4 : ( ), F :AM Fig. 1 Skull of Indarctos zdanskyi sp. nov., holotype F :AM A. right lateral view ; B. frontal view ; C. ventral view

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1 VERTEBRATA PALASIATICA pp figs ) 1 2 ( ) ( ), Indarctos zdanskyi,, p4,p4 M1,m1, m2,, Q , (R. H. Tedford) Frick ( Indarctos) Childs Frick ( ) 1934, 46 - L Indarctos lagrelii I. sinensis, ( Can Llobateres Samos), ( ) Indarctos zdanskyi sp. nov. 1924? Hyaenarctos sp., Zdansky, 26 27, pl. 4 5 F :AM 22345,,, ; ( Zdansky, p , pl. 4 5),,IVPP, > 400mm, > 300mm,, ;, ;, ; p4 ;P4 M2 1/ 4 ; ; P4 M1, ;M2 ( ) ;m1 ;m2 1) ( : ) ( : KZCX22103) :

4 : 279 1 ( ), F :AM 22345 Fig. 1 Skull of Indarctos zdanskyi sp. nov.

2 4 : ( ), F :AM Fig. 1 Skull of Indarctos zdanskyi sp. nov., holotype F :AM A. right lateral view ; B. frontal view ; C. ventral view

280 41 (Otto Zdansky) ;, ; ; ; ( ) 150mm 83mm, 35mm ; ( ) 200mm ;,40mm (25mm) ; ; ; 11mm 11mm ( 1 A) ;, ;, 117mm ; 41.

3 (Otto Zdansky) ;, ; ; ; ( ) 150mm 83mm, 35mm ; ( ) 200mm ;,40mm (25mm) ; ; ; 11mm 11mm ( 1 A) ;, ;, 117mm ; 41. 5mm ; M2 ;, 42mm ;, ;, P4 ; C ( 1 B) ;, ; ;, ; ;,,, 115mm, 52mm ( 1 C ; 2) 80mm, 2 ( ), F :AM Fig. 2 Basicranium of Indarctos zdanskyi sp. nov., holotype F :AM b. bulla ; cf. condyloid foramen ; em. external meatus ;,, mp. mastoid process ; plf. posterior lacerate foramen ; pp. paroccipital process ; sf. stylomastoid foramen ; ss. suprameatal shelf ( 2 plf) 55mm, ( ) 60mm ; ( 2 cf), ( 2 pp), 14mm ; (7 8mm) (15mm) ; ( 2 sf), ; ( 2 mp),,, 33mm ( 2 b),,, ( 2 em), 12mm 25mm, ( 20mm), 46mm ; ( 2 ss) ; (120mm) (25mm),

4 4 : 281 M2 ;, 110mm 110mm M1, (, ), M2 ; M2, ( ) P4 ;, C 1/ 3, I2 42mm ; I3 6315mm ( 1 C) I2 3 I3,, P1 P3, I2, I3, ( ) I3 C 10mm C,, ( ) (P1 P2) P3, P1, P3, P2 P1 P3,,,, ;P1, P3 C P4 40mm P4,, ;M1 P4 ;M2, ( 3),,, m3, p4 13mm ; ; c, p4, p4 ;,, ; ; (shelf) ; ( 3 B) c, p1, p2, p3 ( ) ; p3 ( ) p4 ;m1, ; ;m2 ;m3 F :AM Indarctos Agriotherium, M2 ( Agriotherium M2 ) ( Agriotherium ),P4,M2, ( Agriotherium P4,M2, ),m1 ( Agriotherium ) Samos Indarctos atticus ( Helbing, 1932 ; Thenius, 1949,1959) ; Can Llobateres Indarctos vireti (Crusafont2Pairo and Kurt n, 1976) Indarctos vireti F :AM ( 1 2) ; ( ) ; P4 ;

282 41 3 ( ), F :AM 22345 Fig. 3 Mandibles of Indarctos zdanskyi sp. nov., holotype F :AM 22345 A. right lateral view ; B.

