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The European Descendants of Ursus Etruscus C. Cuvier (Mammalia, Carnivora, Ursidae)

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The European Descendants of Ursus Etruscus C. Cuvier (Mammalia, Carnivora, Ursidae) Boletin Geologico y Minero. Vol. 103-4. Ano 1992 (632-642) GEOLOGIA The European descendants of Ursus etruscus C. Cuvier (Mammalia, Carnivora, Ursidae). Por T. DE TORRES PEREZHIDALGO (') ABSTRACT This paper deals with a review of the Pleistocene Bears origin, evolution, and stratigraphical distribution. U. ruscinensis DEP. could be considered the common ancestor of all the European Pleistocene Bears, as the source of two evolutive lineages: one the more conservative, U. mecliterraneus F. MAJOR, and another the more evolved which starts with U. etruscus G. CUV. and gave origin o two evolutive trends: the today vanished speloid one (U. deningeri v. REICH, and U. spelaeus ROS.-HEIN) and the still living artoid one which is represented today in the true brown bear (U. arctos LIN.). U. minimus DEV.-BOUILL. does not constitute a link between U. ruscinensis and U. etruscus: it is a lateral branch, in the general evolutionary schedule, more evolved than U. etruscus. Recent findings of arctoid Ursidae remains in the Iberian Peninsula, ranging from Lower to the Middle (Upper) Pleis­ tocene, allowed us to think that the general migration southwards of an Asiatic Brown bear population during the Wurm glacial period beginning, superimposed on an authochtonous European Brown bear population (prearctoid): U. prearctos BOULE, which is the probable ancestor of the polar bear [U. maritimus PHIP.). Key words: Ursidae, Phylogeny, Stratigraphy, Pleistocene, Europe. R E S U M E N En este trabajo se revisan el origen, evolucion y distribucion estratigrafica de los osos del Pleistoceno. IS. ruscinensis DEP. puede considerarse como e! ancestro comun: de el derivaria un grupo muy conservador (U. medite- rraneus F. MAJOR)) y otro mas evolucionado U. etruscus G. CUV. que sera el origen de las dos principales lineas evo- lutivas de Ursidos pleistocenos en Europa: la linea espeloide (U. deningeri V. REICH, y U. spelaeus ROS.-HEIN.), muy bien conocida, y la linea arctoide, cuyo representante actual es U. arctos LIN. U. minimus DEV.-DEBOUILL. no constituye un estadio intermedio entre U. ruscinensis y U. etruscus, debe tratarse de un representante lateral dentro del esquema evolutivo general, mas evolucionado que U. etruscus. Hallazgos recientes en la Peninsula Iberica, de osos arctoides con edades que van desde el Pleistoceno Inferior al Medio (Superior), permiten pensar que la gran migracion hacia el sur de parte de la poblacion asiatica de oso pardo, que tuvo lugar a comienzos del Wurm, se superpuso a una poblacion europea autoctona de oso pardo (prearctoide): U. prearctos BOULE, de la cual derivo el oso polar (U. maritimus PHIP.). Palabras clave: Ursidos, Filogenia, Bioestratigrafi'a, Pleistoceno, Europa. 1. FOREWORD cies. To solve this will be the goal of this article. In spite of the fact that there are a very large amount of papers dealing with Pleistocene bears, there remain some gaps in the phylogeny and 2. PHYLOGENY stratigraphical distribution of the different spe- The genus Ursavus could be considered the com­ mon ancestor of the Ursus genus, the first re- (*) Dpto. de Ingenieria Geologica. Escuela Tecnica Su­ presentings of which appeared at the Upper perior de Ingenieros de Minas. Madrid. Pliocene. 12 a: IOUGOCENE! MIOCENE PUOCENE PLEISTOCENE r0. I .., ::l ;l: ~ iii C'l bl MINDEL RISS WURM ~ c: ~ 0 ~ ~ ;l:" ;l: ~ ~ a 0 /; '"C'l ~ "z ~ LOW I MIDDLE I UPPER C'l " t"" "'" t"' U rusclnensis - U. minutus ._ Uminlmus _ U. """",,""""'JS _ •••••••••••••••• \ \ i \ 4-634 T. DE TORRES PEREZHIDALGO The Ursavi were Carnivora with a typical enough with a heavily built skeleton and bunodont-sec- Ursoid dentition: there is a certain cusp dupli­ torial cheek-teeth morphology: it is the Deninger city in molars and the lower carnassials show bear, Ursus deningeri VON REICHENAU, which a cutting-like paraconid, while the metaconid is would be superseded by its descenent: the cave very small and placed backward. They were not bear, Ursus spelaeus ROSEN-MULLER-HEINROTH, too different from true bears, but smaller, like which finally vanished at the end of Wurm with­ the wolverine (Gulo gulo). This genus had a out further descendants. In this species those full Miocene development. morphological characteristics which were poin­ ted in the Deninger bear appeared exagerated. The first relatively well documented bear spe­ cies are Ursus ruscinensis DEPERET and Ursus In the Cromer, and until the Riss, there is a minimus DEVEZE-DEBOUILLET. Their first ap- more conservative evolutionary lineage represen­ peareance was during the Pliocene. ting: U. prearctos BOULE, which retained very close similarities with its ancestor U. etruscus. U. ruscinensis was found in the Ruscinian locali­ In the WCirm this species would be substituted ty of Perpignan (France).) Very close to this by the still living brown bear: Ursus arctos species, if not the same, are Ursus minutus GER- LINNEO. VAIS from the Pliocene of Montepellier (France); U. wenzensis STACH from the Polish site of Although they will not be studied in this paper Weze, of similar age to Montpellier, and Ursus it is interesting to briefly cite a group of small (Protursus) boecki SCHLOSSER from the Plio­ sized Pleistocene bears very closely related to cene lignites of Barot-Kopeck (Hungary) which the U. ruscinensis, also known as Plio narctos in FICCARELLi's opinion (1979) is a U. ruscinen­ group, of which the best known, and probably sis synonymous. the only one representative was U. mediterra- neus FORSYTH-MAJOR, in STEHLIN (1933)'s opi­ Along the Lower ViHafranchian, Villarroya (Spain), nion, which KURTEN and POULI.ANOS (1977) there appeared, for the first time, Ursus etrus- called «european black bear», assuming a nar­ cus G. CUViER, which reached its highest eco­ row parallelism with the Thibet bear Ursus thi- logical development in the Middle ViHafranchian: betanus G. CUVIER. Val d'Arno (Italy), Tegelen (The Netherlands), Saint Vallier (France), La Puebla de Valverde The bear of Achenheim (France) Ursus schertzi (Spain) and a certain number of other minor lo­ DEHM, must be considerated as a U. mediterra- calities. This species disappeared at the Upper neus synonymous, in spite of MOTTL (1951 )'s ViHafranchian, Venta Micena (Spain), and it is opinion that it was an arctoid lineage relation, considered to be the ancest or of the main Euro­ like an intermediate stage between the brown pean Pleistocene bears. bear of Repolust and U. priscus: «Der Braun bar der Repolust Hole also zwischen Ursus Although it is possible that U. minimus and U. schertzi DEHM und Ursus priscus GOLDFUSS etruscus were related to U. ruscinensis, it seems der Steiermark...». difficult that they are both related, in spite of the fact that U. minimus has been considered Others representatives of this group of small an U. etruscus ancestor species, STEHLIN (1933), sized bears, and probably U. mediterraneus sy­ THENIUS (1959), KURTEN (1975), TORRES (1984, nonymous too, are: U. (Plionarcstos) Stehlini 1986). BERZI's analysis (1966) of Gaville U. mi­ KRETZOI from Gombasszog (Hungary) of Riss (?) nimus mandible morphology, and of this author's age, Ursus (Plionarctos) telonensis BON I FAY of own data on unpublished material from Layna, Cimay (France) of Mindel age and Ursus sack- proved that both species were unrelated, dillingensis HELLER from Sackdillinger Hole showing U. minimus mandibular characteristics (Austria)) of doubtfull age (Lower- Middle Pleis­ more evolved than in U. etruscus; FICCARELLI tocene?). (op. cit.) agrees on an early extintion of the former, but thought that this had more carnivo­ The Plionarctos genus can not be employed in rous characteristics that U. ruscinensis. European bear description, mainly because it was employed by FRICK (1926-1929) to describe In the Cromer, for the first time, a bear of un- an Ursus (Tremarctos) ornatus F. CUVIER an­ doubtable speloid characteristics was detected cestor: Ursus (Plionarctos) edensis FRICK, it 14 THE EUROPEAN DESCENDANTS OF URSUS ETRUSCUS C. CUVIER (MAMMALIA, GARNIVORA, URSIDAE) 4-635 was employed later by KRETZOI (1970-1971) as other, more conservative (arctoid)), was com­ «Plionarctosgroup» to describe some small sized posed of U. prearctos and U. arctos, showing a species that he throught similar, ERDBRINK similar, but quite less marked, tendency; U. ma­ (1953), to Ursus (Helarctos) malayanus RAFFLES. ritimus can be explained as result of an insulari- zation period effecting a prearctoid population In short: since the Upper Villafranchian there during an advance of the ice cap period. has been a relativelly net species distribution: some were big sized, U. etruscus derivated, A description and discussion of the six species coexisting with others clearly smaller and clo­ before mentioned will be the central objective sely related to its ancestor U. ruscinensis It of the following. is not necessary to invoque an U. etruscus in- vassion, as was suggested by FICCARELLI (op. cit., p. 1681: "Col Villafranchiano medio-supe- riore VUrsus etruscus invade larga parte dell 'Eu­ rasia. ..». This species was already all over 3. DESCRIPTION AND DISCUSSION since the Lower Villafranchian, although its de- OF THE SPECIES mographical boom took place at the moment Ursus etruscus G. CUVIER 1823 indicated by the author. Synonymous: Ursus cultridens NESTI 1826, There is also a co-existence of two evolutive Thalassarctos etruscus AIRAGHI 1922. lineages apart from the residual presence of As stated before, this species appeared at the U. mediterraneus: since the Cromer and until the Riss it was composed by U. deningeri and Lower Villafranchian (Viliarroya), reached its po­ U. prearctos, since the Riss it was composed of pulation peak at the Middle Villafranchian (Val U. spelaeus and U. prearctos. Later, during the d'Arno, Saint Vallier) and disappeared during the Wurm, U. arctos substituted U. prearctos. The Upper Villafranchian (Venta Micena, Spain). former was an asiatic inmigrant which at the It is anatomically characterised by a notably si­ end of Pleistocene times colonized the whole of zed skeleton when compared with U.
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