Chronological Sequence and Subdivision of Chinese Neogene

ELSEVIER Palaeogeography,Palaeoclimatology, Palaeoecology 116 (1995) 41-70

Chronological sequence and subdivision of Chinese Neogene mammalian faunas * Zhanxiang Qiu and ZhudingQiu Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, Beijing, China Received 21 January 1994; revised and accepted 28 September 1994

Abstract

Six land mammal ages are recognized for the Neogene of East Asia. The ages are based upon updated large and small mammal faunas and replace or redefine the land mammal stages erected by Li et al. (1984). Comparisons and correlations are made to faunas in Europe, Northern and Southern Asia, and North America, In addition, methods of utilizing mammalian faunal data to define and recognize geochronologic ages are discussed.

1. Introduction marine sequences were enhanced. In China systematic research in Neogene stratigraphy began only Chinese Neogene deposits to date have yet to in the late 1970s. Li et al. (1984) conducted an be categorically recognized, in contrast with introductory profile of recent international devel- Europe, where a standard sequence of continental opments and made preliminary correlations of the Neogene sediments and their chronologic subdivi- Chinese Neogene with that of Europe above the sions has been developed. A vast majority of level of stage/age. Research during the past decade Chinese Neogene sediments pertain to continental has gradually refined the correlation precision to facies although some marine sediments are located the stage/age level. These recent results are scat- along the southeast coastal region and in the tered throughout the literature on individual taxo- southwestern corner of the Tarim Basin, Xinjiang nomic descriptions and often out of sight of the Autonomous Region. Consequently, in China, reli- stratigraphers in general. Recently, Z.X. Qiu able chronologic subdivision of Neogene terrestrial (1989) provided a succession of Chinese Neogene sediments is best provided through the application mammal localities with their chronologic subdivi- of fossil mammals, by virtue of their high evolu- sions. Utilizing that text as a foundation, this tionary rates, abilities for far-ranging migration, paper augments the information by the inclusion and high probability for being fossilized. of recent micromammal data to provide a further After the conclusion of World War II, both basis for chronologic subdivision and correlations. Europe and North America made rapid advancements in Neogene research. Traditional subdivisions of Neogene sediments were greatly modified 2. A brief introduction to the Chinese Neogene and the precision for correlation of terrestrial to mammalian local faunas and their succession

* Translated by Will Downs, BilbyResearch Center, Northern This text adopts the terminology "local fauna" Arizona University, Flagstaff,AZ 86011-6013 USA. proposed by H.E. Wood et al. (1941). Woodburne

(1987) later defined the term as: "An assemblage specialized Oligocene fauna. The consensus of of fossil vertebrates of specific taxonomic composi- European work over the past several decades has tion recovered from one or a few sites that are shifted the Aquitanian Stage from the Oligocene closely spaced stratigraphically and geographi- into the Miocene, which pushes the Lower cally." Tedford (1970) interpreted "local fauna" Miocene boundary back to approximately 24 Ma. as a reflection of what is actually and most primar- The Lanzhou taxa Tsaganomys, Leptotataromys, ily discovered with little element of abstraction and Tataromys grangeri are elements frequently and extrapolation. The precise definition of observed in Middle to Late Oligocene assemblages; "fauna" then, is an aggregation of many taxa of however, T. suni is a derived ctenodactylid, the mammals from a few stratigraphic levels, over a type specimen of which was found in the Miocene relatively broad area, with more elements of Xiejia Local Fauna. This situation is very similar abstraction. For brevity's sake, in case no special to that of Aquitanian mammal assemblages in emphasis is needed and no ambiguity may occur, Europe. (2) Two faunas in the former Soviet the generalized term "fauna" is applied. The over- Union, Agysp6 in Kazakhstan and Benara in all arrangements of these ages and their interconti- Georgia, have similar relationships and are recog- nental correlations are summarized in Table 1 and nized as earliest Miocene. The former locality the locations of their local faunas are shown produces taxa including Desmatolagus and in Fig. 1. Tataromys cf. deflexus; the latter locality produces forms including Brontotheriidae, Tsaganomys, 2.1. Earl); Miocene." Xiejia Age Lophiomeryx, and others. These taxa clearly indicate these faunas to be a bit more archaic. If these Lanzhou Local Fauna two faunas are assigned to MN1 (Mammal Locality: North slope of Gaolanshan Mountain, Neogene Zone 1), there is no reason to exclude south of the Lanzhou Railway Station, Gansu the younger Lanzhou fauna from the Miocene. (3) Province. The appearance of Metexallerix, which is much Fossil mammals: Six species total: Metexallerix more advanced than Exallerix from the Middle gaolanshanensis, Tataromys grangerL T. sp., T. Oligocene fauna of Hsanda Gol, Mongolia may sunL Leptotataromys cf. gracilidens, Tsaganomys also favour the assignment. However, with regard altaicus (Qiu and Gu, 1988). to its evolutionary stage, Metexallerix approaches Discussion: Four formations are recognized the earliest North American Brachyerix, the record among the red beds of the Lanzhou Basin accord- of which does not exceed 21 Ma in age. ing to a 1981 report by the Gansu Regional Survey Related faunas: A similar fauna dominated by Brigade: the Xiliugou Fm., Yehucheng Fm., micromammals was discovered in 1982 within the Xianshuihe Fm., and Linxia Fm. The Yehucheng Suosuoquan Formation in the Wulonghe River Fm. is "characterized by abundant gypsum and Valley along the northern borders of the Jungar mirabilite." The aforementioned taxa are reco- Basin, Xinjiang Autonomous Region. Taxa pub- vered from abundantly gypsiferous red clays, lished from this assemblage include Prodistylomys which undoubtedly represent the Yehucheng Fm. xinjiangensis (Wang and Qi, 1989) and The regional survey brigade regards the age of Sinolagomys ulunguensis (Tong, 1989). A prelimi- these deposits as Oligocene. nary report by Y.S. Tong et al. (1987) also includes This paper assigns the age of Early Miocene to the genera Palaeoerinaceus, Tachyoryctoides, this fauna for the following reasons: (1) Its faunal Parasminthus, and Palaeogale. Furthermore, a character resembles the European fauna correlated manuscript by M.C. McKenna suggests a new to the Aquitanian marine stage. Its principal ele- species of Exallerix in this fauna. Currently, the ments consist of advanced or derived Oligocene age of the fauna is generally recognized as forms, yet the fauna lacks typical Miocene immi- Oligocene, although S. ulunguensis is demonstrably grants such as Proboscidea and Anchitherium. more derived than Late Oligocene species of the Consequently, it may be regarded as a derived and same genus. This assemblage is regarded here as Table 1 Subdivision and corellation of Chinese terrestrial Neogene faunas

.,-7,. EUROPE C H I N A Marine Paratethys Continental CONTINENTAL

AGE/ AGE/ IMAMMAL~ DIAGNOSTIC TAXA ~ AGE UNIT AGE Typic~ Related Local Locfl STAGE STAGE AGE ~ (Li, 1984) (Qiu, This Fauna Fauna N < ~' 1989) Text MICRO MAMMALS LARGE MAMMALS N Pliest. Calabrian Biharian Nihewan- Nihewan- Borsodiacldnensis, Mimomys gansunicus Equus sanmeniensis Romanian 18 ian VI ian . . Xiashagou Piacenzian Villafran- 17 Youhean Youhe Mimomysorientalis, Germanomys, Elephasyouheensis chian 16 Mazegou Jingle Daodi Chardinomyslouisi, Pliopentalagus* E" Yushean Chasmaportheteskani, Pliohyaena, ;~ 15 Jinglean V Bilike Mimomyssp. nov.* Chardmamysyusheensis :~ ~ Zanclian Dagian Ruscinian 14 Gaozhuang Hypolagus,*Rhagapodemus Hipparion( Proboscidipparion) pater, 5 m Nyctereutes,Paracamelus Ertemte HartObo Apodemusorientalis, Karnimata, 13 ~ Messinian Orientalomyssimilis, Lophocricetus Procapreolus,Meles, Martes anderssoni grabaui,Microtodon Pontian Turolian 12 Baodean (Loc.Baode 30) JilongQingyang Songshza HipparionFAUNA Baodean/ Huanggang Heterosorex.#Brachyrhizomys tetracharax. Ailurarctosorientalis, Hipparion nagriensis, Tortonian 11 IV Lufengian Lufeng Kowalskia~*Pmgonomys wui,# Yunomys Lufengpithecuslufengensis ,.d Hipparionweihoense, Dinocrocuta, 10 aahe -10 Pannonian Vallesian Bahean Wuzhong Bulong Ningxiatherium Hipparion,*Anchitherium, Acerorhinus Amuwtlsu 9 Tsaidam Xiaolongtan Democwtcetodon,#Plesiod~us,# Pros~phneus* tsaidamensis 8 Platybelodongrangeri,# Oioceros grangerl, Sarmatian ~ ~ Tunggur Etlanggang Plesiodipusleei, Heterosminthus, Alloptox Lm~'anshan gobiensis,Bellatona forsythmajori Listriodon -r- Karamaygay = .~ Serravallian Astaracian 7 ~ III ~ Lengshuigou Alloptoxminor "Hispanotherium,"Paleotragus Sansanosmiluspalmidens, Platybelodon tongxinensis,*Kubanochoerus, Chilotherium* •15 o .- Badenian 6 ['~ ['~ ' Dingjiaergou JiulongkouKoujiacun

Shanwang Xiaodian • Ansomysshanwangensis, Meinia asiaticus, ' Ursavusorientalis, Palaeotapirus ._ Langhian Diatomysshanwangensis xiejiaheensis,Lagomeryx colberti, Palaeomeryxtricornis -& Karpatian 5 .~~ II Anam~orienta~ Megt~itz.to&m aft :)ttnangian Orleanian 4 ~ ~-~ co#onga~i~~ d. brevi~* Dionysopithecusshuangouensis, Semigenetta Sihong Fangshan Puzhen ningensis,Megacricatrk~ * ~r4~ Burdigalian Oq Wuellu huaiheensis,Anchitheriura, Dorcatherium 20 Eggin- ZhangjiapinI Jiaozigou Xishui Tataromyssp.# Gomphotherium,Aprotodon i, burgian 3 t~ i~a.1 1 Xiejiaan Xiejia SuosuoquanShaw Eucricetodonyoungi, Parasminthus Diceratherium,Sinopalaeoceros xiejiaensis 2 siningensis,Tataromys suni, Aquitanian i Egerian Agenian Xi~iaan Sinolagomyspachygnathus Metexallerixgaolamshanensis, Tsaganomys Dzungariotheriumorgosense, "Lophiomeryx" •24 i 1 Lanzhou altaicus,#Tataromys grangeri * First appearance # Last appearance 44 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70

42 • P( •7

8• 33e ~26_14o !o4138ee e23 4o 5o -,._.,L .3o I 36

15 go 25 32 q 27Q e19 11 t e13

Scale ~,J 0 500km • 28 I I ~ 22 f

Fig. 1. Distribution of fossil mammal localities in China. 1 = Lanzhou. 2 = Suosuoquan. 3 = Shawa 4 = Xiejia. 5 = Zhangjiaping. 6 =

Jiaozigou. 7 = Xishui. 8 = Wuertu. 9 = Sihong. 10 = Fangshan. 11 = Puzhen. 12 = Shanwang. 13 = Xiaodian. 14 = Dingjiaergou. 15 =

Koujiaeun. 16 = Jiulongkou. 17 = Lengshuigou. 18 ~ Tunggur. 19 = Erlanggang. 20 = Lingyanshan. 21 = Karamaygay. 22 = Xiaolongtan. 23=Tsaidam. 24=Amuwusu. 25=Bahe. 26=Wuzhong. 27=Bulong. 28=Lufeng. 29=Baode (Loc 30). 30= Qingyang. 31=Jilong. 32=Huanggang. 33=Songshan. 34=Ertemte. 35=Harr Obo. 36=Gaozhuang. 37=Bilike. 38=Mazegou. 39 = Youhe. 40 = Daodi. 41 =Jingle. 42 = Xiashagou. most closely approaching the Lanzhou fauna. The lies the Shawa Fm., which correlates generally to new species of Exallerix determined by McKenna the Suosuoquan Fm. Dzungariotherium orgosense also suggests that it is similar to Metexallerix of (Qiu, 1973) and Lophiomeryx sp. (Qiu, 1965) are Lanzhou but differs from Exallerix of the Hsanda produced from the Shawa Fm. The former is an Gol Fauna. indricothere rhinoceros of a very high evolutionary Along the southern margin of the Jungar Basin grade. Although the Lophiomeryx displays several Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 45 plesiomorphic characters for the genus, it also ments (MN2b) of Europe, and differs greatly from exhibits derived states that greatly surpass other the true rhinoceros Brachypotherium. Middle to species within the genus. It is very possibly a Late Oligocene taxa such as Tsaganomys and new genus. Desmatolagus are absent from the Xiejia Local The assemblages from the three aforementioned Fauna, which differentiates the Xiejia Local Fauna localities very probably represent the earliest from the Suosuoquan Local Fauna. It may thereby Miocene fauna of northwest China. be stated that this is an even more derived assemblage but with an Oligocene complexion. Xiejia Local Fauna Additionally, there are no typical Miocene ele- Locality: One kilometer north of Xiejia Village, ments such as Proboscidea or Megacricetodon, Tianjiazhai, Huangzhong County, Qinghai indicating that the age of the Xiejia Local Fauna Province. is probably younger than the Lanzhou Local Fossil m~amals: 14 species total (Li and Qiu, Fauna but older than the Proboscidea and 1980): Sinolagomys pachygnathus, Eucricetodon Megacricetodon datum in Europe. youngL Parasminthus xiningensis (originally Plesiosminthus), P. huangshuiensis, P. lajeensis, Tataromys suni, Tachyoryctoides kokonorensis, Zhangjiaping Local Fauna Sinopalaeoceros xiejiaensis (originally Oioceros, Locality: Approximately 30 km north of ref. Chen, G.F., 1988a), Diceratherium sp. (origi- Lanzhou, Gansu Province. nally. Brachypotherium), several indeterminate Fossil mammals: Unpublished, but preliminary taxa. identifications include Tataromys, Sinolagomys, Discussion: Material is recovered from tan- Tachyoryctoides, Hyaenodon, Schizotherium, Apro- yellow mudstones bearing sands and gravels with todon, Indricotheriidae, and a suspect piece of secondary gypsum veins. Underlying the fossilifer- proboscidean tusk (Qiu et al., 1990). ous unit is 440 m of tan-red sands and mudstones, Discussion: According to the Gansu Regional and gray-green gypsiferous mudstones. At the Survey Brigade, fossils occur in the yellow sands generic level, the Xiejia fauna is similar to the Late at the base of the Xianshuihe Formation. The Oligocene Taben-buluk fauna at Danghe, Gansu fossil assemblage is notable by the possible appear- Province; however, at the species level all members ance of Proboscidea, with the remaining taxa all are new and morphologically more derived. For remnant Oligocene forms that are very large and example, the species Sinolagomys pachygnathus, morphologically derived compared to their true Parasminthus lajeensis, Tataromys sunL and Tachy- Oligocene counterparts. Most European paleontol- oryctoides kokonorensis are larger, more hyp- ogists believe the Proboscidea appear abruptly at sodont, and maintain a morphologically more European mammal Zone MN4, although some derived dentition than other species of the same recognize this event at MN3b, or at approximately genera. The presence of Eucricetodon youngi per- 19 Ma. In Asia, the earliest record of Proboscidea mits a correlation of the Xiejia fauna to faunas of is from the Bugti Hills of Baluchistan in southern Europe, where this genus appears in the Oligocene Pakistan, but a precise date for this locality is still to Early Miocene. The Xiejia Eucricetodon most not determined. Flynn et al. (1986) described a closely approaches those from the Early Miocene. totally endemic rodent fauna from this locality. The extension pattern of the dental lophs and the By extrapolation they inferred an age of 21-18.3 degree of dental sinus and cingulum reduction Ma for Bugti. Tassy (1990) estimates the appear- suggest an affinity closest to E. aquitanicus and E. ance of Proboscidea in Asia (notably the Indian infralactorensis from the European Mammal Zones subcontinent) to be at 21-23 Ma, and to possibly MN1-3. Macromammals from Xiejia are rare. A predate the first appearance in Europe. Unlike the rhinocerotid is represented by a single fourth meta- situation in Europe, the Zhangjiaping data tarsal, which in size most closely approaches illustrates that, in China, Proboscidea may possibly Diceratherium aginense from the Laugnac sedi- appear earlier than other typical Miocene elements 46 Z. Qiu, Z. Qiu/Palaeogeography. Palaeoclimatoh~gy, Palaeoecology 116 (1995) 41 70

