Cainozoic Research, 5(1-2), pp. 93-102, November 2006
A primitive species of Chilotherium (Perissodactyla,
Rhinocerotidae) from the Late Miocene of the Linxia Basin
(Gansu, China)
Tao Deng
Institute of VertebratePaleontology and Paleoanthropology, ChineseAcademy of Sciences, P. O. Box 643, Beijing 100044, China: e-
mail: [email protected]
Received 1 June 2003; revised version accepted March 2006
skull with its articulated attributed the Late Miocene of Linxia Basin A mandible to Chilotheriumprimigenius sp. nov. from the (Gansu,
is described. This Late Miocene the earliest and in the with China), early species is most primitive genus Chilotherium.Compared the
other Late Miocene of C. →C.wimani→other of Chilotherium is species Chilotherium, a tentative lineageof primigenius→ species pro-
A C. posed. Late Miocene Asian radiationis suggested for this genus, starting fromLinxia primigenius. The origin and dispersal history
ofthe genus Chilotherium are discussed and correlated with major faunal exchanges. The sequence ofthe Late Miocene Hipparion fauna
in the Linxia Basin, which yields the remains described herein, is also discussed.
KEY WORDS: Chilotherium, Late Miocene, Linxia Basin, new species.
of of Introduction cies Chilotherium, leading to the description a new
species.
Late Cenozoic terrestrial deposits from the Linxia Basin
(Gansu, China) have yielded many mammalian fossils. In
these faunas, rhinocerotids are very abundant and species- Systematic palaeontology rich. The rhinocerotids have essentially been collected and
studiedsince the 1980s(Qiu etal, 1987,1990;Qiu & Xie, Family RhinocerotidaeOwen, 1845
1998; Guan, 1988; Guan & Zhang, 1993). Recently, we Subfamily AceratheriinaeDollo, 1885
have collected many new rhinocerotid material (Deng, Tribe ChilotheriiniQiu etal., 1987
2002a, 2003; Qiu et al. 2002). Qiu et al. Genus ChilotheriumRingstrom, 1924 2001b, c, b, ,
(1987)attributed a skull with its articulatedmandiblefrom
the Linxia Basin to Acerorhinus hezhengensis and consid-
ered that it differed from the species ofthe genus Chiloth- Chilotheriumprimigenius sp. nov.
erium. They established a newtribe Chilotheriini, including (Figures 1-2, Tables 1-3)
the genera Acerorhinus and Chilotherium.Their material
was recovered from unnamed Late Miocene deposits at Holotype — HMV 0102, an adultskull with its articulated
Dashengou (Hezheng, Linxia Basin). Deng (2001a) de- mandible. This mandible lacks the posterior part of the
scribed much more materialof Chilotheriumwimani from horizontal ramus, the lower part of ascending ramus and
than and consid- Fugu (Shaanxi, China) Ringstrom (1924), the left anterior corner of symphysis.
ered that C. wimani was the most primitive species of Chi-
lotherium. The species of Chilotheriumform a group con- Etymology — The specific name, primigenius, is a Latin
sidered be related the Acerorhinus and first. This is considered to closely to genus (Qiu word, means the new species
etal., 1987). the most primitive species of Chilotherium.
We have collected thousands ofmammalianfossils in the
Linxia Basin, including carnivores, rodents, perissodactyls, Type locality — Zhongmajia (35° 24' 47.0"N, 103° 24'
proboscideans, artiodactyls, and primates (Deng, 2001b, c, 30.6"E; altitude: 2218 m), 6 km southeast to the Hezheng
2002a, b, 2003; Qiu et ai, 2000, 2001; Wang and Qiu, County seat in Gansu Province, China. The specimen is
2002; An almost skull with its articulated collected from 2003). complete the gray green sandstoneand conglomerate
broken mandible from a small and primitive species of in the bottom of the Liushu Formation. The bed bearing
Chilotherium has been discovered at Zhongmajia in Hez- this fossil is of the early Late Miocene, corresponding to
Its known heng. morphology is different from all the spe- early and middleMN 9. -94-
— The braincase is Diagnosis Ventral surface ofmandibularsymphysis flat frontalbones is slightly erect. very narrow, Chiloth- with exterior walls. The is (different from concave surfaces of other known steep parietal crest slightly pro- erium species); parietal crests very slightly separated, form- jecting sideward to form a brim. The two parietal crests
to ing a sagittal crest different from the broadly separated meet posteriorly form a sagittal crest with the smallest
reach the lat- parietal crests of C. anderssoni, C. habereri, C. samium, C. width of 18 mm, from where they diverge to schlosseri and C. kowalevskii and also different from the eral ends ofthe occipital crest. The posttympanic process is ,
of with smoothanterior and surface. slightly separated parietal crests lacking a sagittal crest thin, a a rough posterior
C. C. wimani C. and C. kiliasi. This sideward, and declines for- xizangensis, , persiae process expands slightly
ward to fuse with the postglenoid process on the upper part
— exists Description Skull: The skull face is high. The premaxil- of the latter. As a result, a pseudoauditory meatus The is thin larybone is retracted into a vertical flake with a sharp ante- between them. postglenoid process longitudi-
incisors absent. Both ends of with transverse section. The interior rior end. The upper are pre- nally, a triangular sur- maxillary bones are in close proximity. In ventral view, the face of the postglenoid process is smooth, and the exterior
surface vertical The end of the premaxillary bone is thin and sharp, and its end is at the has a groove posteriorly.
