Ursus deningeri hercynicus Rode, 1935

Marciszak, Adrian, Kot, Małgorzata, Zarzecka-Szubińska, Katarzyna & Lipecki, Grzegorz, 2025, Ursidae (Carnivora, Mammalia) from Tunel Wielki Cave (southern Poland), Comptes Rendus Palevol 24 (14), pp. 241-302 : 244-260

Ursus deningeri hercynicus Rode, 1935

REFERRED MATERIAL. — The material of U. deningeri , the most abundant carnivore species from TW is represented by almost all skeletal elements ( NISP 641 , MNI 26 ; left/right bones are given in brackets): 4 cranium fr., 5 maxilla fr. (4/1), 15 mandibles (8/7), 19 I1 (10/9), 26 I2 (8/18), 30 I3 (14/16), 19 C1 (11/8), P1 (0/1), 15 P4 (9/6), 30 M1 (17/13), 44 M2 (22/22), 15 i1 (7/8), 26 i2 (15/11), 15 i3 (6/9), 25 c1 (12/13), 3 p1 (3/0), 13 p4 (5/8), 43 m 1 (17/26), 34 m 2 (16/18), 27 m 3 (15/12), scapula (1/0), humerus (0/1), 7 radii (3/4), 5 ulnae (3/2), 6 tibiae (3/3), 2 fibulae (2/0), sternum (1), 4 patellae (2/2), 4 naviculares (4/0), 6 calcanei (0/6), 4 tali (0/4), 2 capitatums (1/1), 2 cuboides (0/2), 3 ectocuneiformes (1/2), 4 pisiformes (3/1), 2 scapholunares (0/2), 3 mc 1 (1/2), 6 mc 2 (1/5), 9 mc 3 (3/6), 7 mc 4 (3/4), 9 mc 5 (6/3), 7 mt 1 (5/2), 5 mt 2 (3/2), 6 mt 3 (3/3), 13 mt 4 (5/8), 7 mt 5 (5/2), 2 mtpd, 5 atlas (5), 8 thoracic (8), cervical (1), lumbar (1), 2 caudale (2), 44 ph 1, 17 ph 2, 20 ph 3, and 2 sesamoides ( Appendix 1). Some bones hold the traces of chewing, gnawing and biting, and documented the interaction with other carnivores. The main factors responsible for the accumulation of bear remains on the site were probably carnivores, accidental falls, and water transport .

EMENDED DIAGNOSIS. — Smaller than the nominotypical subspecies Ursus deningeri deningeri ( von Reichenau, 1904) , often present P3, I3 with a calyx, i1 and i2 with larger and broader crowns and enlarged distoconid, P4 and p4 with morphologically more complicated crowns, M1 with a stronger developed parastyle and lingual cingulum, M2 with a narrower talon, m1 with a longer talonid and triple entoconid, m2 with a broader and longer talonid, and m3 with a shorter trigonid ( Rode 1935; Schütt 1968; Baryshnikov 2007).

DESCRIPTION

Upper incisors

Among 39 I2s from TW, 29 teeth belong to the most ancestral morphotype d with an asymmetrical, narrow crown without a lingual edge nor fossa lunaris ( Fig. 3). The other 10 I1-I2s represent the more advanced morphotype d/p,which is broader, with a weakly developed lingual part, but still without any cusplets on the mesial cingulum. The morphotype d predominates in the sites dated on MIS 19-13; it is characteristic for all I2s from Kozi Grzbiet, Mosbach 2, and Stránska skálá ( Table 1 ). The I2 from TW is metrically comparable to those from the sites dated on MIS 19-15, but with more robust crowns ( Fig. 3). The B/L index for TW is 81.5 (75.6-85.6, n = 10), and it is higher than MIS 19-15 group of U. deningeri (B/L = 77.9, 77.1-78.7, n = 9) ( Table 1 ). The bear crowns of younger populations (MIS 11-9) are larger and broader (B/L = 82.7, 77.8-91.9, n = 15), the ratio of the morphotypes p to the d is higher, and the calyx size increasing ( Table 1 ).

All I3s from the group 1 (MIS 19-13) represent the most ancestral morphotype 0, without calyx ( Table 1 ). Only 2 I3s from Hundsheim hold a very delicate structure resembling a minute calyx. Among 26 I3s from TW, seven specimens represent the more evolved morphotype 1, with a minute to small calyx located on the buccal side, above the apex ( Fig. 4). The rest 21 specimens of I3 from TW were assigned to the morphotype 0 ( Fig. 4). The I3 from TW has a developed lingual edge and fossa lunaris that occurs as a clearly visible pit on the mesial side of the tooth. The cingulum is weak and even absent in some of the specimens ( Fig. 4). In all I3s from TW, the mesial and distal edges delimit a somewhat recessed field with the lingual cingulum. The I3s from TW are larger than those from MIS 19-13, but smaller than I3s from MIS 11-9. A general tendency to increasing in size and broadening of all three upper incisors is observed ( Table 1 ).