and coexist largely with remnant Oligocene taxa. It is regrettable that the precise stratigraphic posi- The Zhangjiaping Local Fauna is hereby tenta- tion of this alleged fauna is unknown. tively correlated to the European MN3 Zone but not MN4. 2.2. Early Miocene." Shanwang Age Related faunas: Several species of mammals were discovered at Jiaozigou, west of Dongxiangxian Sihong Local Fauna and east of Lanzhou, including Dzungariotherium Locality: Encompassing Xiacaowan, orgosense, Paraentelodon macrognathus, a species Songlinzhuang, Shuanggou, Qizui, Zhengji, and of primitive rhinoceros, and a fragment of probos- other localities 20 km southwest of Hongzehu cidean tusk (Qiu et al., 1990). Formerly, Lake, Sihong County, Jiangsu Province. Paraentelodon was known only from the Benara Fossil mammals: Published taxa include-- fauna in Georgia. The Jiaozigou specimen appears Rodentia: Ansomys orientalis (Aplodontidae; Qiu, a bit more derived. Dzungariotherium is also reco- 1987), Parapetaurista tenurugosa, Shuanggouia liui vered from the Shawa Formation of the Jungar (Petauristinae; Qiu and Lin, 1986), Eutamias Basin. Although a proboscidean tusk is recovered sihongensis, Plesiosciurus sinensis (Sciuridae; Qiu from this locality, its precise stratigraphic position and Lin, 1986), Youngofiber sinensis (Castoridae; is still to be verified; hence, the Jiaozigou Local Chow and Li, 1978), Microdyromys orientalis Fauna is not regarded in this paper as a representa- (Gliridae; Wu, 1986); Primates: Dionysopithecus tive fauna for this time period. shuanggouensis ( Li, 1978), Platodontopithecusjian- The Wuertu Local Fauna is located approxi- ghuaiensis (Gu and Lin, 1983); Carnivora: Semigenetta huaiheensis (Viverridae; Qiu and Gu, mately 50 km northwest of Bayanhot, Alxa Zuoqi, 1986), Pseudalurus cf. lorteti (Aeluroidea; Qiu and Inner Mongolia. Fossils are recovered from tan- Gu, 1986); Proboscidea: Rulengchia hueiheensis yellow sandstones overlying the Oligocene (Chow and Zhang, 1983); Perissodactyla: Ulantatal Formation. A total of 13 taxa are Plesiaceratherium shanwangensis (Wang, 1965). recorded (Wang and Wang, 1990) including: Discussion: This assemblage, first discovered in Amphechinus minimus, Amphechinus sp., Sino- 1952, was designated the Xiacaowan fauna. Later lagomys cf. ulungurensis, S. sp., Distylomys qian- discoveries produced elements indicating its lishanensis, Prodistylomys xinjiangensis, Tachy- Miocene affinity (Chow and Li, 1978). Systematic oryctoides sp., Megacricetodon sp., Protalactaga investigation and excavations were conducted in sp., Gomphotherium sp., and indeterminate frag- 1981 throughout the widely distributed 90-m thick mentary material. Sinolagomys ulungurensis and Miocene deposits. The upper section of the sedi- Prodistylomys xinjiangensis are elements found in ments consists of yellow-red clays and gray sandy the Suosuoquan Local Fauna. Tachyoo,ctoides sp. conglomerates, while the lower section is domi- is asserted to be extremely close to T. kokonorensis nated by green clays. Fossil mammals were pro- from Xiejia. The Protalactaga specimen is regarded duced from the green clays and sandy gravels. here as possibly being a derived species of Li (1983) listed 47 species in the assemblage. Parasminthus. Two elements in the local fauna, In addition to the published taxa listed above a relatively primitive gomphothere, and are forms including Lanthanotherium, Primus, Megacricetodon, which first appears in Europe at Allop-tox, Sayimys, Diatomys, Megacricetodon, Zone MN4, suggest the possibility of a general age Democricetodon, Anchitherium, Dorcatherium, correlation to Zhangjiaping or possibly a little Stephanocemas, and more. It is possible that the later, but still rather older than the Sihong Local initial taxonomic list contained some misidentifi- Fauna. cations, whereas Galerix may be a species of Bohlin (1937) described the so-called Gom- Lanthanotherium; Protictitherium may be photherium-Sayimys-Kansupithecus fauna over- Semigenetta; and Palaeotragus may in actuality be lying the Taben-buluk deposits in the Danghel Palaeomeryx. River Valley which may equate to this time period. This fauna is quite different from that of Wuertu, Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 47 for remnant Oligocene taxa are extremely rare, Palaeomeryx tricornis (Qiu et al., 1985), among which only the ctenodactylid genus Sayimys Lagomeryx colberti (Yan et al., 1983). is recognized. The sciurid complex within this Discussion: This assemblage is one of the few fauna characterizes it as closely resembling the classic Miocene faunas discovered in China prior Aliveri fauna (Zone MN4) of Europe. In evolu- to the establishment of the People's Republic. The tionary level of their dentitions, two cricetodontine fossil site has a national as well as international forms are generally equivalent to Megacricetodon reputation for its high taxonomic diversity and collongensis and Demoericetodon brevis of the exquisite preservation, including insect and plant European Zone MN4. Dorcatherium first appears specimens. In 1980 the Department of State in Europe in Zone MN4. Several genera within declared this locality a national monument. Yan the Sihong Local Fauna equate to those from the et al. (1983) suggested the presence of Shanwang Local Fauna, although at the species Anchitherium based on a section of metatarsal, but level they are noticeably more primitive at Sihong. his specimen is hereby reidentified as belonging to Consequently, the Sihong Local Fauna should lie the Tapiridae. between Zhangjiaping and Shanwang, and corre- The Lagomerycidae dominate the Shanwang late to Zone MN4 of Europe. assemblage. Formerly, several species assigned to Related faunas: The Dongxuanguan Fm. at different genera were recognized. However, they Fangshang, Nanjing, Jiangsu Province produces probably all belong to the same species, Lagomeryx taxa including Spanocricetodon ningensis and colberti, which displays a relatively large range of Anchitherium (Chow and Hu, 1956; Li, 1977). In variation in antler morphology. One species of addition, the Puzhen Formation on the north bank suid was recognized: "Hyotherium" penisulus. A of the Yangzi River in the municipality of Nanjing single tooth assigned to Palaeochoerus cf. pascoi produces several species of fossil mammals (Chow (Zhang 1974) should be attributed to the former and Wang, 1964); among them is the unquestion- taxon. There may be more than one species of able Dicrocerus, which in Europe appears at Zone rhinoceros. At least one species of brachypothere MN4. These two localities may generally be is present that very closely approaches regarded as contemporaneous with Sihong, Diaceratherium (a complete skull is housed in the although it is possible that Puzhen is slightly Shandong Provincial Museum). Proboscidea and younger. Chalicotheriidae are also represented in this fauna, the latter represented by several distal parts of limb bones. Their size and morphology most Shanwang Local Fauna closely approach Chalicotherium grande. Locality: Approximately 22 km NE of Linqu The Shanwang Local Fauna shares three micro- County Seat in Shandong Province. mammal genera with Sihong, Ansomys, Fossil mammals: Published fossil mammals Plesiosciurus, and Diatomys. However, the include--Chiroptera: Shanwangia unexpectata Shanwang species are slightly larger and clearly (Young, 1977); Rodentia: Ansomys shanwangensis morphologically derived, as are the Carnivora and (Aplodontidae; Qiu and Sun, 1988), Plesiosciurus Artiodactyla. In Europe, Chalicotherium grande aft. sinensis (Sciuridae; Qiu and Lin, 1986), Meinia and the relatively elaborately antlered Lagomeryx asiatica (Petauristinae; Qiu, 1981), Diatornys shan- first appear in Zone MN5. It is highly probable tungensis (Geomyidae?; Li, 1974); Carnivora: that the Shanwang Local Fauna correlates to this Amphicyon confucianus (Young, 1937), Hemicyon time interval. (Phoberocyon) youngi, Ursavus orientalis,(Qiu, Related faunas: The type specimen of Hemicyon Z.X., 1986), Thaumastocyoninae gen et sp. nov. youngi is derived from the locality of Zhongxiang Qiu et al., 1986; Perissodactyla: Palaeotapirus (Xiaodian) (Chen, 1981). Gu (1980) described a xiefiaheensis ( Xie, 1979), Plesiaceratherium gracile single tooth as Macaca youngi from the same (Young, 1937), P. shanwangensis (Wang, 1974); locality, however this identification is suspected by Artiodactyla: Hyotherium penisulus (Zhang, 1974), Pickford (1987) and E. Delson (pers. comm.), who 48 Z. Q&, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70