forward. The same level with the nasal tip. The nasal notch is narrow and postglenoid process curves temporal condyle
deep, and its bottom is located at the level of M1/P4 is flat and straight, and its interior margin spaces out the boundary. A large infraorbital foramen on the maxillary postglenoid process by a deep groove. The articular surface is with continuous face is located near the bottomof the nasal notch. The dis- behindthe temporal condyle narrow, a tance between the bottomof thenasal notch and the orbit is wide crest on its exterior and posterior margins to form a
transverse flute to the The very short, 55.5 mm. The wide and prominent anterior parallel temporal condyle. margin ofthe orbit is located at the level of the middle of pterygoid bones are high, with a deep valley between them.
M2, and connects downward with the anterior margin of The maxillary tubercle is well developed, with a sharp pos- zygomatic arch. A frontward face is in front of the large terior end. The palatal surface is smooth and narrowly
ofthe is and and round orbit. The postorbital process is well developed arched. The posterior margin palate V-shaped,
its bottom is located the level of the of M3. on the frontal bone, but absent on the the zygomatic bone. at protocone
The supraorbital and lacrimal tubercles are relatively Mandible:There is a long distance between the ascending
is notch between the and m3. The of anterior of the marked. There a postorbital process ramus upper part margin
rounded. The coronoid and supraorbital tubercle, and there is another one between ascending ramus is thin and process the supraorbital and the lacrimal tubercles. The zygomatic is well developed, and strongly curved backward. The
is wide and arch is projecting highly to form a strong facial crest with a condyloid process very transversely narrow right angle, and its anterior end is located at the level of longitudinally, especially narrower interiorly. Near the inte- there is marked P4/M1 boundary. This arch is constricted inward at the rior end of the condyloid process, a con-
is striction. On the of the level of the postorbital process, while its posterior part posterior part condyloid process,
is and with expanded outward as a semicircle. The widest part of the there a narrow deep transverse groove a sharp zygomatic arch is located at the level of the postorbital bottom and a well-developed transverse crest.
the end of the arch, its In view, the mandibular process. On posterior zygomatic top symphysis expands strongly
sideward form shovel. The alveolar is crest posterior upper corner is rounded, and its posterior margin to a margin a
and its anterior is straight and inclined forward. In ventral view, the zygo- with a straight posterior part, part expands
The outward to be close to i2. There is wide matic arch is very thin.The occipital elevationis high. a U-shaped valley occipital surface and lateral margins incline forward obvi- between the two alveolar margins. The posterior margin of
mandibular is located the level of ously. The lower part of the occipital surface is broken. the symphysis at p3/p4 In front view, the alveolar between the The foramen magnum is onion-shaped, with a sharp upper boundary. margin
shallow central second lower incisors is thinand The ventral end. The occipital crest has a wide and uplifted. sur-
and its lateral corners are rounded. The lateral face of the mandibular is broad and flat, on groove, two symphysis nutrient foramina distributed margins ofthe occipital crest are straight and parallel each which there are some large
other. The nuchal ligament depression is wide and shallow. densely and irregularly, four on the right side.
