ΐῒ῏ῑῐ ῍῎῍῎ῌThe Quaternary Research.1 - p. +/3 +1, June ,**2

Early Pleistocene soricomorphs and lagomorphs from the Xiaochangliang site in the Nihewan Basin, Hebei, Northern China

Yingqi Zhang*+, and Yoshinari Kawamura* ,a

This paper systematically describes soricomorph and lagomorph remains collected by a screen-washing procedure from the sediments of the Xiaochangliang site with the age of+4-0 Ma. The soricomorph remains described are assigned to the tribe Soricini in the subfamily Soricinae of the family Soricidae, on the basis of the detailed dental morphology. Among the genera of the tribe known from the Pliocene and Pleistocene of Asia,Sorex has morphological characters well coincident with those of the remains. Owing to their incompletion, the specific determination can not be done, and thus they are assigned merely toSorex sp. All the lagomorph remains belong to the family Ochotonidae. The family comprises two genera,Ochotona and Ochotonoi- des, in the Pliocene and Pleistocene of China. The remains are easily divided into a smaller and larger forms by their striking di# erence in size. The dental morphology and size of the smaller form indicate its allocation toOchotona. The smaller form is compared with many species ofOchotona known from the Pliocene and Pleistocene as well as the present day of China, and is assigned to the extinct species,O. youngi. The larger form is, however, assigned merely toOchotona or Ochotonoides sp., because of its limited specimens with poor preservation.

Keywords : soricomorph, lagomorph, systematics, Early Pleistocene, Xiaochangliang site, Nihewan Basin, China

knowledge on small mammals from the layer I. Introduction had been quite limited, until we washed the The Xiaochangliang site is one of the most sediments stratigraphically equivalent to the important archaeological sites of Early Pleisto- layer with fine-mesh screens in,**/ . Our cene age in China, and is situated in the Nihe- screen-washing successfully produced a large wan Basin about+/* km west o north o west of number of small mammal remains which in- Beijing (see Fig.+ of Zhanget al., ,**2 ). The cluded +* forms identifiable at the generic or artifact layer of the site is intercalated in flu- specific level (Zhanget al., ,**2 ). Among them, vio-lacustrine sediments called the Nihewan soricomorph and lagomorph remains are de- Formation, and was dated as+4-0 Ma by paleo- scribed systematically in this paper. The geo- magnetic method (Zhuet al., ,**+ ). It was logical setting of the layer and the screen- known that the layer had yielded mammalian washing procedure are explained in Zhang et remains in association with artifacts, but theal. (,**2 ).

Received July,+, ,**1. Accepted January +3 , ,**2. * + Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box0.- , Beijing, +***..,China(Presentaddress : Graduate School of Science, Osaka City University.--+-2ῌῌ Sugimoto, Sumiyoshi-ku, Osaka,//2ῌ 2/2/ , Japan). *Dep,artment of Earth Sciences, Aichi University of Education. + Hirosawa, Igaya-cho,Kariya, ..2ῌ 2/.,,Japan. * a Corresponding author:[email protected]. ac. jp 160 Zhang, Y.Q. and Kawamura, Y. June ,**2

face of the crown, the posterior cingulum runs II. Terminology and measuring method along the posterior margin of the crown. It is Almost all the remains described herein are thick and not undulated. The cingulum con- detached teeth. Figure+ shows the terminol- tinues to the ventral face of the crown, and ogy and measuring method used for the teeth forms a knob at the linguo-ventral corner of of soricid soricomorphs, which follow those of the crown. On the lingual face of the crown, Kawamura (+33, , +33- ). Figure , also shows however, it is absent or feeble. The root is long those for the cheek teeth of ochotonid lago- and stout, and extends postero-dorsally. morphs, which mainly follow those of Lopezo M+ : In one of the two specimens described (V Martinez (+320 ). The measurements of the lago- +/-+24- ), the antero-buccal part of the crown is morph teeth were taken parallel to their occlu- lost (-- in Fig. ), but this part is observable in sal surfaces. another specimen (V+/-+24. ). The occlusal out- All the measurements were taken by using a line of the crown is squarish, and is not elon- measurescope (Nikon : MMῌ++ ) with an electric gated linguo-buccally so that W is only little digital counter (Nikon : CMῌ0/ ). larger than L (Table + ). The anterior border of the crown is gently convex anteriorly in occlu- III. Systematic description sal view. The buccal border of the crown is Order Soricomorpha Gregory,+3+* weakly emarginated both in its anterior and Family Soricidae Fischer de Waldheim,+2+1 posterior halves. The V o shaped ridge formed Subfamily Soricinae Fischer de Waldheim,+2+1 by the parastyle, paracone and mesostyle has Tribe Soricini Fischer de Waldheim,+2+1 nearly the same size and shape as that formed GenusSorex Linnaeus,+1/2 by the mesostyle, metacone and metastyle. A Sorex sp. postero-buccal protrusion of the metastyle is (Fig.- ) weak. In the lingual part of the crown, the pro- Referred specimensῌ+ left first upper incisor tocone is well developed, and forms a broad- (V+/-+24+ ) ; + right first upper incisor (V +/-+24, ) ; angled Vo shaped ridge, whose posterior part ,broken right M+, (V +/-+24-ῌ . ) ; + left M (V (postprotocrista) runs posteriorly. This ridge is

+/-+24/) ; + broken left M+ (V +/-+240 ) ; + right separated from the lingual base of the meta-