5 ( ), F :AM Fig. 3 Mandibles of Indarctos zdanskyi sp. nov., holotype F :AM A. right lateral view ; B. crwon view,m1, Indarctos vireti Indarctos atticus Dames 1883 m1 m2 m3, Pikermi,Dames Weithofer 1888, Dames : Hyaenarctos atticus ( Indarctos atticus) Samos,, Samos Indarctos atticus Thenius

6 4 : 283 (1949,1959),F :AM ( 1 2),,F :AM F :AM mm, Samos, 325mm ( 1),F :AM M2 110mm, P4 M2 (95mm) ; Samos, 82mm ( Thenius, 1959, Fig. 6 ), P4 M2 86mm,,F :AM F :AM 22345, p4,p4 M1 m2 Samos F :AM Indarctos atticus Indarctos Indarctos lagrelii (Zdansky, 1924) F :AM ( 1 2) ;P4 M1 ; p4 Indarctos sinensis (Zdansky, 1924 ; Li and Chi, 1964), ( ), p2 (Li and Chi, 1964, Pl., 1) F :AM ( 2), m2, sinensis Samos (Helbing, 1932), Indarctos sinensis Indarctos atticus 1 Table 1 Measurements and comparison of skulls of Indarctos (mm) Indarctos vireti (VP646) atticus lagrelii zdanskyi Crus. & Kurt.,1976 Zd., 1924 F : AM ( Greatest L) 465 (Basilar L) (Palatal L) (Rostral W) 108 (W at postorb. proc. ) 135 (W at mastoid proc. ) 200 (Condylar W) C M2 L 170 P1 (L W) 7. 1? P2 (L W) P3 (L W) P4 (L W) M1 (L W) M2 (L W) 3 (Heavily worn) ?; ; 12? ; ; ;

7 Maragheh Indarctos maraganus (de Mecquenem, 1925), F :AM p2 ; :, p4,,m2 ( Crusafont2Pairo and Kurt n, 1976, p. 13),Maragheh Indarctos atticus : Indarctos salmontanus Indarctos punjabiensis Pilgrim 1913, 1914,, M2 Indarctos punjabiensis Lydekker (1884), P4 M1 P4,, Pilgrim (1932) Agriotherium palaeindicum ( Ind. Mus. No. D 8, Lydekker, 1884, pl. 1), Indarctos,m1 I. punjabiensis,, p3,m1 m2 ( 2), 2 Table 2 Measurements and comparison of mandibles of Indarctos (mm) Indarctos vireti atticus punjabiensis lagrelii zdanskyi Can Llobateres V 2922 Pikermi Samos Maragheh Ind. Mus. D 8 Baode Baode Crus. & Kurt.,1976 Li & Chi, 1964 Weith.,1888 Crus. & Kurt., 1976 Lyd.,1884 Zdan.,1924 F :AM (Total L) (Ramus vert. H) p4 (H ant. p4) m2 (H under m2) p1 L p1 W 7 p2 L p2 W p3 L p3 W p4 L p4 W m1 L m1 W (trig. ) m1 W (tal. ) L/ W (tal. ) m2 L m2 W (trig. ) m2 W (tal. ) L/ W (trig. ) m3 L m3 W (Measured by the present authors according to plates or specimens)

