Chapter 22 Author(s): QIU ZHUDING and LI CHUANKUEI Source: Bulletin of the American Museum of Natural History, :586-602. Published By: American Museum of Natural History DOI: http://dx.doi.org/10.1206/0003-0090(2003)279<0586:C>2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1206/0003-0090%282003%29279%3C0586%3AC%3E2.0.CO %3B2

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Chapter 22

Rodents from the Chinese Neogene: Biogeographic Relationships with Europe and North America

QIU ZHUDING1 AND LI CHUANKUEI2

ABSTRACT Cenozoic terrestrial deposits with a dense fossil record are widespread in China. At least 126 genera of rodents, belonging to 25 families and subfamilies, are known from the Neogene. Geographical distribution of the fossil rodents indicates that zoogeographic differentiation in China was already quite distinct and faunal provinces similar to the present day Palearctic Realm in North China and to the Oriental Realm in South China existed throughout Neogene time. An initial phase in development of the present Oriental region emerged by the early Miocene in southeastern Asia. Faunas in northern China were Holarctic in character and showed greater similarity in composition to Europe than to North America. Apparently, im- migration and dispersal of rodents in the Holarctic Region repeatedly took place via the Bering Landbridge during the Neogene. Interchange of rodents between Asia and Europe tended to gradually increase during the Neogene, whereas it declined between Asia and North America after the late Miocene.

INTRODUCTION 1999). To attain more precise biochronologic The widespread Cenozoic terrestrial de- dating, a combination of biostratigraphic and posits and rich fossil record in China afford paleomagnetic data is possible in some areas an opportunity to study the paleontology of with long stratigraphic sections and rich this era. Research on the Chinese Neogene mammal remains. A chronostratigraphic se- in zoogeography, biostratigraphy, biochro- quence with paleomagnetic data has been ac- nology, and systematic paleontology has ad- complished at Yushe and Lanzhou basin, vanced in the last 20 years due to efforts by mainly through cooperation with American paleontologists from China, Europe, and and Swiss colleagues. Investigations of se- North America. The fossil record has been quences with long composites of successive enriched greatly, and a large number of microfaunas and magnetostratigraphic con- mammalian faunas are recognized in Neo- trol were initiated a couple of years ago in gene deposits. Chinese paleontologists have the Lantian area by a joint expedition from seriated the faunas and developed biochrons Finland and the Institute of Vertebrate Pale- based on correlation with those of Europe. ontology and Paleoanthropology (IVPP) and The Neogene biochronologic framework ini- in Lingtai, Gansu, by Shaohua Zheng and tiated by Chiu (Qiu) and others in 1979 has others. A section with rich late Oligocene been re®ned signi®cantly, with local Neo- through middle Miocene remains in northern gene biochrons continuously modi®ed (Chiu Junggar, Xinjiang, including magnetic stra- et al., 1979; Li et al., 1984; Qiu, 1990; Qiu tigraphy analysis, is under study by Wenyu and Qiu, 1995; Tong et al., 1995; Qiu et al., Wu and others. In addition, the two biotic

1 Professor, Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, P.O. Box 643, Beijing 100044, China. 2 Professor, Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, P.O. Box 643, Beijing 100044, China.

586 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 587

Fig. 22.1. Distribution of Neogene rodent localities in China. , Early Miocene (Xiejian ϩ Shan- wangian): 1, Suosuoquan; 2, Xiejia; 3, Gaolanshan; 4, Zhangjiaping; 5, Gashunyinadege; 6, Wuertu; 7, Shanwang; 8, Sihong (Songlinzhuang, Zhengji, Shuanggou); 9, Fangshan. ⅷ, Middle Miocene (Tung- gurian): 10, Halamagai; 11, Quantougou; 12, Lierpu (Qijia, Danshuilu); 13, Dingjiaergou; 14, Tunggur; 15, Tairum Nor. Ⅵ, Late Miocene (Baodean): 16, Songshan; 17, Bulong; 18, Jilong; 19, Qingyang; 20, Baode; 21, Lantian (Bahe); 22, Amuwusu; 23, Shala; 24, Baogedawula; 25, Ertemte (Harr Obo); 26, Shihuiba; 27, Yuanmou. ᭡, Pliocene (Yushean): 28, Bilike; 29, Jingle; 30, Dingcun; 31, Youhe; 32, Daodi; 33, Zhoukoudian (Cap Travertine); 34, Yinan; 35, Zhaotong; 36, Wushan. ૽, Late Miocene ϩ Pliocene: 37, Yushe (Mahui; Gaozhuang; Mazegou; Haiyan; Jiayucun); 38, Lingtai (Wenwanggou). provinces of northern and southern China prior to the 1980s was based on a handful of have been recognized to persist throughout specimens at a few localities. To date, more Neogene history (Qiu, 1996b; Tong et al., than 40 assemblages of rodents have been 1996). recovered from Neogene deposits of different Progress in research on the Chinese Neo- ages. Figure 22.1 shows the distribution of gene partly should be ascribed to the great major localities of Neogene rodents in China. advances in intensive collection and study of Available collections prove that fossil ro- small mammals, particularly rodents in the dents