 1 0 - M a m m i f e r i della g r o t t a di Reale presso Porto Longone (Isola d'Elba). Mondo sotteraneo, VI, 1-2, 23 p. ELHAI H. 1966 -Deux g i s e m e n t s du Q u a t e r n a i r e moyen: Bruges (S.-W. de la France), Bafiolas (Espagne).
Bulletin de l'Association fran~aise de l'Etude du Quaternaire, 6, 1: 69-78.
ERDBRINK D.P. 1953 - A review of fossil a n d r e c e n t b e a r s of the old world. 320 p., Deventer. FISTANI A. & CRI~GuT-BoNNOURE E. 1993 - D~couverte d'Ursus thibetanus ( M a m m a l i a , Carnivora, U r s i d a e ) dans le site Pleistocene moyen de G a j t a n (Shkoder, Albanie). Geobios, 26, 2: 241-263. FORSYTH MAJOR C.J. 1873 - R e m a r q u e s s u r quelques mammif'eres p o s t - t e r t i a i r e s de l'Italie, suivies de considerations g~n~rales s u r la f a u n e des m a m mif~res post-tertiaires. Atti de la Societa Italiana di Scienze Naturale, XV, 5, 27 p. GUI~RIN C. 1980 Les r h i n o c e r o s ( M a m m a l i a , Perissodactyla) du Miocene t e r m i n a l a u P15istoc~ne sup~rieur en E u r o p e Occidentale. Documents des Laboratoires de Ggologie de Lyon, 79, 3:785-1185. KURTEN B. & POULIANOSN. 1977 - New s t r a t i g r a p h i c a n d f a u n a l m a t e r i a l from P e t r a l o n a cave w i t h special reference to the Carnivora. Anthropos, 4, 1-2: 47-130. RUSTIONI M. • MAZZA P. 1993 - The T i b e t a n - l i k e b e a r from G r o t t a di Reale, Porto Azzuro (Isle of Elba, Italy). Il Quaternario, 6, 1: 35-38. THENIUS E. 1958 - ~ o e r eine Kleinb~iren aus dem Pleistoz~in von Slowenien. Razprave Dissertationes, IV: 333646. VIRET J. 1947 - S n r les U r s i d a e de Bruges (Gironde). Eclogae Geologicae Helvetiae, 40, 2: 356-360. VIRET J. & BALLAND R. 1948 - S u r quelques n o u v e a u x m a m m i f e r e s pl~istoc~nes des "Argiles de Bruges" (Gironde). Comptes-Rendus sommaires de la Socigtg ggologique de France, 12: 238-239. WERNERT P. 1957 - S t r a t i g r a p h i e pal~ontologique et pr6h i s t o r i q u e des s ~ d i m e n t s q u a t e r n a i r e s d'Alsace. Achenheim. Mdmoire du Service de la Carte ggologique d'Alsace Lorraine, 14, 262 p. E. C R I ~ G U T - B O N N O U R E
Museum Requien 67, rue Joseph-Vernet F-84000 Avignon