M,- (V+/-+241 ) ; + left M (V +/-+242 ). cone, and thus the trigon basin opens posteri- RepositoryῌAll the specimens are stored in orly. The hypocone is much smaller and lower the Institute of Vertebrate Paleontology and than the protocone, but shows a well-defined Paleoanthropology (IVPP), Chinese Academy conical shape. It is separated from the postpro- of Sciences, Beijing. tocrista by a shallow valley. The lingual bor- LocalityῌXiaochangliang site, about*4/ km der of the crown is gently emarginated, and a northwest of Guanting, Yangyuan County, He- short cingulum is observed along its central bei Province (IVPP field locality number12**/ ). part. The distocrista extends posteriorly from Geographic coordinates :++.῍῍ .*ῌῌ E, .* +- N. the apex of the hypocone to the postero-lingual HorizonῌUpper part of the Nihewan Forma- corner of the crown, where it forms a feeble tion defined by Wei (+331 ) cusplet. Furthermore it continues to the post- AgeῌEarly Pleistocene ; dated paleomagneti- cingulum, which runs along the posterior cally as+4-0 Ma. border of the crown and reaches the postero- DescriptionῌAll the specimens are secondar- buccal corner of the crown. The posterior emar- ily stained dark brown to brown, so that origi- gination is weak. nal pigmentation of their cusps cannot be de-M, : This tooth can be regarded as a small ver- tected. sion of M+ , but the following minor di# erences First upper incisor : The anterior cusp has are observed. W is almost equal to L (Table+ ). no medial cusplet, and thus this tooth is not The metastyle never protrudes postero-buccally. fissident. The posterior cusp is well developed, The postprotocrista bends at right angles, and and its apex reaches close to that of the ante- extends buccally to the base of the metacone. rior cusp in dorso-ventral level. On the labial The distocrista has no cusplet. ,**2῍ῌ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 161

Fig.+ Terminology and measuring method for the soricid soricomorph teethdescribed here, according to Kawamura (+33,,) +33- bl : base line for measurement, h : height of crown, L : length of crown, W : width of crown.

Fig., Terminology and measuring method for the ochotonid lagomorph cheek teethdescribed here, according to Lopezo Martinez ( +320 ) All the measurements are taken on the occlusal surface of each tooth. L : maximum length of crown, W : maximum width of crown, Wa:width of anterior loph in P- . 162 Zhang, Y.Q. and Kawamura, Y. June ,**2

Fig.- Detached teethofSorex sp. from the Lower Pleistocene of the Xiaochangliang site +. left first upper incisor (V +/-+24+ ), , . right first upper incisor (V +/-+24, ), - . right M+ (V +/-+24- ), . . , left M (V+/-+24/ ), / . right M,- (V +/-+241 ), 0 . left M (V +/-+242 ). +῍῍ , . labial view, - 0 . occlusal view.

M+ : The trigonid is mostly lost in the only rior part of the entocristid. The anterior end of specimen described. The oblique cristid is con- the entocristid is connected to the posterior nected to the posterior face of the protoconid. base of the metaconid.

A well-marked cingulum is observed on theM- : The occlusal outline is also trapezoidal. posterior base of the hypoconid. The postcris- The trigonid is larger than the talonid. A dis- tid extends from the hypoconid to the posterior tinct cingulum starts at the anterior base of the base of the entoconid, but its lingual end is paraconid, extends along the anterior and buc- separated from the entoconid by a narrow cal faces of the crown, and terminates on the groove extending postero-lingually from the posterior base of the hypoconid. Another nar- talonid basin. The entoconid is a conical cusp, row cingulum runs along the lingual face of and separated from the metaconid by a depres- the crown. The talonid is longer than broad. It sion between them. The entocristid is not rec- is not reduced, and shows nearly the same ognized. A narrow cingulum runs from the shape as those of M+, and M . But owing to the postero-lingual base of the entoconid to the lack of the narrow groove between the lingual lingual base of the metaconid. end of the postcristid and entoconid, the talo-

M, : The occlusal outline is trapezoidal. The nid basin becomes a closed depression sur- trigonid is nearly as large as the talonid, but rounded by a continuous ridge formed by the the former is slightly narrower than the latter. oblique cristid, hypoconid, postcristid, entoco- A distinct cingulum extends buccally from the nid and entocristid. The mesoconid is recog- anterior base of the paraconid along the ante- nizable on the oblique cristid. rior face of the crown. The morphology of theMeasurementsῌ The measurements of the talonid is the same as that of M+ except the teeth are given in Table+ . following points. The oblique cristid has aComparison and discussionῌ The specimens weak incision on its central part, so that the described here undoubtedly belong to the fam- mesoconid is recognizable. The entoconid is ily Soricidae. Repenning (+301 ) divided the elongated antero-posteriorly to form the poste- family into such four subfamilies as Heteroso- ,**2῍ῌ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 163

+ Table+ Measurements of the teethofsp.Sorex parastyle in M and extremely reduced M- . As from the Lower Pleistocene of the Xiao- regards the members of Notiosoricini, Repen- changliang site in mm ning (+301 ) gave their morphological charac- ters. According to him, Notiosoricini di# ers in

having a more reduced talonid of M- where the entocristid is lacking. The remaining tribe, Soricini, is therefore com- pared with the present specimens. This tribe comprises such nine genera as Sorex, Drepano- sorex, Deinsdorfia, Petenyia, Blarinella, Dimylo- sorex, Zelceina, Crocidosorexand Oligosorex (Reu- mer,+32. ). Among them, Sorex, Deinsdorfia, Petenyiaand Zelceina have been recorded from the Pliocene and Pleistocene of Asia (Storch et al., +332). Thus the four genera are compared with the present specimens on the basis of the descriptions by Reumer (+32. ). h:height of crown, L : length of crown, W : Deinsdorfia di# ers from the present speci- width of crown. mens in having a less developed posterior cusp of the first upper incisor, stronger posterior emargination in M+, and M , proportionately ricinae, Crocidurinae, Limnoecinae and Sorici- smaller talonid in M, , and strongly reduced nae, with the descriptions of morphological talonid of M- which is usually single-cusped. characters for each subfamily. According toPetenyia is also distinct in having a dorso-ven- him, Heterosoricinae di# ers from the present trally shorter posterior cusp of the first upper specimens in having no posterior emargination incisor, no hypocone in M+, and M , the entoco- + in M , a much thicker buccal cingulum in M, nid situated close to the metaconid in M+ and and M- , and proportionately smaller talonid in M, , and a reduced talonid with a single cusp in