The exterior lateral crest is strong, and a deep vertical de- Upper teeth: The premolars have been worn deeply to
reach the The labialwalls ofthe cheek teeth pression is present between this crest and the lateral margin root. upper are
the nuchal flat. DP 1 has root. The molars also have been worn of the occipital crest. The median crest on liga- a single M3 becomes ment depression connects with the upperend of the fora- to be close to the root. As a result, a quadran-
On the molars, the is constricted; the en- men magnum. The cranial top is smooth and flat, and it is gle. protocone
the widest at the level of the supraorbital tubercle. The trance ofmedian depression is wide, with some pillars; the
and and extend and of and nasals are narrow long, flatly straight. lingual margins protocone hypocone are straight.
is enamel board the The nasal top is convex, and its basehas a marked constric- There a well-developed on posterior
tion. The nasal tip is appreciably truncated and slightly wall of M3. Lower teeth: The first lower incisors absent, and the rough. The nasal lateral margins are drooping markedly are
The nasal second lower incisors with posteriorly and curved inward. ventral surface are very strong tusks a worn, middle of the nasal transversal backward oriented surface. also is convex, so the part lingual
section is lentoid. The connection between the nasal and -95-
Liushu Late Figure 1. Chilotherium primigenius sp. nov. Holotype (HMV 0102), Formation, Miocene, Zhongmajia, Hezheng
County, Gansu Province, China. 1, lateral view of skull. 2, ventral view of skull. 3, dorsal view ofskull. Scale bar = 6 cm. -96-
2. Chilotherium Liushu Formation, Late Miocene, Figure primigenius sp. nov. Holotype (HMV 0102), Zhongmajia, Hezheng
County, Gansu Province, China. 2-1, occipital view of skull. 2-2, dorsal view of mandible. 2-3, lateral view ofthe right side of
the mandible. Scale bar = 6 cm. -97-
The two tusks extend outward and upward markedly, and out to be a Hispanotherium matritense (Villalta & Crusa- their medial also almost The flanges are erect. transverse font, 1955). Similarly, another species, Chilotherium section of the crown of i2 is a round triangle with an inte- ibericum, described by Antunes (1972) fromPortugal, was rior sharp angle, but thatofthe root is an oval. The base of also laterattributed to Hispanotherium matritense(Heissig,
ofi2 crown is 37 mm wide and 26 mm thick. 1975). Borissiak (1915) described anew species Acerath-
The distancebetweenthe tusks is about from relatively large, 80 erium zernowi Odessa, and Kretzoi (1942) created a mm. The lower cheek teethalso are worn deeply. The ante- new genus Acerorhinus for this species. Heissig (1975) rior end of talonid of p2 is sharp and projecting forward. reduced the genus Acerorhinus to the rank of subgenus of
The talonids of and be- p2 p3 are much better developed than the genus Chilotherium, so that Acerorhinus zernowi the On the lower cheek the cin- C. referred trigonids. teeth, lingual came (Acerorhinus) zernowi. Heissig also one gulum is absent, and the V-shaped labial groove is wide species ofAceratherium and two species ofDiceratherium and shallow. from China into this subgenus of Chilotherium, including
C. (A.) hipparionum (Koken, 1885), C. (A.)palaeosinense
Discussion Since established the tsaidamense Ringstrom (1924) ge- (Ringstrom, 1924), and C. (A.) (Bohlin, 1937). nus Chilotherium, twelve species have been described, and Qiu et al. (1987) consideredthat the subgenus Acerorhiuns
19 other species have beenreferred to this genus (Table 4). shouldbe ranked a genus, so above-mentionedfour species
Ringstrom (1924) referred two species from Siwalik into ofthe subgenus Acerorhinusbecame Acerorhinuszernowi, the Chilotherium, i.e. C. (Lydekker, 1884) A. A. and A. tsaidamnesis. In genus blanfordi hipparionus , palaeosinensis, and C. the Acerorhinus has constricted nasal fatehjangense (Pilgrim, 1910). Foster-Cooper fact, genus a strongly
(1934) changed Aprotodon smith-woodwardi, a new genus base and a relatively narrow mandibularsymphysis, which and species established by him in 1915, to Chilotherium are very different fromthe genus Chilotherium.Ringstrom
smith-woodwardi. Heissig (1975) attributed1 C. blanfordi (1924) revised Rhinoceros persiae from Maragha (Iran) and C. fatehjangense to the genus Aprotodon. Qiu & Xie described by Pohlig (1885) into Chilotheriumpersiae. This
C. smith-woodwardito its has the characters of (1997) reassigned original name species typical Chilotherium, espe-
A. smith-woodwardi. There are essential differences be- cially a widely expanded mandibular symphysis. On the
tween Aprotodon and Chilotherium:Aprotodon has a pro- other hand, the M 3 of this species is quadrangular, with a
portionally larger and wider symphysis than Chilotherium', rudimental posterior depression, which is different from
the horizontal ramus of Aprotodon is curved not only in other species of Chilotherium. Most species of Chiloth-
side view (lower border curved) but also in dorsal view erium have been recorded from Neogene deposits of China.