Lower incisors (i1-i3) and canines (C1/c1)

Among 14 i1s from TW, eight specimens represent the most ancestral morph. A, simply build and small, while five others were assigned to the morph. C ( Table 2 ). The latter have larger and broader crowns, with an enlarged distoconid, and thickened mesial and mesio-buccal margins. The moph. A is the only occurring in the group 1 ( MIS 19-13). The i1 from TW is large and robust, and its size exceeds the dimensions of the i1 from most of the sites the material from which was used for comparison ( Table 2 ). The relatively large crown of the i2 is divided into two (in most specimens) or three (in a few teeth) parts, and its apex is oriented vertically and slightly distally, being oval-shaped in occlusal view ( Fig. 5; Table 2 ).

Among 25 i2s from TW, 10 represent the most ancestral morph. d, with a simply build crown, divided into two parts ( Table 2 ). The other 10 i2s from TW belong to the morph. d/s, with a weakly developed mesioconid, smooth distal surface, and without any cusplets. Finally, two specimens were assigned to the tricuspid morph. s, with a large mesioconid and a double distoconid ( Fig. 5; Table 2 ). The i2 from TW represents a higher evolutionary level as compared to the group 1 ( MIS 19-13). Among six i2s from Kozi Grzbiet, five specimens represent the morph. d, while only a single – d/s. In Hundsheim (n = 5) and Stránska skálá (n = 4), the morph. d/s is represented only by a single tooth. The i2 from the group 1 ( MIS 19-13) is also slightly smaller than that from TW, which in turn is smaller than i2 from the group 2 ( MIS 11-9). In the group 2, the number of more evolved morphotypes (d/s and s) is higher than that in TW ( Table 2 ). Among 15 specimens of i3 from TW, nine were assigned to the morph. B, a double-crowned, well-developed distoconid distinct from the protoconid, and a well-developed sulcus medialis, while six others belong to B/C ( Fig. 6). The i3 from the group 1 ( MIS 19-13) is smaller and evolutionarily more ancestral, although a high variability is inherent to this group ( Table 2 ).

Among 37 canines from TW, majority of the specimens is more or less worn, and therefore it is not possible to take their precise measurements. Among them, large and robust male canines are easily distinguishable from the smaller and narrower female canines. In addition, the lower canines are moderately curved distally and weakly flattened bucco-lingually, while the upper canines are straighter. All canines are characterised by an elongated and massive root, oval-shaped in cross-section. The crown is quite short (as compared to the root), blunt, with a poorly developed, very thin inner enamel crest running through its entire length.

P4

Among 15 P4s from TW, eight specimens represent the ancestral morph. A, with a simple morphology, without any cusplets such as protoloph and metaconule ( Fig. 7; Table 3 ). Seven other P4s from TW are more evolved, with a distinct metaloph or a small metaconule ( Fig. 7; Table 3 ). Comparison of P4s from TW with those from other Middle Pleistocene bear palaeopopulations showed that they do not differ metrically ( Table 3 ), but the TW material represents a slightly higher evolutionary level. Among the P4s from the group 1 ( MIS 19-13), teeth with a small metaloph or metaconule can be found, but the morph. A predominates in the sample. Among nine P 4 specimens from Kozi Grzbiet, only a single tooth represents A/B, while the others – A ( Table 3 ).

p4

The morphology of p4 from TW is highly variable, and metrically most of them are quite small ( Fig. 8; Table 4 ). Four specimens from TW belong to the ovoid morphotype B2, with a prominent protoconid, moderate metaconid edge running distally, conical paraconid located mesio-lingually and a small but distinct hypoconid shifted disto-lingually. Two other p4s from TW represent the morphotype D2, with a well-developed paraconid and metaconid of variable size, and a small cusplet between them. One specimen was recognised as representing the morphotype D3, with a small entoconid ( Fig. 8; Table 4 ). Two p4s from TW belong to C2, similar to B2, but with a small metaconid and a strong hypoconid shifted mesio-lingually. Three others were assigned to C3, with a small entoconid located between the paraconid and the hypoconid ( Fig. 8; Table 4 ).

Ancestral morphotypes of p4 (B1 and C1) are the most common in the group 1 ( MIS 19-13), where also a very ancestral p4 is recorded ( Table 4 ). These single-cusped teeth without para- and metaconid closely resemble the morph. b1, typical for arctoid bears. Metrically, p4s from TW do not differ from those in bear palaeopopulations of both groups, also the width to the length ratio (B/L) does not show any particular differences. Only the material from Biśnik Cave shows slightly higher dimensions ( Table 4 ). The p4 from TW is already not so ancestral as those from the group 1 ( MIS 19-13), but it cannot by any means be considered to be advanced. The morphodynamic index of P4/p4 shows that the TW material is grouped alightly above the populations from the group 1 ( MIS 19-13) ( Fig. 9). Simultaneously, the TW index is clearly lower than that of the group 2 ( MIS 11-9) and U. deningeri from TW holds an intermediate position between both groups ( Fig. 9).