believe it may represent a suid. The age of this Koujiacun Formation at Lantian, Shaanxi, where, locality probably corresponds to Shanwang. at the same locality, K. gigas and Bunolistriodon intermedius are produced. These three suids are 2.3. Middle Miocene." Tunggur Age also produced from the Dingjiaergou Local Fauna (J. Yeh, pers. comm.). Chow et al. (1978) corre- Dingjiaergou Local Fauna lated the Koujiacun Formation with the Tunggur Locality: Approximately 15km southeast of Formation and believed both are a little later in Tongxin, Ningxia Islamic Autonomous Region. age than Dingjiaergou. Chow et al. did not base Fossil mammals: Published taxa include Alloptox their correlations upon the fossil evidence from gobiensis (Wu et al., 1991), Pliopithecus sp. (Qiu the locality of Koujiacun itself, however, but from and Guan, 1986), Percrocuta primordialis (Qiu, a locality designated Gaopo on the other bank of z.x et al., 1988), Platybelodon tonxinensis (Yeh the Bahe River which produced Platybelodon et al., 1986), and Kubanochoerus lantienensis (Qiu, grangeri, Lagomeryx complicidens, and antelopine Z.X., et al., 1988). teeth. These taxa are probably even closer to the Discussion: Tongxin County is famous for its Tunggur fauna (Young, 1964). But, Koujiacun abundant production of "dragon bones." It is and Gaopo are situated on opposite banks of the possible that the fossils produced here encompass Bahe River, and the lithologic characters of the several late Tertiary stratigraphic horizons. The two localities are extremely difficult to correlate most abundant fossils are derived from the yellow directly, for Koujiacun is dominated by yellow sands in the vicinity of Dingjiaergou Village, where lithologies and the Gaopo by red lithologies the fauna is represented by Platybelodon and (Zhang et al., 1978). Whether or not these are Kubanochoerus. In addition to the aforementioned different facies of the same stratigraphic unit is five published taxa, the fauna includes Sansanosmi- extremely difficult to determine. Based on the fossil lus, Gobicyon, Hemicyon, Stephanocemas, Eotra- mammals, the Koujiacun (or Koujiacun gus, Turcoceros, Aceratheriinae, Elasmotheriinae, Formation stratotype) could correlate to the and Chalicotheriidae. Dingjiaergou sediments, both of which are clearly Although this local fauna is still under study, older than the Tunggur Locality. there is no doubt that it closely matches the The first reference to the Jiulongkou locality in Belomechetskaya Fauna in Belorussiya, Russia. Cixian County, Hebei Province, was by Young Platybelodon and Kubanochoerus dominate both (1937). Later, C.K. Hu (1959) described the Russian and Chinese assemblages with their Maerotherium cf. brevirostris from this locality. In associated taxa either identical or similar in nature. 1976, Chen and Wu described sixteen species from The Russian fauna is produced from sediments this locality, the principal taxa consisting o~ that grade into the marine Tshokrakian Stage, Carnivora: Percrocuta hebeiensis, Sansanosmilus which, on the basis of invertebrates, may be corre- palmidens; Perissodactyla: Dicerorhinus cixia- lated to a lower member of the Badenian Stage nensis, Plesiaceratherium gracile, Chilotherium sp.; (but not the lowest member which is in the Artiodactyla: Palaeomeryx sp., Turcocerus jiulong- Tarhanian Stage) at the center of the Paratethys kouensis, T. robustus, and T. stenocephalus. sequence. The Badenian may be directly correlated Percrocuta hebeiensis is almost identical to P. to the Mediterranean Langhian Stage by the miocenica from Europe, and may only be appearance of the marine taxon Praeorbulina glom- differentiated at the subspecies rank (Howell and erosa, with the basal boundary age at 16.8 Ma. Petter, 1985). The saber tooth cat-like carnivore This comprises the lower boundary of the Middle cannot be distinguished from Sansanosmilus from Miocene in Europe. Consequently the age of the Zone MN6 of Europe. The chalicothere from Dingjiaergou Local Fauna is regarded generally as Cixian is also similar to Chalicotherium grande of 16 Ma. Zone MN5 (Zapfe, 1979), although current obser- Related faunas: The type specimen for vations suggest the Cixian form is slightly more Kubanochoerus lantienensb is derived from the derived by its slightly larger size, the rapidly Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 49 increased posterior depth of the mandible, and its from Lengshuigou, whereas Dingjiaergou and relatively expanded angular lobe. Furthermore, the Koujiacun produce the bunodont Kubanochoerus. Cician Dicerorhinus is fundamentally consistent Additionally, the morphology and size of the elas- with Lartetotherium sansanense. All of these ele- motheres from Lengshuigou are a bit more derived ments indicate that Cixian Local Fauna very than those from Dingjiaergou (work in progress). closely approaches the European Zone MN6 fauna According to Zhang et al. (1978) no direct relation- at Sansan, France. Certain distinct discrepancies, ship exists between the two stratotypes of however, may be noted between this assemblage Lengshuigou and Koujiacun. Their age determin- and the Dingjiaergou Local Fauna. Although the ations are determined solely upon paleontological two faunas share the carnivore Sansanosmilus, evidence. By eliminating interference from other other taxa such as Kubanochoeros, Platybelodon, localities and utilizing only the paleontological and Elasrnotherium, found in such abundance at characteristics from each of these two localities, Dingjiaergou, are conspicuously absent at Cixian. the age of the Lengshuigou Formation is younger Furthermore, there is relatively abundant material instead of older than the Koujiacun Formation. of Turcocerus, but no horns have been recovered, and only a pair of protuberances have been noted Tunggur Local Fauna on a skull. G.F. Chen (1988a) interpreted this Locality: On the western and northern bound- feature to indicate that bovoids had not yet formed aries of a 20-40 m mesa outside Erlian, Nei (Inner) horns. Bovoid horn-cores are noted from the Mongol Autonomous Region. The northwest Turkish locality of Candir (Zone MN5) in addition corner of the mesa is approximately 60 km from to Dingjiaergou. It is extremely difficult to deter- Erlian. mine if the discrepancies between Dingjiaergou Fossil mammals: Approximately 30 species have and Jiulongkou local faunas are due to temporal been published with another approximately 30 or ecologic factors; the latter, however, appears species more recently discovered (Qiu, Z.D., 1988). more plausible. Discussion: With the exception of the Chinese Hipparion faunas, this assemblage of fossil mam- Lengshuigou Local Fauna mals is one of the most well known Miocene Locality: Western foot of Lishan Hill, approxi- faunas in China. It has been the object of a large mately 20 km east of the ancient capital of Xian, amount of research, and represents the last stage Shaanxi Province. of the Anchitherium fauna on the Eurasian conti- Fossil mammals: Selenolophodon spectabilis nent, or approximately Zone MN8 in the European (Zhang and Zhai, 1978), Hispanotherium lin- mammal chronology. tungense (Zhai, 1978), Listriodon lishanensis, Related faunas: The Shaping Formation in Palaeomeryx sp., Turcocerus lishanensis (Li and Fangxian County, Hubei Province, produces Wu, 1978). Tesselodon fangxianensis, Anchitherium aureli- Discussion: This paper contradicts the original anense, Turcocerus noverca and Listriodon robustus deductions regarding the stratigraphic relation- (Yan, 1978). The last three species are typical ships of the Lengshuigou and Koujiacun forma- elements of the Tunggnr fauna. Tesselodon is a tions (Chow, 1978; Zhang et al., 1978) by relatively derived and hypsodont elasmothere, recognizing the later deposition of the Lengshuigou more advanced than those at Dingjiaergou or Formation. A taxonomic comparison of the fossils Lengshuigou. This fauna is extremely close to from these two localities reveals that the Tunggur. Lengshuigou assemblage seems to be more derived. The Karamagay Formation exposed along the Research undertaken by Ye and Jia (1986) deter- banks of the Ulungurhe River, at the northern mines the evolutionary level of Selenolophodon margin of the Jungar Basin, Xinjiang Autonomous spectabilis to lie between Platybelodon tonxinensis Region, has produced the following published from Dingjiaergou, and Platybelodon grangeri taxa--Rodentia (Wu, 1988): Sinomylagaulus hala- from Tunggur. Genuine Listriodon is recovered magaiensis (Mylagaulidae), Atlantoxerus junggar- 50 Z. Qiu. Z. Qiu/Palaeogeography. Palaeoclirnatology, Palaeoecology 116 (1995) 41-70

ensis, A. giganteus (Sciuridae), Amblycastor Similar paleontological data are recovered from tunggurensis (Castoridae); Proboscidea (Chen, Lierbao, Minhe, Qinghai Province (Qiu et al., G.F., 1988b): Gomphotherium cf. shensiense. 1981). Apparently, the so-called Xianshuihe fauna Platybelodon sp., Zygolophodon junggarensis, is spread extensively throughout the Qinghai and Carnivora (Qi, 1989): Amphicyon ulungurensis, Gansu Provinces. Its chronological determinations Ictitherium cf. gaudryi," Perissodactyla (Ye, 1989) and correlations require further work. At the very Anchitherium cf. aurelianense, Brachypotherium sp., least, a portion of the fauna should equate to Chilotherium sp.; Artiodactyla (Ye, 1989): Tunggur. Kubanochoerus sp., Stephanocemas aft. thomsoni, The Lingyanshan sand quarry 20 km north of Dicrocerus grangerL Lagomeryx sp., Palaeomervx Nanjing, Jiangsu Province, has produced sp., Eotragus halamagaiensis, and Turcocerus nov- Tetralophodon sp., Acerorhinus sp., and Hyo- erca. A vast majority of the taxa listed above are therium cf. palaeochoerus. Twenty meters above either identical to, or slightly more primitive than the fossil unit lies a basalt dated 12.17 Ma (Shao the Tunggur taxa. Ictitheriurn gaudryi, a typical et al., 1987). This stratigraphic unit is broadly element in the Hipparion faunas, has never been equivalent to Tunggur. recovered from older sediments. Reidentifications The Xiaolongtan Local Fauna of Kaiyuan, of the original text figures in Qi (1989) reveals: Yunnan Province, has long been considered to fig. 2-1 may be a fragment of mandible from represent the last stage of the Anchitherium fauna Gobicyon, and figs. 2-2 and 2-3 may be Miohyaena. and consequently correlative to Tunggur. It con- The former taxon is a Tunggur element, while the tains the noted primate Dryopithecus keiyuanensis, latter is a genus recovered from Europe, Turkey, Hipparion is absent, and many other elements of and from stratigraphic units equivalent to the assemblage correlate to the Chinji fauna from Tunggur. In summary, the Karamagay fauna may the Siwaliks of Pakistan. After Barry et al. (1982) be generally correlated with Tunggur, although it defined the upper limit of the Chinji fauna at appears to be slightly older. approximately 9.5 Ma based upon paleomagnetic Another Chinese Miocene fauna is represented stratigraphy, complications arose regarding the by the Xianshuihe fauna, Yongdeng, Gansu age of the Xiaolongtan Local Fauna. Dong (1987) Province. The assemblage was collected by J.G. listed eleven species in the Xiaolongtan Local Andersson and is currently housed at the Fauna: the most prominent are Sivapithecus sp. Paleontologiska Institutionen in Upsala, Sweden. (= Dryopithecus keiyuanensis), Tapirus cf. yunna- Research on the specimens has been undertaken nensis, Propotamochoerus parvulus, Dicorypho- by Young (1927) and Pearson (1928), with later choerus sp., Listriodon sp., Tetralophodon xiaolong- revisions conducted by Schaub (1930). Among the tanensis, Gomphotherium cf. macrognathus, and taxa recorded are Protalactaga grabauL Hetero- Zygolophodon chinjiensis. He recognized the suids smithus orientalis, Plesiodipus leeL Para- as being derived and possibly correlative to the cricetulus schaubL Kubanochoerus gigas, and suids of the Siwalik Nagri Formation. G.F. Chen Gomphotherium. This fauna may be directly corre- (1986) suggested that the Xiaolongtan suids were lated with Tunggur, although it is impossible to misidentified, and proposed the possibility of a verify whether all the specimens were derived from relatively younger faunal nature. This suggestion the same stratigraphic unit. The Gansu Regional raises the problem of whether a direct correlation Survey Brigade describes the Xianshuihe should be made between the Yunnan faunas and Formation as an extensive package of yellow sands the Siwalik sequence on the Indian subcontinent. and gravels exceeding 300 m in thickness that Although the Yunnan mammalian faunas repre- contain fossil mammals. In fact, however, since sent dense tropical forest assemblages, there are the Early Miocene Zhangjiaping fauna is produced many European elements that are dissimilar to the from this formation, it is probable the Xianshuihe segregated Siwalik elements. The Lufeng Local fauna was derived from different stratigraphic Fauna may be correlated to Zone MNll of levels, the latest of which is equivalent to Tunggur. Europe. The Xiaolongtan Local Fauna clearly Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 51 differs from Lufeng, is noticeably older in nature, Paleontology: In excess of 20 species (Liu and would more appropriately be equivalent to et al., 1978). Zone MN8. Discussion: At Lantian, this fauna is derived from gray interbedded sandy conglomerates and yellow mudstones that unconformably underlie 2.4. Late Miocene: Baode Age (North China) typical "Hipparion red clays." A vast majority of the taxa are commonly found in the Hipparion Tsaidam Local Fauna faunas of North China, although some species are Locality: Lake Tosonhu (Tosson nor) region at clearly distinct from the younger Baode fauna, the northeastern margin of the Tsaidam Basin, such as H. weihoense and H. chiaL and more Qinghai Province. closely approach H. primigenium from the Fossil mammals: Nearly 20 species (Bohlin, Vallesian of Europe. Absent from the Bahe assem- 1937). blage is the hyaena Adcrocuta commonly found in Discussion: Bohlin neglected to publish a the Baode fauna. Instead, Dinocrocuta is present. regional geologic description or detailed account Z.X. Qiu et al. (1988) believe this latter genus to of the fossil localities. It is generally believed that be a significant element of the Vallesian Age. the fossils were recovered from the Youshashan Related faunas: The locality of Wuzhong, Fm. at the northern boundary of the Tsaidam Ningxia Autonomous Region, produces Hipparion Basin (Li, 1984). The fauna is characterized by weihoense, Tetralophodon cf. exoletus, Acerorhinus coexistence of elements of the Anchitherium fauna tsaidamensis, and Qurliqnoria cheni (Qiu, 1987), and the earliest Hipparion fauna. Cervids belonging and is considered similar in age to Bahe. Another to the older fauna include Stephanocemas, fauna similar in nature is derived from the Bulong Lagomeryx, and "Dicrocerus"; elements of the Basin, Biru County, northern Tibet (Zheng, 1980). younger fauna include Hipparion and Tetra- Although Hipparion xizangense differs from the lophodon, in addition to locally derived and species in the Bahe Local Fauna, it is still more autochthonous bovids in the fauna such as primitive than the Hipparion from the Baode Qurliqnoria, Olonbulukia, and Tsaidamotherium. If Fauna. Moreover, Chilotherium tanggulaense from it is possible to document this assemblage as being this fauna is clearly comparatively primitive, and derived from a single stratigraphic unit, it would a species of Dinocrocuta is present, indicating its thereupon undoubtedly constitute the earliest age is probably older than Baode. occurrence of the Hipparion fauna in China. Related faunas: A predominantly micromammal Baode mammalian fauna fauna was recovered in 1976, 1983, and 1986 from Locality: The most fossiliferous and earliest dis- Amuwusu, approximately 8 km west of Jurh, Inner covered locality is Locality 30 at Daijiagou, Baode, Mongolia Autonomous Region. Macrofossils Shanxi Province. The fauna represented here is include Anchitherium and Hipparion, while 18 extensively spread over North and West China, species of micromammals are recorded (Qiu, including Tibet, and is particularly concentrated 1988). Among the taxa are elements shared with in the Loess Plateau region. Tunggur including "Protalactaga," Demo- Fossil mammals: Extremely diverse taxonomi- cricetodon, and Plesiodipus, in addition to species cally, with several hundred species recorded. associated with the Hipparion fauna including Discussion: This fauna is undoubtedly the most Sinocastor and Prosiphneus. significant Cenozoic fossil assemblage due to its taxonomic diversity, quantity of specimens, and range of distribution. The productive stratigraphic Bahe Fauna unit is referred to as the Hipparion red clays, with Locality: On the northern slope of the Bailuyuan a multitude of localities found along the Yellow loess plateau, southern bank of the Bahe River, River in the Shaanxi-Shanxi Province boundary Lantian, Shaanxi Province. region, within Henan Province, and those noted 52 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41- 70

