The first record of “spelaeoid” bears in Arctic Siberia Andrei V. Sher a, Jacobo Weinstock b,c, Gennady F. Baryshnikov d, Sergey P. Davydov e, Gennady G. Boeskorov f, Vladimir S. Zazhigin g, Pavel A. Nikolskiy g,* For a long time, “spelaeoid” (cave-bear-like) bears, Ursus (Spelearctos) spp., were believed to be almost purely European animals. Their geographic range has recently been extended to the east, in southern Siberia, Transbaikalia, Kirghizia, Mongolia and Korea. Two unexpected new findings, presented here in detail, significantly change existing views on the distribution of cave bears; both were found in North- Eastern Siberia, far beyond the Arctic Circle, more than 1500 km to the north-east of the previously accepted range. One of the fossils is a mandible, found near the town of Cherskiy at 68.73N, 161.38E. The analysis of local geology and accompanying mammal fossils suggests that it comes from the Olyorian Fauna (Early to early Middle Pleistocene). Morphologically, the Cherskiy mandible is closest to Ursus savini, a small middle Pleistocene cave bear from the British Cromer Forest-bed Formation, but differs in having a slightly more advanced dentition, and thus it is described as a new subspecies Ursus savini nordostensis. Another newly recognized fossil of the “spelaeoid” bear is an astragalus found at the Oskhordokh site at 67.54N, 135.67E, on a large gravel bar on the right bank of the Adycha River. This specimen is attributed to Ursus cf. deningeri. The paper also presents an interesting example of the interaction between classical and “molecular” palaeontology. The new finds significantly change existing ideas on the ecology and evolution of cave bears, some of the most remarkable members of the extinct Pleistocene megafauna.
Torres P.H. 1988a. Osos (Mammalia, Carnivora, Ursidae) del Pleistocene Ibérico: Filogenia; Distribución estratigrafica y geográfica. Estudio anatomico y metrico del craneo. Boletín Geológico y Minero
ABSTRACT Stratigraphical distribution, geographical dispersion and morphological and biometric studies of ¡berian pleistocene bears, are the main topics of this paper. Geographieal distribution is poorly known in U. etruseus, U. mediterraneus and U. prearetos. U. deningeri mainly appeared in the Central part of Spain. U. spelaeus appeared in three distribution areas: Mediterranean, Cantabrian and Central. U. aretos colonizated the whole of the Iberian Peninsula. Stratigraphical distribution: U. etruseus from the Upper Pliocene until the GÜnz. U. prearetos Cromer-Upper Riss. U. deningeri Mindel-Lower Riss. U. spelaeus Upper Riss-Later Würm 111. In the Iberian Peninsula U. arctos appeared at the Lower Würm. Skull: Sex dimorphism and ontogenic changes are studied. The general shape of the cave bear skull varies between two extreme morphologies: dolichocephalic and brachycephalic. The last is absent in U. etruseus and U. arctos. In the cave bear skull, the facial structure is characteristic: step and frontal inflation. The morphology of the brain roof is flattened in U. spelaeus and concave in U. arctos. The olfactory lobe is big in the bown bear and smaller in the cave bear. The cave bear skull is heavy, although pneumatized, and its condyles are bigger and oriented in a different way than in U. arctos. The zigomatic archs are oriented up and backwards in U. spelaetls and directly backwards in U. arctos. In the cave bear skull appear as relatively bigger: basilar and basifacial length, dental and cheek tet`h length. Facial length and zygomatic width are smaller. The facial shortening of the cave bear skull does not affect thc basilar and palatal regions oa.upm.es/3972/2/TORRES_ART_1988_06.pdf Observations on the Ursus gr. spelaeus remains from the Pocala cave (Trieste, Friuli Venezia Giulia, N. Italy) Mario Rossi1 & Giuseppe Santi2 3. MATERIAL AND METHODS 10 skulls, 9 of which are stored into the Geological and Paleontological Museum of the University of Padua and 1 which is preserved in the Museo di Storia Naturale of Milan, 28 P4, 38 P4 and 21 M1 compose the samples under study. The parameters of the measurements of the skulls and teeth are those codified by Torres (1988a, b,c, d), and are presented in Table 1.