M1

Morphologically, the M1 from TW shows the predominance of ancestral features, closely resembling those from the group 1 ( MIS 19-13), although with a number of more progressive characteristics ( Fig. 10; Table 5 ). Among 21 M1s from TW, nine teeth have a smooth internal slope of the paracone, (morph. 0), while the next 12 specimens have a moderately thick edge running from the paracone apex to its base, and accompanying with one or two smaller thin pillars reaching the half of its height (morph. 1) ( Table 5 ). Internal slope of the metacone is smooth in 10 of 24 teeth (morph. 0), while 14 others represent the morph. 1 ( Fig. 10; Table 5 ). The small metastyle is shallow and elongated, without any distal edge and is strongly associated with the distal cingulum. The mesocone is undivided. The shallow and small metastyle is weakly developed, with a smooth internal surface. The protocone is a low rectangular cusp, on the internal wall of which there are a few thin and sharp pillars and ribs. In a few M1s from TW, these pillars and ribs occur also on the boundary between the protocone and the mesocone, and in one tooth they are present even on the internal slope of the mesocone. In this respect, they represent various morph. B (2 B1, 9 B2, 6 B3) and one specimen morph. C1. One to three small cusplets, accompanying with the delicate and thin enamel, are mostly situated on the talon surface. Most of them represent the morph. B (19 among 23), while the talon field in four specimens is almost smooth, with a few weak pillars. The talon field is also separated from the weakly developed distal cingulum. Cingulum crest around the crown is almost absent. The lingual cingulum in nine M1s represents the morphotype 1.5, and is situated mostly on the lingual slope of the protocone. In the other nine specimens representing the morph. 2, the lingual cingulum collared the lingual slope of the protocone and terminated in the indentation between the mesocone and hypocone ( Fig. 10; Table 5 ).

Metrically, the M1s from TW are moderately large, being slightly larger than those from the group 1 ( MIS 19-13) ( Table 5 ). They are morphologically similar, although the features mentioned above are stronger developed. Among others, the parastyle in most M1s from TW is larger and more strongly separated from the paracone. The lingual cingulum is more strongly developed. In half of the M1s from TW, the lingual cingulum collared the entire protocone (morph. 1.5), while in the other half it starts from the mesial wall of the protocone and reaches the bounduary between the mesocone and the hypocone. In the M1s from the group 1 ( MIS 19-13), the lingual cingulum is weaker, and represents morph. 1, more strongly developed only on the mesocone and protocone base. Morphotypes 1.5 and 2 also occur, but they are much rarer than in the TW material ( Table 5 ).

M2

The mesial ridge of the M2 paracone in eight of 29 teeth is in contact with the mesial ridge of the protocone, forming a smooth junction (A3). The next 11 teeth represent the morph. B (5 B1, 5 B2, 1 B3), with an additional branch running on the mesio-lingual slope of the parastyle and mesial ridge of the parastyle, that formed a group of 4-5 cusplets like structures that collared the mesial wall of the trigon ( Fig. 11; Table 6 ). The 26 from 28 M2s yield two or three pillars on the internal wall of the paracone, which started from the base and reaching the half of its height (B). In two M 2 specimens, the internal wall of the paracone is smooth, without any pillars and ribs ( Fig. 11; Table 6 ). Among 27 M2s from TW, 13 specimens represent the morphotype A (2 A, 7 A1 and 4 A2), where the mesostyle complex forms a double arch ( Fig. 11; Table 6 ). The next 13 teeth belong to the morphotype B (8 B, 4 B1, B2), with the distal arm of the paracone curved considerably centrally, where it loses connection with the mesial arm of the metacone and forms a separate arch. The metaloph complex is less variable (single A/B), the 26 other specimens belong to B. Among them, there are seven teeth representing the morphotype B2, with a few (5-7) small cusplets in disordered forms located between the protocone and the metacone, and the metaloph complex is pushed more distally. The next four specimens represent B1, where a few (5-7) cusplets running in quite order from the metacone to the hypocone. Finally, five teeth belong to the morphotype B3, where the metaloph complex is located between the metacone and the protocone.

All M2s (n = 29) have the posteroloph complex uniform in shape (morph. 1), where between the metastyle and hypocone, there are a few (4-6) small cusplets, which do not form any regular structure. The talon field is moderately developed, with 15 M2s from TW representing A/B, where the surface is partially smooth and partially covered by a few (5-8) small cusplets. The next 16 teeth belong to B, where the entire talon field is covered with numerous small cusplets. The talon field is collared by a thick wall of the distal cingulum, which forms numerous small cusplets, and, except one tooth, all the M2s from TW represent B. Among 25 M2s from TW, metastyle and posthypocone in 23 are present but weakly developed into a small and low cusp. The metastyle and posthypocone are absent in three other specimens. Most of M2s from TW (n = 23) have the internal wall of the protocone with a few, usually 3-4, pillars and ribs starting from the internal base of the protocone but not reaching the apex.