8 4 : 285 Zdansky? Hyaenarctos sp. (Zdansky, 1924,p , pl. 4 5) (Andersson Lok. 13), M2 Indarctos M2 ( ) M2, F :AM 22345,, M2, ( Zdansky, 1924, Pl. 5) 115mm, P4 M2 (95mm) ; F :AM mm, 95mm ; Samos ( Thenius, 1959, fig. 6) 82mm 86mm, M2 P4 M2, (Zdansky, 1924, Pl. 4), F :AM 22345, P4 M1 P4 M1 Indarctos zdanskyi,zdansky Indarctos lagrelii Indarctos sinensis, Zdansky P4 (Zdansky, 1924, Pl. 6 7),? Hyaenarctos sp. P4, Agriotherium ( ) Childs Frick, Childs Frick Edward Pedersen Lorraine Meeker A NEW SPECIES OF INDARCTOS FROM BAODE, CHINA QIU Zhan2Xiang 1 R. H. Tedford 2 (1 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Science Beijing ) (2 American Museum of Natural History New York 10024) Key words Baode, Shanxi, late Miocene, Indarctos Summary While studying the Carnivora material from the Yushe area a few years ago, Tedford noticed a well preserved Indarctos skull from Baode, China, in the Frick Collection housed in the AMNH. The skull in association with its mandibles and atlas was sent to Frick by one of his agents, Mr. Liu Hsi Ku, in Qiu, while visiting to the AMNH in 2001, together with Tedford, studied this specimen and found that it belonged evidently to a new species of the genus Indarctos, widely diffe2 rent from the two known species in China. So far more or less complete skulls have been known only from two European localities ( Can Llobateres and Samos). No complete skull has ever been described in China. In view of its importance for our understanding of the Chinese Indarctos, it is described here.

9 Indarctos zdanskyi sp. nov. 1924? Hyaenarctos sp., Zdansky, 26 27, pl. 4 5 Holotype F :AM 22345, almost complete skull in association with its mandibles, from Hip2 parion beds at Zhaojiagou of Baode County, Shanxi Province, China, now housed in the AMNH. Referred specimen A very poorly preserved skull from Chengouwan of Xin an County, Henan Province, China, described by Zdansky in 1924 as? Hyaenarctos sp. Diagnosis Indarctos of large size : basilar length of skull > 400mm, and total length of mandible > 300mm. Skull with high cranium, sagittal crest high, plate2like ; mastoid process very robust, strongly extended laterally, surpassing the suprameatal crest which lies anterior to it ; external meatus very long, tubular, its length > the bulla width, choana very narrow and long. Mandible high and robust, posterior border of symphysis located slightly anterior to p4. Cheek teeth relatively small, P4 M1 shorter than 1/ 4 of the basilar length ; ante2carnassial premolars strongly reduced in size and number ; P4 longer than M1, with large parastyle and double2cusped protocone ; m1 m2 wide, talonid wider than trigonid in m1, and trigonid wider than talonid in m2. Etymology The species name is dedicated to Otto Zdansky, who already knew some particular features of this form in Brief description and comparison The occipital surface is triangular in form, with its upper angle very sharply pointed, the sagittal crest is plate2like, 35mm high. The mastoid process is extended downward and externally, widely exposed on the occipital surface. The paroccipital process is small, separated from the mastoid one by a wide and shallow depression. In lateral view (Fig. 1 A), the profile of the skull is rather straight. The sagittal crest starts at the level of the glenoid process. The zygomatic process of the temporal bone stretches anteriorly to the base of the postorbital process, 117mm long. The posterior border of the anterior end of the zygomatic arch lies at the middle of the M2. The infraorbital foramen lies above the posterior part of the P4. On ventral side ( Fig. 1 C ; Fig. 2), the basioccipital bone is very wide. The condyloid foramen lies 14mm posterior to the posterior lacerate foramen. The stylomastoid foramen lies lateral to the posterior lacerate foramen. The mastoid and paroccipital processes form a large depression on the ventral side. The external meatus is long, tube in form, its length is longer than the width of the bulla, ventral surface of which is almost flat, elevated only at its medial border. The postglenoid foramen lies at the medial border of the postglenoid process. No alisphenoid canal can be observed. The choana is very long (120mm) and narrow (25mm). The temporal fossa, seen from the ventral side, is very large (110mm 110mm). The posterior palatal foramen lies at the level of the middle of the M1. Of the incisors only right I2 I3 and left I3 are preserved. All the incisors and molars are heavily worn. The wear facet on the I2 is horizontal, whereas there are two wear facets on the I3 : one is horizontal, the other faces postero2laterally. Perhaps due to wear, no secondary cusplets or cingulum can be observed on the incisors. The I3 C diastema is 10mm long. C is almost vertically positioned, with two wear facets ( horizontal and antero2internal) and weak posterior crest. On the right side, there are two small oval alveoli (for P1 and P2) and a P3 ; on the left side there are P1 and two alveoli for the P3. P1 and P3 are of the same morphology, both small, low crowned, with a convex labial side, and flat lingual one. P1 slightly converges anteriorly. P3 is almost transversely positioned. The P4 has a large parastyle and large protocone, probably double2cusped. The M1 is worn to its root. But evidently, it is shorter than P4. The postero2lingual cingular shelf ( talon ) of M2 is not particularly large. The outline of the M2 crown is roughly triangular. The mandible is very robust ( Fig. 3). Its lower border is markedly convex, with marked chin anteriorly and a hook2like angular process posteriorly. The posterior border of the symphysis extends almost to the p4. There are three mental foramina on each side, the posterior2most one located under the posterior root of the p4.