M- . Crocidurinae is also distinct in having a M- .Zelceina is distinguishable from the pre- less developed posterior cusp in the first upper sent specimens by its larger size, posterior incisor, broader occlusal outline of M++, elongat- emargination of M and M being weaker, tet- ing bucco-lingually, much stronger posterior rahedric hypocone of the same teeth, and + emargination in M , and a much reduced talo- strongly reduced talonoid of M- . nid in M- . Limnoecinae is di# erent in having The remaining genus,Sorex, bears the main a proportionately shorter talonid in M+ and characters well coincident with those of the much reduced talonid in M- . Thus the mem- present specimens. They are as follows : The bers of the remaining subfamily, Soricinae, are first upper incisor is fissident or not fissident, compared with the present specimens. and has the posterior cusp being well-devel- Reumer (+32. ) revised the tribal classification oped or moderately developed. M+, and M has of Soricinae by Repenning (+301 ), and recog- a squarish outline, the posterior emargination nized such seven tribes as Soricini, Blarinini, being moderate or weak, and the hypocone

Notiosoricini, Soriculini, Beremendiini, Ambly- being well-difined and normal in shape. M, and coptini and Allosoricini. His descriptions indi- M- have the entocristid, and the talonid of M- is cate that Soriculini and Beremendiini are di# er- not reduced. The present specimens are there- ent from the present specimens in having a fore assignable to Sorex. fissident first upper incisor. Blarinini and Allo- As stated by Reumer (+32. ),Sorex is a very soricini also di# er in having no entocristid in large genus with many species. The present

M,- and M , and a reduced talonid in M - . Ambly- specimens are too incomplete to carry out any coptini is easily distinguishable from the pre- specific determination within the genus, and sent specimens by its much better developed thus they are assigned merely toSorex sp. 164 Zhang, Y.Q. and Kawamura, Y. June ,**2 herein. The hypoflexus is relatively wide and shallow, and is usually filled with little cementum. The Order Lagomorpha Brandt,+2// lingual wall of the hypocone protrudes more Family Ochotonidae Thomas,+231 lingually than that of the protocone. The pos- GenusOchotona Link,+13/ terior wall of the posterior loph is nearly Ochotona youngi Erbajeva and Zheng,,**/ straight. The paraflexus is a long and narrow (Figs../῍ ) fold, which is initiated posteriorly to the para- Ochotona nihewanica: Zheng and Cai,+33+ : Contr. XIII cone and terminates lingually to the metastyle. INQUA, Inst. Vert. Paleont. Paleoanthrop. Acad. Sinica, The metacone surrounded by the paraflexus +*,ῌ +*.. remarkably protrudes out on the occlusal sur- Ochotona youngiErbajeva and Zheng,,**/ : Acta Zool. face. The enamel thickens on the lingual walls Cracov., .2A, +**ῌ +*-,. +*2 of the metacone, protocone and hypocone. Referred specimensῌ+right P, (V +/-+34+ ) ; / PorM.+ : Detached P.+ and M of ochotonids left P-- (V+/-+34,῍῍ 0 ) ; 2 right P (V +/-+341 +. ) ; are generally indistinguishable from each other 2left P.+ or M (V +/-+34+/῍ ,, ) ; . right P .+ or M owing to their morphological similarity. Thus (V+/-+34,-῍ ,0 ) ; + left M, (V +/-+34,1 ) ; , right detached upper cheek teeth determined merely , .+ M (V+/-+34,2῍ ,3 ) ; + right mandible with P. as P or M are described here ( 3῍ +- in Fig. . ). and M+- (V+/-+34-* ) ; . left P (V +/-+34-+῍ -. ) ; , In these teeth, the crown is composed of two right P- (V+/-+34-/῍ -0 ) elliptical lophs elongating bucco-lingually in Repository, locality, horizon and ageῌThe occlusal view, which have nearly the same size same asSorex sp. described above. and shape. The lingual ends of the lophs are Descriptionῌ always pointed, while their buccal ends vary P, : This tooth is represented only by one from being pointed to blunt. The anterior wall specimen (+. in Fig. ). It is shaped as a single- of the anterior loph is slightly convex anteri- lobed curved shaft, which is convex antero- orly. The hypoflexus is a narrow and long lingually. Its occlusal surface has an elliptical cement-filled fold without any crenulation on outline elongating bucco-lingually, and shows its anterior and posterior enamel walls. It ex- a simple pattern. The paraflexus, the only fold tends buccally to reach near the buccal wall of observed, separates the protocone from the met- the crown, which forms a shallow buccal fold. acone. The anterior wall of the protocone is The posterior wall of the hypoflexus, in other arranged in the same bucco-lingual line as that words the anterior wall of the posterior loph, of the metacone. Neither hypoflexus nor meta- protrudes out on the occlusal surface. The flexus is observed on these cusps. The para- posterior wall of the posterior loph is nearly flexus is filled with cementum, and extends straight. The enamel layer thickens on the postero-buccally from the anterior wall of the anterior walls of the anterior and posterior crown somewhat lingual to its center, and lophs, while it is interrupted on the buccal wall reaches near the posterior wall of the crown. of the crown. The posterior wall is very slightly convex pos-M, : The morphology of this tooth is the teriorly. The enamel layer is thin, and its di# er- same as that of P.+ or M described above, ex- entiation in thickness is not remarkable. cept for the presence of a process on the poste- P- : This tooth is shaped as a bilobed curved rior wall of the posterior loph (+. in Fig. . ). shaft, which is convex lingually, as P.+ , M and This process is pointed posteriorly or postero- M, are. But it clearly di# ers from these cheek lingually, and is set considerably lingual to the teeth in having the Uo shaped paraflexus inside center of the posterior wall. A shallow but the crown (,2῍ in Fig. . ). The protocone is remarkable fold is formed between the process coalescent with the paracone to form the ante- and the lingual end of the posterior loph. The rior loph, whose anterior wall is slightly con- bucco-lingual width of the posterior loph is vex anteriorly. The anterior loph is much somewhat smaller than that of the anterior shorter than the posterior loph formed by the loph. coalescence of the hypocone and metastyle.Mandible : The mandible is represented only ,**2῎῍ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 165

