Rend. Fis. Acc. Idncei s. 9, v. 5:17-26 (1994)

Paleontologia. - - The fossil bear from Sen&e (Southern France). Nota di PAUL MAZZA e MARCO RusvIoNI, presentata (*) dal Socio A. Azzaroll.

ABSTRACT. - - The bear from Sen~ze is here referred to the black bear stock of Eurasia. The coherency of the faunal assemblage is discussed, in the light of the occurrence of late Pliocene and several Late Villafranchian elements, among which Hippopotamus cf. antiquus. The stratigraphical position is also controversial. In accordance with this, the black bear from Sen~ze could represent either the last occurrence of this stock in Europe, or an isolated presence during the Early Pleistocene.

I~v worms: Ursus; Plio-Pleistocene; Eurasia.

RIASSUrqTO. - - Lbrsofossile di Sen&e (Francia meridionale). L'orso di Sen&e ~ qui fiferito al ceppo degli orsi neri dell'Eurasia. Viene messa in discussione la coerenza dell'associazione faunistica per la presenza di alcuni elementi tardo pliocenici e di alcuni tardo villafranchiani, fra i quali Hippopotamus cf. antiquus. Anche la posizione stratigrafica ~ controversa. Alia luce di ci6, l'orso di Sen~ze potrebbe rappresentare l'ultima testimonianza di questo gruppo in Europa oppure una presenza isolata nel Pleistocene inferiore.

HISTORICAL

Seneze is a hamlet situated in the central depression of a volcanic structure. This locality has been known for a long time for its rich fauna of fossil vertebrates.

Schaub (1943) examined the fossiliferous site providing a full revision of the fauna. His list included Ursus etruscus.

In 1947 Jung published the complete list of mammals from Sen~ze, passed to him by J. Piveteau. This list includes Ursus arvernensis in the place of Ursus etruscus. Furthermore Jung reported the occurrence of an unspecified specimen of hippopota- mus, which he attributed to ,Hippopotamus Major>>, in the collections of the Facultd des Sciences de Clermont.

Bout (1960) made a further revision of the site. He recognized two fossiliferous levels, one with fragmental remains (<<dep(3t du pente>>), the other with complete skeletons (,maar>>), perhaps of different age, on the basis of the characters of the site and of the occurrence of Hippopotamus major, which is an anomalous element in the faunal context of Sen~ze. In the faunal list published by Bout, the bear was again quoted as Ursus 8truscus.

Bout (1970) revised his previous study emphasizing four successive phases of deposition. The lowermost level is made of pyroclastic rocks and lavas; the following term is represented by <<... a maar with explosion breccia accumulated on the western side of the cone of scoria>); the third level is a lacustrine deposit which provided complete skeletons cited by Bout ( 1960); the fourth level, which caps the local sequence, is a slope deposit with fragmentary bone remains.

(*) Nella seduta det 18 giugno 1993.

1 8 P. M A Z Z A - M. R U S T I O N I

Prevot and Dalrymple (1970) dated the Sen~ze fauna of the <<maar>~ at a time younger that 2.3 Ma and found a normally magnetized subchron below the fossiliferous beds of Sen~ze, which Opdyke et al. (1983) referred either to the Reunion or to the Olduvai Subchron.

Heintz et al. (1984) published an updated list of the fauna, which is reported in a section below.

Mein (1979, 1990) placed the Sen6ze fauna, together with several others, such as those from Mas Rambault, Brielle, Ostramos 8, Betfia XIII Tasso, etc., in the Quaternary, at a time following that of this MN17 mammal zone.

Replying to Opdyke et al. (1983), Repenning (1983) precised that the youngest lava flow underlying the oldest fossiliferous lacustrine deposits dates to 2.3 Ma; that the latter are reversely magnetized and thus referable to the Matuyama Chron; and that the normaUy magnetized beds present in the lake deposits may be correlated with the Reunion Subchron (which dates 2.14 +2.01 Ma).

In their general revision of the paleomagnetism of several French sites, Thouveny and E. Bonifay (1984) dated the Sen6ze fauna, at most, to 2.01 Ma, but made no mention about the two fossiliferous levels identified by Bout (1960, 1970).