in the Shaanxi-Gansu Province region. Additional the relatively derived cervid Procapreolus, and lithologies producing this fauna include several other elements. basins in southeastern Shanxi that contain "purple fluviolacustrine facies"; the locality of Luwangfen, 2.5. Late Miocene." Lufeng Age (South China) Henan Province, and the Duodaoshi Formation of Jingrnen, Hubei Province where there are pro- Lufeng Local Fauna ductive fossiliferous marls. The numerous and Locality: Shihuiba, Lufeng County, Yunnan diverse depositionai environments that produce Province. this fauna may be due to ecologic conditions or Fossil mammals: Published taxa include temporal factors. From the perspective of the Scandentia: Prodendrogale yunnanica (Qiu, 1986); faunal characteristics, the assemblage should at Rodentia: Platacanthomys dianensis, Typhlomys least be correlated to the European MNII-12 primitivus, 72. hipparionum (Qiu, 1989), Brachy- zones, and may even be extended to include por- rhizomys nagrii, B. cf pilgrimL B. tetracharax tions of MN10 and MN13. (Flynn and Qi, 1982; Qi, 1986); Lagomorpha: Alilepus longisinuosus (Qiu and Han, 1986); Primate: Sinoadapis carnosus, Laccopithecus robus- Ertemte local fauna tus (Wu and Pan, 1985), Lufengpithecus lufengensis Locality: Approximately 4km southeast of (Wu, 1987); Carnivora: Aelurarctos orientalis (Qiu Huadexian County Seat, Inner Mongolia and Qi, 1989); Artiodactyla: Yunnanotherim sim- Autonomous Region. plex, Dorcabune progressus (Han, 1986). Fossil mammals: Over 70 species of mammals Discussion: This locality represents the most have been discovered (Schlosser, 1924: fossiliferous and taxonomically diverse late Fahlbusch, 1983). Tertiary locality in South China. Over 100 species Discussion: This Chinese late Tertiary assem- have been discovered from lignitic sandy mud- blage is predominantly composed of micromam- stones less than 10 m in thickness (Qi, 1985). A mals. It was discovered in 1919 by the Swedish majority of the paleontological data is still under paleontologist J.G. Andersson, after which over study. Flynn and Qi (1982) assigned an age of 8 30 species were published by Schlosser (1924). Ma, or perhaps younger, to the locality based on Later, revisions were conducted upon Schlosser's the correlation of rhizomyid rodents to the Dhok diagnoses by Miller (1927) and Schaub (1930, Pathan sediments of the Siwaliks of Pakistan. 1934, 1938). In 1980 extremely successful excav- Most recently Z.D. Qiu and Storch (1990) have ations were conducted under the auspices of a revised the diagnosis of a murid rodent originally joint German-Chinese cooperative program. called Parapodemus to a new species of Schlosser believed the age of the fauna to most Progonomys, a genus that first appears in the closely approach the European locality of Polgardi, Vallesian of Europe and extends only into the Hungary (Zone MN 13). Storch (1987) recognized Early Turolian (Zone MN11). Consequently, the the dental morphology of Apodemus as Lufeng fauna should probably be correlated as approaching A. primaevus and A. gudrunae from such. the European Late Turolian (e.g. zones MN 12-13). The evolutionary level of the Ertemte Karnimata is believed to lie between K. darwini 2.6. Pliocene: Yushe Age and K. huxleyi within the Selenoportax interval zone, 5-7 Ma more or less, in the Siwaliks of Gaozhuang Fauna Pakistan. The condition of Occitanomys and Locality: Approximately lOkm southeast of Micromys are similar. Relatively derived elements Yushe County seat, Shanxi Province. are also represented among the macromammals, Fossil mammals: Still under study. with the carnivores approaching the extant genera Discussion: Pliocene mammalian faunas in China Meles and Martes, in addition to the presence of have customarily been chronologically constrained Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 53 by Yushe Zone II and the Jingle Age. The Yushe Ertemte is the locality of Har Obo which was Zone II assemblage, however, was not represented formerly equated with Ertemte. Two genera found by any definitive single or several localities, nor at this locality, Rhagapodemus and Hypolagus, was it represented by a single stratigraphic unit, appear relatively late in geologic history, both but instead was established by Teilhard de Chardin occurring in the Early Pliocene. Furthermore, the on the basis of composite bioevolutionary levels. average size of Ochotona here is larger than at Some specimens were recovered from relatively Ertemte and the premolar structure is more com- low stratigraphic positions, while others were reco- plex (Qiu, 1987). These factors suggest that the vered from higher sites. The Jingle Age was erected age of Har Obo is slightly younger than Ertemte on the basis of several mammalian taxa recovered and should correlate to the base of the Gaozhuang west of the village of Hefengcun, Jingle County, Formation. Shanxi Province, where Elephas (the age of which The Bilike local fauna lies 45 km northeast of cannot exceed 3 Ma) and several clearly more Ertemte and is predominantly a microfauna shar- primitive elements are represented. Most recent ing many genera with both Ertemte and Har Obo. publications (Zhou, 1988) have still not deter- However, there are also genera absent from Bilike mined whether the fauna is derived from a single but present at the other two localities including stratigraphic unit or whether it is a composite Anatolomys, Microtodon, Lophocricetus, Microto- from different stratigraphic levels. It is apparent scoptes, and others. Moreover, Mimomys occurring from investigations conducted in the Yushe Basin here constitutes the most primitive morphology of that the general complexion of the fauna produced the genus in China to date. Also appearing at this from the central section of the Yushe Group differs locality are numerous new insectivores such as markedly from the standard Hipparion fauna. The Beremendia, Drepanosorex, and Desmana. From Gaozhuang sediments consist of an approximately the perspective of its general complexion it most 450 m thick set of interbedded gray-green clays closely approaches the German Gundersheim and yellow ferruginous sandstones, and for which fauna (MN15), and equates generally to the late the nomenclature was erected by Qiu, Jia, and stage of the Gaozhuang Formation. Guo (pers. comm.). More formal recognition of this nomenclature was undertaken by Qiu et al. (1987a). From 1988 to 1991 American and Chinese Mazegou Fauna paleontologists conducted biostratigraphic re- Locality: 12 km west of Yushe County seat in search including paleomagnetic stratigraphy of Shanxi Province in an east-west trending arroyo the region. It was determined that the entire between the villages of Zhaozhuang and Gaozhuang Formation fell within the Gilbert Baihaicun. reversed paleomagnetic chron constraining its age Fossil mammals: Data are still under study. to 5.2-3.4 Ma. At the base of the Gaozhuang Discussion: The faunal nomenclature was Formation are first appearances including derived from the Mazegou Formation, which con- Nyctereutes, Paracamelus, Stegodon zdanskiL sists of a set of interbedded yellow sands and Pliohyaena pyrenaica orientalis, Chasmaporthetes, purple-red clays that gradually phase to predomi- Hipparion (Proboscidipparion) pater, and H. nantly argillaceous sediments. The formation is (Plesiohipparion) houfenense. More typical approximately 200 m thick. Paleomagnetic data Hipparion faunal elements are absent including illustrates that its age falls within the Gauss polar- Cervavitus novorossae, Adcrocuta, lctitherium, and ity chron, with an age constraint of 3.4-2.5 Ma. Chleuastochoerus, among others. Detailed organ- Many elements of the fauna are a continuation of ization of data from the Gaozhuang Formation is the Gaozhuang, with first appearances including currently in progress. These data illustrate this Ananeus, Ochotonoides, Mimomys orientalis, rock unit to correlate to zones MN14 + 15 in the Prosiphneus paratingi, and Chardinomys louisi. The European chronology. top of the Mazegou Formation produces Elephas Related faunas: Approximately 3 km north of but, to date, not Equus. This faunal age equates 54 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70 generally to the European Early Villafranchian, or Fossil mammals: Over forty species described Zone MN16. principally by Teilhard and Piveteau (1930). Related faunas: Xue (1981) described a locality Discussion: As opinions vary, no decision has from the lower reaches of the Youhe River at been reached regarding the concept of the Nihewan Weinan, Shaanxi Province which produced Elephas fauna, its age constraints, its relationship to the youheensis, Hipparion houfenense, Sus subtriqueta, lower Quaternary boundary, and the regional cor- Cervavitus sp., Nyctereutes sinensis, Ochotonoides relation of stratigraphic cross-sections. cf. complieidens, and Mimomys youhenicus. H. Consequently, a discussion on the late Tertiary houfenense was later reidentified as /4. relationship of Nihewan is required, which entails (Proboscidipparion) pater (Qiu, Z.X. et al., 1987). the following subjects: (1) A definitive concept of Later an additional species of Mimomys was added the Nihewan fauna is required. It is proposed here to the fauna, M. orientalis (Zheng and Li, 1986). to restrict the concept to the widely recognized It does not differ greatly from the entire Mazegou fauna described by Teilhard and Piveteau (1930). assemblage, although it probably relates more The characteristics of discoveries made recently closely to the upper part of the formation. are still ambiguous, and to assign all the mammals Most recently, older fossils have been discovered found in the Nihewan region to a single assemblage beneath the Nihewan local fauna (in the restricted would increase complications leading toward a sense) within the Nihewan vicinity. The localities faunal consensus. (2) A clarification of Nihewan are distributed along both banks of the Huliuhe River in the southeast Nihewan Basin. The fossil faunal characteristics is required. The question mammals from the Dongyaozitou locality were arises whether the 1930 descriptions reflect a single published by Tang (1980) and Tang and Ji (1983). assemblage, or a mixture of taxa from different Cai (1987) erected the Daodi Formation here and faunas. The Nihewan faunal elements are hereby subsequently described additional taxa (1989). recognized as falling into the following category Several species of large mammals from based upon recognition of their evolutionary posi- Dongyaozitou are more primitive than the same tion. The principle taxa equate generally to Zone taxa shared with the Nihewan fauna (in its MN18 of the European mammal age, represented restricted sense). The Daodi fauna is predomi- by localities such as Seneze and Le Coupet, and nantly a micromammal assemblage including the which postdate Zone MN17 localities such as St. described lagomorphs Hypolagus schreuderi, Vallier. This assemblage includes Pliohyaena licenti Pliopentalagus nihewanensis, Ochotona cf. lagrelii. (Qiu, Z.X., 1987), Megantereon nihowanense, Canis and O. erythrotis. Cai's identifications illustrate chiliensis, Eucladoceros boulei, and Elaphurus bifur- many genera shared with the Bilike fauna, such as catus. Although Myospalax tingi cannot be directly Beremendia, Prosiphneus, Nannocricetus, and compared to a counterpart in Europe, its evolu- Sminthoides, but with additional first appearances tionary level is noticeably higher than the including Eucastor, Germanomys, Chardinomys, Myospalax and Prosiphneus species from lower and Rattus. The faunal age is clearly intermediate stratigraphic positions, and hence belongs with between Bilike and Nihewan (in the restricted this category. Several remnant taxa from earlier sense). stages are present in this fauna, including The Mazegou fauna is hereby selected to repre- Nyctereutes sinensis and Paracamelus gigas, in sent a biochronologic unit due to the in-depth addition to several derived or specialized forms stratigraphic research that constrains its age from older stages, the ages of which are not easily (Tedford et al., ms). determined. These include Hipparion (Probo- 2.7. Late Pliocene to Early PleBtocene: Nihewan scidipparion) sinense and Postschizotherium. All Stage (restricted sense). these factors illustrate that the main body of the fauna may be correlated to Zone MN18, although Xiashagou Local Fauna the fact that older elements have infiltrated the Locafity: Approximately 3km northeast of assemblage may not be disregarded. (3) It is Nihewan, Yangyuan, Hebei Province. necessary to resolve the stratigraphic positions that Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 55 produce Nihewan taxa. Based on information from dard, regardless of whether the inception of Weilong Huang (pers. comm.), the main body of the Nihewanian occurs at 1.9, 2.5, or 3.0 Ma, at specimens described by Teilhard and Piveteau were least a portion of this interval falls within the collected from the green clays and yellow sands in Neogene. Xiashagou Arroyo. However, in the introduction by Wang et al. (Chen, M.N, 1988), it is stated that the Xiashagou stratigraphic cross-section is a 60- 3. A discussion of several related problems m-thick complex composed predominantly of yellow sands and green clays. Two methods may 3.1. The function of terrestrial sediments and be utilized to correlate this section of deposits. mammals in the subdivisions of international Shufang Wang utilized paleomagnetic data to con- stratigraphic units clude that the inception of deposition in Xiashagou began in the Kaena reversed polarity subchron The majority of Cenozoic and pre-Cenozoic (approximately 3.0 Ma) and terminated in the international stratigraphic subdivisions is based on Olduvai normal polarity subchron (approximately marine sediments. But terrestrial sediments and 1.6 Ma). Qiang Wang and Jingzhe Wang recog- their faunas also greatly influence the recognition nized four ostracod complexes in the late Cenozoic of geologic time periods, such as the Tertiary- sediments of the Nihewan region. Ostracod com- Quaternary boundary. In the nineteenth century plex II is present in the first stratigraphic unit, of the lower Quaternary boundary was recognized Xiashagou, while the remaining overlying strati- through glacial periods, paleoanthropological graphic units contain ostracod complex III. data, and the so-called E-L-E (Elephas-Leptobos- Complex III is also distributed throughout strati- Equus) event. More recently, controversy concern- graphic units 22-43 in the relatively complete ing the lower Quaternary boundary has been exposures noted in Hongyanangou Arroyo, a sec- caused by research on terrestrial sediments and tion that has been determined to lie predominantly mammalian paleontology. This is exemplified by in the Matuyama magnetic polarity interval. paleoanthropological discoveries in the 1970's Through these interpretations, the base of the which promoted hypotheses that would extend the Xiashagou section is probably 2.5-3.0 Ma or lower Quaternary boundary further back in time. thereabouts. This deduction is in agreement with More recently, it has been proposed that the base the fossil mammal data wherein the main faunal of the Quaternary in China should correspond to body correlates to Zone MN18 but is associated the inception of loess deposition. Further influence with even older taxa that are identified from sedi- of mammalian paleontology upon the subdivision ments 1.9-2.5 Ma. (4) The relationship between of epoch (series) levels was confirmed in the 1930s the Nihewan Age and the lower boundary of the when it became fashionable to subdivide the Quaternary requires discussion. The lower bound- Neogene on the basis of horse evolution. ary of Zone MN18 in Europe is placed at 1.92 Consequently, the Miocene, Pliocene, and Ma. The main body of the Nihewan fauna lies at Pleistocene boundaries in Eurasia were recognized approximately 1.9 Ma but with the data from by the appearances of Anchitherium, Hipparion, Xiashagou the lower boundary of Nihewan sedi- and Equus respectively. mentation may extend to 2.5-3.0 Ma. Most recent Trends and developments led by modernized data from the marine sediments in Europe con- stratigraphic methods and techniques encourage strain the lower Calabrian boundary at the Vrica late Cenozoic research to gradually clarify the section, where continuous sedimentation is recog- following three complications: (1) Epoch (series) nized, in stratigraphic unit e, 30 m above the rank subdivisions based on mammalian and conti- inception of the Olduvai normal polarity event. nental stratigraphy differ greatly from those based Consequently, the lower boundary of the on marine stratigraphy. Mammalian biostratigra- Quaternary in these marine sediments is estab- phy previously recognized the lower boundaries of lished at 1.6 Ma. Utilizing this datum as a stan- the Miocene, Pliocene, and Pleistocene at 20, 12, 56 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70 and 3.4-2.4 Ma, while marine lower boundaries sions would require an alternate timetable to be for the same geochrons are currently recognized acknowledged by a multitude of stratigraphers, a at 24, 5, and 1.6 Ma (2). Although regional situation that would seemingly require lengthy and geologic synchrony may be recognized, there is time-consuming negotiations to accomplish. nearly a total absence in the geologic record of Consequently, it is hereby proposed that the synchronous global geologic events, with the marine stratigraphic subdivisions serve as the stan- exception of paleomagnetic events. Global geologic dard formal technical vocabulary to represent con- events, presumably, are the foundation for interna- tinental biostratigraphic correlation at the tional stratigraphic subdivisions, as exemplified by stage/age level (Table 1). the research upon global sychrony of glacial devel- Based upon the discussions above, international opment. Boundaries, subdivided on the basis of epoch (series) and subepochs (subseries) should be lithostratigraphy, and which are themselves based recognized as follows: upon biostratigraphy and chronostratigraphy, are (1) The inception of the marine Aquitanian frequently discordant with those of geochronologic Stage is established as the beginning of the units. An example of this lies with the stratigraphic Neogene with a radiometric date of 24 Ma. position of the Lanzhou local fauna, which lies in Previous concepts for the initiation of the Miocene, the upper section of the Yehucheng Formation, such as a radical change of mammalian faunal where the Oligocene-Miocene boundary is arbitrar- complexion or the appearance of Anchitherium in ily recognized. But arbitrary boundaries frequently the so-called Burdigalian Age, should be aban- provoke revisions of subdivisions and correlations doned. In Europe this boundary line is represented which may be rejected by stratigraphers. (3) The as the base of MN1 (Paulhiac); in the former precision and range of marine stratigraphic corre- Soviet Union beneath the Kazakhstan Agyspe lation are greatly elevated due to the application Fauna; and in China the boundary is recognized of micropaleontology. Consequently, the flourish- beneath the Lanzhou local fauna. ing development of marine stratigraphic studies (2). The inception of the Middle Miocene has expanded proportionally to late Cenozoic marine Langhian Stage is established with a date marine geology as a whole. of 16.8 Ma. In the terrestrial sequence in Western Advancements in marine stratigraphy have over- Europe this occurs at Zone MN6 (represented by shadowed the study of terrestrial stratigraphy and disturbed the establishment of late Cenozoic the locality of Sansan, France). In the Paratethys international stratigraphic units. It is believed here region the base of the Badenian Stage is recognized that international continental epochs (series) and as the Middle Miocene boundary. In Russia, this subepochs (subseries) should abide by traditional boundary is represented by the Belomechetskaya principles by applying marine stratigraphic subdi- Fauna. In China recognition of the Middle visions as their standard, regardless of their Miocene had been rather confused, with both the impractical time distribution. For historical Sihong and Shanwang Local Faunas considered reasons, several geologic units erected on the basis representative. But currently, these two faunas are of marine stratigraphy are extremely unreasonable reassigned to the Early Miocene, and the earliest when applied to the terrestrial realm. For example, Middle Miocene faunas now recognized include the 19-m.y. history of the Miocene epoch far the Dingjiaergou and Jiulongkou local faunas. exceeds the mere 3-m.y. range of the Pliocene. (3). The inception of the Late Miocene Furthermore, terrestrial stratigraphy and mamma- Tortonian Age is established with a radiometric lian subdivisions may attain the precision of date of 12 Ma. In the past this subdivision was marine correlations through recent advances in based on erroneous marine correlations and the micromammalian paleontology and paleomagnetic evolution of horses. The former lower Pliocene stratigraphy; however, the erection of a new boundary was based upon the appearance of the nomenclature to replace the former is not easily Palearctic Hipparion fauna, an event that should accomplished, for the replacement of these subdivi- now be emended to occur in the Late Miocene. Z. Qiu, z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 57