in this country are as abundant and di- last 20 years. Knowledge of fossil rodents verse as in Europe and North America. Fig- 588 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279 ure 22.2, a correlation chart, serves as a ly Miocene assemblages of eastern China (Li framework for evaluating faunas. Subdivi- et al., 1983). It is composed of quite a num- sion of the Chinese Neogene mammalian ber of rodents and other mammals either par- ages and intercontinental correlation of ro- ticular to the present Palearctic region or dis- dent assemblages in the correlation chart are tributed over the Oriental region and tropical/ mainly based on Li et al. (1984); Qiu and subtropical areas today. The fauna is closely Qiu (1995); Tong et al. (1995); and Qiu et related to the early Miocene Li Mae Long al. (1999). In this paper, we review the fossil fauna of Thailand. record of Chinese Neogene rodents and are The Third Central Asiatic Expedition or- pleased to dedicate this effort to Dr. Richard ganized by the American Museum of Natural H. Tedford for his contributions to the de- History initially investigated the Tunggur velopment of Chinese Neogene paleontology fauna. The site was re-collected by Chinese and his works that inspire great respect. paleontologists in 1986, who recovered 21 rodent taxa to be added to the so-called Pla- MAJOR CHINESE NEOGENE RODENT tybelodon fauna. This is the most diverse and FAUNAS abundant middle Miocene fauna known in China and all of Asia (Stirton, 1934, 1935; Neogene rodent localities are centered Wood, 1936; Li, 1963; Qiu, 1996a). largely in northern and northwestern China, The Amuwusu fauna (19 taxa) contains ei- scattered in southwestern areas and the area ther Tunggurian relict forms or very primi- between the Yangtze River and the Huai Riv- tive Baodean elements (Qiu and Wang, er, but so far not known in southeastern and 1999), and is considered an earliest late Mio- northeastern parts of China. A set of local cene fauna in China. rodent faunas in central Inner Mongolia and The Shihuiba fauna from the Lufeng hom- the middle part of the Yellow River valley inoid locality, including 19 species of ro- characterize very well the history of Chinese dents, is the best represented late Miocene Neogene rodents in North China. The best representative and highly signi®cant Neo- fauna in South China. It exhibits a quite dif- gene local faunas are the Suosuoquan fauna, ferent composition from that of the contem- Xiejia fauna, Sihong fauna, Tunggur fauna, porary faunas of North China and is obvi- Amuwusu fauna, Shihuiba fauna of Lufeng, ously Oriental in character (Qiu et al., 1985). Yushe faunas, Ertemte fauna, Wenwanggou The Ertemte fauna was ®rst discovered in faunas of Lingtai, Bilike fauna, Daodi fauna, 1919 and studied by Schlosser in 1924. Re- and Wushan fauna. collection in 1980 added a rodent fauna with The Suosuoquan fauna has been variously up to 32 forms and made it the richest fauna considered late Oligocene (Tong et al., 1995) among the numerous late Miocene rodent as- and early Miocene (Qiu and Qiu, 1995; Qiu semblages in North China (Schlosser, 1924; et al., 1999). It is here referred to the earliest Fahlbusch et al., 1983). It re¯ects a typical Neogene fauna of China because the nine ro- temperate steppe or forest±grassland envi- dents associated with other mammals are de- ronment, like that of the present day Palearc- rived species of the genera present in latest tic province. Oligocene faunas, such as at Taben-buluk. The Yushe faunas (containing Mahui, The Xiejia fauna well represents the early Gaozhuang, Mazegou, and Haiyan assem- Miocene in northwestern China, and contains blages) and Wenwanggou fauna, spanning survivors of endemic Oligocene forms, but from about 6±7 Ma to 2 Ma, have long com- with de®nitely advanced species characters posites of successive rodents and magneto- (Li and Qiu, 1980). The species of Paras- stratigraphic control (Flynn, 1993, 1997; minthus in this fauna shows more derived Flynn et al., 1995, 1997; Zheng and Zhang, morphology than that of Suosuoquan. 2000). They have the potential to provide a The Sihong (Xiacaowan or Hsiatsaowan) key reference for the late Neogene faunas of fauna, consisting of 17 species of rodents North China. from the Songlinzhuang, Zhengji, and The Bilike fauna, containing 30 rodents, Shuanggou sites, represents the very few ear- shows strong similarities with the Ertemte 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 589