Only three M2s have a single, similarly developed pillar. The lingual cingulum shows a low evolutionary stage, with most teeth (24 of 32 specimens) representing the morph. 1. It is characterised by the cingulum running through the base of the protocone and terminating between the protocone and the hypocone. In eight other M 2 specimens, the lingual cingulum is better developed and ends near the top of the hypocone (1.5) ( Fig. 11; Table 6 ). Metrically, the M2 from TW is moderately large, and in average it is slightly larger than those from Koněprusy C 178 or Mosbach 2, but smaller than M2 from Hundsheim ( Table 6 ). Any particular differences between populations have not been found. Only the distal to the mesial breadth ratio shows a tendency to decreasing, with the narrowing of the distal part of the crown. In this index, the M2 from TW is closer to the group 2 ( MIS 11-9), than those from the group 1 ( MIS 19-13) ( Table 6 ).

m1

The main cusps of the m1 from TW are connected to each other by a crest, which follows the outline of the tooth. In occlusal view, the tooth is elongated, and has moderately expanded talonid. The mesial margin of the trigonid is rounded, the lingual one is straight, while the buccal margin is almost straight, with a gentle concavity at the level of the protoconid. The broader talonid has a straight lingual margin, strongly convex buccal and rounded or blunt distal margins. The trigonid is moderately broad and long and clearly distinguishing from the wide and long talonid. The apex of the triangular and low paraconid inclines slightly mesially. Situated behind, rectangular protoconid is the highest and largest cusp, with almost with the vertical axe of the cusp almost perpendicular to the tooth occlusal plane. Both main cusps are separated by a wide, V-shaped valley. A relatively high, elongated and rectangular metaconid is situated after the metastylid. The lingual half of the talonid is occupied by entoconid, square-shaped to rectangular and relatively low cusp, with well developed entoconid 2. In TW, 15 of 23 m 1 have a single metastylid (1), rather it is double or triple (2, n = 7) ( Fig. 12; Table 7 ). Among 28 m 1 from TW, the entoconid is sometimes double (A2, n = 7), and the most often triple (A3, n = 19), and rarely quadruple (A4, n = 4) ( Table 7 ). On the internal slopes of the entoconids occur a few moderately developed ribs and pillars, reached half of the entoconid height (B in 22 of 23 teeth). Only in one tooth pillars and ribs are thicker and stronger developed (C). The entyphoconid is present in all 23 m 1, but in 13 it is weakly developed and not separated from the hypoconid (A/B). In the next 10 m 1, well-developed entyphoconid is separated from the hypoconid by a thick ridge (B).

The talonid cusps are closely situated, and in five of 24 m 1 from TW its distal margin is very compact and narrow, like that in U. arctos arctos , without any entyphoconid or hypoconulid. However, a slightly broader morphotype 2, with entyphoconid and small hypoconulid dominated (n = 19) ( Fig.12; Table 7 ). Metrically, the m1 from TW does not differ from the group 1 ( MIS 19-13). Slow increasing in size is observed since MIS 11 ( Table 7 ). There are no particular differences in breadth proportion between different populations. The m1 from TW has proportionally slightly longer talonid, and the L ta/L tr index is closer to the group 2 ( MIS 11-9) than to the group 1 ( MIS 19-13). The trend to shortening trigonid with simultaneous lengthening of the talonid starts to be well visible since MIS 11, and it is well recognised in the m1 from Biśnik Cave ( Table 7 ).

m2

The m2s from TW strongly vary in size, but they are within the range of variation of U. deningeri ( Fig. 13; Table 8 ). The mesial and distal margins are gently rounded or blunt, the lingual margin is straight, moderately concave in the middle part, in the transition between the trigonid and the talonid. The buccal margin of the trigonid is slightly rounded, while that of the talonid is moderately expanded and rounded. In some teeth, the lingual margin is almost straight ( Fig. 13). Occlusal surface is moderately complicated, less than in U. spelaeus , but definitely stronger than in the group 1 ( MIS 19-13). Among 25 m 2s, nine specimens have a simple metalophid complex developed as a straight, thick edge (morph. A). The dominant is B (n = 16), with the entprotoconid located on the internal slope of the protoconid. The internal field of the trigonid in eight m 2 specimens holds only a few weak furrows and pillars (morph. 1). Among them, the dominant is the morphotype 2 (n = 17), with one to three small but prominent cusplets and one or two thick furrows and pillars.

Most of the m2s from TW (24 of 26) have the metastylid complicated by the presence of small cusplets. As was showed by Rabeder (1999), the presence of smaller cusplets (usually 3 mesial, 1 paterial and 2 distal) is more ancestral state, and the occurrence of one large cusp is a more derived one. Only in two m2s, the metastylid is double, with two larger cusplets (mesial and distal). The mesolophid complex is moderately developed, still ancestral, with dominant morphotype B (22 of 24 teeth). In two specimens, the mesolophid is developed as a short and moderately thick ridge, not connected with the distal slope of the protoconid (A). The enthypoconid is variably developed, with the dominant morphotype B (20 among 27), where it is prominent, thick, triple or quadruple, separated from the hypoconid by a wide groove. The four m 2 specimens represent the morph. A/B, with the moderately developed enthypoconid not separated from the surrounded hypoconid. The other three teeth represent the morph. B/C, where a very strongly developed enthypoconid is still not divided. In half of the m 2 specimens from TW (16 of 30), the hypoconulid is absent, while it is well marked in 14 other teeth, being situated on the internal slope of the hypoconid ( Fig. 13; Table 8 ).