+. right P, (V +/-+34+ ) ,. left P- (V +/-+34, ) -. left P- (V +/-+340 ) .. right P- (V +/-+342 ) /. right P- (V +/-+34+* ) 0. right P- (V +/-+34++ ) 1. right P- (V +/-+34+, ) 2. right P- (V +/-+34+. ) 3. left P.+ or M (V +/-+34+1 ) +*. left P.+ or M (V +/-+34+2 ) ++. left P.+ or M (V +/-+34+3 ) +,. left P.+ or M (V +/-+34,* ) +-. right P.+ or M (V +/-+34,0 ) +.. left M, (V +/-+34,1 )

+/. left P- (V +/-+34-+ )

+0. left P- (V +/-+34-, )

+1. left P- (V +/-+34-- )

+2. left P- (V +/-+34-. )

+3. right P- (V +/-+34-/ )

,*. right P- (V +/-+34-0 )

,+. right M+ (V +/-+34-* )

Fig.. Cheek teethofOchotona youngi from the Lower Pleistocene of the Xiaochangliang site (occulsal pattern) by a poorly preserved horizontal ramus, whose part of the crown, and has nearly the same lingual and buccal faces are smooth (Fig./ ). length as the protoflexid which extends pos- The buccal face is perforated by several small tero-lingually. These two folds separate the foramina, among which the most posterior one anteroconid from the remaining part of the corresponding to the posterior mental foramen crown, but a very narrow bridge connects the is the largest, and opens probably below the dentine field of the former with that of the occlusal position of the talonid of M, . The latter. The bottom of the paraflexid is posi- maximum height of the ramus preserved (H in tioned somewhat posterior to that of the proto- Fig./ ) measures /40. mm along its buccal face. flexid. The anteroconid varies from trapezoi-

P- : This tooth is shaped as a curved trigonal dal to rectangular shapes with corners rounded prism which is convex buccally. Thus the in various degrees. No additional fold is ob- occlusal surface has a triangular outline with served on the antero-lingual and antero-buccal rounded corners, and shows a relatively com- walls of the anteroconid. The buccal end of the plicate pattern formed by one lingual and two protoconid protrudes more buccally than that buccal folds (+/ῌ ,* in Fig. . ). These folds are of the anteroconid. The hypoflexid extends filled with cementum. Among them, the para- postero-lingually, and is somewhat longer than flexid extends postero-buccally to the central the protoflexid. The hypoconid remarkably 166 Zhang, Y.Q. and Kawamura, Y. June ,**2

MeasurementsῌThe measurements of the cheek teeth are given in Table, . Comparison and discussionῌThe lagomorph remains obtained from the Xiaochangliang site are easily divided into two forms by striking di# erence in size. The smaller form comprising much more specimens than the larger one is described here. The smaller form has the fol- lowing dental characters : The Uo shaped para- flexus is present in P- . The hypoflexus lacks enamel crenulation and reaches near the buc- cal wall of the crown in P.+ , M and M , . The lingual ends of the anterior and posterior lophs are pointed in the same teeth. The posterior process of the posterior loph is present in M, . These characters indicate that the smaller form is referable to the family Ochotonidae. Two ochotonid genera,Ochotona and Ocho- Fig./ Right fragmental mandible of Ochotona youngi tonoides, have been known from the Pliocene from the Lower Pleistocene of the Xiaochang- and Pleistocene of China.Ochotonoides di# ers liang site (V+/-+34-* ; buccal view) fromOchotona in its larger size and more com- H:maximum height of the horizontal ramus plicate pattern of P- . The size of the smaller measured along the buccal face. form falls into the range of small species of

Ochotona, and its P- pattern is simpler than that elongates antero-buccally, and protrudes buc- ofOchotonoides. The smaller form is, therefore, cally far beyond the buccal end of the protoco- assignable to Ochotona. nid. The metaconid and entoconid are indistin- Ochotona species from the Pliocene and Lower guishable from each other and form a large Pleistocene of China are represented by O. la- dentine field continuous to the protoconid and greli, O. minor, O. nihewanica, O. plicodenta, O. hypoconid, because the lingual wall of the den-magna, O. youngi, O. lingtaica and O. gracilis. tine field is almost straight, and no metaflexid Among them,O. lagreli is well known by abun- is recognized. The posterior wall of the dentine dant materials from the uppermost Miocene field is also straight, and the postero-lingual and lowermost Pliocene of Inner Monglia, corner of the field is nearly right-angled. The which were first described by Schlosser (+3,. ), enamel layer thickens on the buccal walls of and subsequently re-examined in detail by Qiu the protoconid and hypoconid as well as on the (+321 ). On the basis of the description by Qiu lingual wall of the metaconid. (+321 ),O. lagreli is easily distinguishable from