Fig. 22.2. Correlation of Chinese Neogene biochrons with those of Europe and North America. 590 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279 fauna and represents, on the whole, a sort of ctenodactylids occur. Aralomys is known modernized Ertemte fauna (Qiu and Storch, from Kazakhstan and is represented by quite 2000). a number of specimens from Suosuoquan, The Daodi fauna is a younger Neogene as- which were previously reported as Tachyor- semblage with 13 well-represented rodents yctoides obrutschewi and T. pachygnathus and one of the best records of a late Pliocene (Qiu et al., 1999). Li and Qiu (1980) de- small mammal community in North China scribed Tachyoryctoides kokonorensis from (Cai, 1987). Xiejia (early Miocene) and indeterminate The Wushan fauna from cave deposits of species of this genus were reported at Hala- Central China consists of 30 rodents. It was magai, Zhangjiaping, Dingjiaergou, Wuertu, considered Pleistocene in age by Zheng in and Gashunyinadege (Qiu et al., 1999; Qiu 1993. The assemblage represents the youn- and Wang, 1999). It is clear that the two gen- gest Neogene fauna, that is, late Pliocene era are survivors from Oligocene faunas, and rather than Pleistocene in age, if the age in- endemic to central and northeastern Asia. terpretation of 2 Ma for the locality is correct They coexisted with and disappeared simul- (Huang and Fang, 1991). taneously with ctenodactylids in the Mio- cene. MAJOR GROUPS OF CHINESE TSAGANOMYIDAE: Tsaganomys is the only NEOGENE RODENTS Neogene genus of Tsaganomyidae, and it ®rst appeared in the early Oligocene in the One hundred and twenty-six genera of ro- Mongolian plateau. Tsaganomys cf. altaicus dents within 25 groups (families and subfam- collected from Gaolanshan (previously Lan- ilies) and representing Protrogomorpha, zhou) represents the last record of the family Sciuromorpha, Myomorpha, and Hystrico- (Qiu et al., 1999). morpha are known in the Chinese Neogene. DIATOMYIDAE: Diatomys shantungensis, Figure 22.3 lists major groups of Neogene collected from Shanwang and Sihong, is the rodents with their temporal and geographical only Chinese species of this family. Diato- distribution, and relationships to Europe and mys was ®rst considered indeterminate to North America. family and questionably referred to Geomy- CTENODACTYLIDAE: Ctenodactylidae ¯our- oidea (Li, 1974). Some students transferred ished in central and northeastern Asia during it to the family Pedetidae (McKenna and the Oligocene, but declined greatly in the Bell, 1997). We follow P. Mein and L. Gins- early Miocene and made their last occurrence burg (1997) in their de®nition of a new fam- in the early middle Miocene of Dingjiaergou. ily for this genus plus probably Fallomus. Six genera, Tataromys, Yindirtemys, Prodis- Diatomys is also known from Thailand, Pak- tylomys, Distylomys, Prosayimys, and Sayi- istan, and Japan (Kato and Otsuka, 1995), mys, grouped in three subfamilies, Tataro- and appears to be a kind of wet/warm-adapt- myinae, Distylomyinae, and Ctenodactyli- ed animal distributed in a tropical or sub- nae, have been reported mainly in the Mon- tropical area. golia±Xinjiang areas and northern edge of APLODONTIDAE: Aplodontid rodents are Qinghai±Tibet Plateau. All the genera except quite well known in Holarctic Oligocene de- Sayimys can be found in Oligocene deposits posits in Europe and North America, but are (Wang, 1977) at low abundance except for represented in China by scarce materials of the common Yindirtemys. Survivors migrated four genera. These are Promeniscomys, Hap- southwestward through Asia to the Mediter- lomys, Prosciurus, and Ansomys, of which ranean area and North Africa after the early only Ansomys survived into the Miocene Miocene, and were affected by change of en- (Rensberger and Li, 1986; Qiu, 1987; Wang, vironment caused by uplift of the Himalayas. 1987). Ansomys may have evolved from a See Wang's (1997) signi®cant revision of the Prosciurus-like ancestor, and made its ®rst family. appearance in the early Miocene. Ansomys TACHYORYCTOIDIDAE: Two genera of this orientalis was described from Sihong and A. family, Aralomys and Tachyoryctoides, have shanwangensis from Shanwang (Qiu, 1987; been recognized in the same regions where Qiu and Sun, 1988), and questionable An- 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 591 Fig. 22.3. Biogeographic relationships of major groups of Neogene rodents found in China. 592 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279 somys was reported from Gashunyinadege, 1998). All the taxa, except Youngo®ber, can Tunggur, Amuwusu, and Shala of Inner be found in the European and North Ameri- Mongolia (Qiu and Wang, 1999). In addition, can Neogene. Youngo®ber was a massive the aplodontine Pseudaplodon is known form also known from Mizunami, Japan from the Mio/Pliocene of Ertemte and Harr (Tomida and Setoguchi, 1994) as well as Si- Obo, Inner Mongolia, and represents the last hong. The occurrence together with Diato- record of this family in Asia (Fahlbusch et mys in both places suggests existence of a al., 1983). An upper and a lower premolar connection between the Chinese mainland from Amuwusu, similar to those of Menis- and the Japanese islands during the early comys of North America, may represent a Miocene. An indeterminate genus from Shi- meniscomyine rodent in the Old World. huiba and ''Castor'' from Zhaotong, Yunnan, MYLAGAULIDAE: Mylagaulids are quite di- represent the southernmost records of bea- verse in North America, and also known vers in China, even in the Old World. As- from Zaisan Basin, Kazakhstan. Sinomyla- sociated with the beavers at Shihuiba are gaulus halamagaiensis described by Wu some small