taxonID	type	description	language	source
B475EB14FFF172439B82FAC1FA858E87.taxon	description	Description of the Bulgarian bear m a t e r i a l f r o m M i s h i n K a m i k. The MK bone sample exhibits an impressive robustness. All the skulls are from very old individuals, the teeth extremely worn, and canines absent. The skulls are robust and very wide, with massive zygomatic arches (Tab. 1); with a strongly domed and bilobated frontal portion, especially in males (Text-fig. 1); the temporal crests are prominent and the sagittal crest is deep. In most cases the sagittal crest (as in many other cave bears) seems longitudinally split (or double) because of the not fully fused temporal lines on its dorsal surface from which the crest is formed (Pl. 1, Figs 1 – 3). The choanae are narrow, the muzzle is rather short, the palate is concave in the area of M 2; the nasal aperture is oblique in lateral view; the rostral end of the nasal bones is at the level of the mesial surface of P 4. The glabella of the MK specimens is deep – varying from 7.85 to 15.4 mm (11.41 mean). Two of the skulls (Nos. FM 3111, FM 3385) have a less domed (more oblique) frontal profile, practically without bilobation; frontals with weaker zygomatic processes and respectively a slightly narrower frontal width; weaker sagittal and transversal crests and less pronounced postorbital constriction. One of them (No. FM 3111) is the smallest of all the skulls from this location (Pl. 1, Fig. 3). We consider these two skulls to be from females. The tooth morphology remains rather unclear due to the degree of wear. Nevertheless, when considering the P 4 outlines it is possible to state that the protocone is placed relatively centrally, close to the paracone-metacone border and that the paracone and metacone are in line and not at an angle with respect to each other (Pl. 2, Fig. 6); these features could also be observed in some better preserved isolated teeth. Only one isolated M 1 has a preserved occlusal morphology (Pl. 2, Fig. 7). It is archaic, with a limited number of swellings and cusplets, the middle longitudinal valley is narrow, wider only on the talonid. The mesocone is single; the cingulum is developed on the labial side from the protocone to the distal end of the hypocone. Two M 2 have a preserved occlusal surface. Each of them has different morphology – one with a more complex morphology and with well-developed cingulum, the other with more simple morphology and without a cingulum (Pl. 2, Figs 8, 9). The hemi-mandibles are well preserved in the MK sample (Pl. 2, Figs 1 – 5, Tab. 2). Only one hemi-mandible has an alveolus for p 2; another has molars only (FM 3331; Pl. 2, Fig. 3), p 4 is not developed at all, and this must be an anomaly. Some of the hemi-mandibles have a preserved p 4 and it is possible to observe its morphology (Pl. 2, Figs 11 a – 15 b). All of the specimens have a paraconid and metaconid more or less of a similar size, situated close to each other. In some cases, there is an additional cuspid of the same size behind them, and all p 4 have smaller additional cusplets (morphotypes D 2, D 2 / D 3 after Rabeder 1989, Rabeder and Tsoukala 1990). The rather simple m 1 morphology is without constriction between the talonid and the trigonid and with relatively more archaic entoconid (Pl. 2, Fig. 10). On the mandibles the preserved m 1 are excessively worn, but in all cases the archaic outline is clearly visible. For all teeth measurements see Table 3. The postcranials are also robust (Pl. 4, Figs 1 – 9, Tabs 4 – 8). The lateral epicondyle of the humerus is enlarged, the lateral condyle of the femur distally protruding, the distal segments of the limbs (ulna, radius and especially the tibia) very shortened. The postcranials of males and females could be distinguished because of the different sizes.	en	Spassov, Nikolai, Hristova, Latinka, Ivanova, Stefanka, Georgiev, Ivan (2017): First Record Of The “ Small Cave Bear ” In Bulgaria And The Taxonomic Status Of Bears Of The Ursus Savini Andrews - Ursus Rossicus Borissiak Group. Fossil Imprint 73 (3 - 4): 275-291, DOI: 10.2478/if-2017-0015
B475EB14FFF172439B82FAC1FA858E87.taxon	materials_examined	Comparison of the Bulgarian bear material from Mishin Kamik Compared to Ursus arctos, the MK bear differs in the robustness of the skull, by the completely different proportions, by the strongly domed frontals, the massive zygomatic arches and the better developped temporal crests, the larger muzzle with very wide surface for the incisors, much deeper palatal bone; visibly narrower choanae, the more complex cheek teeth morphology, absence of upper and lower anterior premolars (they very rarely persist, and are reduced in size). The nasal aperture is more oblique in lateral view and the nasal bones are more caudally situated: their rostral end is at the level of the mesial surface of P 4 (in U. arctos it is normally at the level between I 3 and C 1). The postcranial elements are also more robust, the distal segments of the limbs are significantly shortened. The noted features, as well as the extremely speleoid morphology of p 4 (see Wagner