M+ : The mandible described above (Fig./ ) the present form in having much larger size, bears the only M+ worthy to describe here. the posterior mental foramen positioned more

This tooth is composed of the trigonid and anteriorly, and such features of P- as the hypo- talonid with rhombic to rhomboid shapes in flexid shallower and the bottom of the proto- occlusal view (,+ in Fig. . ). The trigonid is as flexid opposite to that of the paraflexid. large as the talonid, but is higher than theO. minor was first discriminated as a new talonid. The antero-buccal wall of the trigonid subspecies ofO. lagreli by Bohlin (+3., ) in the is weakly concave to form the protoflexid. The remains assigned toO. lagreli by Schlosser hypoflexid is somewhat deeper than the meta- (+3,. ). The subsequent study on abundant re- flexid. The posterior wall of the talonid is mains by Qiu (+321 ) elevated the subspecies to slightly convex posteriorly. The enamel layer a full species,O. minor. The data given by Qiu thickens on the posterior walls of the trigonid (+321 ) discriminateO. minor from the present and talonid, as well as their buccal ends. form in the following features :+ ) size usually ,**2῍ῌ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 167

Table, Measurements of the cheek teethofOchotona youngi from the Lower Pleistocene of the Xiaochangliang site in mm

L : maximum length of crown, W : maximum width of crown, Wa : widthofanterior lophinP.- smaller, although its size range partially over- by Zheng and Cai (+33+ ) almost overlap the size laps that of the present form in some cheek ranges for the corresponding cheek teeth of the teeth,, ) the protoflexid and paraflexid of P- present form. Thus the specimens described shallower, so that the bridge between their by these authors probably belong to the same bottoms is broader,- ) the metaflexid or a weak species as the present form, namely O. youngi enamel fold on the postero-lingual wall usually as mentioned below. present in P- . O. plicodenta, O. magna, O. youngi, O. lingtaica O. nihewanicawas described by Qiu (+32/ ) as and O. gracilis were recently described as new a new species on the basis of the holotype only, species by Erbajeva and Zheng (,**/ ). The which comprises a skull in association with left following comparisons are based on the data and right mandibles from the Lower Pleisto- given by these authors. cene of the Danangou section in the NihewanO. plicodenta occurs from the Upper Miocene Basin, about+/ km south o south o east of the to Pliocene of Lingtai in Gansu. This species is , Xiaochangliang site, and was originally allo- larger than the present form in P and P- , while cated to “O. lagrelii minor ” by Zheng (+32+ ). it is smaller in M, . Additionally it is nearly as Judging from the descriptions of the authors, large as the present form in P- . This species is,

O. nihewanica di## ers from the present form in however, di erent in having such features in P- having the hypoflexus and metaflexus in P, , as plication on the anterior wall of the para- and the metaflexid in P- , although there are no flexid, the metaconid elongated more anteri- di# erences in the remaining cheek teeth, and orly, the paraflexid deeper, and the hypoconid althoughO. nihewanica is as large as larger shorter. specimens of the present form. Additionally,O. magna is obtained from the Lower Pleisto- Zheng and Cai (+33+ ) described .- specimens cene of the Danangou section. This species is , including one P- (but no P ) from the Danangou much larger than the present form, and di# ers section and allocated them toO. nihewanica. in having larger anteroconid, deeper paraflexid

Their figure for P- (0,# in Fig. ) is di erent from and shorter hypoconid. that of the holotype, but is well coincident withO. youngi occurs from the Lower Pleistocene

P- of the present form in shape and size. Moreo- of the Danangou section as well as that of ver, the measurements of the remaining teeth Majuangou which is very near to the Xiao- 168 Zhang, Y.Q. and Kawamura, Y. June ,**2 changliang site. The morphology and size of Beijing are referred toO. cf. koslowi by Pei this species are well coincident with those of (+3-0 ).O. cf. koslowi from this locality seems to the present form, which leads to the allocation be considerably larger than the present form, of the form toO. youngi (compare Fig.. and and to have the posterior mental foramen posi- Table, of the present paper with Figs. -.2 , , tioned more posteriorly on the mandible, al- and Tables0 , 1 of Erbajeva and Zheng, ,**/ ). though the information on the cheek teeth is O. lingtaica known from the Pliocene of Ling- quite limited. Remains from the Upper Cave of tai is almost as large as the present form. It Zhoukoudian are allocated toO. daurica by Pei di# ers in P- with relatively larger anteroconid ( +3.* ). Judging from his description, O. daurica with a rhomboid shape, and with shorter hypo- from this locality di# ers from the present form conid, but the di# erences are not so clear. in having much larger size, and shorter hypo-

O. gracilis known also from the Pliocene of conid and noticeable metaflexid in P- . Remains Lingtai has roughly the same size as the pre- from Jinniushan are assigned to O. hyperborea sent form, but di# ers in having narrower P- , by Zhanget al. ( +33- ). Their description indi- which is resulted from shorter hypoconid. It cates thatO. hyperborea from this locality dif- also di# ers in having shorter paraflexid in the fers from the present form in having a squarish , same tooth, whose bottom is opposite to that of outline of P and remarkable metaflexid in P- , the protoflexid. although the cheek teeth of the former are as