(both extremities deflectoutward), which is seldom seenin Ringstrom (1924) described three new species ofChiloth-
rhinocerotids; Aprotodon has semi-molariformpremolars, erium and one revised species described by Schlosser
while Chilotherium fully molariform(Qiu & Xie, 1997). (1903) from the Hipparion fauna in northern China, but
Matthew (1929) revised Rhinoceros sivalensis var. inter- Heissig (1975) merged C. planifrons into C. anderssoni,
medius described by Lydekker (1884) from Siwalik into C. and synonymized C. wimaniwith iC. habereri. However, C.
intermedium, and Heissig (1975) placed C. intermediumin wimani has a narrow surface between the parietal crests, a
Subchilotherium. the new subgenus Heissig (1989) raised concave cranial profile, and well-developed parastyle folds
the Subchilotherium the the the cheek which characters subgenus to genus rank, so on upper teeth, can obviously
species became S. intermedium.The mandibularsymphysis be distinguished from C. habereri, so C. wimaniis a valid of Subchilotheriumis differentfrom narrow, and the widely separate species (Deng, 2001a, c, 2002a). Tung et al.
expanded one ofChilotherium.As a result, there is not any (1975) showed that the surface between the parietal crests real species of Chilotherium in the Siwalik faunas. of C. fenhoense is wider than that of C. anderssoni. How-
Ringstrom (1924) referred four species of Aceratherium ever, the widthbetweenthe parietal crests ofChilotherium
C.from Samos (Greece) into Chilotherium, including is variable (Deng, 2001a, c), and the larger width of C.
samium (Weber, C. schlosseri falls 1905), (Weber, 1905), C. fenhoense within the rangeof variation of C. anders-
wegneri (Andree, 1921), and C. angustifrons (Andree, soni. From Late Miocene deposits ofTibet, Ji etal. (1980)
1921). At the same time, he referred Teleoceras poticus described a new species of Chilotherium, C. xizangensis, and Aceratherium kowalevskii from Odessa (Ukrain) into and Zheng (1980)addedanother species. C. tanggulaense.
Chilotherium C. and as ponticum (Niezabitowski, 1913) C. The widely expanded mandibularsymphysis of C. xizan- kowalevskii (Pavlow, 1913). Heissig (1975) merged C. gensis underlinesits attributionto Chilotherium.The dental
ponticum weeneri and C. — into C. schlosseri, and attributed characters of C. tanggulaense show that it should be re-
C. angustifrons to C. kowalevskii. Therefore, only C. ferred into Subchilotherium. Heissig (1975) revised Rhi-
samium, C. schlosseri and C. kowalevskii are consideredto noceros pygmaeus described by Ringstrom (1927) only be valid snecies of Chilotheriumfrom Samos and Odessa. based upon limb bones from Guide, Qinghai (previously Geraads & Koufos (1990) described a new species, Kansu) into Chilotherium (Subchilotherium) pygmaeum.
Aceratherium kiliasi from (Greece). Because Because Subchilotheriumhas Pentalophos been raised as a genus. C.
its mandibular is is referred to symphysis broadly expanded, Heissig pygmaeum now as S. pygmaeum. Schlosser
(1999) attributed A. kilaisi to Chilotherium kiliasi. (1903) described a new species Rhinocerosbrancoi based
of Zbyszewsky (1952) described a new species. Chilotherium upon a specimen the cheek teeth with complicated
quintanelensis from Portugal, but this species later turned enamel plications from northern China. Rretzoi (1942) -98-
created a new genus for this species, i.e. .Shansirhinus, but concave ventral surface of the mandibular symphysis. Node 5,
Heissig (1975) renamed it as Chilotheriumbrancoi. More broadly separated parietal crests, weak lingual cingulum, low
discoveries from occipital surface, robust paroccipital slender postglenoid China prove that the genus Shansirhinus process, well-developed postorbital weak is valid.Chilotherium yunnanensisfrom Yunnan described process, process, supraorbital thin orbital tubercle, upper margin, narrow nasal bone, low and by Tang et al. (1974), C. cornutum from Shanxi (Qiu & rounded braincase, high crown ofthe cheek teeth, strong constric- Yan, 1982), and C. tianzhuensis from Gansu (Zheng, 1982) tion ofthe protocone, and comparatively large P2. Other species should be referred into Shansirhinus as well. of include Chilotherium C. anderssoni, C. habereri, C. samium, In conclusion, nine species of Chilotherium are valid: C. C. schlosseri, C. kowalevskii, C. xizangensis, C. persiae, and C. samium C. schlosseri C. and kiliasi , kowalevskii. C. from , kiliasi. Europe, C. anderssoni, C. habereri, C. wimani and iC. xi- ,