mammals con®ned to the present (1988) from Halamagai Formation, Junggar tropical or subtropical areas, such as tree Basin, represents the only record of this fam- shrew, mole shrew, spiny dormouse, and ily in China. bamboo rats (Qiu et al., 1985). Xu (1994) SCIURIDAE: We recognize 23 genera of presented a thorough review of castorid ro- sciurid rodents, with Sihong, Ertemte, Shi- dents found in North and East China. huiba (Lufeng), and Wushan being relatively EOMYIDAE: Eomyids are another cosmo- diverse and common squirrel faunas (Qiu et politan group of fossil rodents, but they are al., 1985; Qiu and Lin, 1986; Qiu, 1991; poorly diversi®ed with limited material in Zheng, 1993). An undiagnosed new species China. Two genera, Keramidomys and Lep- of Palaeosciurus from Suosuoquan repre- todontomys, have been determined from ear- sents the oldest record of the family (Qiu et ly Miocene (Gashunyinadege) through early al., 1999). Palaeosciurus, Eutamias, Sciurus, Pliocene (Harr Obo) localities, and these usu- Tamiasciurus, Atlantoxerus, Heteroxerus, ally occur together in the Miocene deposits Sciurotamias, Miopetaurista, Pliopetaurista, of North China (Zheng and Li, 1982; Fahl- Petinomys, Albanensia, Pteromys, and Hy- busch et al., 1983; Qiu, 1996a; Qiu and lopetes show af®nities either to Europe or to Wang, 1999). Keramidomys failed to persist North America, whereas Sinotamias, Pros- into the Pliocene. An undetermined genus of permophilus, Tamiops, Callosciurus, Dre- this family from Shihuiba is a bunodont momys, Parapetaurista, Shuanggouia, Ple- eomyid, which shares dental similarities with siosciurus, Meinia, and Belomys are restrict- Eomys or Pentabuneomys of Europe, or with ed in geographical distribution to Asia. Gen- Adjidaumo of North America (Qiu, 1994). erally, sciurids were not so diverse as they GLIRIDAE: Since the ®nding of Myomimus were in Europe and North America during at Ertemte, dormouse material has been Neogene time, and ¯ying squirrels are not as found at several localities, one after another. common as they are in European sciurid fau- Nevertheless, only three genera, Microdyro- nas. Sciurids found in China demonstrate ap- mys, Miodyromys, and Myomimus, are rec- parently ecologic provinciality throughout ognized, and specimens of the ®rst two taxa the Neogene, with dominance of ground are rather rare (Wu, 1985, 1986; Qiu, 1996a; squirrels in the north, and of tree and ¯ying Flynn et al., 1997; Wu et al., 1998; Qiu and squirrels in the south. Wang, 1999). All three genera are commonly CASTORIDAE: Eight genera of beavers, known in Europe, but do not occur in South Youngo®ber, Anchitheriomys, Steneo®ber, China. Neither Microdyromys nor Miodyro- ?Hystricops, Castor, Dipoides, Trogonther- mys survived into the Pliocene. ium, and Eucastor, have been recovered from PLATACANTHOMYIDAE: Platacanthomys and Sihong, Tunggur, Halamagai, Amuwusu, Yu- Typhlomys, con®ned to the present Oriental she, Ertemte, Bilike, and Wushan (Li, 1963; region, are known from Shihuiba (Qiu, 1989) Chow and Li, 1978; Zheng, 1993; Xu, 1994; and Yuanmou recently. Zheng (1993) de- Qiu, 1996a; Flynn et al., 1997; Wu et al., scribed three species of Typhlomys from 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 593

Wushan. An isolated tooth from Sihong, as- (Li and Qiu, 1980). The indeterminate spe- signed to Neocometes, represents the third cies previously reported as Eumyarion sp. genus of platacanthomyid rodents found in based on a lower molar from Tunggur has China. The latter is also known from Thai- been transferred to Gobicricetodon (Qiu, land. Remains of these animals are not found 1996a). in North China. CRICETODONTINAE: Five genera in this ZAPODIDAE: The nine genera representing group, Megacricetodon, Democricetodon, this family in North China are Litodonomys, Primus, Spanocricetodon, and Paracricetu- Parasminthus, Heterosminthus, Eozapus, lus, have been recovered from the early and Protozapus, Sinozapus, Plesiozapus, Sicista, middle Miocene. Megacricetodon and De- and Lophocricetus. Litodonomys and Paras- mocricetodon had a large geographic distri- minthus are known only from early Miocene bution and made their ®rst appearance in ear- localities, such as Suosuoquan and Xiejia, ly Miocene (Sihong, Wuertu, Gashunyinad- whereas Heterosminthus is mainly middle ege) as in Europe. They seem to have thrived Miocene, Quantougou, Dingjiaergou, and in North China in the middle Miocene. Wes- Tunggur, for example (Li and Qiu, 1980; sels (1996) argued that Megacricetodon did Qiu, 1996a; Qiu et al., 1999). Remains of the not occur in Pakistan and that all material other forms derive mainly from the late Mio- from the Indian subcontinent previously as- cene and Pliocene deposits. Eozapus, Sicista, signed to Megacricetodon should be allocat- and Lophocricetus are common members of ed as the myocricetodontine Sindemys. The the Ertemte and Harr Obo faunas (Qiu, 1985; tiny cricetodontine Primus is known from the Fahlbusch, 1992). Specimens of Protozapus early Miocene of Pakistan, and similar spe- reported from Wenwanggou and Plesiozapus