The m2s from TW differ from those of the group 1 ( MIS 19-13) in a more complicated occlusal surface, more strongly developed metalophid and mesolophid complexes, and in a broader inner talonid field. Dimensions of m2s and their particular indexes do not differ among bear palaeopopulations recorded from different sites ( Table 8 ). The size of the m2 is quite variable, and the TW palaeopopulation is characterised by a small size of the m2. The mean length of the m2 from Hundsheim (28.66 mm), Koněprusy C 178 (29.56 mm), Kozi Grzbiet (29.27 mm) and Stránska skálá (28.35 mm) is higher than that of TW (27.56 mm), while only that from Mosbach 2 (26.97 mm) has a smaller value. Increasing in size is observed in the specimens which come from the layers 19ad-19 of Biśnik Cave ( MIS 10-9) and later. It was also found a tendency to shortening and narrowing the trigonid and elongating and broadening the talonid. In all these ratios, the m2 from TW holds an intermediate position between the group 1 ( MIS 19-13) and the group 2 ( MIS 11-9) ( Table 8 ).

m3

In outline of the m3 from TW, the morphotype B predominates (20 of 22); it has an irregular shape, broad and expanded trigonid, long and wide talonid, and distinctly marked concavity on the buccal side of the boundary between the trigonid and talonid ( Fig. 14; Table 9 ). Two others represent a more evolved morphotype C, oval-shaped, with less marked buccal concavity and sharper distal margin of the crown. The morphology of the protoconid complex is quite variable, and among 20 m 3s from TW the most common are morphotypes B (1 B2, 1 B3, 7 B4, and 6 B5), a strongly developed mesolophid of which is connected with the protoconid. The next four teeth have anmesolophid (A4) extending disto-lingually. Most of the m3s (n = 15) hold one small but prominent cusplet on the internal wall of the metaconid (morph. B). Five other specimens have two small cusplets, and one tooth also has a thin pillar (morph. C). All m3s from TW hold still quite simple, but already present hypoconid, from which running 1-3 (B2, n = 9) or 4-6 thin (B3, n = 10) short ridges and pillars. In most of the m3s (22 of 24), the centrolophid is present as a series of small and low cusplets running disto-buccally from the metaconid base to the buccal slope of this cusp (B) ( Table 9 ). In two m3s, the centrolophid is similarly developed, but additionally it is connected with the mesolophid by a thin and sharp ridge (C). The entoconid is always present, usually (19 of 24) occurring in a series of quite large and low cusplets (4-7), arranged one behind the other (B). In five others, the entoconid is similarly developed, but 1-2 among these cusplets are enlarged (C). The stage of development of the talon field is variable, and in nine teeth this structure is quite smooth, sometimes the are one or two gentle pillars (B). In most specimens (n = 14), the talonid field holds one to three moderately large cusplets (C), while this structure in a single m3 is more complicated and holds six additional cusplets (D) ( Fig. 14; Table 9 ).

The m3 from TW is moderately large and broad and matches in this respect other Middle Pleistocene populations. No particular differences between the m3 from TW and those from the group 1 ( MIS 19-13) and the group 2 ( MIS 11-9) have been found ( Table 9 ). A tendency to the lengthening of the talonid is observed in the course of time. In the trigonid length to the total length (L tr/L m3) index, the m3 from TW matches the value of the group 2 ( MIS 11-9), and the trigonid is shorter than that in the group 1 ( MIS 19-13) ( Table 9 ).

POSTCRANIAL MATERIAL

Long bones of U. deningeri from TW are numerous but preserved only as fragments and do not give any reliable information. Similarly, differential carpals and tarsals are mostly damaged and represent moderately large and quite robust animals. Among 73 metapodials, only 24 specimens are complete enough to take most of the measurements, including the total length ( Table 10 ). Even more numerous phalanges, with the total of 75 (41 ph 1, 16 ph 2 and 18 ph 3), mostly complete, belong, however, to not fully immature animals, and their biochronological or taxonomical validity is restricted ( Table 10 ). Phalanges of fully adult specimens, similarly to long bones, also represent large and robust animals. Among elements of the postcranial skeleton, the most valuable information came from metapodials, which were also commonly found in TW.

Metrically, the metapodials from TW resembled those from the group 1 ( MIS 19-13) and the group 2 ( MIS 11-9). Their size falls within the size variability of equal bones, while K-index and P-index are variable. Sometimes, these indexes for the TW material are higher, smaller or comparable, therefore it is not possible to find one general trend ( Table 10 ). A more in-depth statistical analysis was omitted because of the small number of metapodial bones. Concluding analysis of the postcranial material, because of the incompleteness of most of their elements, and not enough metapodials, the results obtained have a limited value. No substantial differences between the TW material and that from both groups were found in morphology of the postcranial bones. The variability of particular features is so high that it is impossible to establish any characteristics in this matter and should rather be treated as a case of intraspecific variability.

GENERAL REMARKS

All analyses confirmed the assignment of the described material to Ursus deningeri and showed that it resembles that recovered from the other Middle Pleistocene localities dated at MIS 19-13 like Hundsheim, Koněprusy C 178, Kozi Grzbiet, Mosbach 2, and Stránska skálá. However, we found also a set of features, which suggested a higher evolutionary level of these bones and teeth supports a younger age for TW bears, closer to MIS 13-12. Most of these characteristics were found during the study of the teeth, while postcranial bones showed a rather limited biochronological and taxonomical value.Among the most important features, there are the higher proportion of evolutionarily more advanced morphotypes, I3 with a calyx, i1 and i2 with larger and broader crowns and enlarged distoconid, P4 and p4 with morphologically more complicated crowns, M1 with a stronger developed parastyle and lingual cingulum, M2 with a narrower talon, m1 with a longer talonid, m2 with a proportionally narrower and shorter trigonid and a broader and longer talonid, and m3 with a shorter trigonid.