Additionally, Cai (+323 ) described remains of large as those of the latter except P- . Ochotonafrom the Pliocene localities near the Many species of Ochotona are known in pre- Danangou section, which were first reported by sent China. The main species are O. thibetana Cai (+321 ). Cai ( +323 ) allocated them to “O. cf. (including O. cansus ), O. roylei, O. macrotis, O. lagrelii,” O. minor and O. erythrotis. Unfortu- erythrotis (including O. gloveri ), O. kamensis, O. nately, Erbajeva and Zheng (,**/ ) did not dis- daurica, O. curzoniae, O. koslowi, O. ladacensis, cuss the relationship among these forms andO. pallasi, O. alpina and O. hyperborea. The pre- their new species mentioned above, but the sent form is compared with these species on allocation by Cai (+323 ) needs to be revised. At the basis of the descriptions of Gureev (+30. ) any rate, “O. cf. lagrelii ” by Cai (+323 ) di # ers and Erbajeva ( +322 ). Among the species, O. from the present form in having much larger macrotis, O. erythrotis, O. daurica, O. cruzoniae, size and the posterior mental foramen posi-O. koslowi, O. ladacensis, O. pallasi and O. al- tioned more anteriorly. On the other hand,O. pina are larger than the present form, and have minorand O. erythrotis by Cai (+323 ) are more the posterior mental foramen positioned more similar to the present form, but further discus- posteriorly (below M- or more posteriorly). O. sion cannot be done owing to the shortage ofroylei and O. hyperborea di# er from the present the data. form in having larger size, and shallower proto-

Ochotona species have also recorded from the flexid and paraflexid in P- so that the bridge Middle and Upper Pleistocene of China. Erba- between their bottoms is broader. Further- jeva and Zheng (,**/ ) described a new species, more in P- , the hypoconids of these species are O. zhangi, from the Middle Pleistocene of Xiao- shorter than that of the present form. O. kamen- xishan in Shandong. This species is largersis is much larger than the present form, and is than the present form, and also di## ers in P- di erent in having shallower protoflexid and which has relatively larger anteroconid, nar- paraflexid in P- which are opposite at their rower paraflexid and protoflexid, and some- bottoms, and in having shallower hypoflexid what shorter hypoconid. also in P- .O. thibetana is as large as the present Besides Xiaoxishan, several fossil localities form, but di# ers in having the posterior men- of Middle and Late Pleistocene age yield re- tal foramen positioned more posteriorly, and mains ofOchotona which have been mostly in having P- whose paraflexid is longer and assigned to such extant species asO. koslowi, extends posteriorly, and whose hypoconid is O. dauricaand O. hyperborea. Remains from shorter. The present form is, therefore, distinct Locality- of Zhoukoudian (ῌ Choukoutien) in from all the extant species compared. ,**2῎῍ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 169

Summarizing the comparison and discussion are much larger than the corresponding teeth mentioned above, the present form di# ers from ofO. youngi described above. such Pliocene and Early Pleistocene extinctP- : P- is represented by the damaged tooth species asO. lagreli, O. minor, O. nihewanica, O. showing a shape of a curved shaft which is plicodenta, O. magna, O. lingtaicaand O. gracilis, convex lingually. Its posterior loph is lost. In as well as the Middle Pleistocene extinct spe- occlusal view, the anterior wall of the anterior cies ofO. zhangi. It is also di# erent from the loph is slightly convex anteriorly. The para- Middle and Late PleistoceneOchotona remains flexus is a Uo shaped narrow and long fold which assigned or compared to such extant species as is filled with cementum. The metacone sur- O. koslowi, O. dauricaand O. hyperborea, and rounded by the paraflexus protrudes out on the from the main extant species now distributed occlusal surface. in China. On the other hand, the present formPorM.+ : The tooth determined merely as P. is coincident with the Early Pleistocene spe- or M+ is described here. The tooth is shaped as cies,O. youngi, in morphology and size, and a bilobed curved shaft which is convex lin- thus it is referable toO. youngi. gually. In occusal view, the anterior and poste- According to Erbajeva and Zheng (,**/ ), O. rior lophs show nearly the same elliptical shape youngi occurs from Layers.,- to of the Danan- elongating bucco-lingually, and have nearly gou section. The geological ages of the layers the same size. The lingual ends of the lophs are are obscure within the Early Pleistocene, be- pointed, while the buccal ends are blunt. The cause chronological data obtained by non- anterior wall of the anterior loph is slightly faunal dating methods including paleomagnet- convex anteriorly. The hypoflexus has no cre- ism are lacking. On the basis of the faunal nulation, and extends buccally to reach near comparisons, Caiet al. (,**. ) concluded that the the buccal wall of the crown. The posterior boundary between Layers01 and , and that wall of the posterior loph is slightly convex between Layers+2 and +3 were inferred to be posteriorly. The enamel layer becomes thick +42and *412 Ma, respectively. But their chro- on the anterior walls of the anterior and poste- nology is quite doubtful especially in the upper rior lophs, where it protrudes out on the occlu- part of the section, because mammalian re- sal surface. mains are few in Layers+3 to ,1 which lackMeasurementsῌ The width of the anterior reliable time markers. Therefore we believe loph (Wa) measures,*3 . mm in P- , while the that the record ofO. youngi from Layer,- by length of the crown (L) measures , . +, mm in P. Erbajeva and Zheng (,**/ ) does not always or M+ . indicate its occurrence in the Middle Pleisto-Discussionῌ The two teeth are assigned to cene. In the paleomagnetically dated sections, the larger form of lagomorphs from the Xiao- the latest record ofO. youngi is dated as+400 changliang site. This form has the U o shaped Ma at Majuangou (its age by Zhuet al., ,**. ) paraflexus in P- , and lacks enamel crenulation except for the Xiachangliang site described of the hypoflexus in P.+ or M , where the hy- here (+4-0 Ma). Thus the occurrence ofO. poflexus reaches near the buccal wall of the youngi from the Xiaochangliang site is the lat- est in the reliably dated records.O. youngi had survived at least until+4-0 Ma in northern China.