This fauna is now regarded as characteristic for malian taphonomy that confront investigators. this time period. Hence, resolution of the lithostratigraphic discrep- (4). The currently recognized Pliocene lower ancies mentioned above has caused North boundary lies at the end of the marine Messinian American and Western European biostratigraphers Stage and is provided with a radiometric date of to progress along two different paths. In North 4.8 or 4.9 Ma (paleomagnetic stratigraphers rec- America lithostratigraphic emphasis is placed upon ommend this boundary be recognized at approxi- the study of deposits filling the multitude of well- mately 5.2 Ma by an interpolation of the developed terrestrial basins, resulting in the subdi- paleomagnetic time scale to the base of the Thvera vision of numerous lithologic members, forma- paleomagnetic subchron (D. Kent, pers. comm.), tions, and groups. and which may be universally recognized in both Biostratigraphically, the concept of "zone" is marine and terrestrial sediments (N. Opdyke, pers. thereby not employed and a more generalized comm.). This boundary is represented by the category, the Mammal Age, is applied. In the late European mammal faunas at the beginning of the Tertiary of North America there are six of these Ruscinian Age. In China, this is represented by ages: Arikarean (partly Oligocene), Heming- the Gaozhuang Fauna in the Yushe Basin. fordian, Barstovian, Clarendonian, Hem- (5). The inception of the Pleistocene, or phillian, and Blancan. In practice, these ages corre- Quaternary, marine Calabrian stage is provided spond to "age" geochronologicaUy, but North with a radiometric date of 1.6 Ma. The European American stratigraphers do not utilize the mammal zone representing this period is in the European marine "stage" which is equivalent to upper section of Zone MN18. In China, it is the chronostratigraphic designation "age". The represented in the upper part of the Nihewan cause behind this discrepancy is due to strati- Formation. graphic units contained within these ages being radically different, such that the concept of stage 3.2. Problems concerning terrestrial sediments in for basin lithologies differs markedly from the subdivisions of regional stratigraphic units. concept of stage in the marine realm. Consequently, North American methodology uti- Facies changes within terrestrial sediments are lizes geochronologic units and lithostratigraphic generally more abrupt than those in marine sedi- units: member, formation, and group. Strictly ments, and these terrestrial lithostratigraphic units speaking, the nature of the North American land are frequently very limited in their surficial distri- mammal age differs from that of a true geologic bution. Frequently, a single basin will contain a age, for the land mammal age is based on an set of local stratigraphic subdivisions, as witnessed amalgamation of faunas from different biogeo- in North America, where up to 100 formations are graphic provinces. The practical application of this present in 15 separate large-scale late Tertiary type of temporal subdivision, then, is not foolproof basins. Furthermore, the quantity of fossil mam- and is merely a tool to be applied to chronology. mals is vastly inferior to that of invertebrates, It is extremely difficult to conduct direct correla- mammalian distribution is frequently very local- tions on the basis of faunas. ized, and only under exceptional circumstances is Stratigraphic work in Western Europe is it possible to recognize mammalian phyla within extremely difficult due to the numerous well-known a continuous stratigraphic section. Even more mammalian faunas that are recovered from caves exceptional circumstances lie in the recovery of or fissure deposits. Therefore, at the inception of fossil mammals from cave or fissure deposits, for investigation, emphasis is placed upon bioevolutio- after the termination of an excavation, the limited nary perspectives. Much of the chronologic pockets of sediment cannot be replicated for future nomenclature applied to European terrestrial stra- study. Currently, there are no sufficiently quantifi- tigraphy is derived from the marine nomenclature, able stratigraphic methods to apply to the unique as these terrestrial workers frequently lack strati- characteristics of radical facies changes and mam- graphic type-sections on which to base their efforts. 58 Z. Qiu, z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70

Consequently, numerous difficulties have arisen is inconvenient in practical application, and will from this application of marine stage nomencla- become more so with the passage of time. ture. Another consideration is the long-standing Mammalian ages are only mammalian zone com- history and comparatively high degree of mamma- plexes which are similar in character. In a vast lian paleontological research in Europe. In the majority of circumstances ages fail to correspond 1970s biochrons were adopted (those of P. Mein, to homologous chronostratigraphic units, or that Mammal Neogene Zones 1-17), with each "zone" is to say, lithologic entities. based upon several nearly contemporaneous faunal To date, research upon Chinese Neogene ter- complexes. Strictly speaking the concept of these restrial stratigraphy is still in a relatively early mammalian zones and the concept of a genuine stage of development, as it has not yet attained "biozone" are not equivalent, for a biozone repre- the degree upon which formal mammalian zones sents a record of fossils in rock, while mammalian are established. At this time, it is possible to faunas represent abstract spans of time. provide a chronologic sequence incorporating only Consequently, mammalian faunas do not consti- the more significant mammalian faunas and local tute a single continual body in the rock record. As faunas, and to correlate this sequence with the zones MN 1-MN 17 does not compose a complete European mammal zones (MNI-17). and unified lithologic body, it does not represent the entire range of the late Cenozoic, and merely 3.3. Chronologic subdivisions and characteristics of constitutes several unconnected and unmatched the Chinese Neogene mammal faunas sequential points in time. From the aforementioned discussions it is appar- It is possible to recognize six mammalian com- ent that in North America and Western Europe, plexes by merging each Chinese late Tertiary fauna both a theoretical and practical approach to solve and local fauna into an aggregation based upon subdivision problems in late Tertiary terrestrial degree of similarity. These complexes also repre- sediments remain elusive. Perhaps, in the final sent six phases of Neogene mammalian evolution analysis, the current foundations for late Tertiary in China. terrestrial subdivisions in North America and Xiejiaan: This age is represented by three diach- Europe may not be satisfactory for solving these ronous local faunas encompassing Lanzhou to geological problems and a new foundation should Zhangjiaping (with the possible inclusion of be erected upon mammal zonations with corre- Wuertu, or at least a portion of it) in addition to sponding stratotypes. From a theoretical aspect other included faunas. The diagnostic characteris- then, this mammal zone would be equivalent to tic of this complex includes the predominance of an assemblage zone, the composition of which remnant Oligocene genera and species. During this would be preserved in general consistency. stage many of these remnant genera evolved to However, its corresponding chronostratigraphic produce more derived species. The entire assem- unit, i.e. chronozone, would not be required to be blage may be viewed as a specialized yet impover- restricted or consistent lithologically. A stratotype ished Oligocene fauna. The Oligocene faunas of could differ in lithologic character from other China bear a clearly autochthonous complexion contemporaneous localities but its characteristic that is preserved in the Xiejiaen mammalian com- suite of mammals should be consistent. Zonation plex. The Proboscidea (Gomphotheriidae), which nomenclature may be selected from biologic, geo- immigrate from Africa, appear toward the end of graphic, or geologic nomenclature. In Western this complex, in addition to a small number of Europe the application of a sequence of MN characteristic Miocene elements such as the numbers is utilized; however, its disadvantage is Cricetodontinae. This condition is similar to that the difficulty in establishing additional emended in Europe, but is not identical. In Europe it is only zones. To date, there are seventeen late Tertiary zones MN1 and 2 that are recognized as impover- European mammal zones, six of which have been ished yet derived Oligocene faunas. Zone MN3 emended with an alphabetical suffix a or b. This represents the appearance of relatively more typical Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 59

Miocene elements, including comparatively forest phase, and the latter grassland plateau phase derived carnivores like Ursavus, the antlered cer- recognized as the Vallesian and Turolian ages. In vids, Procervulus and Stephanocemas, and the China these distinctions are not clear. Although horse Anchitherium. In Europe Zones MN3-5 are the earliest Tsaidam fauna and latest Ertemte incorporated into the Orleanian Age. In China, fauna are typical Baode grassland plateau assem- the principle components of the Zhangjiaping blages, they do contain some discrepancies, such fauna are still remnant Oligocene elements. as the addition of dense-forest taxa, including Shanwangian: This age is represented by the cervids. It is quite possible that forest taxa repre- combination of local faunas from Sihong and sent a relatively ephemeral transition phase that Shanwang. Remnant Oligocene genera and species altered from a grassland plateau to a different kind within the assemblage are nearly extinct, and of ecological system. To recognize this variety of replacing them are a large number of elements assemblages as individual units, however, would commonly observed in the Miocene of Europe be inappropriate. including Lanthanotherium, Microdyromys, Demo- The age of the Lufeng fauna from South China cricetodon, Megacricetodon, Amphicyon, Hemi- rests within the Baodean; however, its composition cyon, Ursavus, Semigenetta, Pseudaelurus, Anchi- is clearly unlike any contemporaneous fauna in therium, Dorcatheriurn, Palaeomeryx, Lagomeryx North China, and more closely approaches the and others. This assemblage represents the first ecological model reflected in contemporaneous relatively large-scale late Tertiary faunal inter- faunas of South Asia, as they share numerous change on the Eurasian continent. taxa. Apparently, there are even more European Tunggudan: This age is an agglomeration of taxa in this fauna than previously thought three diachronous local faunas from Dingjiaergou (research in prep.). This fauna, then, is charac- to Tunggur, in addition to other associated faunas. terized ecologically by its similarity to the modem The assemblage represents the continued develop- Oriental faunal region. ment of the Shanwangian complex, but with the Yushean: This age is represented by the faunas addition of several elements characteristic to Asia, at Gaozhuang and Mazegou. It is a continuation including Plesiodipus, Heterosminthus, Bellatona, of the Baode complex but with many additional Platybelodon, Chilotherium, Kubanochoerus, Turco- dense forest taxa represented by the appearance ceras, and others. Several forms appear in central of a large number of cervids. The other initial Asia that are adapted to a grassland plateau genera and species have become more derived, habitat such as the dipodid Paralactaga and the developing even more toward dense forest habitats. lagomorph Alloptox. This time period displays frequent interchanges on Baodean: This age encompasses five diachronous the Eurasian continent, while the faunal relation- faunas from the Tsaidam Basin to Ertemte, Inner ships between Asia and North America are also Mongolia, in addition to other related localities. clearly strengthened. Immigrating from Europe are The mammalian assemblage constitutes the the genera Mimomys, Germanomys, and so-called "Hipparion fauna" phase on the Eurasian Pliopentalagus. Immigrating from North America continent. Among the micromammals, the are Hypolagus, the Camelidae, and Canidae. Muridae appear and rapidly evolve and radiate to Concurrently, several genera emigrated from Asia various regions, while the extant Cricetinae replace to North America, including Agriotherium, the archaic Cricetodontinae. As many as 100 Eomellivora, and Plesiogulo. species of macromammals, including many forms Nihewanian: This age consists of the Nihewan of Hipparion, were adapted to grassland plateau fauna in its restricted sense. The lower section of habitats as represented by certain species of hyae- this complex falls within the late Tertiary, while nids, saber-toothed cats, Tetralophodon, Zygo- the upper boundary lies outside of the scope of lophodon, Chilotherium, Chleuastochoerus, and var- this text and will not be discussed further. From ious species of antelopines and bovines. In Europe, the perspective of the total faunal body complex- this phase may be divided into the early dense- ion, there does not appear to be any great differ- 60 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclirnatology, Palaeoecology 116 (1995) 41-70