There is also another method to check the taxonomic position of the particular bear palaeopopulation as was shown on the material from the Early Pleistocene Austrian site Deutsch Altenburg ( Rabeder et al. 2010). This grouping is based on the functional morphology attempt on two main components. The first is the strong enlargement of the cheek teeth and increased plumpness of the metapodials in spelaeoid bears. The second trend is the remaining of the ancestral cheek teeth and elongation and narrowing of the extremities in arctoid bears ( Rabeder et al. 2010). The relation between the dentition and extremities can be shown best by means of the total lengths of the teeth and metapodial bones. In order to include also palaeopopulations with a low number of individuals, like in the case of metapodials from TW, the arithmetic means of all measured lengths of all cheek teeth used in this study (P4/P4, M1/m1, M2/m2 and m3) were correlated to those of the total length of all measurable metapodial bones. For this purpose, all values had to be standardised, and the means of U. s. ingressus from Gamssulzenhöhle served as a such a standard. As expected, the TW bear group is placed together with other deningeroid and spelaeoid bears on a scatter-plot ( Fig. 15).

Tables

TABLE 1. — Measurements and morphological indices of the upper incisors of Ursus deningeri Richenau, 1904, Ursus spelaeus deningeroides Mottl, 1964, and Ursus spelaeus ingressus (Rabeder,Hofreiter,Nagel & Withalm,2004) from GamssulzenhÖhle (GS) as a standard.Abbreviations:JB, Biśnik Cave;KZ, Kozi Grzbiet; RP, RepolusthÖhle; SS, Stránska skálá; TW, Tunel Wielki Cave; V2, Vértesszőlős 2.

		I1-I2 L			I1-I2 B			Fossa lunaris/cingulum cusp index
	MIS	M	Min-Max	N	M	Min-Max	N	M
KZ	17	8.9	7.3-11.2	16	8.3	6.5-9.6	16	3.2/4.3
SS	19-17	9.1	7.9-11.6	9	8.6	7.6-9.3	9	11.1/11.1
TW	13-12	9.5	8.4-12.4	39	8.9	6.9-10.9	39	12.8/20.5
JB	10-8	9.7	7.9-12.3	43	10.1	8.5-11.5	43	16.3/26.7
RP	8-7	8.8	7.1-10.5	74	10.1	8.8-12.3	74	19.7/36.4
GS-S	3	10.0	8.2-12.3	59	11.4	9.5-13.6	59	150.9/168.8
		I3 L				I3 B
	MIS	M	Min-Max	N	M	Min-Max	N	Calyx index
KZ	17	12.6	10.9-13.9	6	9.9	8.9-10.3	6	0
SS	19-17	13.1	12.2-15.3	17	11.7	11.2-12.7	17	14.7
TW	13-12	13.3	10.7-14.6	26	11.1	9.9-12.2	26	30.8
V2	11	13.2	11.5-15.3	9	11.4	10.3-12.5	9	38.9
JB	10-8	13.7	11.8-16.5	24	12.0	10.5-14.3	24	43.4
RP	8-7	13.6	–	30	–	–	–	50.0
GS-S	3	13.9	–	46	–	–	–	100.0

TABLE 3. — Measurements and morphological indices of P4 of Ursus deningeri von Reichenau, 1904, Ursus spelaeus deningeroides Mottl, 1964, and Ursus spelaeus ingressus (Rabeder,Hofreiter,Nagel & Withalm,2004) from GamssulzenhÖhle (GS) as a standard. Abbreviations: HU, Hunas; JB, Biśnik Cave; KZ, Kozi Grzbiet; M2, Mosbach 2; RP, RepolusthÖhle; SS, Stránska skálá; TW, Tunel Wielki Cave; V2, Vértesszőlős 2.

	Age		P4 L		P4 Bp		Bp/L	Index	Stand.
	MIS	M	Min-Max	M	Min-Max	M	Min-Max	M	M	N
KZ	17	18.3	16.5-19.8	12.4	11.1-13.7	67.6	63.1-73.2	5.6	2.2	9
SS	17	18.9	16.7-20.8	13.8	12.3-15.5	72.1	65.1-75.5	14.3	5.6	7
M2	15-13	18.6	16.3-21.2	13.9	12.4-16.9	74.2	65.3-83.9	19.2	7.5	13
TW	13-12	18.3	15.5-19.9	13.0	11.2-14.2	71.3	65.2-80.7	23.3	9.1	15
V2	11	18.4	16.5-20.5	12.9	11.2-14.2	70.3	66.6-75.9	31.8	12.4	11
JB	10-8	18.4	15.9-20.2	12.8	10.7-14.2	70.7	63.8-77.8	31.3	12.3	40
RP	8-7	–	–	–	–	–	–	32.4	12.7	105
HU	7	–	–	–	–	–	–	39.9	15.6	56
GS	3	–	–	–	–	–	–	255.7	100.0	123

TABLE 4. — Measurements and morphological indices of the p4 of Ursus deningeri von Reichenau, 1904, Ursus spelaeus deningeroides Mottl, 1964, and Ursus spelaeus ingressus (Rabeder, Hofreiter, Nagel & Withalm, 2004) from GamssulzenhÖhle (GS) as a standard. Abbreviations: HU, Hunas; JB, Biśnik Cave; KZ, Kozi Grzbiet; RP, RepolusthÖhle; SS, Stránska skálá; TW, Tunel Wielki Cave; V2, Vértesszőlős 2.

