Ursus deningeri VON REICHENAU, 1904

Wagner, Jan, Jiangzuo, Qigao, Lenardić, Jadranka Mauch & Liu, Jinyi, 2017, Šandalja I (Croatia) And, Fossil Imprint 73 (3 - 4), pp. 533-544 : 537-538

Ursus deningeri VON REICHENAU, 1904

Text-figs 1e, f , 6a , 7a

1975 Ursus cf. deningeri REICHENAU ; Malez, pp. 186, 188, 189, 198.

1996 Ursus cf. deningeri ; Bosinski, p. 55.

2000 Ursus [of similar stage of evolution as bears from Vallonnet and Pirro]; Spassov, p. 108. (partim)

2002 Ursus [of similar stage of evolution as bears from Vallonnet and Pirro]; Spassov, p. 234. (partim)

2003 Ursus [of similar stage of evolution as bears from Vallonnet and Pirro]; Spassov, p. 214. (partim)

2011 Ursus sp. ; Kahlke et al., p. 1376. (partim)

2012 U. cf. deningeri ; Wagner et al., pp. 50, 51.

2015 transitional stage in Ursus etruscus-U. deningeri lineage; Vislobokova and Agadjanian, p. 655. (partim)

2016a U. etruscus-U. deningeri ; Vislobokova and Agadjanian, p. 195. (partim)

2016b U. cf. deningeri ; Vislobokova and Agadjanian, p. 196.

N o t e. As p4 sin. assigned here to U. deningeri , was originally determined by Malez (1975) as U. etruscus , all the references of U. etruscus should be repeated also in synonymy of this species; but we see it as somewhat redundant and we refer here to the synonymy of U. t. mediterraneus in this case.

M a t e r i a l. Mesial fragment of m2 dex. (specimen G;

Text-fig. 1e ) and p4 sin. (specimen H; Text-fig. 1f ).

D e s c r i p t i o n. The mesial fragment of m2 dex. (specimen G) is distally limited by breakage extending from the distal part of Prd-complex (in front of contact with Hyd) to the distal arm of End 2. The tooth is only slightly worn. There is a very well developed central constriction. The cusps are well developed and strong. EPrd and EMed are rather low, but clearly separated from Prd and Med. There are well developed secondary cusps and swellings in the anterior trigonid basin and well developed and diversified swellings and crests are also present in the distal part of the trigonid.

p4 sin. (specimen H) is almost complete (only the lingual crown base is damaged) and only slightly worn. Except the dominant Prd, Pad, Med, Hyd and End are well developed. With the exception of these main cusps, a smaller secondary cusp is developed between Pad and the mesial arm of Prd. The distal arm of Prd is segmented by 2 transverse grooves.

D i s c u s s i o n. The mesial fragment of m2 was already determined as Ursus cf. deningeri by Malez (1975). Although fragmentary, this tooth has the characters typical for spelaeoid bears, e.g., well developed central constriction, EPrd and EMed forming independent cusps or a highly diversified inner field in the tooth (i.e., not only the presence of crests in the distal part of the trigonid, but also their further divisions, partly forming by small cusps). The combination of these characters is typical for spelaeoid bears and is more advanced compared to U. etruscus or even U. arctos (cf. Wagner and Čermák 2012). The state of diversification of the inner field seems to be greater than in U. deningeri from the beginning of the Middle Pleistocene, i.e., from the Late Biharian ( Text-fig. 6 ). Although we assigned this tooth to U. deningeri , it ought to be noted that similar morphology can also be found in some specimens of Ursus gr. spelaeus. The p4 sin. was assigned by Malez (1975: 186) to U. etruscus , but the tooth is more evolved than in this species ( Text-fig. 7 ), where the cusps, other than Prd (and sometimes Pad), are only weakly developed, if not missing. A similar situation is in U. thibetanus , in which this tooth is usually even less developed. Even in the type specimen of U. “ karabach ” (= U. t. mediterraneus ; see Baryshnikov 2010) from Azykh (= Azokh) cave (Nagorno-Karabakh), which has larger and more robust lower teeth than is usual in Asiatic black bear, the p4 is much simpler (although Prd, Hyd and End are present, all of them are only very slightly developed; see Text-fig. 7e ). In U. arctos this tooth has a not so diversified occlusal surface (at least in most cases). Moreover, relatively mesially located Med is typical character for spelaeoid bears, distinguishing them from brown bears. On the other hand, compared to the Late Pleistocene U. gr. spelaeus, this tooth is rather simple in construction (Pad and Med are smaller). We therefore assigned it to U. deningeri .

Figures

Text-fig. 1. Dental material of ursids from the locality Šandalja I (Croatia) in occlusal view. U. t. mediterraneus – a: m2 dex. (specimen A), b: m2 dex., c: m3 dex. (both specimen B), d: distal fragment of m1 dex. (specimen C); U. deningeri – e: mesial fragment of m2 dex. (specimen G), f: p4 sin. (specimen H).

Text-fig. 6. m2 of U. deningeri from Šandalja I (a: specimen G; 1– occlusal, 2 – lingual view) compared with Late Biharian U. deningeri from C 718 cave (b: NM-R 9740, c: NM-Ra 129; both occlusal view). Specimens coated by ammonium chloride; a reversed.

Text-fig. 7. p4 of U. deningeri from Šandalja I compared with other bear species. a: Šandalja I (specimen H; 1 – occlusal, 2 – lingual view), b: U. etruscus, Casa Frata (private collection), c: U. etruscus, Olivola (IGF 4605), d: U. deningeri, C 718 cave (NM-Rv 20003), e: U. t. mediterraneus, Azykh cave (ZIN 32549) (all in occlussal view). Specimens coated by ammonium chloride; d reversed.