Departement du Rhone - Museum, Lyon NEW TAXA OF ALPINE CAVE BEARS (URSlDAE, CARNIVORA) Gemot RABEDERt, Michael HOFREITER2, Doris NAGEL1, and Gerhard WITHALM' ABSTRACT Rt':sUMt Morphological and metrical differencesbetween seve­ Lesdifferences morphologiques et metriqucs entre plu­ ral alpine cave bear associations of the same age are sieurs populations d'ours des cavernes alpins de meme large enough to suspect more than one evolutionary age sont suffisamment importantes pour qu'on puisse line. The contemporaneous age of several morpholo­ supposer I'existence de plusieurs lignees evoiutives dis­ gically distinct forms is confinned by radiocarbon tinctes. Les datations par Ie mdiocarbone confinnent la dating. Moreover analyses of fossil DNA indicate at coexistence de fonnes morphologiquement differentes. least three different lineages within the cavebear De plus les analyses sur I'ADN fossile montrent group. For two of these groups, both morphological qu'existent au moins trois lignees d'ours des cavernes. and genetic data show evidence of reproductive isola­ Pour deux de ces groupes, les donnees morphologiques tion. Thus, we suggest that Ursus spelaeus comprised comme les donnees genetiques prouvent des isolats de at least two different species. Ursus illgressus n. sp. reproduction. Aussi nous suggerons qu' Ursus spelaeus occurred in the eastern parts of the Alpine region and correspond au moins a deux especes differentes. Ursus in the Dinarids of Slovenia and Croatia. For two other ingressus n. sp. est repandu dans les Alpes orientales et morphological forms, subspecific status with Ur.HI$ dans les Alpes Dinariques de Siovenie et Croatie . Pour spe/aeus fadillicus n. ssp. in the Dolomites and Ursu$ les deux autres fonnes morphologiques, nous propo­ $pelaeus eremlls n. ssp. in the Totes Gebirge is sug­ sons la distinction de deux sous-especes, Ursus spe­ iaeus ladinicus n. ssp. dans les Dolomites et Ursus spe­ gested, as they are genetically relatively close and no data about their reproductive relationship is yet laeus eremus n. ssp. dans Ie Massif des Totes Gebirge : available. elJes sont en effet relativement proches genetiquement mais il n'existe pour J'instant aucune donnee sur Icur croisement eventuel. Mots-cl6i : Keywords phylogeny, eYolWion, uuonomy, cave bear. phylogenie. evolulion, /(uOIlOlllie, ours des DNA-analysis cal'emes, ADN. INTRODUCTION on the excavation activity was intensified under the leadership of Gernot Rabeder; excavation campaigns The study of the Alpine bear caves has a - in some cases lasting for many years - were carried out in the Ramesch Knochenhohle, the Brieglersberg long tradition aI lnstitute of Palaeontology at the cave, the Gamssulzen cave, the Brettstein cave and in University of Vienna. It began more than 80 years ago under the direction of Othenio Abel in the the Lieglloch (all of them in the Totes Gebirge). DrachenhOhle of Mixnitz, continued under Kurt Other excavations were undertaken in the Ehrenberg in the Schreiberwand cave on the Hartelsgraben cave (Gesause), the Herdengel cave Dachstein, the Merkenstein cave near Baden, the and (he Schwabenreith cave near Lunz am See bear cave of Wind en, and finally in the Salzofen cave (Lower Austria), in the Nixloch near Losenstein­ in the Totes Gebirge as well as in the Schlenken­ Ternberg in Upper Austria, the Sulztluh cave in the Durchgangshohle in !.heOstcrhom group. From 1980 R1itikon (Switzerland), the Conturines cave in the 'lnstitutfUr PaJaootologie dcr Universitill Wien, lMa;< Plaock-Gesellsc:haft, evo Inslilut rur lutionlire Anthropologie, leipzig Cahiers scicntifiques - Departement du Rh6ne· Museum, Lyon, Hors sene n02 (2004) - p. 49-67. II fig., 5 tabl., 2 pI. Actes du 9< Symposium international sur I'ours des cavernes, Entremont-Ie-Vieux (Savoie, France), septcmbre 2003 ,,°2 (2004) 49 Cahiers scienlifiques/ Hors serie Centre de Consermtiol! er d'£tude des Collectiolls Dolomites (Italy) as well as in the Potacka zijalka in lution in this range of time (hypothesis of saltation) the Karawankes (Slovenia). Under O. Abel qu.estions or a bear population on a high level of evolution about the morphology of function and the life style of advanced from a so far unknown area to the eastem the cave bear became imponant as well as the reasons Alps (hypothesis of immigration):' for their extinction (ABEL & KYRLE 1931), while Another high alpine small fonn was discovered in the K. Ehrenberg was above all fascinated by the rela­ Dolomites. In the years 1998 to 200 I a large number tionship between bear and palaeolithic man (keyword: of fossils was excavated from the Conturines cave, cave bear cUlt). In the last two decades due to consi­ situated at an altitude of nearly 2800 m high that were derably advanced excavation methods we could go at first assigned to a smaller cave bear relative. deeper into questions of evolution and fine stratigra­ Because of the mixture of archaic attributes (small phy. because now statistically relevant numbers of fos­ dimensions. P3 in over 25% of the skulls still existing, sils are available and the possibility of