Ochotonaor Ochotonoides sp. (Fig.0 ) Referred specimensῌ+right P- ( +/-,*4+ ), + .+ left P or M (+/-,*4, ). Fig.0 Detached cheek teethoforOchotona Ochotonoi- Repository, locality, horizon and ageῌThe des sp. from the Lower Pleistocene of the Xiao- same asSorex sp. described before in this paper. changliang site (occlusal pattern) DescriptionῌThe two teeth described here +. right P-.+ (V +/-,*4+ ), , . left P or M (V +/-,*4, ). 170 Zhang, Y.Q. and Kawamura, Y. June ,**2 crown, and the lingual ends of the anterior and All the lagomorph remains show the mor- posterior lophs are pointed. These characters phological characters of the family Ochotoni- indicate the allocation of the larger form to dae. Almost all of them are isolated teeth. Ochotonidae, as in the case of the smaller form. They are easily divided into a smaller and Ochotonid genera known from the Pliocene larger forms by their striking di# erence in size. and Pleistocene of China are represented by Only two ochotonid genera,Ochotona and Ocho- Ochotonaand Ochotonoides, which are indistin- tonoides, are known from the Pliocene and guishable from each other by the morphology Pleistocene of China. The dental morphology of P-.+ and P or M , but species ofOchotona are and size of the smaller form coincide with generally much smaller than those ofOcho- those of Ochotona, which has a large number of tonoides.Most species of Ochotona are, there- species in the Pliocene and Pleistocene as well fore, distinguishable fromOchotonoides by size as the present day of China. The comparisons di# erence. On the basis of the measurements with theOchotona species indicate that the given by Qiu (+321 ), and Erbajeva and Zheng smaller form is referred toO. youngi very re- (,**/ ), however, larger individuals ofOchotona cently described as a new extinct species. The lagreli,the largest species of Ochotona in the occurrence of this species from the Xiaochang- Pliocene and Pleistocene of China, are as large liang site is the latest in the reliably dated asOchotonoides complicidens, the only repre- records, and thus this species had survived at sentative ofOchotonoides in the same age and least until+4-0 Ma in northern China. The area. The two teeth described here probably lie larger form is referred merely toOchotona or within this overlap of the size ranges of the twoOchotonoides sp., because it is very few in speci- species, although the measurements of the pre- men number, and its morphological informa- sent form are not su$ cient owing to the poor tion is limited. preservation and shortage of material. Thus Acknowledgments they cannot be determined whether to be Ocho- tonaor Ochotonoides. We thank Professor Shaohua Zheng from the IV. Summary and conclusion Institute of Vertebrate Paleontology and Paleo- anthropology, Chinese Academy of Sciences The detailed systematic descriptions given and Professor Baoquan Cai of Xiamen Univer- in this paper have brought a su$ cient under- sity for their helpful discussion. We also ac- standing of the soricomorph and lagomorph knowledge the great convenience provided for remains from the sediments of the Xiaochang- the present study from Professor Shusaku Yoshi- liang site dated paleomagnetically as+4-0 Ma. kawa and the graduate students belonging to All the soricomorph remains are isolated teeth Natural History of Anthropogene Laboratory, which show morphological characters of the Graduate School of Science, Osaka City Univer- family Soricidae. The morphology of the teeth sity. The present study was supported by the is well coincident with that of the tribe Soricini National Science Foundation of China (Grant in the subfamily Soricinae, but is distinguish- No..*.1,*+, ) and the Major Basic Research Pro- able from those of the other subfamilies as well jects (,**0 CB 2*0.** ) of MST (Ministry of Sci- as those of the other tribes of Soricinae. Among ence and Technology of the People’s Republic the genera of Soricini, such four genera as of China). Sorex, Deinsdorfia, Petenyiaand Zelceina, are known from the Pliocene and Pleistocene of References Asia, and thus they are compared with the Bohlin, B. (+3., ) A revision of the fossil Lagomorpha present remains. The comparison reveals that in the Palaeontological Museum, Upsala. Bulletin the remains are allocated toSorex. The re- of the Geological Institution of the University of Upsala,-* , ++1ῌ +/. . mains are, however, too incomplete to carry Cai, B.Q. (+321 ) A preliminary report on the Late Plio- out any specific determination within the ge- cene micromammalian fauna from Yangyuan and nus. The remains are therefore assigned to Yuxian, Hebei. Vertebrata PalAsiatica,,/ , +,.ῌ +-0 . Sorex sp. (Cῌ E) ,**2῏῎ 0 Soricomorphs and lagomorphs from the Xiaochangliang site 171