TIlE MAMMALS FROM SENI~ZE

The updated faunal list provided by Heintz et al. (1974) is the following:

Archidiskodon meridionalis Equus stenonis senezensis Equus bressa~us Dicerorhinus etruscus Sus strozzii Croizetoceros ramosus minor ~(Cervus~ phlisi philisi Eucladoceros senezensis senezensis Libralces gallicus Gazellospira torticornis Procamptoceros brivatense Megalovis latifrons Gallogoral meneghinii Pliotragus ardeus Leptobos etruscus Macaca cf. florentina (~Dolichopithecus~ arvernensis Nyctereutes megamastoides Vulpes alopecoides? Canis sp. Ursus etruscus Hyaena perrieri

THE FOSSIL BEAR FROM SENI~ZE (SOUTHERN FRANCE) 19

Euryboas lunensis Acinonyx pardinensis Homotherium sainzelli Megantereon megantereon.

In this list the bear material is attributed to Ursus etruscus and no mention is made about the hippopotamus reported by Jung (1947) and Bout (1960).

Concerning the hippopotamus, Heintz et al. (1974) stressed that <<Le genre Hippopotamus n'existe dans aucun des gisements &udi&>>.

Faure and G@rin (1979) and Faure (1983, 1985) excluded, the presence of hippopotamus material from Sen~ze as well as from any other ViUafranchian site of France. The authors stressed that the hippopotamus material believed to come from Sen~ze should actually come from a nearby locality, Chambezon, also located in the Haute-Loire area.

In a recent visit at the Facultd de Science de l'Universit~ Claude Bernard, Lyon I, we had the opportunity to observe a first phalanx of a manus of Hippopotamus, FSL 211082 (fig. 1: 1A, 1B), which is reported to come from Domeyrat (Sen~ze)fide the registers of the collection; unfortunately, the specimen was erroneously mistaken for a horse element and referred to Equus sp. We ignore if this is the specimen cited by Jung. It has a polished surface, clear evidence of abrasion by transportation.

The measures of the hippopotamus phalanx are the following (in mm):

greatest length: 75.7; physiological length: 59.2; proximal breadth: 47.4; proximal depth: 46.6; breadth of the diaphysis: 38.5; depth of the diaphysis: 30.3; distal breadth: 41.6; distal depth: 28.7.

We believe that the specimen may be attributed to Hippopotamus cf. antiquus. The occurrence of this definitely Pleistocene element of the fauna opens the

problem of the possible occurrence of two distinct faunas at Sen~ze. This point was already raised by Azzaroli et al. (1988). Besides the hippopotamus, other species speak for an early Pleistocene age. Libralces (now Cervalces) gallicus has been recorded in East Anglia in an early Pleistocene faunal context. A small equid may be identified with the early Pleistocene Equus stehlini and a small dog with Canis arnensis of the same age. This may also be the case of Equus bressanus and Megalovis latifrons.

THE BEAR FROM SENI~ZE

The bear material that is preserved at the Facultd de Science de l'Universitd Claude Bernard de Lyon I is represented by a right fragment of facial skeleton with mandible, FSL 210966 (fig. 1: 1A-D).

i

Fig. 1. - 1: Hippopotamus sp., first phalanx of the manus; A, dorsal view (about nat. size); B, lateral or medial view (about nat. size). 2: Ursus minimus-thibetanus; A, fragmentary right maxillary, lateral view (about 1/2 nat. size); B, fight molar row, occlusal view (about nat. size); C, incomplete right marld/ble, lateral view (about

1/2 nat. size); D, right lower dentition, ocdusal view (about nat. size).