10 4 : 287 All the lower incisors are absent. On the right side there is a large oval alveolus for the p1, a dumbbell2shaped alveolus for the p2 and a very small alveolus for the p3. On the left side there is only one alveolus for the p3 (with some remnant of root). The P4 is relatively small, more convex on lingual side. The m1 is a typical Indarctos carnassial, wider at its posterior half. The trigonid is wider than talonid in m2. Comparison and discussion Indarctos zdanskyi is distinguished from all the other known species of the genus by being larger in general size ( Tables 1 2). As far as the skull is concerned, its comparative study is very limited, since skulls are only known in 2 European species ( I. vireti and I. atticus ) among the Indarctos species. Indarctos zdanskyi can be clearly distinguished from these 2 species by the skull characters stated above in the diagnosis. As far as the teeth are concerned, the distinction between I. zdanskyi and the other known species is also clear. In I. lagrelii the premolars are much less reduced than in I. zdanskyi : the p4 is very large and both upper and lower 2 nd and 3 rd premolars are still large in size and two2rooted ; the P4 is shorter than M1. Indarctos sinensis is represented only by mandibles and lower teeth. The horizontal ramus of I. sinensis is low, almost as low as the anterior part of the ramus under the premolars (probably due to age), and its m1 m2 are less widened. In general the morphology of the lower teeth of I. sinensis is almost the same as those of the European I. atticus. They may well be conspecific. The mandible discovered at Maragheh in Iran and described as I. maraganus is also very large, and close to that of I. zdanskyi in size. However, the posterior border of the symphysis is situated anterior to the p3, all the premolars are less reduced, p4 remains very large, and the molars are not particularly widened ( Table 2). If sexual and individual variation in size is fairly large as Crusafont2Pairo and Kurt n (1976) thought, the Maragheh mandible may well be a large male of I. atticus. There are 2 species of Indarctos described from Indo2Pakistan : I. salmontanus and I. punjabiensis. The former is characterized by having enlarged talon and better developed secondary cusplets on its M2, and the latter by having a little developed parastyle and compressed protocone in the P4. Therefore, both are different from I. zdanskyi in tooth morpholgy. The mandible bearing the catalogue number Ind. Mus. No. D 8, which was first described by Lydekker ( 1884) as belonging to Hyaenarctos punjabiensis, but later transferred by Pilgrim ( 1932) to Agriotherium palaeoindicum, may well belong to Indarctos, in view of its lack of premasseteric fossa and the double2cusped entoconid in its m1. In size this mandible is very close to that of I. zdanskyi. However, the horizontal ramus is very low in this mandible, and the molars are narrow (Table 2). It seems inappropriate to refer them to one and the same species. It is interesting to note that the badly crushed skull from Andersson s Lok. 13 (Chengouwan, Xin an County, Henan) described by Zdansky (1924, p , pl. 4 5) as? Hyaenarctos sp. shows great similarity with F :AM In size they are comparable. The distance between the posterior border of M2 and the posterior border of the glenoid fossa in the Henan skull is 115mm (measured from the plate). This same distance in F :AM is 110mm, while it is only 82mm in the Samos skull of I. atticus (Thenius, 1959, fig. 6). The P4 M2 length is 95mm in both the Henan skull and F : AM 22345, while it is 86mm in the Samos skull, longer than the distance between M2 and glenoid fossa. In the Henan skull and F :AM the distance between the M2 and the glenoid process is far longer than the P4 M2 length. In the Henan skull the sagittal crest is also very high and plate2like, the P4 is longer than M1 and the talon of M1 is small. All this shows that the Henan skull is to be included in the new species, Indarctos zdanskyi. Acknowledgements It seems appropriate when describing a Baode fossil from the Frick Collection, to acknowledge the work of Liu Hsi Ku, who while employed by Childs Frick, worked diligently during the turbulent years of making a large collection of fossil mammals from