zangensis from China, and C. from Iran. All nine persiae The shows that C. shares cladogram primigenius many species have a strongly concave ventral surface of the man- synapomorphies (Fig. 3, node 3) with the other species of dibular while C. symphysis, primigenius has a relatively Chilotherium, but not with the primitive chilothere flat ventral surface. Chilotheriumanderssoni, C. habereri, Acerorhinu and otherrhinocerotids whole: fused as a post- C. samium C. and C. kowalevskii have , schlosseri, broadly tympanic process with the postglenoid process, low parietal separated parietal crests (Schlosser, 1903; Weber, 1905; crest, narrow zygomatic arch without a projection ofpos- Pavlow, 1913; Ringstrdm, 1924). On the other hand, C. terodorsalangle, gradually narrowing cranialdorsal surface has that primigenius very poorly separated parietal crests before the orbit, ofthe mandibular strong broadening sym- form a sagittal crest backwards. Chilotheriumxizangensis , physis, well-developed secondary structures on the cheek C. wimani, C. persiae, and C. kiliasi have the slightly sepa- teeth, comparatively small P2, and upturned medial flanges rated al, parietal crests (Ji et 1980; Ringstrom, 1924; of i 2 are apomorphies and mean that C. primigenius is truly 2001a, Geraads Koufos, not Deng, c; & 1990), but they do member a of the genus Chilotherium. Meanwhile, some form a sagittal crest. The surface between the crests parietal apomorphic characters ofthe otherspecies ofChilotherium ol C. wimani is and this has very narrow, species a ten- thick are lacking: posttympanic process, obviously sepa- dency to form a sagittal crest. The M 3 of C. persiae is rated parietal crests, wide nasal notch, long distance be- unique by its quadrangular occlusal surface in all known tween the bases ofi 2 and p2, concave ventral surface ofthe of species Chilotherium, so the C. primi- triangular M 3 of mandibularsymphysis, which also characterize the “C. wi- genius can be easily distinguished from i C. persiae. Conse- mani”-other Chilotherium species clade” (containing C. quently, C. primigenius is a new species different from anderssoni, C. habereri, C. samium, C. schlosseri, C. other species of Chilotherium. The relatively small size kowalevskii, C. xizangensis, C. persiae, and C. kiliasi)” (Table 1-3) is characteristic ofskull, mandible and teethof node (Fig. 3, 4), are absent in this species. Consequently, C. It is much smaller than C. primigenius. wimani, but a C. primigenius is the sister group ofother species of Chi- little larger than C. xizangensis (Ji etal, 1980). lotherium. The second dichotomy within the genus Chi- Characters C.checked on skull, mandible and teeth of lotherium (Fig. 3, node 5) sets C. wimani as the sister primigenius allow us to investigate the relationships ofthis group ofthe remaining species ofChilotherium.These lat- new species using cladistic analyses (Fig. 3). ter species are defined by markedly separated parietal
crests, weak lingual cingulum, low occipital surface, robust
slender paroccipital process, postglenoid process, well-
developed postorbital process, weak supraorbital tubercle,
thin orbital nasal upper margin, narrow bone, low and
rounded braincase, high crown of the cheek teeth, strong
constriction of the protocone, and comparatively large P2.
Chilotheriumprimigenius and C. wimani are distinct from
other species of Chilotherium. Only cranial and dental
characters have been discussed so far, because postcranial
remains attributed C. to primigenius are entirely unknown.
It is impossible to define the characteristics of the skull at
the basal node of the genus Chilotherium.