Cai, B.Q. (+323 ) Fossil lagomorphs from the Late Plio- (The Netherlands) and Hungary. Scripta Geologica, cene of Yangyuan and Yuxian, Hebei. Vertebrata1- , +ῌ +1- . PalAsiatica,,1 , +1*ῌ +2+ , pl. + . (C῍ E) Schlosser, M. ( +3,. ) Tertiary vertebrates from Mon- Cai, B.Q., Zhang, Z.Q., Zheng, S.H., Qiu, Z.D., Li, Q. and golia. Palaeontologia Sinica, Ser. C,+ , fasc. + , +ῌ +-- . Li, Q. (,**. ) New advances in the stratigraphic Storch, G., Qiu, Z.D. and Zazhigin, V.S. (+332 ) Fossil study on representative sections in the Nihewan history of shrews in Asia. Wo´jcik, J. M. and Wol- Basin, Hebei. Professional Papers of Stratigraphy san, M. (eds.) Evolution of shrews :3-ῌ +,* , Mammal and Palaeontology,,2 , ,01ῌ ,2/ . (C῍ E) Research Institute, Polish Academy of Sciences. Erbajeva, M.A. (+322 ) Pikas of the Cenozoic. ,,- p, Wei, Q. ( +331 ) The framework of archaeological geol- Nauka. (R) ogy of the Nihewan Basin. Tong, Y.S., Zhang, Y.Y., Erbajeva, M.A. and Zheng, S.H. (,**/ ) New data on Wu, W.Y., Li, J.L. and Shi, L.Q. (eds.) Evidence for Late Mioceneo Pleistocene ochotonids (Ochotonidae, evolutionῌ Essays in honor of Prof. Chungchien Lagomorpha) from North China. Acta Zoologica Young on the hundredth anniversary of his birth : Cracoviensia,.2 A, 3-ῌῌ ++1 . +3- ,*1 , China Ocean Press. (C῍ E) Gureev, A.A. (+30. ) Fauna of the USSR. Mammals, Zhang, S.S., Wang, C.L., Li, Y.S., Cui, D.W., You, Y.Z., Vol.- , Pt. +* , Lagomorpha. ,10 p, Nauka. (R) Han, D.F., Ye, X.K., Hou, L.H., Zheng, S.H., Yu, Q.L., Kawamura, Y. (+33, ) Descriptive terminology for teeth Wang, P.F., Xia, Y.M. and Chen, W.Y. (+33- ) Com- of Japanese Quaternary insectivores. Bulletin of prehensive study on the Jinniushan Paleolithic site. Aichi University of Education (Natural Science), Memoirs of Institute of Vertebrate Palaeontology .+, ,/ῌ .- . (J῍ E) and Palaeoanthropology, Academia Sinica,+3 , +ῌ Kawamura, Y. (+33- ) Method of tooth measurements +0- , pls. +ῌ +0 . (C῍ E) for Japanese Quaternary insectivores. Bulletin of Zhang, Y.Q., Kawamura, Y. and Cai, B.Q. (,**2 ) Small Aichi University of Education (Natural Science), mammal fauna of Early Pleistocene age from the .,, -+ῌ ., . (J῍ E) Xiaochangliang site in the Nihewan Basin, Hebei, Lopezo Martinez, N. ( +320 ) The mammals from the northern China. The Quaternary Research (Dai- Lower Miocene of Aliveri (Island of Evia, Greece). yonkio.12+3, Kenkyu), ,ῌ . VI. The ochotonid lagomorphAlbertona balkanica Zheng, S.H. (+32+ ) New discovered small mammals in n. gen. n. sp. and its relationships. Proceedings of the Nihowan bed. Vertebrata PalAsiatica,+3 , -.2ῌ the Koninklijke Nederlandse Akademie van We--/2 , pl. + . (C῍ E) tenschappen, Ser. B,23 , +11ῌ +3. . Zheng, S.H. and Cai, B.Q. ( +33+ ) Micromammalian fos- Pei, W.C. (+3-0 ) On the mammalian remains from sils from Danangou of Yuxian, Hebei. Contribu- Locality- at Choukoutien. Palaeontologia Sinica, tions to the XIII INQUA, Institute of Vertebrate Ser. C,1 , fasc. / , +ῌ +,+ . Paleontology and Paleoanthropology, Academia Pei, W.C. (+3.* ) The Upper Cave fauna of Choukou- Sinica : +**ῌ +-+ , Beijing Scientific and Technologi- tien. Palaeontologia Sinica, New Ser. C,+* , +ῌ +*+ . cal Publishing House. (C῍ E) Qiu, Z.D. (+32/ ) A new ochotonid from Nihewan bed Zhu, R.X., Ho # man, K.A., Potts, R., Deng, C.L., Pan, Y. of Yuxian, Hebei. Vertebrata PalAsiatica,,- , ,10ῌ X., Guo, B., Shi, C.D., Guo, Z.T., Yuan, B.Y., Hou, Y.M. ,20. (C῍ E) and Huang, W.W. (,**+ ) Earliest presence of hu- Qiu, Z.D. (+321 ) The Neogene mammalian faunas of mans in northeast Asia. Nature,.+- , .+-ῌ .+1 . Ertemte and Harr Obo in Inner Mongolia (Nei Mon- Zhu, R.X., Potts, R., Xie, F., Ho# man, K.A., Deng, C.L., gol), China.ῌῌ0 . Hares and pikas Lagomorpha : Lepo- Shi, C.D., Pan, Y.X., Wang, H.Q., Shi, R.P., Wang, Y. ridae and Ochotonidae. Senckenbergiana lethaea, C., Shi, G.H. and Wu, N.Q. (,**. ) New evidence on 01, -1/ῌ -33 . the earliest human presence at high northern lati- Repenning, C.A. (+301 ) Subfamilies and genera of the tudes in northeast Asia. Nature,.-+ , //3ῌ /0, . Soricidae. Geological Survey Professional Paper, /0/, +ῌ 1. . (C῍ E) in Chinese with English abstract or summary, Reumer, J.W.F. (+32. ) Ruscinian and early Pleistocene (J῍ E) in Japanese with English abstract, (R) in Rus- Soricidae (Insectivora, Mammalia) from Tegelen sian 172 Zhang, Y.Q. and Kawamura, Y. June ,**2

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