"FIE F O S S I L B E A R F R O M S E N I E Z E ( S O U T H E R N F R A N C E ) 21

)readth

19

17

15

13 * x x

11 i i

14.5 15.5 16.5

.F i rs t upper molar

A

g a ~ ~. o a Ao

x " ~ o xv ~a ~ , A ~ x ~ o gm~'a-- ~o A

o o t ' " x

x x ~ x x

x x a

I I I I I I I I

17.5 18.5 19.5 20.5 21,5 22.5 23.5 24.5

length

o ,#

25.5

Im o U.aro tos x U.mln . - th lbe t . �9 Nlhewsn |

! 8 e n # z e ~ U . e t r u e c u s o U. sf f . e t r u s o u 8 I

Fig. 2. -Scat tcr diagram length vs. breadth of the first upper molar.

breadth 25

23

21

19

17

15

13

11 21

S e c o n d upper mo lar

o

@o

o oo o o o

,'g oo ~ go% 0o o• o oo o

o

xXx x x~ x ~ o o • ~ @ ~ x ~ ~ ox

x x xX OXx x

x x u

I B I I l I i I I I

23 25 27 29 31 33 35 37 39 41

length

o o

m

I

43 45

a U .arotos x U .mln . - th ibe t .

t, U . e t r u s c u s o U. off . e t r u s o u s

~" 8 e n # z e

Fig. 3. - Scatter diagram length vs. breadth of the second upper molar.

22 P . M A Z Z A - M . R U S T I O N 1

The facial fragment is represented by the maxillary and part of the zygomatic arch (fig. 1: 1A). The third incisor and the first and second molar are still preserved. The mandible is represented by most of the horizontal ramus and by part of the basal portion of the ascending ramus.

The measurements of the specimens are the following (in mm):

total length of the upper molar row: 52.8; length and breadth of 13/: 12.8, 9.8; length and breadth of M1/: 21.6, 16.7; length and breadth of M2/: 30.6, 17.1; total length of the lower cheek teeth (P/4 - M/3): 76.5 (?); total length of the lower molar row: 63.7; length and breadth of P/3: 5.8, 4.2; length and breadth of length and breadth of length and breadth of

P/4:12.4 (?), 7.3; M/l: 25 (?), 10.6; M/2: 23.4, 14;

length and breadth of M/3: 15.4, 13.5; diastema canine-P/4: 38.2; diastema P/l-P/2: 3.1; diastema P/2-P/3: 6.3; diastema P/3-P.4: 7.9; height of the horizontal ramus under M/l: 47.2; height of the depth of the horizontal ramus between P/4 and M/l: horizontal ramus under M/3: 56.8; depth of the horizontal ramus between P/4 and M/l: 17.3; depth of the horizontal ramus between M/2 and M/3: 20.8.

On morphological grounds, the specimen is referable to the Ursus minimus- thibetanus group sensu Mazza and Rustioni (1994); we therefore agree with Jung (1947), and through him with J. Piveteau, in attributing the bear from Sen~ze to a black bear rather than to an etruscan bear. The most significant characteristics of this fossil are its particularly short muzzle, the persistence of the anterior premolars (P1-P3) which are separated by fairly wide diastemas, the slightly prominent dental cusp(-id)s, the weak styles of the first upper molar, the second upper molar with relatively short, weak talon, the lower teeth rather small and compressed latero-medially, the lower carnassial tooth with the typical <<zig-zag>> longitudinal groove, stressed by Erdbrink (1953) as typical of the Asian black bears, and the mandible with short and massive horizontal ramus and deep masseterine fossa. The mandible also shows a pachyostosis at level of the third lower molar, which is particularly evident, in occlusal view, laterally to M/3. This feature is also present in all the U. thibetanus specimens that we have studied; it is not mentioned in the list of characters of the U. minimus-thibetanus group reported by Mazza and Rustioni (1994) because it is generally obliterated in badly preserved fossil specimens.

T I t E F O S S I L B E A R F R O M S E N I ~ Z E ( S O U T H E R N F R A N C E ) 23

breadth

First lower molar

14

12

10

8

o

o w o o ~ o A

A

o

o ~ ~ ~ . ~ ~ 1 7 6 o o xo ~ x ~ , x

x x ~ 1 7 6 x x • x X x

xX x x x x x x x

x x

6 I t I I I I I I I I I I

15.5 16.5 17.5 18.5 19,5 20.5 21,5 22.5 23,5 24.5 25.5 26.5 27.5 28.5

length W

o U.arctos x U.mln.-thlbet. �9 8en6ze |

I U.etrusoue o U. aff. etrusous

Fig. 4. - Scatter diagram length z's. breadth of the first lower molar.

breadth

Second lower molar

19

17

15

13

11

o o

o

oo o o o

~ x x X x ~ , ~ x , ,

x x x ~

x x x x

9 I I I I I I I I I I I I 1

16.5 17.5 18.5 19.5 20.5 21.5 22.5 23.5 24.5 25.5 26.5 27,5 28.5 29.5

length

o U.atoto8 x U.mln.-thlbet. o Plrro i

m

I 8en6ze ~ U.etrusous o U. 8ff. etruacua

Fig. 5. - Scatter diagram length vs . breadth of the second lower molar.