ence between it and the Yushean. Possibly the a single locality to include faunas from three most notable event within the complex is the chronologic levels. Furthermore, the upper bound- appearance of Equus. In order to justify the ary of this age does not coincide with the upper Nihewanian as an individual unit, careful consider- boundary of the European Aquitanian, which only ation must be provided to noticeable geologic encompasses zones MN1-2. Zone MN3 correlates events that occurred between the Mazegou and to the Zhangjiaping Local Fauna with a general Nihewan (depositional hiatuses, inception of loess complexion that unifies it with the older Chinese deposition, climatic deterioration, etc.). Careful localities. Consequently, it is hereby regarded as consideration must also be provided to historical part of the same age. and conventional factors, as similar phenomena (3) Chronologic adjustments have been made may be observed in both Europe and North upon the positions of several faunas within the America. In Europe the faunas of the Ruscinian Shanwangian and Tunggurian ages. For instance, the Jiulongkou and Lengshuigou local faunas have and Villafranchian ages do not differ greatly, while been transferred from the Shanwangian into the in North America, after the Hipparion faunas, or Tunggurian Age; the Zhongxiang (Xiaodian) from the Blancan stage upward to the Olduvai Local Fauna has been transferred from the normal paleomagnetic subchron, the faunas Tunggurian to Shanwangian Age, etc. appear consistent. (4) The Bahean Age has been eliminated. The The six complexes mentioned above are agglom- Baode Age now represents the age of the Hipparion erates based upon mammalian characteristics. faunas in its entirety. In China this fauna is These complexes are regarded both implicitly and chronologically imprecise and does not correlate categorically as equivalent to the mammalian ages exactly with that of Europe. utilized currently by North American and (5) The Yushean Age replaces the former European workers. It is proposed here that the six Jinglean Age and incorporates a portion of the complexes be regarded as representing Chinese former Youhean age. The precise characteristics Neogene mammal ages. The Lufeng fauna, how- of the Jingle Local Fauna are still vague. It is ever, should be regarded as exclusive and indepen- proposed here that the nomenclature be elimi- dent from the North China ages. In 1984 Li et al. nated, as the stratigraphy is well developed in the proposed a mammalian subdivision chart and Yushe region, fossil mammals are abundant, and nomenclature based upon the data as then recog- diagnostic faunal characteristics are more explicit. nized. However, Li et al.'s chart was fundamentally In the future, it will be possible to erect stratotypes subordinated to European subdivisions. Currently, of chronozones for mammal zones in this region. several distinguishing factors are identified in the (6) The subsequent use of the classical concep- historic development of Chinese Neogene mam- tion for Nihewanian is regarded as follows: The mals that differ from those of both North America lower boundary is established generally at 2.5-3.0 and Europe. Subdivisions, therefore, should be Ma. In this manner it includes the late Youhean erected to conform to the actual conditions in Age erected by Li et al. Because the Youhean has China. A new subdivision chart is hereby proposed now been subsumed by the Yushean and that derives from the chart of Li et al. with the Nihewanian, it has lost its value as a formal body. following amendments and alterations: From another aspect, early Nihewanian is within (1) The new correlation chart displays the chro- the realm of the Tertiary Period. If the upper nologic sequence of mammalian faunas within boundary of the Nihewanian is established later each age. In general it is possible to correlate the than 1.6 Ma, the Tertiary-Quaternary boundary main body of the sequence to the European would fall within the Nihewan Age. mammal Neogene (MN) zones. Chinese mammalian zones are not formally erected here; further Acknowledgements refinement of temporal precision will later replace these transition phase age units. The authors and translator hereby express their (2) The Xiejiaen Age has been expanded from gratitude for the opinions of colleagues Richard Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 61

A. Tedford, Lawrence J. Flynn, Chuankui Li, Eutamias sihongensis Renjie Zhai, Shaohua Zheng, Guoqin Qi, and Plesiosciurus sinensis Sciurinae gen et sp. indet. Qinqi Xu, as numerous beneficial recommenda- Youngofiber sinensis tions were adopted into the text. Appreciation is Microdyromys orientalis particularly expressed to S. David Webb and Sayimys sp. Everett H. Lindsay for important suggestions, Rhizomyidae gen. et sp. indet. corrections, and modifications adopted into the Diatomys cf. shantungensis text. cf. Cricetodon sp. Megacricetodon sp. Democricetodon sp. 2primus sp. 2Neocometes sp. Appendix 1--Faunal list for the major reference Cricetidae gen. et sp. indet. fauna (~paper in press; 2determined by specimen, LAGOMORPHA waiting for description) Alloptox sp. CARNIVORA ?Amphicyon sp. Lanzhou Fauna Ursidae gen. et sp. indet. INSECTIVORA Metexallerix gaolanshanensis Proputorius sp. RODENTIA Semigenetta huaiheensis Tataromys grangeri Pseudaelurus cf. lorteti T. suni Mustela sp. T. sp. ?Protictitherium sp. PROBOSCIDEA Leptotararomys cf. gracilidens Tsagannomys altaicus Rulengchia huaiheensis Xiejia Fauna PERISSODACTYLA RODENTIA ?Anchitherium sp. Atlantoxerus sp. Plesiaceratherium shanwangensis Eucricetodon youngi ARTIOCACTYLA Plesiosminthus xiningensis Suidae gen. et sp. indet. P. huangshuiensis Pecarichoerus sp. P. lajeensis Dorcatherium orientale Tataromys suni Micromeryx sp. T. sp. Dicrocerus sp. Tachyoryctoides kokonorensis ,4mphimoschus sp. LAGOMORPHA Stephanocemas sp. Leporidae gen. et sp. indet. Lagomeryx sp. Sinolagomys pachygnathus Palaeotragus sp. S. cf pachygnathus CETACEA CARNIVORA Delphinus sp. Mustelidae gen. et sp. indet. PRIMATES PERISSODACTYLA Dionysopithecus shuangouensis ?Diaceratherium sp. Platodontopithecus jianghuaiensis ARTIODACTYLA Hominoidea indet. Sinopalaeoceros xiejiaensis Shanwang Fauna Sihong Fauna CHIROPTERA INSECTIVORA Shanwangia unexpectula Lanthanotherium sp. RODENTIA Crocidosorex sp. ,4nsomys shanwangensis CHIROPTERA Plesiosciurus aft. sinensis Myotis sp. Meinia asiatica Vespertilionidae gen. et sp. indet. Diatomys shantungensis RODENTIA CARNIVORA Ansomys orientalis Amphicyon confucianus Parapetaurista tenurugosa Hemicyon youngi Shuanggouia lui Ursavus orientalis 62 Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70

Thaumastocyoninae gen. et sp. indet. Selenolopodon spectabilis PROBOSCIDEA PERISSODACTYLA Gomphotherium sp. Hispanotherium lingtungensts PERISSODACTYLA ARTIODACTYLA Palaeotapirus xiejiaheensis Listriodon lishanensis Chalicotherium sp. Palaeotragus sp. Plesiaceratherium gracile Stephanocemas sp. P. shanwangensis Palaeomeryx sp. Diaceratherium sp. ?Micromeryx sp. ARTIODACTYLA Turcocerus lishanensis Hyotherium penisulus Tunggur Fauna Palaeomeryx tricornis qNSECTIVORA Lagomervx eolberti Mioechinus? gobiensis Dingjiaergou Fauna M.? sp. 2INSECTIVORA Erinaceinae gen. et sp. indet. Erinaceidae gen. et sp. indet. Proscapanus sp. Talpidae gen. et sp. indet. Yanshuella sp. Soricidae gen. et sp. indet. Quyania sp. 2CHIROPTERA Desmanella sp. Chiroptera gen. et sp. indet. Talpidae gen. et sp. indet. 2RODENT1A Mongolosorex qiui Sayimys sp. Soricinae gen. et sp. indet. Tachyorvctoides sp. Soricidae gen. et sp. indet. Atlantoxerus sp. ICHIROPTERA Steneofiber sp. Chiroptera gen. et sp. indet. ?Leptodontomys sp. RODENTIA Prodryomys sp. 1A nomys? sp. Heterosminthus orientalis 1Eutamias aft. ertemtensis Protalactaga grabaui Sinotamias primitivus Paralactaga sp. LAtlantoxerus sp. Megacricetodon sp. Anchitheriomys tungurensis Democricetodon sp. Monosaulax tunggurensis LAGOMORPHA tHystricops? sp. Alloptox gobiensis Leptodontomys lii 1 CARNIVORA L. aft. gansus 1 Tongxinictis primordialis Keramidomvs fahlbuschi I Gobicyon sp. 1Microdyromys wuae 1 HemicTon sp. l Miodyromys sp. Sansanosmilus sp. t Heterosminthus orientalis PROBOSCIDEA Protalactaga grabaui 1 Platybelodon tongxinensis P. major 1 PERISSODACTYLA 1Gobicricetodon flynni j Caementodon tongxinensis ~G. robustus 1 Chalicotherium sp. iG. sp" 1 ARTIODACTY LA 1Plesiodipus leei Kubanochoerus lantienensis tP. progressus ~ Stephanocemas sp. Megacricetodon sinensis ~ Eotragus sp. M. pusillus a Turcoceros sp. Democricetodon lindsayi ~ PRIMATES D. tongi ~ Pliopithecus zhanxiangi LAGOMORPHA Lengshuigou Fauna ~Desmatolagus? moergenensis LAGOMORPHA Alloptox gobiensis Alloptox minor Bellatona forsythmajori Tsaganolagus wangi CARNIVORA PROBOSCIDEA Gobicyon macrognathus Gomphotherium shensiensis Pseudarctos sp. Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 63

Hemicyon teilhardi Soricinae gen. et. sp. indet. Amphicyon tairumensis Yunoscaptor scalprum Leptarctus neimonguensis Talpinae gen. et sp. indet. Melodon sp. CHIROPTERA Mionictis sp. Pteropidae gen. et sp. indet. Martes sp. Hipposideridae gen. et sp. indet. Tungurictis spocki Myotis sp. Percrocuta tungurensis Epthsicus sp. Metailurus mongoliensis Pipistrellus sp. Machairodus sp. Plecotus sp. PROBOSCIDEA RODENTIA Platybelodon grangeri Tamiops sp. Serridentinus gobiensis Sciurotamias sp. Zygolophodon sp. Callosciurus sp. PERISSODACTYLA Dremomys sp. Anchitherium gobiensis cf. Albanensia sp. Chalicotherium brevirostre Hylopetes sp. Chalicotheriidae gen. et sp. indet. Castoridae gen. et sp. indet. Rhinocerotidae gen. et spp. indet. Platacannthomys dianensis ARTIODACTYLA Typhlomys primitivus Listriodon mongoliensis T. hipparionum Kubanochoerus sp. Leptodontomys sp. Stephanocemas thomsoni Eomyidae gen. et sp. indet. Dicerocerus grangeri Brachyrhizomys nagrff Dicerocerus sp. B. cf pilgrimi Micromeryx sp. B. tetracharax Lagomeryx triacuminatus 1Kowalskia hanae Euprox sp. Progonomys yunnanensis Palaeotragus tungurensis Yunomys wui Turcocerus grangeri Hystrix sp. T. noverca LAGOMORPHA Tsaiflam Fauna Alilepus longisinuosus CARNIVORA CARNIVORA ?Ictitherium sp. Ursavus sylvestris PROBOSCIDEA U. sp. ?Tetralophodon sp. lndarctos sinensis PERISSODACTYLA L sp. Acerorhinus tsaidamensis Ailurarctos lufengensis Hipparion sp. Martes cf. palaeosinensis ARTIODACTYLA M. sp. ?Stephanocemas sp. Mustelinae gen. et. sp. indet. Lagomeryx tsaidamensis Eomellivora wimani ?Dicrocerus sp. Melinae gen. et sp. indet. Eostylocerus sp. Proputorius lufengensis Qurliqnoria cheni P. sp. Tossunnonia pseudibex Sivaonyx bathygnathus Tsaidamotherium hedini Lutra sp. Lufeng Fauna Lutrinae gen. et sp. indet. SCANDENTIA Mustelidae gen. et sp. indet. Prodendrogale yunnanica Viverra sp. INSECTIVORA Viverrinae gen. et sp. indet. Lanthanotherium sanmigueli Paradoxurinae gen. et sp. indet. Hylomys aft. suillus Viverridae gen. et sp. indet. Heterosorex wangi Ictitherium gaudryi Anourosorex oblongus I. sp. Blarinella sp. Epimachairodus fires Sorex sp. Pseudaelurus sp. 64 Z Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70

Felis sp. ?Ly~Taena dubia PROBOSCIDEA Machairodus palanderi Gomphotherium sp M. tingii Serridentinus sp. Metailurus major Zygolophodon lufengensis M. minor PERISSODACTYLA PROBOSCIDEA Hipparion spp. Tetralophodon exoletus Chalicotherium salinum PERISSODACTYLA C. sp. Dicerorhinus orientalis Tapirus sp. Chilotherium habereri Chilotherium sp. C. planifrons Aceratherium sp. C anderssoni ARTIODACTYLA ,4 eerorhinus palaeosinensis Hyotherium sp. Sinotherium lagrelii H. cf. palaeochoerus Hipparion (Hipparion) hippidiodus Lophochoerus lufengensis H. (H. ) dermatorhinum Potamochoerus spp. H. (H.) fossatum Suidae gen. et sp. indet. H. (H. j plocodus Dorcabune progressus tL ( Cremohipparion ) Jorstenae Yunnanotherium simplex ARTIODACTYLA Moschus sp. Chleuastochoerus stehlini Dicrocerus sp. Microstonyx major Metacervulus cf. simplex Propotamochoerus hyothericides Metacervulus spp. Cervocerus novorossiae Muntiacus cf. nanus Procapreolus latifrons M. spp. Palaeotragus mierodon Cervidae gen. et sp. indet. P. cf. eoelophrys Selenoportax sp. Samotherium sp. Bovidae gen et. sp. indet. Honanotherium schlosseri PRIMATES Urmiatherium intermedium Sinoadapis carnosus Plesiaddax depereti S. shihuibaensis Tragocerus spectabilis Laccopithecus robustus Gazella paotehensis Lufengpithecus lufengensis G dorcadoides Baode Fauna (Loc. 30) G. altidens RODENTIA ?Tragoreas lagrelii Sinocastor zdanskyi ?T anderssoni CARNIVORA ?72 palaeosinensis? Simocyon aft. primigenius Sinotragus wimani Indarctos lagrelii Paraprotoryx minor L sinensis Palaeoryx sinensis Sinictis dolieognathus Ertemte Fauna Mustela palaeosinensis INSECTIVORA Proputorius minimus Erinaceus mongolieus Plesiogulo brachygnathus Erinaceidae gen. et sp. indet. Lutra aonychoides Quyania chowi Parataxidea sinensis Yanshuella primaeva P. crassa Talpinae gen. et sp. indet. Melodon majori ?Anourosorex sp. ?M. ineertum Neomyini gen. et sp. indet. Promephitis cf. maeotica Crocidura kormosi Eomellivora wimani Blarinella sp. nov. Ietitherium sinense Alluvisorex sp. I. gaudryi Sorex spp. Thalassietis wongii Soricinae gen. et sp. indet. Hyaenictitherium hyaenoides CHIROPTERA Aderocuta variabilis Chiroptera gen. et sp. indet. Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 65