cies occur in the equivalent age Sihong fau- from Amuwusu are inadequate and need fur- na. Spanocricetodon from the Shanwangian ther identi®cation. Only Eozapus and Sicista of eastern China (Fangshan) is also known persist to the present day. from Pakistan and Thailand. Young (1927) DIPODIDAE: Neogene dipodid rodents described Paracricetulus schaubi from Hsien found include Protalactaga, Paralactaga, Shui Ho (Quantougou) based on a fragmen- Sminthoides, Brachyscirtetes, and Dipus. The tary upper jaw with a damaged M1. Addi- earliest Neogene record of this family is an tional material has been collected from the indeterminate species of Protalactaga rep- resented by an upper molar from Wuertu fau- type locality, and further study will deepen na (Qiu et al., 1999); that it possibly is a understanding of this genus. Cricetodon it- derived species of Parasminthus cannot be self, which is known from the Miocene of excluded. Protalactaga occurs usually in the Europe and western Asia, seems not to ap- middle Miocene of North China, such as in pear in eastern Asia. Records of ``Criceto- the Quantougou, Tunggur, and Dingjiaergou don'' from Suosuoquan, Dingjiaergou, and a faunas. These rather primitive jumping mice similar genus from Sihong are misidenti®ed failed to survive into the late Miocene. The (vide infra). other genera are known from the late Mio- GOBICRICETODONTINAE: Gobicricetodontine cene and Pliocene (the indeterminate species rodents are relatively large-size cricetids with of Paralactaga from Dingjiaergou in a pre- mesodont and bunolophodont cheek teeth, vious faunal list may be Protalactaga ma- and include two genera, Plesiodipus and jor). Sminthoides is quite common in late Gobicricetodon. Plesiodipus is known from Neogene deposits, whereas Brachyscirtetes several middle and early late Miocene local- and Dipus are relatively scarce. The ques- ities of North China (Lierpu in Xining Basin, tionable Dipus from Amuwusu probably rep- Quantougou, Tunggur, and Amuwusu). It resents the ®rst record of the three-toed jer- was thought to include ancestry of siphnei- boa. Both zapodids and dipodid rodents are nes, which ¯ourished later in northeastern found only in North China. Asia (Qiu et al., 1981). Qiu (1996a) de- PARACRICETODONTINAE: Eucricetodon scribed two species of Gobicricetodon from youngi from Xiejia is the only representative Tunggur. Previously reported Cricetodon sp. of this group found in the Neogene of China from Suosuoquan, Halamagai, Dingjiaergou, 594 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279 and Amuwusu should be referred to this ge- atavus and Anatolomys teilhardi from Ertem- nus. te, Harr Obo, and Bilike of central Inner CRICETINAE: Cricetines rapidly replaced Mongolia. They are very common in the Er- the archaic cricetids and diversi®ed since the temte and Harr Obo faunas, but rare in Bi- late Miocene. Including Wushan in the late like. Baranomyine rodents seem to be re- Pliocene, 11 genera of this group, Sinocri- placed by the arvicolines that arise abruptly cetus, Nannocricetus, Kowalskia, Neocrice- during the early Pliocene in this region, and todon, Bahomys, Cricetulus, Cricetinus, Al- persist into the Pleistocene only in eastern locricetus, Phodopus, Chuanocricetus, and Europe. Amblycricetus, have been recognized in the SIPHNEINAE: Siphneines are a group of ro- late Neogene. Sinocricetus and Nannocrice- dents derived probably from a Pleisodipus- tus from Shala in North China and Kowal- like ancestor and endemic to central and skia from Shihuiba in South China represent northeastern Asia. They made their ®rst oc- the ®rst appearance of this subfamily (Bao- currence in the very early late Miocene dean age). They are quite common in the late (Amuwusu) and had a rapid Pliocene radia- Miocene and early Pliocene faunas (Ertemte, tion in the great land mass. They are com- Yushe, Bilike, Wenwanggou, etc.). Neocri- monly known in the late Neogene faunas of cetodon is known only from the late Miocene North China (see table 22.1) and are poten- of Yushe (Schaub, 1934; Flynn et al., 1997). tially very useful in biochronological corre- Bahomys, a form of cricetid with rather com- lations. On the basis of the presence or ab- plex occlusal structure, was ®rst described sence of molar roots, Teilhard and Young from the Pleistocene of Lantian (Chow and (1931) assigned these animals to two gen- Li, 1965), and recently reported from the Pli- eraÐthe rooted Prosiphneus and rootless Si- ocene of Wenwanggou (Zheng and Zhang, phneus (ϭ Myospalax). Zheng (1994, 1997), 2000). The six other genera occur in the Yu- however, based on patterns of their occipital shean Pliocene (Yushe Basin, Wenwnggou, shield and corresponding skull features, and Wushan). Differential diagnosis of these grouped them into three subfamilies, the con- taxa is not so clear, and de®nition and real- vex-skulled Prosiphneinae, the ¯at-skulled location of late Neogene cricetines remains Myospalacinae, and the concave-skulled Me- undone. In addition, three damaged teeth sosiphneinae, comprising together 10 genera. from Shala have been