11 the classic Baode sites developed by Zdansky in the early 1920 s. Liu s collection resides in the American Museum of Natural History, New York, and represents the 2 nd most important collection (the 1 st being that in Uppsala) from Baode outside of the People s Republic of China. The material described herein was expertly prepared in 2001 by Edward Pedersen, and the photos illustrating it were made by Lorraine Meeker, both of the AMNH staff. References Crusafont2Pairo M, Kurt n B, Bears and bear2dogs from the Vallesian of the Vall s2pened s Basin, Spain. Acta Zool Fenn, 144 : 1 29 Dames W, Βber das Vorkommen von Hyaenarctos in den Pliozgn2Ablagerungen von Pikermi bei Athen. Sitz2Ber Gesel naturf Freunde, 8 : 1 8 De Mecquenem R, Contribution a l tude des fossiles de Maragheh. Ann Paleont, 13 : , 14 : Helbing H, Βber einen Indarctos2Schgdel aus dem Pontien der Insel Samos. Abh Schw Paleont Ges, 52 : 1 18 Li C K( ), Chi T( ), Pontian mammals from Wenquan, Sinkiang. Vert PalAsiat ( ), 8 (3) : (in Chinese with English summary) Lydekker R, Siwalik and Narbada Carnivora. Paleont Ind, 10(2) : Pilgrim G E, Further description of Indarctos salmontanus Pilgrim, the new genus of the bear from the Middle Siwaliks, with some remarks on the fossil Indian Ursidae. Rec Geol Surv Ind, 44 : Pilgrim G E, The fossil Carnivora of India. Paleont Ind, N S, 18 : Thenius E, Βber die Gehgrregion von Indarctos (Ursidae, Mammalia). Sitz ; str Akad Wiss, Math2naturw Kl, Abt 1, 158(9 10) : Thenius E, Indarctos arctoides (Carnivora, Mammalia) aus dem Pliozgn ; sterreichs nebst einer Revision der Gattung. Neues Jahrb Geol Palgontol, Abh, 108(3) : Weithofer K A, Beitr ge zur Kenntnis der Fauna von Pikermi bei Athen. Beitr Pal Str2Ung, 4 : Zdansky O, Jungtertgre Carnivoren Chinas. Pal Sin, C, 2(1) : 1 149

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~ ~ ~ 33 4 2014 467 478 Studies in the History of Natural Sciences Vol. 33 No. 4 2014 030006 20 20 N092 O6-092 A 1000-1224 2014 04-0467-12 200 13 Roger Bacon 1214 ~ 1292 14 Berthold Schwarz 20 Luther Carrington