Ancient DNA and dating of cave bear remains from Niedzwiedzia Cave suggest early appearance of Ursus ingressus in Sudetes Mateusz Baca a,*, Pawe1 Mackiewicz h,*, Anna Stankovic b,e , Danijela Popovic c , Krzysztof Stefaniak d , Kinga Czarnogórska e , Adam Nadachowski f , Micha1 Ga˛siorowski g , Helena Hercman g , Piotr Weglenski b,c a Center for Precolumbian Studies, University of Warsaw, Krakowskie Przedmiescie 26/28, 00-927 Warsaw, Poland b Institute of Biochemistry and Biophysics, Polish Academy of Science, Pawinskiego 5a, 02-106 Warsaw, Poland c Centre of New Technologies, University of Warsaw, Miecznikowa 1, 02-096 Warsaw, Poland dDepartment of Evolutionary Biology and Ecology, University of Wrocław, Sienkiewicza 21, Wrocław 50-335, Poland e Institute of Genetics and Biotechnology, University of Warsaw, Pawinskiego 5A, 02-106 Warsaw, Poland f Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31-016 Cracow, Poland g Institute of Geological Sciences, Polish Academy of Sciences, Twarda 51/55, 00-818 Warsaw, Poland hDepartment of Genomics, Faculty of Biotechnology, University of Wrocław, Przybyszewskiego 63/77, 51-148 Wrocław, Poland
abstract The migration of Ursus ingressus from Southern to Western Europe, where it replaced native forms of Ursus spelaeus, is well documented. However, its occurrence in Central and Eastern European countries is unexplored. We have characterized the cave bear population inhabiting the Sudetes Mountains during the Late Pleistocene. DNA analyses were performed on 39 cave bear specimens from the Niedzwiedzia Cave. Ten successfully yielded the 254-bp sequences of an mtDNA control region. Phylogenetic analyses allowed assignment of nine samples to the U. ingressus haplogroup and one to brown bear (Ursus arctos). The mtDNA haplotypes of U. ingressus formed a single cluster and differed from haplotypes obtained for cave bears from other excavation sites in Europe. Uraniumethorium and radiocarbon dating of selected samples has shown that U. ingressus appeared in the Sudetes region at least 80,000 years ago. This finding may suggest that the appearance of U. ingressus in this area resulted from migration that occurred earlier than the colonization of the Alps and Swabian Jura. It is possible that migration of U. ingressus proceeded along the Carpathian and Sudetes arc. It is widely accepted that two main forms of cave bear, Ursus ingressus and Ursus spelaeus, inhabited Europe during the Late Pleistocene (Rabeder, 1995; Rabeder et al., 2000, 2004a, 2004b; Rabeder and Hofreiter, 2004; Hofreiter et al., 2004b) (Fig. 1). However, the taxonomical position of these forms is still under debate: they are considered either as separate species (Rabeder, 1995; Rabeder and Hofreiter, 2004; Rabeder et al., 2004a, 2004b; Hofreiter et al., 2004b) or as U. s. spelaeus and U. s. ingressus, subspecies of U. spelaeus (Baryshnikov and Puzachenko, 2011). U. spelaeus has been further divided into ‘classic’ U. s. spelaeus and two high-Alpine forms with reduced body size, U. s. eremus and U. s. ladinicus (Rabeder et al., 2004b, 2008). U. spelaeus remains have been found in Spain, France, Germany, Belgium, Italy, and Austria, while U. ingressus inhabited mostly Eastern Europe and has been discovered in Romania, Slovenia, Ukraine, Czech Republic, Slovakia, and Greece but also in Switzerland, Austria, and Germany. Divergence of these two forms took place 173e414 ka (Knapp et al., 2009). The place of U. ingressus origin remains unknown: however, the basal position of haplotypes from the Romanian site Pes¸ tera cu Oase in phylogenetic trees suggests Southeastern Europe (Baca et al., 2012). It has been proposed that the westward migration of U. ingressus started ca. 60,000 years ago (Rabeder and Hofreiter, 2004; Münzel et al., 2011). The presence of U. ingressus was genetically confirmed in several sites in Austria (Gamssulzen, Nixloch, and Herdengel caves), and its first appearance has been dated to ca. 50,000 years ago (Rabeder and Hofreiter, 2004; Hofreiter et al., 2004a, 2004b). U. ingressus cohabited with Alpine forms of U. spelaeus for some time and ultimately replaced it. Further migration of U. ingressus to the Swabian Jura ca. 32,000 years ago probably also resulted in the final replacement of U. spelaeus ca. 28,000 years ago (Hofreiter et al., 2007; Münzel et al., 2011). We previously reported two aDNA sequences of this species from Niedzwiedzia Cave in Poland, moving its range northwards (Baca et al., 2012). The taxonomical affiliation of these samples was confirmed by phylogenetic and morphometric analyses. One sample was found to be over 49,000 years old, indicating an earlierthan-expected migration of U. ingressus into the Sudetes region. To verify this interesting hypothesis and to assess the genetic diversity of cave bears from this region, we have carried out aDNA analyses on a larger sample and applied uraniumethorium dating of bone collagen to obtain the precise age of the studied samples.
"...Interestingly, one sequence obtained from a III metatarsal (KLEVI_22) excavated in Niedzwiedzia Cave clusters with sequences of brown bear (U. arctos) (Fig. 3). This positioning confirms previous biometrical and morphological analyses indicating that this bear species also inhabited this cave in the Late Pleistocene (Wiszniowska, 1989; Wiszniowska et al., 1996; Bieronski et al., 2009)..."