24 P . M A Z Z A - M . R U S I I O N I

b r e a d t h

Third lower molar

19

17

15

13

11

9

7 11.1

x

x x

x • ~

x • •

I I I

12.5 13.5 14.5

o o

x & O o

x xX~Ox o = oo o~,~,

x x~, x

0 0

0

a 0

OD~]

I I I I I I I I I

15.5 16.5 17.5 18.5 19.5 20.5 21.5 22.5 23.5 24.5

l e n g t h

k U.aretos x U.mln.-thibet. = Nlhewan |

I ~' Sen~)ze ~ U.etruscua 0 U. aff. etruscua

Fig. 6. - Scatter diagram length vs . breadth of the third lower molar.

The specimen from Sen~ze shows some peculiarities that single it out from other Pliocene black bears of Europe. It is a relatively large-sized form for the group and its dental proportions fall between the ranges of the typical black bears and those of Ursus arctos, as shown by the scatter diagrams here included (figs. 2-6). This f~lrtherly supports the hypothesis that brown bears may have originated from the black bear stock in the course of the Pliocene (Mazza and Rustioni, 1994).

CONCLUDING REMARKS

The probable occurrence of two s levels at Seneze raises a number of problems. On the basis of the results of the studies of Opdyke et al. (1983), Repenning (1983) and Thouveny and E. Bonifay (1984), the lower fossiliferous beds of the site are negatively magnetized and dated to about 2.01 Ma; but the age of the more recent specimens is uncertain. A further difficulty is represented by the fact that we do not know the stratigraphic position of the bear material. Bout (1960) informed that the older fossiliferous beds yielded complete skeletons, while the younger ones are characterized by fragmental remains; the bear fragments may thus come from the latter. In this case, however, we should correlate the black bear with the Late Villafranchian elements, Canis arnensis, Equus bressanus, possibly E. stehlini and Hippopotamus cf. antiquus, an

THE FOSSIL BEAR FROM SENI~ZE (SOUTHERN FRANCE) 25

association in which black bears have never been recorded before. Moreover, an Early Pleistocene occurrence of black bears in Europe would be stratigraphicaUy isolated from both the Late Pliocene ones of France and Italy and the Middle Pleistocene ones of Germany.

On the other hand, if the bear comes from the Pliocene sediments, it may be one of the last representatives of the black bear stock in Europe before its new, Mid-Pleistocene occurrence at Mosbach. This would apparently fill in the gap in the distribution of the European bears hypothesized by Rustioni and Mazza (1993). In this case, the demise of black bears could be seen as an ecological displacement or an evolutionary replacement by primitive etruscan bears (Ursus aff. etruscus, sensu Mazza and Rustioni, 1994) which made their appearance in Eurasia approximately at this time.

ACKNOWLEDGEMENTS

We are deeply indebted to Prof. C. Gudrin, Dr. A. Prieur and M.me M. Sirven for granting us access to the collections of the Facult~ de Science de l'Universite Claude Bernard de Lyon I. We also thank Prof. A. Azzaroli, Prof. G. Ficcarelli and Prof. D. Torre for critically reading the manuscript and for the proficuous discussions.

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26 p . M A Z Z A " M . R U S T I O N I

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P. Mazza: Museo di Geologia e Paleontologia Universit~ degli Studi di Firenze

Via G. La Pira, 4 - 50121 FW,~NZE

M. Rustioni: Dipartimento di Scienze della Terra Universit~ degli Studi di Firenze

Via G. La Pira, 4 - 50121 FW,~NZE