Chilotheriumprimigenius is the most primitive species of Figure 3. Phylogenetic relationships of Chilotherium primigen- Chilotherium described far. This so species shares many ius sp. nov. exclusive characters with the other species of Chilotherium. Node Node Acerorhinus. 1, Outgroup. 2, genus Node 3, fused For its known characters. C. primigenius constitutes the with the low posttympanic process postglenoid one, parietal crest, ancestral morphotype of the genus Chilotherium. More- narrow zygomatic arch without a projection ofposterodorsal an- over, it implies that the itself is older. The locality gle, gradually narrowing cranial dorsal surface before the orbit, genus with C. primigenius corresponds to the early Late Miocene strong broadening ofthe mandibular symphysis, well-developed
(early and middle MN 9). But a Middle Miocene is structures on the cheek teeth, comparatively small P2, upturned origin
For some doubtfulremains of medial flanges ofi2, flat labial wall ofuppercheek teeth. Node 4, probable. example, Chiloth-
thick posttympanic process, obviously separated parietal crests, erium sp. were reported from Halamagai (Xinjiang, China)
wide nasal distance between the bases of i2 notch, long and p2, by Chow (1957) and from Jiulongkou (Hebei, China) by -99-
Chen and Wu (1976). The described material from Ha- References lamagai is only a right M1. This molar has a sharply de- marcated parastyle fold, and its protocone is not con- Andree, J. 1921. Rhinocerotiden aus dem Unterpliocan von
Samos. 3: 189-212. stricted. As a result, this molar should belong to Ac- Palaeontologische Zeitschrift, Antunes, M.T. 1972. Un Rhinocerotide, Chilotherium erorhinus instead of Chilotherium (Deng, 2000; Deng & nouveau ibericus Boletim do Museu Laboratorio n. sp. e Mineralogico Downs, 2002). Tong et al. (1990) also reported that there e Geoldgico daFaculdade de Ciencias, 13: 25-33. are some questionable fossils of Chilotherium, but they did Borissiak, A. 1915. Mammiferes fossiles de Sebastopol, II. Trudy not describe these fossils. The referred material from Jiu- Geologicheskago Komiteta, Novaja Seria, 137: 1-45. longkou comprises the anteriorpart of a young skull, the Bohlin, B. 1937. Eine tertiare Saugetier-FaunaausTsaidam, Pa- left maxillary bone of another young skull, and a P2, but laeontologia Sinica, Series C, 14; 1-111. have of Chilotherium. At the they no diagnosis moment, Chen, G.F. & Wu, W.Y. 1976. Miocene mammalian fossils of Linxia Basin is with old- Zhongmajia in the the locality the Jiulongkou, Ci Xian district, Hebei. Vertebrate Palasiatica, est undubiousremains ofthe genus Chilotherium. Because 14: 6-15, the most primitive species ofChilotherium, C. primigenius Chow, M.C. 1957. Notes on some mammalian fossils from the
Late Cenozoic of Vertebrate 1: 33-41. and C. wimani, originated in China, an East Asian origin Sinkiang. Palasiatica,
Deng, T. 2000. A new species of Acerorhinus (Perissodactyla, for this genus is suggested. Heissig (1989) consideredthat Rhinocerotidae) from the Late Miocene in Fugu, Shaaxi, Chilotherium emerged in the early Middle Miocene of China. Vertebrata Palasiatica, 38: 203-217. South Asia with Subchilotherium from the lower Siwalik Deng, T. 2001a. New materials ofChilotherium wimani (Perriso- series. The first appearance of C. wimani was necessarily dactyla, Rhinocerotidae) from the Late Miocene of Fugu, later than that of C. primigenius, and must precede that of Shaanxi. Vertebrata Palasiatica, 39: 129-138. the other species of Chilotherium.In Zhongmajia locality Deng, T. 2001b. New remains ofParelasmotherium (Perissodac- of the Linxia Basin, the bed bearing C. wimani remains is tyla, Rhinocerotidae) from the Late Miocene in Dongxiang, bed located above the containing C. primigenius. The ear- Gansu, China. Vertebrata Palasiatica,39: 306-311.
liest records of the Chilotheriumoutside the Linxia genus Deng, T. 2001c. Cranial ontogenesis of Chilotherium wimani
Basin are in western China: C. habereriis present in Bahe (Perissodactyla, Rhinocerotidae). In: Deng, T. & Wang, T.
the Annual the (Lantian, Shaanxi) (Liu et al., 1978) at location attributed (eds). Proceedings of Eighth Meeting of Chi-
nese Vertebrate China Ocean to late MN 9-10 (Qiu et al., 1999). Society of Paleontology, Press,