referred to Microto- Zheng's proposal may be reasonable, but in- cricetus, but this taxon awaits more material complete knowledge of skulls and great mo- for con®rmation. lar similarity in various species make it dif- GERBILLINAE: Pseudomeriones is the single ®cult to allocate the isolated teeth that usu- published representative of this group, and ally occur in deposits. In addition, evolution- occurs frequently in the late Miocene and ary relationships of the diverse extant Pliocene of North China (Qingyang, Yushe, Myospalax species with various siphneines Ertemte, Wenwanggou, Bilike, and Ningxi- remain to be solved. According to Zheng's an). Two species of the genus, at different de®nition, the Neogene siphneine genera are stages of evolution, have been recognized Myotalpavus, Prosiphneus, Chardina, Me- (Teilhard, 1926; Zhang, 1999). sosiphneus, Pliosiphneus, Episiphneus, and MICROTOSCOPTINAE: Microtoscoptine ro- Youngia. The ®rst two appeared in the late dents have a Holarctic distribution. Micro- Miocene, whereas the others are Pliocene. toscoptes, ®rst named by Schaub in 1934, ARVICOLINAE: High-crowned rodents evi- represents this subfamily in China. Fahl- dently increased in abundance in northeast- busch (1987) described the additional mate- ern Asia since the late Miocene. Like the mi- rial from Ertemte and Harr Obo in detail. crotoscoptines, baranomyines, and siphnei- More recently remains of this animal have nes in the late Miocene, arvicolines show a been reported from Shala that may represent Pliocene radiation. Aratomys, Mimomys, the earliest record of the genus (Qiu and Germanomys, Hyperacrius, Villanyia, Mi- Wang, 1999). crotus, Clethrionomys, and Eothenomys are BARANOMYINAE: Two taxa can be referred the genera of this group found in the Plio- to Baranomyinae. These include Microtodon cene of China. The oldest certain arvicoline 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 595

TABLE 22.1 Faunal List for the Major Reference Faunas 596 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279

TABLE 22.1 (Continued) 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 597 is Aratomys from the early Yushean, which cromys occur in Ertemte and Harr Obo. In dominates the Bilike fauna (Qiu and Storch, addition to these elements, Chardinomys and 2000). Zheng and Li (1986, 1990) reviewed Huaxiamys are often found in the Pliocene, the Mimomys of China. Since then, more ma- as at Yushe, Bilike, Wenwanggou, and Daodi terial has been recovered from Yushe, Wen- (Jacobs and Li, 1982; Wu and Flynn, 1992; wanggou, and Daodi. Germanomys is re- Cai and Qiu, 1993; Flynn et al., 1997; Qiu ported from Yushe, Daodi, and Jingle. The and Storch, 2000; Zheng and Zhang, 2000). later arvicolines include the poorly known Huaxiamys, Chardinomys, and Allorattus Hyperacrius, Villanyia, and Microtus from were apparently restricted in geographical Dachai, and Clethrionomys and Eothenomys distribution to the Pliocene of North China. from Wushan (Zheng and Li, 1990; Zheng, Storch (1987) reported Rhagapodemus from 1993). Harr Obo. Saidomys is known from the late RHIZOMYIDAE: Fossil rhizomyids in eastern Pliocene of Afghanistan and occurs in the Asia are not so diverse as in the Siwaliks of equivalent age fauna of Daodi. Further, Pakistan and India. An indeterminate rhizo- Zheng (1993) determined Mus, Hapalomys, myid from Sihong (also Shanwang) repre- Chiropodomys, Vernaya, Leopoldamys, Ni- sents the earliest record of the family in Chi- viventer, and Wushanomys from Wushan, na. Only subfamily Rhizomyinae is certainly which represent the earliest occurrence of recognized in the Neogene deposits. They are these genera. primitive species of Rhizomys (Brachyrhizo- HYSTRICIDAE: Porcupines are quite com- mys) from Lufeng and Yuanmou, and more mon in the Pleistocene faunas of South Chi- advanced Rhizomys (Brachyrhizomys) shan- na. Neogene hystricids are represented by a sius from Yushe. Flynn (1993) added a spe- single genus, Hystrix, with limited speci- cies of Rhizomys to the Miocene fauna of mens. An indeterminate species from Shi- Yushe. Flynn and Qi (1982) and Qi (1986) huiba is the oldest record of the genus so far described three species of Brachyrhizomys known (Qiu et al., 1985). Porcupines extend- from Lufeng, which occur together at about ed northward to Yushe in distribution during 8 Ma in Pakistan. A species reported previ- the Pliocene (Flynn et al., 1997). Zheng ously as Brachyrhizomys hehoensis from Bu- (1993) reported H. subcristata and H. magna long, Tibet, is considered congeneric with from Wushan, which herald the common por- Pararhizomys (Jacobs et al., 1985). Parar- cupines of the Pleistocene. hizomys from Fugu, Shanxi, is poorly known and, if truly a rhizomyid, represents the ZOOGEOGRAPHIC REGIONS OF northernmost distribution of this family in CHINESE NEOGENE RODENTS China. Rhizomys occurs also in the late Pli- ocene fauna of Wushan (Zheng, 1993). Distributions of extant land vertebrates MURIDAE: Murid rodents appeared in Chi- demonstrate clearly two different zoogeo- na much later than in the Indian subconti- graphic