	Age		L		B		B/L	Index	Stand.		P4/p4 stand.
	MIS	M	Min-Max	M	Min-Max	M	Min-Max	M	M	N	M
KZ	17	14.4	13.3-15.7	9.2	8.4-9.9	64.3	59.3-69.8	10.6	5.4	9	–
SS	17	14.7	13.4-16.7	7.7	6.5-8.2	63.7	46.4-69.1	18.9	9.5	5	–
M2	15-13	14.9	12.4-17.1	10.2	8.1-12.5	68.6	53.6-83.3	21.9	11.1	31	–
TW	13-12	14.1	12.3-16.3	8.8	7.4-10.1	62.6	57.1-69.3	22.9	11.6	12	20.8
V2	11	14.3	12.2-16.6	8.9	7.9-11.2	63.7	57.9-70.5	45.9	23.2	8	–
JB	10-8	14.5	11.8-17.9	9.5	8.1-11.6	66.1	56.9-76.9	62.1	31.3	44	–
RP	8-7	–	–	–	–	–	–	64.1	32.3	71	20.2
HU	7	–	–	–	–	–	–	74.4	37.5	45	24.2
GS	3	–	–	–	–	–	–	198.2	100.0	97	100.0

TABLE 7. — Measurements and morphological indices of m1 of Ursus deningeri von Reichenau, 1904. Abbreviations: C 718, Koněprusy C 178; HH, Hundsheim; JB, Biśnik Cave; KP, Koněprusy; M2, Mosbach 2; TW, Tunel Wielki Cave; ZH, Za Hájovnou Cave.

	Age		1			2			3			6			7
	MIS	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N
C 718	17	27.8	24.5-31.5	24	17.4	16.4-19.3	23	10.3	8.9-12.0	23	10.2	8.6-11.8	24	12.9	11.5-14.9	25
KP	17	27.0	24.2-31.3	20	17.0	15.2-19.5	21	10.1	8.9-11.7	27	10.0	8.8-11.4	23	12.5	11.3-13.7	26
M2	15-13	27.4	25.3-31.2	19	17.4	16.6-18.8	9	9.9	8.9-10.7	9	10.2	8.6-11.8	24	12.9	11.5-14.9	25
HH	15-13	27.1	24.9-30.1	7	17.7	16.2-19.2	5	10.3	9.4-11.6	7	10.7	9.8-11.4	7	12.8	11.6-14.0	9
TW	13-12	27.5	25.1-30.7	20	17.1	15.2-19.1	21	10.8	8.8-12.0	26	10.9	9.6-12.6	24	13.2	11.9-14.7	29
ZH	11	28.0	25.6-30.9	15	17.4	14.9-19.1	14	10.4	9.2-12.5	18	10.6	9.5-11.5	14	13.3	11.9-15.0	21
JB	10-8	28.4	25.4-32.6	39	16.7	15.6-19.7	39	12.8	10.7-14.8	39	11.5	9.7-13.7	39	13.6	11.6-14.6	39
	Age		2/1			3/2			6/1			7/1			6/7
	MIS	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N
C 718	17	62.5	56.9-66.9	23	59.2	54.3-62.2	23	36.6	33.8-43.4	24	46.6	44.4-52.2	24	85.4	74.7-95.9	24
KP	17	63.0	55.8-67.4	20	59.4	58.6-60.0	21	36.9	33.3-40.1	20	46.4	42.7-49.5	19	84.4	77.9-93.2	23
M2	15-13	62.4	57.1-66.4	19	61.7	58.7-66.9	9	37.7	34.1-40.5	19	48.0	44.8-54.1	18	83.6	81.7-94.6	18
HH	15-13	62.3	55.6-65.3	5	62.5	57.9-65.8	5	38.8	37.2-41.4	6	46.8	44.6-49.5	7	84.9	80.4-93.3	6
TW	13-12	62.1	57.3-64.9	19	64.6	58.8-74.7	19	39.9	36.1-43.9	24	48.3	44.7-50.5	25	82.8	73.1-95.8	24
ZH	11	62.1	58.2-64.6	14	65.1	58.5-72.5	14	37.8	34.7-40.9	13	48.0	45.9-50.7	15	80.0	73.4-85.6	13
JB	10-8	61.6	52.1-68.9	39	69.7	64.7-82.9	39	40.5	37.3-43.9	39	47.8	44.2-51.6	39	84.8	77.8-93.9	39

TABLE 8. — Measurements and morphological indices of m2 of Ursus deningeri von Reichenau, 1904. Abbreviations: C 718, Koněprusy C 178; HH, Hundsheim; JB, BiŚnik Cave; KP, Koněprusy; KZ, Kozi Grzbiet; M2, Mosbach 2; TW, Tunel Wielki Cave; ZH, Za Hájovnou Cave.