RODENTIA Palaeotragus microdon Pseudaplodon asiaticus Axis speciosus Eutamias ertemtensis Procapreolus rutimeyeri Sciurus sp. P. latifrons Sinotamias gravis Paracervulus brevis Spermophilus orientalis Moschus grandaevus Pliopetaurista rugosa Gazella sp. Petinomys auctor Gaozhuang Fauna Sinocastor anderssoni INSECTIVORA Dipoides cf. majori Soriculus praecursus Myomimus sinensis Blarinini gen. et sp. indet. Leptodontomys gansus Yanshuella primaeva Sicista sp. Desmana kowalskae Eozapus similis Scaptochirus sp. Lophocricetus grabaui RODENTIA L. pusillus 2Eutamias cf. ertemtensis Paralactaga anderssoni 2Tamiasciurus sp. Brachyscirtetes wimani 2Sinotamias sp. Sminthoides fraudator Pliopetaurista rugosus Sinocricetus zdanskyi Sinocastor anderssoni Nannocricetus mongolicus aft. Dipoides majori Kowalskia neimengensis Hystrix sp. K. similis Myomimus sp. Microtodon atavus Sminthoides fraudator Anatolomys teilhardi Kowalskia spp. Pseudomeriones abbreviatus Nannocricetus mongolicus Prosiphneus eriksoni Allocricetus sp. Microtoscoptes praetermissus Prosiphneus truncatus Apodemus orientalis P. eriksoni Orientalomys cf. similis P. praetingi Karnimata hipparionum Germanomys sp. Occitanomys pusillus Mimomys sp. Micromys chalceus Apodemus qiui LAGOMOPHA Micromys chalceus Alilepus annectens M. tedfordi Ochotona lagreli aft. Karnimata hipparionum O. minor Chardinomys yusheensis CARNIVORA Huaxiamys primitivus Meles suillus H. downsi Promephitis alexejevi LAGOMORPHA Martes anderssoni Ochotona lagreli M. sp. Alilepus sp. Ictitherium aft. hipparionum Trischizolagus sp. Viverridae gen. et sp. indet. Hypolagus sp. Hyaena sp. CARNIVORA Machairodus sp. Agriotherium sp. Felis sp. Ursus sp. PROBOSCIDEA Plesiogulo brachygnathus Mastodon sp. Martes sp. PERISSODACTYLA Thalassictis sp. Hipparion richthofeni Chasmaporthetes kani H. sp. Pliohyaena pyrenaica orientalis Sinohippus zitteli Metailurus sp. Chilotherium habereri Nyctereutes tingi ARTIODACTYLA N. sinensis Propotamochoerus hyotherioides Canis sp. Honanotherium sp. PROBOSCIDEA 66 Z Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70

Mammut borsoni Ursus sp. Anancus sinensis Meles sp. Sinomastodon intermediu,~ Chasmaporthetes sp. PERISSODACTYLA Pliohyaena sp. ILL (Hipparion ) platyodus Croeuta sp. H. (Plesiohipparion ) houJenense Metailurus sp. H. (Proboscidipparion ) pater Machairodus sp. H. (Baryhipparion ) insperatum Homotherium sp. H. (Cremohipparion ) licenti Lynx sp. Dicerorhinus orientalis Civettictis sp. Acerorhinus sp. ,'~rctereutes sinensis ARTIODACTYLA Canis sp. Sus erymanthius Vulpes baihaiensis Paracamelus sp. PROBOSCIDEA Moschus sp. Stegodon sp. Cervocerus novorossiae Ananeus sp. Paracervulus cf. killgusi Sinomastodon intermedius Procapreolus sp. Archidiskodon sp. Axis speciosus HYRACOIDEA Cervavitus demissus Postschizotherium sp. Metacervulus sp. PERISSODACTYLA Muntiacus cf. lacustris H. (Hipparion) platyodus Gazella blacki H. (Proboscidipparion) pater cf. Protoryx yushensis H. ( Baryhipparion ) insperatum cf. Tragoceras laticornis Dicerorhinus sp. Mazegou Fauna ARTIODACTYLA INSECTIVORA Sus sp. cf. Erinaceus sp. Paracamelus sp. cf. Blarinoides sp. nov. Palaeotragus sp. Peisorex pliocaenicus Axis sp. Yanshuella primaeva Muntiacus sp. Scaptochirus sp, Dama sp. RODENTIA Rusa sp. Sciurus sp. Gazella blacki Tamiasciurus sp. Antilospira licenti Pliopetaurista rugosus Sinorvx cornucopia Sinocastor sp. Caprini gen. nov. Dipoides sp. Megalovis sp, Hystrix sp. Lyrocerus sata Myomimus sp. Xiashagou Fauna Smmthoides fraudator INSECTIVORA Rhizomys shansius Erinaceus cf. dealbatus Allocricetus sp. RODENTIA ?Cricetul~ sp. Alactaga cf. annulata Prosiphneus praetingi Foungia tingi Germanomys sp. Borsodia chinensis Mimomys cf. orientalis Hystrix sp. Apodemus zhangwagouensis LAGOMORPHA Mieromys tedfordi Oehotonoides complieidens Chardinomys nihewanieus CARNIVORA LAGOMORPHA Vulpes chikushanensis Ochotonoides complicidens Nyctereutes sinensis Oehotona sp. Canis chiliensis A6lepus annectens Urn'us etruscus Hypolagus schreueri Pliohyaena licenti CARNIVORA Chasmaporthetes cf. ossiJiTgus Agriotherium sp. Crocuta honanensis Z. Qiu, z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41-70 67

Lutra licenti Chen, G.F., 1988a. The phylogenetic status of Oioceros from Mustela pachygnatha the Early and Middle Miocene of China. Vertebr. PalAsiat., Meles chiai 26(3) (in Chinese). Lynx sp. Chen, G.F., 1988b. Middle Miocene Proboscidea from the Acinonyx pleistoceneus North Bank of the Ulungur He River, Jungar Basin, Megantereon nihowanensis Xinjiang Autonomous Region. Vertebr. PalAsiat., 26(4) Homotherium cf. crenatiders (in Chinese). PROBOSCIDEA Chen, G.F. and Wu, W.Y., 1976. Miocene Mammalia from Palaeoloxodon namadicus (Probably from overlapping Jiulongkou, Cixian, Hebei Province. Vertebr. PalAsiat., deposits) 14(1) (in Chinese). PERISSODACTYLA Chen, M.N. (Editor), 1988. Research on the Nihewan Deposits. Dicerorhinus yunchuensis Ocean Press, Peking (in Chinese). Coelodonta sp. Chow, M.C., 1978. Tertiary mammalian faunas from the Elasmotherium sp. Lantian Region of Shaanxi Province. In: Collected Papers Nestoritherium sp. of Paleontology and Stratigraphy 7. Geol. Publ. House, Postschizotherium chardini Peking (in Chinese). Proboscidipparion sinense Chow, M.C. and Hu C.K., 1956. The discovery of Miocene Equus sanmeniensis mammals from Fangshan, Nanjing. Vertebr. PalAsiat., 4(4) Equus teilhardi (in Chinese). ARTIODACTYLA Chow, M.C. and Wang, B.Y., 1964. Miocene vertebrates from Sus cf. lydekkeri Xiacaowan. Sihong, and Puzhen, Nanjing, Jiangsu Province. Paracamelus gigas Vertebr. PalAsiat., 8(4) (in Chinese). Cervulus bohlini Chow, M.C. and Li, C.K., 1978. "The Xiacaowan System," Elaphurus bifurcatus "Trogontherium," and "The Wei River Transition Phase"-- Eucladoceros boulei Corrections of historical misunderstandings. J. Stratigr., Cervus elegans 2(2) (in Chinese). Gazella sinensis Chow, M.C. and Zhang, Y.P., 1983. Stegotetrabelodon in G. subgutturosa China. Vertebr. PalAsiat., 21(1) (in Chinese). Spirocerus wongi Dong, G., 1987. Further discussions upon the character and S. peii age of the Xiaolongtan mammalian fauna, Kaiyuan, Yunnan Antilospira robusta Province. Vertebr. PalAsiat., 25(2) (in Chinese). Ovis shantungensis Fahlbusch, V., Qiu Zhuding and Storch, G., 1983. Neogene ?Budorcas sp. mammalian faunas of Ertemte and Har Obo in Nei Mongol, Bison paleosinensis China--Report on field work in 1980 and preliminary results. Sci. Sin. B, 26(2). Flynn, L.J. and Qi Guoqin, 1982. Age of the Lufeng, China, hominoid locality. Nature, 298. References Flynn, L.J., Jacobs, L. and Cheema, I.U., 1986. Baluchimyinea, a new ctenodactyloid rodent subfamily from the Miocene of Barry, J.C., Lindsay, E.H. and Jacobs, L.L., 1982. A biostrati- Baluchistan. Am. Mus. Nov., 2841. graphic zonation of the middle and upper Siwaliks of the Gabunia, L.K., 1964. La faune de mammif6res de l'Oligoc~ne Potwar Plateau of North Pakistan. Palaeogeogr. du Benera. Metcniereba, Tbilisi. Palaeoclimatol. Palaeoeeol., 37: 95-130. Gabunia, L.K., 1973. The Belomechetskaya fauna of fossil Bi, X.X. , 1981. An Early Pleistocene mammalian fauna and vertebrates. Metcniereba, Tbilisi, p. 93. its stratigraphic position from Weinan, Shaanxi Province. Gansu Provincial Investigative Brigade, 1984. The Tertiary Vertebr. PalAsiat., 19(1) (in Chinese). System of Gansu Province. In: The Geology of Gansu. Bohlin, B., 1937. Eine Tertiare S~tugetier-Fauna aus Tsaidarn, Gu, Y.M., 1980. A Pliocene tooth of Macaca from Zhongxiang, Sino-Swed. Exped. Pal. Sin., C, 14(1). Hebei Province. Vertebr. PalAsiat., 18(4) (in Chinese). Cai, B.Q., 1987. Late Pliocene Micromarnmals from Yangyuan Gu, Y.M. and Lin, Y.P., 1983. The first discovery of and Yuxian, Hebei Province. Vertebr. PalAsiat., 25(2) Dryopithecus from Sihong, Jiangsu Province. Acta (in Chinese). Anthropol. Sin., 2(4) (in Chinese). Chen, G.F., 1981. Amphicyon from the late Tertiary of Han, D.F., 1986. Tragulidae from the Lufeng hominoid Zhongxiang, Hubei Province. Vertebr. PalAsiat., 19(1) locality. Acta Anthropol. Sin., 5( 1) (in Chinese). (in Chinese). Howell, F.C. and Petter, G., 1985. Comparative observations Chen, G.F., 1986. A new species of Listriodon from the Middle on some Middle and Upper Miocene hyaenids, Genera: Miocene of Xinan, Henan Province. Vertebr. PalAsiat., Pererocuta Kretzoi, Allohyaena Kretzoi, Aderocuta Kretzoi 24(4) (in Chinese). (Mammalia, Carnivora, Hyaenidae). Geobios, 18(4), 68 Z. Qiu, Z. Qiu/Palaeogeography. Palaeoclimatology, Palaeoecology 116 (1995) 41 70