provinces in China. A line roughly nent. Progonomys and Yunomys associated from the Qin Ling Mt. to the Huaihe River with hominoids and Brachyrhizomys from separates the northern Palearctic Realm from Shihuiba, Lufeng, are considered the oldest the southern Oriental Realm. Distribution murids so far known in China (Qiu and and changes of the fossil rodents indicate that Storch, 1990), but an undescribed Progono- similar regions existed during the Neogene. mys from Lantian, Shaanxi, may represent a The numerous localities, especially in North murid appearing earlier than those of Lufeng China, record high faunal diversity and long- (Zhaoqun Zhang, personal commun.). The term stabilities of faunas through Neogene murid group has really ¯ourished since the time. latest Miocene. At least 19 genera of this In the northern region, the history of ro- group have been recognized, with the Lu- dents is characterized by extinction of archa- feng, Ertemte, Harr Obo, Bilike, Yushe, ic elements and the establishment of the Mu- Wenwanggou, Daodi, and Wushan faunas be- ridae and other semiarid or arid-adapted mu- ing the most productive. Apodemus, Orien- roids, Dipodidae, Zapodidae, and Cricetidae, talomys, Karnimata, Occitanomys, and Mi- as major elements in the faunas. Early Mio- 598 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279 cene faunas are dominated by groups that of the area. The fauna includes an extinct originated in the Oligocene or earlier, Cten- family (Eomyidae), two eurytopic families odactylidae, Tachyoryctoididae, Tsagano- (Sciuridae and Muridae, although most of the myidae, Sciuridae, Paracricedontinae, and Sciuridae live in tropical±subtropical zones), Zapodidae. New elements of the fauna are two Holarctic or Palearctic groups (Castori- Cricetodontinae, Gliridae, and Eomyidae. By dae and Cricetinae), and three families (Pla- the middle Miocene in this region rodents en- tacanthomyidae, Rhizomyidae, and Hystrici- ter a new stage: archaic groups, except for dae) con®ned to the present Oriental region Zapodidae, decline greatly or become ex- or tropical±subtropical regions. Associated tinct, and living groups dominate Neogene with the rodents are some small mammals faunas since then. Cricetodontinae and Gob- endemic to the Oriental region (Tupaiidae, icricetodontinae replace Paracricetodontinae, Echinosoricinae) or mainly distributed in advanced members of Zapodidae ¯ourish, tropic or subtropical areas (Pteropidae and and Dipodidae ®rst appear. By the late Mio- Hipposideridae). Thus, the late Miocene Shi- cene most archaic groups disappear. An ini- huiba fauna is quite different in composition tial diversi®cation of Cricetinae ends the from those faunas in the North, and re¯ects dominance of Cricetodontinae and Gobicri- a tropical or subtropical mosaic forest envi- cetodontinae of the middle Miocene. The ronment. Geographical distribution and char- northern area in this period was where the acters of faunas seem to suggest distinct zoo- high-crowned cricetids (Microtoscoptinae, geographic differentiation of animals be- Baranomyinae, and Siphneinae) were highly tween South and North China during the late diversi®ed and radiated with the ®rst appear- Miocene. Palearctic and Oriental regions ance of Gerbillinae and Muridae. As in the similar to those of the present day are fairly middle Miocene, those groups established in clear in China at that time. the early and middle Miocene, such as Inadequate collection in South China Eomyidae, Gliridae, Dipodidae, and ad- makes it dif®cult to reconstruct the evolu- vanced genera of Zapodidae, continue as im- tionary history of the Chinese Oriental portant elements in late Miocene faunas. The Realm. However, composition of the early Pliocene faunas continued the diversity of Miocene Sihong and Shanwang faunas in the the late Miocene with the appearance of ar- East likely indicates that a predecessor of the vicolines. Eomyidae, Microtoscoptinae, Bar- Oriental Province developed before the late anomyinae, and Aplodontidae disappeared Miocene, as the later Shihuiba fauna con- early in the Pliocene. Among the 12 groups ®rms. The Sihong fauna contains nine groups surviving to the present day, Castoridae, Za- of rodents, of which ®ve become extinct or podidae, Dipodidae, Gliridae, Cricetinae, disappear in China (®g. 22.3). Among the Gerbillinae, and Siphneinae are particular to, four persistent groups, Castoridae and Gliri- or mainly distributed over the Holarctic, Pa- dae are Holarctic or Palearctic, whereas Pla- learctic, or Nearctic regions and live today in tacanthomyidae and Rhizomyidae are re- the Chinese Palearctic region. Family Aplo- stricted to the Oriental Region. Apparently, dontidae occurs in the western coastal areas the Sihong fauna shows both Holarctic and of North America, families Rhizomyidae and Oriental components. In addition to Diato- Hystricidae (occurring only as far north as myidae, Rhizomyidae are known from Shan- Shanxi during the Pliocene) are con®ned to wang (Li Fenglin, personal commun.). Dia- the present tropical or subtropical areas of