radiometric il primitive, m2-trigonidtal primitive) and progressive dating gives totally different age classifications attributes (12, ml-and m2-enthypoconid on a high (DOPPF.s & RABEDER 1997). level of evolution. also m2 -mesolophid, M2-metha­ It was K. EHRENBERG (1929) who nOliced that the cave loph and above all P4) the Conturines cave bear must bear remains from different caves differ in size. based have an exceptional position. (RABEDER, 1999) : on the variability of the metric values. In the first des­ "'... this leads to the conclusion that this high alpine cription of the very small bear form from form was derived probably from a late Middle Schreiberwand cave he created the term "hochalpine Pleistocene U. dellillgeri-group... A genetic connec­ Kleinform" (high alpine pygmy form). The reduction tion with the high alpine cave bears of the nonh Alps of the dimensions was seen as an adaptation to life in ... can be excluded". the high Alpine region (shon summers, long winters). The work on the evolutionary statistics of metapodial Other authors explained the on average smaller bones done by G. WtTHALM (2001) and on tibiae by dimensions as sex-specific differences: the smaller M. FROEMEL (2001) clarified the exceptional position females would have preferred the higher situated of the high alpine cave bears also in respect to the caves for hibernation.This hypothesis,jokingly called postcranial skeleton. The more developed first meta­ "Pascha-theory" was taken to ad absurdum quite carpal- and metatarsal bones as well as the relatively recently (RABEOER, 2(01). longer and more slender tibiae are interpreted as an The excavation in the Ramesch bone cave from 1979 adaptation to tife in high alpine regions. to 1984 was the key to modern cave bear research in Now we can discuss the relationship between the the Alps. The until then unique site in an undisturbed, other cave bear faunas and the three aforementioned more than two meters thick profile of sediment which groups. Taking the paleontological data we can accept made it possible to ask questions about metric and with utmost probability that the bear from morphological details. The big difference to the Gamssulzen cave inhabited the following caves: remains from the Drachenhohle near Mixnitz leads at Nixloch near Losenstein-Ternberg, Lieglloch first to the conclusion that the "high 'alpine small (Austria), Schnurenloch (Switzerland), Potacka zijal­ foml" occurs also in the Ramesch bone cave (HtLLE & ka. Mokriska jama (Slovenia). Vindija (Croatia); the RABEDER, 1986). The taxonomic status of this fonn of bear from Ramesch cave is also documented in the the high alpine region could not be bener argued at Salzofen cave, whereas the bear from Conturines cave that time. But the conclusions drawn from the excava­ is so far only known from this cave in the Dolomites tion in the Gamssulzen cave brought a change. of South Tyrol. Finally, numerous cave bear faunas I.n the Totes Gebirge, two cave bear forms that were cannot be slotted into this scheme, for example the unequal in size and at extremely different levels of faunas of the Brieglersberg cave and the Brettstein evolution, lived contemporaneously for many thou­ cave (Totes Gebirge) or the caves of easl Switzerland, sands of years and apparently they did not interbreed. e.g. the Drachenloch near Vatlis and the Sulzfluh Already by then the possibility of a formal separation cave. (Ramesch bear and Gamssulzen bear as two different With the analysis of ancient DNA we have now a species) was taken into consideration (hypothesis of completely new possibility to investigate the phyloge­ speciation). netic relationship of glacial animals. It is well known Two hypotheses were presented (RABEDER, G., for a long time that in the sediments of caves organic 1995:81) : "There are abnormal differences of evolu­ molecules like collages are conserved much longer tion also to other absolute dated bear faunas that can than in fossils from open sites because of the constant only be explained that either there was an erratic evo- temperature and humidity in caves. This applies also CaMers scie!JIijiqlles Nors sirie / 11°2 (2004) 50 Departement du RhOne - Museum, Lyon to DNA. Although the preservation of PCR amplifia­ the University of Vienna and the Institute of ble genetic material differs from cave to cave, DNA Evolutionary Anthropology of the Max Planck­ analysis is possible on bones up to about 100.000 Gesellschaft in Leipzig - led to the investigation of years old. various aspects of the phylogeny and taxonomic posi­ The cooperation between palaeontology and genetics tion of cave bears. - in this case between the Institute of Palaeontology of Fig 1 - Location map of the type localities: GS Gamssulten cave, RK - Ramesch KoochenhOh1e. CU - Conturines cave (Italy) and ZoolithenhOhle (Germany). LOCATION OF THE TYPE LOCALITIES P4/4-lndex The so-called P4/4-index, the geometric mean from the index of the lower jaw-P4 and the upper jaw-P4, seems to be by far the best parameter to investigate the MATERIAL AND METHODS morphological level of evolution.