. Discussion Paleogenetic characterization of the cave bear in Europe is still incomplete. While the distribution and phylogeography of cave bear haplotypes in Western Europe are well documented, the north and northeastern parts of its range are poorly characterized. Our analyses of aDNA from Niedzwiedzia Cave, which is located north from the Carpathian arc, at least partially fill this gap. Phylogenetic analysis of 12 DNA samples obtained from remains excavated from distinct excavation profiles of different geological ages indicated that they undoubtedly belong to the U. ingressus group; an exception was one sample that was clearly affiliated with the brown bear. No sequences related to U. spelaeus were identified, suggesting that U. ingressus was the only form of cave bear inhabiting the cave for at least 40,000 years. The population of U. ingressus from Niedzwiedzia Cave appears quite divergent from other European samples. These findings plus the age of samples (up to ca. 80,000 BP) indicate an early separation of this population from other European U. ingressus. Afterwards, the range of the Carpathians may have played the role of an insulating barrier because no evidence of gene flow has been identified between populations from both sides of this mountain range. The presence of only the U. ingressus haplotype and the lack of U. spelaeus during the Early Weichselian in the Sudetes differ from other central European sites located in the Austrian Alps and Swabian Jura, to which migration of U. ingressus from Southeastern Europe was associated with replacement of U. spelaeus (Rabeder and Hofreiter, 2004; Rabeder et al., 2008; Münzel et al., 2011). It was proposed that U. ingressus was ecologically better adapted to continental environments and could outperform U. spelaeus (Baryshnikov and Puzachenko, 2011). Morphological studies of cave bears from Alpine caves revealed improved masticatory performance of U. ingressus in comparison to U. spelaeus, which allowed for more efficient food intake and processing (Rabeder et al., 2008). Greater morphological variability of U. ingressus also suggests its better adaptational abilities (Rabeder et al., 2008). The earliest known samples of Alpine U. ingressus came from Gamssulzen Cave and are dated to 47,300 BP (Bocherens et al., 2011), i.e., from the beginning of high climate fluctuations during MIS 3. This context may suggest that because of better adaptation skills, U. ingressus could successfully compete with U. spelaeus during sudden and severe climate oscillations. It cannot be excluded that these adaptations also facilitated its migration north to the Carpathians, where U. spelaeus was not reported. According to dating of specimens from Niedzwiedzia Cave, U. ingressus appeared in the Sudetes at least 20,000 years earlier than in the Austrian Alps, during MIS 5a. In this context, it is interesting to consider if potential differences in dietary habits between U. ingressus and U. spelaeus contributed to their different distribution. It has been assumed that the cave bear was generally adapted to a herbivorous diet. The herbivory of the cave bear is well supported by many craniodental features (see e.g., Kurtén, 1976; Mattson, 1998; Rabeder et al., 2000; Grandal-D’Anglade and López González, 2004), the relatively thic dental enamel cap (Mackiewicz et al., 2010), several adaptations in enamel structure (Wiszniowska et al., 2010), and numerous stable isotope studies (e.g., Bocherens, 1990; Bocherens et al., 1994, 2006; Nelson et al., 1998; Vila-Taboada et al., 1999; Fernández-Mosquera et al., 2001). Isotope analyses of samples with confirmed haplotype affiliation based on DNA showed that both U. spelaeus and U. ingressus were vegetarians (Bocherens et al., 2011; Bon et al., 2011; Münzel et al., 2011). Interestingly, studies of cave bears from two closely located caves in Austria showed that U. spelaeus eremus and U. ingressus probably consumed different plant types from different habitats (Bocherens et al., 2011). Recent analyses based on large isotopic data sets revealed that the cave bear isotopic profile does not differ from that of the grizzly bear (U. arctos horribilis), accustomed to a diversified diet that varies from vegetarian to carnivorous (Robu et al., 2013). Moreover, samples from Southeastern European (Carpathian) sites, probably related to U. ingressus, showed higher values of the nitrogen stable isotope, which suggests a greater abundance of animal protein in their diet. In agreement with that, studies of U. ingressus samples from Greece indicated that this bear was mainly vegetarian but with a variable component of what was likely aquatic animal protein (Dotsika et al., 2011). Therefore, it cannot be excluded that U. ingressus was more flexible than U. spelaeus in its dietary habits, which facilitated its migration, especially to colder ecozones, e.g., northwards from the Sudetes and the Carpathians, as our studies have indicated 5. Conclusions Phylogenetic analyses of specimens from Niedzwiedzia Cave in the Sudetes showed that this cave, located north of the Carpathian arc, was inhabited for at least 40,000 years by only one taxon of cave bear, U. ingressus. Dating of some samples (ca. 80,000 BP) indicates that the appearance of U. ingressus in the Sudetes took place earlier than in Western Europe. In agreement, the haplotypes from Niedzwiedzia are divergent from those from other Western European sites. The Niedzwiedzia Cave population was probably isolated from other Western European cave bear populations because the identified haplotypes were not detected in other excavation sites. www.smorfland.uni.wroc.pl/uploads/Main/1-s2.0-S1040618213006538-main.pdf Skeleton of cave bear Ursus speleaus in the the pavilion of the Bear Cave Kletno,Poland. upload.wikimedia.org/wikipedia/commons/a/ae/2015_Jaskinia_Niedźwiedzia_w_Kletnie,_szkielet_niedźwiedzia_jaskiniowego_05.JPG upload.wikimedia.org/wikipedia/commons/6/69/2015_Jaskinia_Niedźwiedzia_w_Kletnie,_szkielet_niedźwiedzia_jaskiniowego_02.JPG