tomyidae, Platacanthomyidae, Cricetodonti- South China, and families Sciuridae and Mu- nae, and Rhizomyidae are also recorded in ridae are widespread across the Old World. Thailand (Mein and Ginsburg, 1997). Apparently, the rodent faunas in the North Among nonrodent taxa, primates and Echi- are distinctly Holarctic in character and re- nosoricinae, which generally live in tropical ¯ect a relatively arid temperate steppe or for- to subtropical forests, are present in both Si- est±grassland environment. hong and Thailand. This implies that an ini- The southern Chinese record in general is tial zoogeographic province similar to the not well documented, but the Shihuiba fauna present Oriental region developed during the of Lufeng well represents the late Miocene early Miocene in southeastern Asia from 2003 QIU AND LI: RODENTS FROM THE CHINESE NEOGENE 599 southeastern China (northward up to Shan- that the interchange of rodents between Asia dong peninsula) to Indo-China. and Europe increased after the middle Mio- cene. In contrast, dispersal of rodents be- BIOGEOGRAPHIC RELATIONSHIPS tween Asia and North America reached its WITH EUROPE AND NORTH AMERICA peak during the late Miocene, and then de- clined. Low levels continued into the Pleis- Of the 25 groups of Chinese Neogene ro- tocene and possibly re¯ected frequent inter- dents shown in ®gure 22.3, 8 have close re- ruption of land connection between the two lationships neither to Europe nor to North continents and shifts of global climate. America. They are mainly endemic to north- In spite of the close relationships to Eu- eastern Asia. Of the others, 16 show af®nity rope, the Chinese Neogene rodent faunas dif- with Europe and 10 with North America. fer signi®cantly. Even since the middle Mio- Among the 126 recognized genera, 44 (35% cene, European faunas contain few elements of the total) are congeneric with European restricted presently to eastern Asia: few high- representatives and only 12 (nearly 10%) are crowned cricetids (temperate forest±grass shared with North American faunas. Appar- lands) and no dipodids adapted to arid ently, the Chinese Neogene rodent faunas steppe. Some European families, such as Tri- display stronger af®nities with those of Eu- lophomyinae, Calomyscinae, Spalacinae, and rope than with North America. Anomalomyinae never reached China. This Rodent taxa shared among the Eurasian probably indicates that the interchange of ro- and North American continents seem to in- dents in the Neogene between Asia and Eu- dicate that interchange in the Holarctic re- rope took place mainly among Palearctic gion had taken place via the Bering Land- groups. Differentiation of faunal composition bridge during the Neogene. However, high might possibly result from different environ- endemism of early Miocene faunas implies mental preferences for these rodents, and dif- that migration happened mainly after the ear- ferent ecotypes or biotic subprovinces within ly middle Miocene. The groups that were ac- the pre-Palearctic region as exist today. Eu- tive in this event are Aplodontidae, Sciuri- rope must have been more forested and dae, Castoridae, and Eomyidae. Families and damper than the Palearctic Chinese regions subfamilies showing af®nities with North during the Neogene. America are also found in Europe, except for Mylagaulidae (®g. 22.3). The greater simi- larities of the Chinese Neogene rodent faunas ACKNOWLEDGMENTS to those of Europe than North America might This work was made possible through the have resulted from the ®ltering action of the efforts of scientists and technicians from Bering Landbridge. It is clear that those el- China, Europe, and North America. The au- ements distributed in tropical to subtropical thors are grateful to Drs. Zhanxiang Qiu and areas, like Platacanthomyidae and Rhizo- Wenyu Wu from IVPP and Dr. Xiaoming myidae in the Oriental region and Erethizon- Wang from Long Island University, for thor- tidae in southern North America, were un- ough review of the manuscript and investing able to withstand the rigors of the bridge. patient effort in offering many improvements North American groups such as the Floren- to the text. Many thanks are given to Dr. L. tiamyidae or even the successful Geomyidae Flynn from Harvard University for critical and Heteromyidae, and the later Sigmodon- reading and correcting of the manuscript. tinae obviously were ®ltered out by the Be- ring Landbridge. The number of genera shared with Europe REFERENCES or North America varied at different times. Cai, B.Q. 1987. A preliminary report on the late Those shared with European faunas are 8 in Pliocene micromammalian fauna from Yan- the early Miocene, 8 in the middle Miocene, gyuan and Yuxian, Hebei. Vertebrata Pal- 24 in the late Miocene, and 25 in the Plio- Asiatica 25(2): 124±136. [In Chinese with En- cene; with North America these ®gures are glish summary] 3, 5, 7, and 4, respectively. This may reveal Cai, B.Q., and Z.D. Qiu. 1993. Murid rodents 600 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 279

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