of Beijing, 101-112. In Europe, the first appearance Chilotherium is in Penta- Deng, T. 2002a. Limb bones of Chilotherium wimani (Perisso- lophos (Greece): C. kiliasi (Geraads & Koufos, 1990) is dactyla, Rhinocerotidae) from the Late Miocene ofthe Linxia present at the location attributed to MN 11-12 (Heissig, Basin in Gansu, China. Vertebrata Palasiatica, 40: 305-316. 1999). Hence the dispersal of the genus Chilotherium to- Deng, T. 2002b. The earliest known woolly rhino discovered in wards Europe can be consideredto begin with the arrival of the Linxia Basin, Gansu Province, China. Geological Bulletin
C. kiliasi. This event is situated between the muridevent at of China, 21: 604-608. 9.6 Ma and Gazella event at 6.9 Ma (Made, 1999). During Deng, T, 2003. New material of Hispanotherium matritense the MN 10-13 least four of Chilotherium Zones, at species (Rhinocerotidae, Perissodactyla) from Laogou of Hezheng
were present in South and East Europe. County (Gansu, China), with special reference to the Chinese
The of the Linxia fauna Middle Miocene elasmotheres. Geobios 36: 141-150. stratigraphical position Hipparion ,
containing abundant fossils of Chilotherium has always Deng, T. & Downs, W. 2002. Evolutionof the ChineseNeogene
Rhinocerotidae and its to climatic variation. Acta been a problem. Fang et al. (1997) arranged the chrono- response GeologicaSinica,16: 139-145. logical position of the Linxia Hipparion faunas and deter- Fang, X.M., Li, J.J., Zhu, J.J., Chen, H.L. & Cao J.X. 1997. Ab- there obvious mined theirpaleomagnetic ages. But was an solute dating and division of the Cenozoic strata ofthe Linxia first of error in this sequence, i.e. the appearance Hipparion Basin in Gansu. Chinese Science Bulletin, 42; 1457-1471. in the Linxia Basin at 15.4-12.5 Ma (Middle Miocene). Foster-Cooper, C. 1934.The extinct rhinoceroses ofBaluchistan. According to our recent fieldobservations and correlations, Philosophical Transactions of the Royal Society ofLondon, the Late Miocene Hipparion faunas in the Linxia Basin can Series B, 223: 569-616. be divided into four levels. The bed the containing speci- Geraards, D. & Koufos, G. 1990. Upper MioceneRhinocerotidae of C. the first men primigenius corresponds to Hipparion (Mammalia) from Pentaloph-1, Macedonia,Greece, Palaeon-
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Measures C. primigenius C. wimani C. xizangensis
3 Distance between nasal tip and occipital crest 390 502.8502.8 (483-528) -
4 Distance between nasal tip and bottom of nasal notch 133.5 148.5 (132-162) -
5 Minimal width of braincase 72 84.9(74-91.5)84.9 (74-91.5) -
6 Distance between crest and 203 257.9 (246.5-267) - Distance between occipital postorbital process 203 257.9 (246.5-267)
7 Distance between occipital crest and supraorbital tubercle 225.5 264.9(214.5-289.5)264.9 (214.5-289.5) -
8 Distance between occipital crest and lacrimal tubercle 233.5 320.3 (299.5-332) -
9 Distance between nasal notch and orbit 55.5 68.1 (60-81) 54
1414 Distance between nasal tip and orbit 186 210.8 (198.5-223) -
15 Width of occipital crest 114 151.5(140.2-168.4) -
16 Width between mastoid ~145-145 195.5 - processes 195.5 (184-219)
17 Minimal width between parietal crests 1818 44.9(29-67)44.9 (29-67) -
18 Width between 147 ~124-124 Width postorbital processes 147 140.3(133.3-148)
19 Width between supraorbital tubercles 158 157.7 (147-164.3) -120
20 Width between lacrimal tuberclestubercles 152 160.4(143-184.8) -
21 Maximal width between zygomatic arches 228 262.7 (248-280) -
22 Width of nasal base 83 88.1 (75-98) 60
23 Height of occipital surface 113 134(120-158) -
25 Cranial height in front ofP2 117 141.2(121-158) -
26 Cranial height in front ofMl 138 168 (139-192) 137
27 Cranial height in frontfront ofM3 165 172(155.5-189) 132
28 Palatal width in frontof P2 42 44.3 (37.5-55.5) -
29 Palatal width in front of Ml 25 43.7(27-57)43.7 (27-57) -
30 Palatal width in front of M3 28 56.9(45-72)56.9 (45-72) -
ofthe skull of and with other of Ten Table 1. Measurements (mm) Chilotheriumprimigenius sp. nov. comparisons species Chilotherium.
specimens of C. wimani were measured. Numbers in front of measuring method in Table 1 correspond with those ofGuerin (1980,
table 1, p- 47).
Measures C. primigenius C. wimani C. xizangensis
I1 Length -340 471.6(455-485)471.6 (455-485) --
9 Distance between ramuses in front of ml 29 62 (54-69) 33
10 Distance between ramuses in front of m3 30 77.3 (65-88.3)(65-88.3) 39
II11 LengthLength of symphysis 96.5 117.5(110-126)117.5 (110-126) 108108
14 Transverse diameterof condyle 7676 90.5 (89-92) “
of Table 2. Measurements (mm) ofthe mandible Chilotheriumprimigenius sp. nov. and comparisons with other species ofChilotherium.
Five specimens ofC. wimani were measured. Numbers under measurements correspond with those ofGuerin (1980, table 3, p. 52).