and Čermák 2012) separate the bears from Mishin Kamik from the arctoid lineage and place them in the cave bear lineage. The tooth morphology demonstrates a number of archaic features (see description). In this respect the Mishin Kamik bear is more similar to U. deningeri than to the large and more evolved cave bears from the U. spelaeus-U. ingressus group. This similarity is expressed also by the more or less similar size (length) of the skulls of some U. deningeri populations (SH and PTR for example) (Text-figs 2 – 4); the mean glabella depth is similar to that of U. deningeri from SH and PTR (García et al. 2007, Santos et. al. 2014). A comparison with U. deningeri suessenbornensis (considered by some authors, especially in older works, as a separate species) is practically impossible, as the type material is extremely scarce (see Baryshnikov 2007). 180 300 350 400 450 500 Condylobasal length 300 350 400 450 500 Condylobasal length On the other hand the skull shape and proportions differs from U. deningeri s. str. from the early Middle Pleistocene. In general, U. deningeri has a less wide (with some exceptions) skull (Text-fig. 2). The MK bear has abruptly and strongly elevated frontals and demonstrates the evolved profile of the U. spelaeus-U. ingressus group: The stepped forehead forms a well developed dome, with a slight depression on the parasagittal line. The CT scan comparison (Pl. 3) revealed several specific profile peculiarities of the Mishin Kamik skulls (Pl. 3, Figs 3, 4), the features and the intraspecific variability in the development of the frontal sinuses as well as the thickness of the skull bones. The sinuses are large, they significantly extend backward and may occupy more than half of the brain cavity to about 2 / 3 of its length (Pl. 3, Figs 3, 4). The occipital region in the area of the external occipital protuberance is very thick. In this respect the MK skulls show similarity with the scanned specimens of U. spelaeus from Spain (García et al. 2006, 2007, Santos et al. 2014) (Pl. 3, Figs 1, 2). The basioccipital is also thick (7.6 mm and 8.1 mm respectively) and most probably pneumatized as in U. spelaeus. In spite of its clearly smaller size, in width the MK skulls may measure the same or even exceed the relative width of the skulls of the U. spelaeus-ingressus gr. On the other hand their tooth morphology is less evolved (see description). Comparison with U. spelaeus ladinicus RABEDER et al., 2004 The skull of this small, high alpine cave bear U. s. ladinicus or U. ladinicus from Conturines cave, is similar in skull size (Rabeder et al. 2004) to the MK bears. But all other proportions are clearly different: the zygomatic width is much larger in the MK sample, while the check teeth length is shorter (Text-figs 2, 4) and the mandibles are deeper (Text-fig. 5). Comparison with Ursus deningeroides MOTTL, 1964 and other taxa from large cave bear group This bear from the Repolust cave in Austria was described by Mottl (1964) as a very primitive and small U. spelaeus. The locality is from the late Middle Pleistocene, with an estimated age of about 200 ka (ca. 223 ka for the lowermost level) (Döppes and Rosendahl 2009). The skulls are considerably larger and relatively narrower than in the MK bears (Text-fig. 6 a). They are however as significantly 70 300 350 400 450 500 550 Maximal SNƲOO length U. spelaeus ZLH U. deningeri PTR Regression line U. s. ladinicus CT U. sa. rossicus KRD U. spelaeus ZLH U. deningeri MOS U. spelaeus PTR U. sa. rossicus MK U. spelaeus PTR U. sa. rossicus KRD MK 230 250 270 290 310 330 350 Maximal mandibular length domed as in MK, but with a clearly slanting caudally neurocranial profile (see Mottl 1964: pl. 2, fig. 1 and pl. 3, fig. 1). The muzzles are longer in general (Text-fig. 6 b). About 60 % of specimens retain anterior premolars (P 3 / p 3 and P 1 – 2 / p 1 – 2) and m 1 is to some degree more archaic than in MK. The mandible length of the Repolust cave bears are between 245 – 313 mm, and the depth under p 4 varies from 50 – 71 mm. The MK mandibles are shorter (with maximal value 294 mm), while the depth under p 4 varies from 54.29 – 71.7 mm, indicating a greater depth. The mandible depth between m 2 and m 3 is also greater in MK – 64 – 78 versus 50 – 75 mm for the Repolust cave bears. A cave bear of small size is described from Einhornhöhle (Middle Pleistocene, Germany). There are only mandibles, no skulls are preserved (Schütt 1968); the p 4 and m 1 morphology is similar to that in the MK sample. The main difference in m 1 morphology is the more complex metaconid and more acute paraconid in MK (sensu Grandal d’Angle and López-González 2004). The MK mandibles (with maximal length from 240 to 294 mm and mean value 269 mm) are smaller on the whole in relation to the Einhornhöhle specimens (260 – 319 mm, mean 280 mm). At the same time they are deeper: the depth under p 4 varies from 54.29 to 71.7 mm (mean 62.8 mm) for MK and for the Einhornhöhle bears from 46 to 75 mm (mean 58 mm). A small cave bear (Ursus spelaeus parvilatipedis TORRES, 1991) was described from the Late Pleistocene of Troskaeta cave, Spain (Torres et al. 1991). The mean value for the maximal mandibular length is 280.9 mm vs. mean 270 260 width 250 Zygomatic 240 230 220 300 320 340 360 380 400 a Basilar legnth 190 180 170 length 160 Rostral 150 140 130 120 300 320 340 360 380 400 b Basilar length value of the height behind m 1 – 65.2 mm. These mean values are similar in size to the MK bears (267 mm vs. 63 mm respectively), but more data are necessary for a reliable comparison and conclusions. In comparison with Ursus kudarensis praekudarensis (BARYSHNIKOV, 1998) from the Middle Pleistocene of the Southern Caucasus (Baryshnikov 1998), the skulls from MK are smaller (Text-figs 3, 4), but their teeth morphology is more complicated; in U. d. praekudarensis p 4 are narrower and more archaic; the m 1 is also more archaic.	en	Spassov, Nikolai, Hristova, Latinka, Ivanova, Stefanka, Georgiev, Ivan (2017): First Record Of The “ Small Cave Bear ” In Bulgaria And The Taxonomic Status Of Bears Of The Ursus Savini Andrews - Ursus Rossicus Borissiak Group. Fossil Imprint 73 (3 - 4): 275-291, DOI: 10.2478/if-2017-0015
B475EB14FFF172439B82FAC1FA858E87.taxon	description	The bear from MK is very similar to Ursus rossicus (U. sa. rossicus) from the type locality, Krasnodar (Borissiak 1930, 1932), with a probable age at the very end of the 85 290 340 390 440 490 Maximal length width Distal 30 250 300 350 400 450 Maximal length 200 220 240 260 280 300 320 340 360 Maximal length Middle Pleistocene or the Eemian (Baryshnikov 2007). The skulls from Krasnodar are robust, clearly domed (Borissiak 1932, Baryshnikov 2007) and similar to the MK bear in proportions. Their zygomatic width is large, similar to the sample from MK (Text-fig. 2). On P 4 the protocone is in a central position between the labial cusps, but the tooth is to some extent more evolved than in U. deningeri, as it also seems to be in the MK sample. The lower teeth morphology are also similar: p 4 has a well-developed paraconid and metaconid, and many additional cusplets; m 1 is archaic with a very weak constriction only at the middle part. The postcranials are smaller, about the size of the female specimens from MK (Text-figs 8 – 10). Some small differences could however be seen: the upper cheek teeth row is shorter (Text-fig. 3). The mandibles have a slightly shorter depth than in the MK sample but the sample of U. sa. rossicus is very limited in number (Text-fig. 6). Comparison with U. savini ANDREWS, 1922 from Bacton Wagner and Čermák (2012) include in U. savini only the specimens from Bacton s. str. (Forest Bed locality). The exact provenance of the only known skull is unclear. The size and proportions of the MK sample compared to the Bacton mandibular material are very similar (Text-fig. 10), while some differences exist in the tooth morphology. In the MK sample the p 4 morphology is more evolved to some extent, with a well-developed metaconid and paraconid and with many additional cusplets and swellings; m 1 is similar in morphology, but with a better developed enthypoconid (sensu Rabeder 1999) and with more cusplets on the talonid. The only known skull from Bacton is a larger size (maximal length is 432 mm) and has a relatively short cheek teeth row in relation to its size, but the absolute length of the cheek teeth row (length of 82.3 mm) (M. Pacher, pers. comm. in 2017) is comparable to that in the specimens from MK. It is difficult to say without more data if this skull belongs to a different species (U. deningeri (? )) or to U. savini from the type locality. It is noteworthy, however, that the specimen had a short muzzle (considering the photo kindly sent by A. Stewart) and strongly domed frontals, which is not diastema the at Height 130 150 170 190 c 1 - m 3 length Regression line 160 170 180 190 200 210 Palatal length typical for U. deningeri, especially at such an early stage in evolution of this species. Comparison with U. “ rossicus ” uralensis VERESHCHAGIN, 1973 from Kizel cave The small bear from Kizel cave (Ural, Russia) has several significant similarities in skull size and proportions to the MK bear, as well as to the typical U. sa. rossicus from Krasnodar (Text-figs 2, 3; see also: Vereshchagin and Baryshnikov 2000, Baryshnikov 2007). The metric comparison (Textfig. 11; for comparison see also the photos in Vereshchagin 1973: fig. 9) shows however, that U. sa. rossicus has a wider skull, and in particular the muzzle, and that in this respect U. uralensis is similar to some extend, in our opinion, to the large cave bears of the U. spelaeus-kanivetz / ingressus group. The bear from Kizel is rather young in geologic age (around 40 000 B. P.). After several opinions it seems to be genetically related more or less to U. ingressus (Pacher et al. 2009, Stiller et al. 2014), but new palaeogenomic data are in contradiction with this view (Barlow 2017).	en	Spassov, Nikolai, Hristova, Latinka, Ivanova, Stefanka, Georgiev, Ivan (2017): First Record Of The “ Small Cave Bear ” In Bulgaria And The Taxonomic Status Of Bears Of The Ursus Savini Andrews - Ursus Rossicus Borissiak Group. Fossil Imprint 73 (3 - 4): 275-291, DOI: 10.2478/if-2017-0015