	Age	1			2			3			4			5			6			7
	MIS M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N
KZ	17 29.2	26.5-31.5	8	15.8	13.9-17.4	8	17.8	15.8-19.5	7	13.2	10.8-15.7	10	11.4	10.1-12.5	9	16.7	15.4-18.4	7	17.3	15.9-19.2	8
C 718 17 28.3		25.4-31.1	16	15.2	12.5-17.5	17	17.4	15.9-20.0	16	13.3	11.9-14.9	16	10.9	9.5-13.0	15	15.5	13.3-17.7	16	17.1	14.8-18.8	16
KP	17 29.5	24.9-32.8	15	15.7	13.5-17.3	15	18.4	15.7-20.7	15	13.8	10.5-15.8	16	11.1	9.1-12.1	17	16.0	14.7-17.9	17	17.2	16.2-18.5	15
M2	15-13 26.9	24.7-30.2	40	15.0	12.7-17.4	15	16.7	15.0-17.8	14	11.8	9.5-14.4	14	10.2	8.8-11.8	14	15.8	13.7-18.1	39	16.7	13.9-19.6	39
HH	15-13 28.6	26.1-32.1	8	15.0	13.3-17.7	9	17.9	15.5-20.0	6	13.4	11.7-15.0	8	10.9	9.5-12.1	6	16.1	14.6-17.7	9	17.2	15.8-18.9	8
TW	13-12 27.5	24.0-31.6	25	15.7	13.5-18.6	28	17.2	14.5-19.4	28	12.9	9.4-14.4	32	10.3	9.7-13.7	35	16.1	13.9-18.8	29	16.7	13.2-19.1	31
ZH	11 27.8	24.7-31.2	19	14.8	12.6-17.0	20	17.2	15.0-19.6	20	13.0	10.5-15.6	19	10.6	9.5-12.3	19	16.4	14.2-19.0	21	16.9	14.6-19.6	20
ZH	9 27.9	26.4-29.8	7	14.7	13.0-17.0	8	17.3	15.6-19.2	7	13.2	12.2-13.7	7	10.7	10.3-11.0	6	16.6	14.3-18.2	9	16.5	15.4-18.0	6
JB	10-8 29.1	26.1-35.5	41	17.0	15.3-19.9	41	17.3	15.5-20.9	41	12.1	10.6-15.6	41	11.9	10.3-14.6	41	17.2	14.2-19.2	43	17.4	14.7-20.3	47
	Age	2/1			4/1			4/2			5/3			6/1			7/1			6/7
	MIS M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N	M	Min-Max	N
KZ	17 58.2	50.2-60.2	8	40.8	39.6-42.4	8	61.9	59.6-73.5	8	60.5	58.5-64.1	8	56.9	51.1-63.9	7	59.3	54.9-66.7	8	97.1	94.4-100.1 8
C 718 17 53.4		48.3-58.5	16	42.1	40.5-47.9	16	62.4	58.6-66.3	16	62.6	59.7-65.1	16	54.7	50.2-62.8	16	60.1	57.2-67.4	15	96.8	91.7-99.8	15
KP	17 53.3	48.4-57.4	15	43.1	41.4-48.8	15	62.8	58.1-67.4	15	60.3	57.9-66.3	15	54.6	51.2-59.1	15	58.6	55.6-62.7	13	96.1	90.9-97.9	13
M2	15-13 56.6	50.6-63.1	14	44.8	40.9-49.8	14	63.9	57.5-69.4	14	61.1	58.7-66.2	14	59.2	53.4-65.5	38	62.4	54.1-69.8	38	96.2	87.5-97.1	38
HH	15-13 53.2	50.8-55.9	8	44.7	40.5-48.8	9	63.2	58.1-68.7	8	60.9	61.3-66.9	6	56.7	54.5-59.2	7	61.0	57.5-64.4	7	96.6	91.9-97.8	7
TW	13-12 56.7	52.1-63.4	25	46.4	36.4-49.7	25	65.2	59.8-69.5	28	64.8	49.8-68.9	28	57.9	51.7-65.4	25	60.2	47.4-68.3	25	93.2	90.9-99.1	29
ZH	11 53.1	48.5-60.2	19	46.9	37.5-45.6	7	66.8	60.6-68.4	7	64.6	57.3-66.1	6	59.3	53.2-62.9	18	60.9	54.7-67.7	18	93.6	90.2-97.7	20
ZH	9 52.3	47.5-58.1	7	47.8	36.9-46.1	7	67.3	62.5-69.1	7	67.8	57.3-66.2	6	58.3	54.2-61.2	7	58.8	55.0-61.9	5	93.9	89.5-95.6	6
JB	10-8 50.5	45.2-55.4	41	49.6	40.5-54.9	41	71.1	65.5-81.0	41	68.6	66.6-69.6	41	58.9	54.0-67.9	41	62.4	55.3-66.9	41	90.7	84.5-94.1	41