Teeth Ml M2M3p2M2 pM33 p2 p p3 4 p4 mlml mm2 2 m3m3
C. primigeniusprimigenius
L 29 33.5 36.5 19.5 26 28.5 26.5 32 37.5
W 51 51 45 13 21 24 26 29 25
C. wimani
L 38.4 48.7 48,7 23.9 29.3 35.7 39.6 46.4 46.9
W 59.5 61 56.8 17.4 20 25.8 27.7 28.428,4 25.9
C. xizangensis
L 41.2 41 35.6 25.2 29 32 36.4 39.2 40
W 50.8 48.5 41.8 18.2 24.3 22.1 23.4 20.9 18.9
Table 3. Measurements ofthe cheek teeth of Chilotherium and with other of Chiloth- (mm) primigeniussp. nov. comparisons species
erium. L = Length; W =Width; the data of C. wimani are mean values (N =5). - 102-
Known species of iChilotherium Revised species Distribution
C. blanfordi (Lydekker.(Lydekker. 1884) Aprotodon blanfordi (Lydekker,(Lydekker, 1884) Siwalik
C. fatehjangensefatehjangense (Pilgrim,(Pilgrim, 1910) Aprotodonfatehjangense (Pilgrim, 1910)1910) Siwalik
C. smith-woodwardi (Foster-Cooper, 1915) Aprotodonsmith-woodwardi Foster-Cooper, 19151915 Siwalik
C. ihericumibericum Antunes, 1972 Hispanotherium matritense (Prado, 1863) Portugal
C. quintanelensis Zbyszewski,Zbyszewski, 1952 Hispanotherium matritense (Prado, 1863) Portugal
C. zernowi (Borissiak, 1915)1915) Acerorhinus zernowi (Borissiak, 1915)1915) Odessa
C. palaeosinensepalaeosinense (Ringstrom, 1924) Acerorhinuspalaeosinensispalaeosinensis (Ringstrom, 1924) China
C. hipparionumhipparionum (Koken, 1885) Acerorhinus hipparionumhipparionum(Koken,(Koken, 1885) China
C. tsaidamensetsaidamense (Bohlin, 1937) Acerorhinus tsaidamensistsaidamensis (Bohlin, 1937)1937) China
C. intermediumintermedium (Lydekker,(Lydekker, 1884) Subchilotherium intermedium (Lydekker,(Lydekker, 1884) Siwalik
C. tanggulaensetanggulaenseZheng, 19801980 Subchilotherium intermedium (Lydekker,(Lydekker, 1884) China
C. Subchilotherium China pygmaeum (Ringstrom, 1927) pygmaeum (Ringstrom, 1927)
C. brancoi (Schlosser, 1903) Shansirhinusbrancoi (Schlosser,(Schlosser, 1903) China
C. yunnanensisTang et al.,al 1974 Shansirhinus brancoi (Schlosser, 1903) China
C. tianzhuensisZheng, 1982 Shansirhinus ringstromi Kretzoi, 1942 China
C. comutumcornutum Qiu & Yan, 1982 Shansirhinus ringstromi Kretzoi, 1942 China
C. samiumsamium (Weber, 1905) C. samiumsamium (Weber, 1905) Samos
C. schlosserischlosseri (Weber.(Weber, 1905) C. schlosserischlosseri (Weber,(Weber. 1905) Samos
C. ponticum (Niezabitowski, 1912) C. schlosseri (Weber. 1905) Odessa
C. wegneri (Andree, 1921) C. schlosserischlosseri (Weber. 1905) Samos
C. kowalevskii (Pavlow, 1913)1913) C. kowalevskii ( Pavlow, 1913)1913) Odessa
C. angustifronsangustifrons (Andree,(Andree, 1921) C. kowalevskiikowalevskii (Pavlow, 1913)1913) Samos
C. kiliasi (Geraads & Koufos, 1990) C. kiliasikiliasi (Geraads & Koufos.Koufos, 1990) Pentalophos
C. anderssoniRingstrom, 1924 C. anderssonianderssoni Ringstrom, 1924 China
C. planifrons Ringstrom, 1924 C. anderssonianderssoni Ringstrom, 1924 China
C.fenhoensisC. fenhoensis Tung etet at., 1975 C. anderssonianderssoni Ringstrom, 1924 China
C. habereri (Schlosser,(Schlosser, 1903) C. habereri (Schlosser, 1903) China
C. gracile Ringstrom, 1924 C. habereri (Schlosser, 1903) China
C. wimani Ringstrom, 1924 C. wimani Ringstrom,Ringstrom. 1924 China
C. xizangensis JiJii etal., 1980 C. xizangensis Ji et ai,al., 1980 China
C. persiaepersiae (Pohlig, 1885) C. persiae (Pohlig, 1885) Maragha
Table 4. Revision of the Chilotherium 4. Revision ofthe genus Chilotherium