Hu, C.K., 1959. Ancylopoda from North China. Vertebr. Whyte (Editor), The Palaeoenvironment of East Asia from PalAsiat., 1(3): 125-132 (in Chinese) the Mid-Tertiary, Proc. 2nd Conf. Centre Asian Stud., Univ. K6hler, M., 1987. Boviden des Tt~rkischen Mioz~ins Hong Kong. (K/inozoikum und Braunkohlen der TOrkei, 28). Paleontol. Qiu. Z.D., 1989. Platycanthomyidae (Rodentia) from the Evol., 21. Lufeng hominoid locality. Vertebr. PalAsiat., 27(4) (in Li, C.K., 1974. Miocene Rodentia from Lanqu, Shandong Chinese). Province. Vertebr. PalAsiat., 12(1) (in Chinese). Qiu, Z.D, Li, C.K. and Wang, S.J., 1981. Miocene mammals Li, C.K., 1977. Miocene Cricetidae from Fangshan, Nanjing. from the Xining Basin, Qinghai Province. Vertebr. PalAsiat., Vertebr. PalAsiat., 15( 1) (in Chinese). 192 (in Chinese). Li, C.K., 1978. A Miocene hylobatid from Sihong, Jiangsu Qiu, Z.D., Han, D.F., Qi, G.C. and Lin, Y.F., 1985. Province. Vertebr. PalAsiat., 16(3) (in Chinese). Micromammals from the Lufeng hominoid locality. Acta Li, C.K. and Qiu, Z.D., 1980. Early Miocene Mammalia from Anthropol. Sin., 4(1 ). the Xining Basin, Qinghai Province. Vertebr. PalAsiat.. Qiu, Z.D. and Lin, Y.P., 1986. The Middle Miocene Vertebrate 18(3) (in Chinese). Fauna from Xiacaowan, Sihong, Jiangsu Province. 5. Li, C.K., Qiu, Z.D. and Wu, W.Y., 1984. Chinese Neogene: Sciuridae. Vertebr. PalAsiat., 24(3) (in Chinese). Subdivision and Correlation. Vertebr. PalAsiat., 22(3) Qiu, Z.D. and Han, D.F., 1986. Lagomorpha from the Lufeng (in Chinese). hominoid locality. Acta Anthropol. Sin., 5( 1) (in Chinese). Li, C.K., Lin, Y.P., Gu, Y.M., Hou, L.H., Wu, W.Y. and Qiu, Qiu, Z.D. and Sun, B., 1988. Recent discoveries of micromam- Z.D., 1983. A Middle Miocene vertebrate fauna from mals from Shanwang, Shandong Province. Vertebr. PalAsiat., Xiacaowan, Sihong, Jiangsu Province. Vertebr. PalAsiat., 26( 1 ) (in Chinese). 21 (4) (in Chinese). Qiu, Z.D. and Storch, G., 1990. New murids (Mammalia; Li, Y.T., 1984. The Tertiary of China. Geol. Publ. House, Rodentia) from the Lufeng hominoid locality, Late Miocene Beijing (in Chinese). of China. J. Vertebr. Paleontol., 10(4). Li, Y.Q. and Wu, W.Y., 1978. Miocene Artiodactyla from Qiu, Z.X., 1965. First Discovery of Lophiomeryx in China. Lantong and Lantian, Shaanxi Province. Prof. Pap. Stratigr. Vertebr. PalAsiat., 9(4) (in Chinese). Paleontol., 7 (in Chinese). Qiu, Z.X., 1973. Indricotheriinae from the Jungar Basin of the Liu, D.S., Li, C.K. and Zhai, R.J., 1978. Pliocene vertebrates Xinjiang Autonomous Region. Vertebr. PalAsiat., 11 (2) from Lantian Shaanxi Province. Prof. Pap. Stratigr. (in Chinese). Paleontol., 7 (in Chinese). Qiu, Z.X., 1987. Die Hyaeniden aus dem Rescinium und Mein, P., 1975. R6sultats du Groups de Travail Vert6br6s. In: Villarfranchium Chinas. MOnchn. Geowiss. Abh., A, 9. J. Senes (Editor), Report on Activity of the RCMNS Qiu, Z.X., 1989. The Chinese Neogene mammalian biochronol- Working Groups. Reg. Comm. Med. Neogene Stratigr. ogy--Its correlation with the European Neogene mammalian Miller, G.S., 1927. Revised determinations of some Tertiary zonation. In: E.H. Lindsay, V. Fahlbusch and P. Mein mammals from Mongolia. Pal. Sin. C, 5(2). (Editors), European Neogene Mammal Chronology (NATO Pearson, H.S., 1928. Chinese fossil Suidae. Pal. Sin. C, 5(5). ASI Ser. A: Life Sci., 180). Plenum, New York, 658 pp. Pickford, M., 1987. Revision des suiformes (Artiodactyla, Qiu, Z.X., Yan, D.F., Jia, H. and Sun, B., 1985. A preliminary Mammalia) de Bugti (Pakistan). Ann. Paleontol., 73(4). investigation upon Palaeomeryx from Shanwang, Shandong Qi, G.Q., 1985. Preliminary diagnoses of the Carnivora from Province. Vertebr. PalAsiat., 23(3) (in Chinese). the Lufeng hominoid site. Acta Anthropol. Sin., 4(1) Qiu, Z,X., Yan, D.F., Jia, H. and Wang, B.Z., 1985. The first (in Chinese). discovery of an Ursavus dentition from Shanwang, Shandong Qi, G.Q., 1985. A general stratigraphic description of the Province. Vertebr. PalAsiat., 23(4) (in Chinese). Lufeng hominoid site. Acta Anthropol. Sin., 4(1) (in Qiu, z.x., Yan, D.F. and Sun, J.H.B., 1986. A new discovery Chinese). of large Ursidae from Shanwang, Shandong Province. Qi, G.Q., 1986. Rhizomyidae from the Lufeng hominoid site, Vertebr. PalAsiat., 24(3) (in Chinese). Yunnan Province. Acta Anthropol. Sin., 5(1). Qiu, Z.X. and Guan, J., 1986. The discovery of a primate Qi, T., 1989. Middle Miocene Carnivora from the Altai vicinity upper molar from Tongxin, Ningxia Autonomous Region. of Xinjiang Autonomous Region. Vertebr. PalAsiat., 27(2) Acta Anthropol. Sin., 5(3) (in Chinese). (in Chinese). Qiu, Z.X. and Gu, Y.M., 1986. The Middle Miocene Qiu, Z.D., 1981. A new genus of Miocene Sciuridae from mammalian fauna from Xiacaowan, Sihong, Jiangsu Lanqu, Shandong Province. Vertebr. PalAsiat., 19(3) (in Province. 3. Two species of Carnivora. Vertebr. PalAsiat., Chinese). 24(1) (in Chinese). Qiu, Z.D., 1986. A fossil tupaiid from the Lufeng hominoid Qiu, Z.X., Huang, H.L. and Guo, Z.H., 1987. Chinese locality. Vertebr. PalAsiat., 24(4) (in Chinese). hipparionine fossils. Palaeontol. Sin. N. S. C, 25 (in Chinese Qiu, Z.D., 1987. The Middle Miocene vertebrate fauna from with English summary). Xiacaowan, Sihong, Jiangsu Province. 7. Aplodontidae. Qiu, Z.X., Yeh, J. and Jiang, Y.J., 1987. Several species of Vertebr. PalAsiat., 25(4) (in Chinese). Bahe Stage mammals from Wuzhong, Ningxia Autonomous Qiu, Z.D., 1988. Neogene micromammals of China. In: P. Region. Vertebr. PalAsiat., 25(1) (in Chinese). Z. Qiu, Z. Qiu/Palaeogeography, Palaeoclirnatology, Palaeoecology 116 (1995) 41-70 69

Qiu, Z.X. and Gu, Z.G., 1988. A Middle Tertiary mammal Shanwang, Lanqu, Shandong Province. Vertebr. PalAsiat., locality from Lanzhou, Gansu Province. Vertebr. PalAsiat., 9(1) (in Chinese). 26(3) (in Chinese). Wang, B.Y. and Wang, P.Y., 1990. The discovery and Qiu, Z.X., Yeh, J. and Cao, J.X., 1988. A new species of significance of an Early Miocene mammalian fauna from Percrocuta from Tongxin, Ningxia Autonomous Region. Wuertu, Zuoqi, Alashan, Inner Mongolia. Sci. Bull., 35(8) Vertebr. PalAsiat., 26(2) (in Chinese). (in Chinese). Qiu, Z.X., Yeh, J. and Huo, F.C., 1988. Description of a Wang, B.Y. and Qi, T., 1989. The discovery of a new genus of Kubanochoerus skull from Tongxin, Ningxia Autonomous Distylomyidae from the Xinjiang Autonomous Region. Region. Vertebr. PalAsiat., 26(1 ) (in Chinese). Vertebr. PalAsiat., 27( 1) (in Chinese). Qiu, Z.X., Xie, J.Y. and Yan, D.F., 1988. The discovery of a Wood II, H.E., Chaney, R.W., Clark, J., Colbert, E.H., Jepsen, skull of Dinocrocuta gigantea. Vertebr. PalAsiat., 26(2) G.L., Reeside Jr., J.B. and Stoch, C., 1941. Nomenclature (in Chinese). and correlation of the North American continental Tertiary. Qiu, Z.X., Xie, J.Y. and Yan, D.F., 1990. Several species of Bull. Geol. Soc. Am., 52. Early Miocene Mammalia from Dongxiang, Gansu Province. Woodburne, M.O. (Editor), 1987. Cenozoic mammals of Vertebr. PalAsiat., 28(1) (in Chinese). North America--Geochronology and biostratigraphy. Univ. Schaub, S., 1930. Fossil Sicistinae. Abh. Schweiz. Palaeontol. Calif. Press. Ges., 49. Wu, R.K., 1987. A revision of the large primates from Lufeng. Schaub, S., 1934. Uber enige fossile Simplicidentata aus China Acta Anthropol. Sin., 6(4) (in Chinese). und der Mongolei. Abh. Schweiz. Palaeontol. Ges., 54. Wu, R.K. and Pan, Y.R., 1985. A new genus of adapid primate Schaub, S., 1938. Terti~tre und Quart~ire Murinae. Abh. from the Miocene of Lufeng. Acta Anthropol. Sin., 4(1) Schweiz. Palaeontol. Ges., 61. (in Chinese). Schlosser, M., 1924. Tertiary vertebrates from Mongolia. Pal. Wu, R.K. and Pan, Y.R., 1985. Preliminary observations on Sin. C, 11. the cranium of Laccopithecus robustus from Lufeng, Yunnan, Shanxi Provincial Investigative Brigade, Third Division, 1976. with reference to its phylogenetic relationship. Acta Early Pliocene Stratigraphy of Luwanfen, Xinxiang, Henan. Anthropol. Sin., 4( 1) (in Chinese). Vertebr. PalAsiat., 142 (in Chinese). Wu, W.Y., Ye, J. and Zhu, B.C., 1991. On Alloptox Storch, G., 1987. The Neogene mammalian faunas of Ertemte (Lagomorpha, Ochotonidae) from the Middle Miocene of and Harr Obo in Inner Mongolia (Nei Mongol), China. 7. Tongxin, Ningxia Hui Autonomous Region, China. Vertebr. Muridae (Rodentia). Senckenbergiana Lethaea, 675 (6). PalAsiat., 29(3) (in Chinese). Tang, Y.J., 1980. Early Pleistocene Mammalia from Yuxian Wu, W.Y., 1986. The Middle Miocene vertebrate fauna from County, Hebei Province and their significance toward Xiacaowan, Sihong, Jiangsu Province. 4. Gliridae stratigraphic subdivision. Vertebr. PalAsiat., 18(4) (in (Mammalia, Rodentia). Vertebr. PalAsiat., 24(1) (in Chinese). Chinese). Tang, Y.J., 1983. An Early to Late Pleistocene transitional Wu, W.Y., 1988. Middle Miocene Rodentia from the northern mammal fauna from Yuxian, Hebei Province. Vertebr. margin of the Jungar Basin. Vertebr, PalAsiat., 26(4) PalAsiat. 21 (3) (in Chinese). (in Chinese). Tassy, P., 1989. The Proboscidean datum event: How many Wu, W.Y., Ye, J. and Zhu, B.C., 1991. On Alloptox Proboscideans and how many events? In: E.H. Lindsay, V. (Lagomorpha, Ochotonidae) from the Middle Miocene of Fahlbusch and P. Mein (Editors), European Neogene Tongxin, Ningxia Hui Autonomous Region, China. Vertebr. Mammal Chronology. Plenum, New York, 658 pp. PalAsiat., 29(3) (in Chinese). Tedford, R.H., 1970. Principles and practices of mammalian Xie, W.M., 1979. The first discovery of Dryopithecus in China. geochronology in North America. Proc. N.A. Paleontol. Vertebr. PalAsiat., 17(2) (in Chinese). Conv. Yan, D.F., Qiu, Z.D. and Meng, Z.Y., 1983. Middle Miocene Teilhard de Chardin, P. and Piveteau, J., 1930. Les mammif6res stratigraphy and fossil mammals from Shanwang, Shandong fossiles de Nihowan (Chine). Ann. Pal., 19. Province. Vertebr. PalAsiat., 21(3) (in Chinese). Tong, Y.S., 1989. A new species of Sinolagomys (Lagomorpha; Yeh, J., 1989. Middle Oligocene Artiodactyla from the northern Ochotonidae). Vertebr. PalAsiat., 27(2) (in Chinese). margin of the Jungar Basin. Vertebr. PalAsiat., 27(1) Tong, Y.S., 1987. Paleovertebrates and Stratigraphy of the (in Chinese). Xinjiang Autonomous Region. Petroleum Developmental Yeh, J. and Jia, H., 1986. Platybelodon from the Miocene of Patterns and Geographic Evolution of the Jungar Basin. Tongxin, Ningxia Autonomous Region. Vertebr. PalAsiat., Inst. Vertebr. Paleontol. Paleoanthropol., Acad. Sin. (in 24(2) (in Chinese). Chinese). Young, C.C., 1927. Fossile Nagetiere aus Nord-China. Pal. Wang, B.Y., 1985. Zapodidae (Rodentia, Mammalia) from the Sin., C, 33. lower Oligocene of Qujing, Yunnan, China. Mainzer Young, C.C., 1937. On a Miocene mammalian fauna from Geowiss. Mitt., 14. Shantung. Bull. Geol. Soc. China, 172. Wang, B.Y., 1965. A new species of rhinocerotid from Young, C.C., 1964. The discovery and significance of 70 Z Qiu, Z. Qiu/Palaeogeography, Palaeoclimatology, Palaeoecology 116 (1995) 41 70

Lagomervx from Lantian, Shaanxi Province. Vertebr. Province, and Kaiyuan, Yunnan Province. Vertebr. PalAsiat., PalAsiat., 8(4) (in Chinese). 12(2) (in Chinese). Young, C.C., 1977. Regarding the Anura and Chiroptera from Zhang, Y.P., Huang, W.P., Tang, Y.J., Ji, H.X., You, Y.Z., Lanqu, Shandong Province. Vertebr. PalAsiat, 15(1) (in Tong, Y.S., Ding, S.Y., Huang, X.S. and Zheng, J.Z., 1978. Chinese). The Cenozoic of the Lantian Region. Inst. Vertebr. Paleontol. Zapfe, H., 1979. Chalicotherium grande (Blainv.) aus der Paleoanthropol. Acad. Sin. Spec. Pap., 14 (in Chinese). mioz~inen Spaltenfilllung von Neudorf an der March, Zheng, S.H, 1980. A Hipparion Fauna from the Bulong Basin, Tschechslowakei. Berger, Wien. Biru, Tibet. In: Paleontology of Tibet, Collected Works on Zhai, R.J., 1978. A Primitive Elasmothere from the Miocene Research on the Qinghai-Tibetan Plateau. I. Science Press of Lantong, Shaanxi Province (Coll. Pap. Paleontol. Stratigr., (in Chinese) 7). Geol. Publ. House, Beijing (in Chinese). Zheng, S.H. and Li, C.K., 1986. Mimomys in China. Vertebr. Zhang, X.T. and Zhai, R.J., 1978. Miocene Proboscidea from PalAsiat., 24(2) (in Chinese). the Lantian Region of Shaanxi (Coll. Pap. Paleontol. Zhou, S.Y., 1988. Pliocene micromammals from Jingle, Shanxi Stratigr., 7). Geol. Publ. House, Beijing. Province, and a discussion of the age of the Jingle Formation. Zhang, Y.P., 1974. Miocene Suidae from Lanqu, Shandong Vertebr. PalAsiat., 26(3) (in Chinese).