Journal of Geosciences, Osaka City University

Vol. 52, Art. 1, p. 1-10, March, 2009

A Preliminary Revision of the Extinct Voles of Mimomys and its Alliesfrom China and the Adjacent Area with Emphasis

on Villanyia and Borsodia

Yoshinari KAWAMURA] and Yingqi ZHANG2

I Department of Earth Sciences, Aichi University of Education, Kariya, Aichj Prefecture 448-8542,

Japan. E-mail: yskawmm@auecc.aichi-edu.ac.jp2 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box

643, Beijing, China. E-mail: arvicolid@gmail.com

Abstract

Mimomys and its allies constitute a group of extinct voles with rooted molars, which is important

for biostratigraphic and evolutionary studies of small mammals. Their remains have commonlyoccurred from the Pliocene and Early Pleistocene sediments of China and Transbaikalia. They m·e

referred to four genera such as Promimomys, Mimomys, Villanyia and Borsodia. Taking controversial

taxonomy of these genera into consideration, we have presented a key to the genera and taxonomjc

notes with the specific composition of each genus in order to clarify the generic distinctions. Among

the four genera, we focus on Villanyia and Borsodia and discuss their chronospatial distributions in

China and Transbaikalia. During the later part of the Pliocene, the more archaic genus Villanyia was

replaced by the more advanced genus Borsodia, although Villanyia survived somewhat later in

southern China than in Transbaikalia.

Key-words: Vole, Mimomys, Promimomys, ViLlanyia, Borsodia, Pliocene, Early Pleistocene, China,

Transbaikalia.

1. Introduction

Extinct voles of Mimomys and its allies constitute a

group adapted to grass-eating habits, which distributes

mainly in the Holm·ctic Region from the Early Pliocene to

the Middle Pleistocene. Reflected by the habits, they are

characterized by high-crowned prismatic molars, but still

retain roots in the molm·s. In China, they have commonlyoccurred from the Pliocene and Early Pleistocenesediments and are considered to have great importance in

biostratigraphic and evolutionary studies of small

mammals in these periods.The first recognition of these voles in China was made

by Kormos (1934) who allocated the remains alreadydescribed as "Arvicolide gen. ind." by Teilhard de Chardin

and Piveteau (1930), to Mimomys. After this work, the

voles were recorded from several localities in northern

China (Young, 1935; Research Group for the Huanghe

Elephant, 1975; Zheng, 1976; Xue, 1981). On the basis of

the knowledge hitherto obtained, Zheng and Li (1986)

published a comprehensive revision of the voles from

China. This work has greatly advanced the subsequent

studies of these voles, and was followed by a number of

studies which described remains of the voles from manylocalities in China (Zong, 1987; Zheng and Cai, 1991;

Zheng, 1993; Qiu and Storch, 2000; Qiu et al., 2004; Cai et

al., 2008). However, these studies described the remains

from each locali ty, and no comprehensive work on thevoles from China has been published since Zheng and Li(1986). Moreover in the previous works, the generic

taxonomy of the voles is rather obscure especially in the

2 Revision of the Extinct Voles of Mimomys and its Allies

Mimomys

distinction among Mimomys, Villanyia and Borsodia.

Thus a new revision on the voles is required on the basis of

the data accumulated after Zheng and Li (1986).

Recently, a new species of Villanyia was described

from southern China by Zhang, Kawamura and Jin (2008)

who provided a generic diagnosis of Villanyia and its

differential characters from Borsodia. But they did not

describe the distinction among Mimomys, Villanyia and

Borsodia sufficiently in order to avoid deviation from the

purpose of the paper. Here we present a preliminary

revision of Mimomys and its allies including their generic

distinction and the chronospatial distribution of Villanyia

and Borsodia on the basis of the data already published.

A more comprehensive revision of the voles will be

published in the near future to include new and

unpublished data from several localities in China that are

not treated herein. The present paper also emends the

chronological confusion in Zhang, Kawamura and Jin

(2008), which was caused by the controversies on the age

of the Pliocene / Pleistocene boundary (whether ca. 1.8 Ma

or ca. 2.6 Ma), because the boundary is strongly related to

the chronological comparisons of Villanyia and Borsodia

between China and the adjacent countries.

2. Generic taxonomy with a key

We here define Mimomys and its allies as the voles

with the following dental characters (Figs. 1-3): The

Promimomysa

sinuous line

Villanyiaa

molars are rooted. M3 has one or two triangles between the

anterior and posterior loops, while M[ has three triangles

between the anterior loop (usually forming the anteroconid

complex) and posterior loop. They comprise four genera:

Promimomys, Mimomys, Villanyia and Borsodia. A key to

the genera is given below.

Molars lower in crown height; sinuous line weakly undulated;

M3 usually without LRA3 (unknown in China); anterior loop

of M[ small and simple, generally lacking LRA4 and BRA3,

and not forming the anteroconid complex. .. ........ Promimomys

- Molars higher in crown height; sinuous line ascending high

aside of the anteroconid complex and posterior loop in M I; M3

with LRA3; anterior loop of M 1 larger and complicated, with

LRA4 and BRA3, and forming the anteroconid complex. .. ... 2

2 Cementum developed except in very primitive species; enamel

negative type (Mimomys-type) in differentiation, or

undifferentiated (in primitive species); Ml and M2 with two or

three roots; M3 generaIJy with an enamel island (but two in

primitive species, and no island in advanced species); posterior

loop ofM3 broader; M[ with the Mimomys angle and an enamel

island except in advanced species Mimomys

-Cementum absent; enamel negative type in differentiation or

undifferentiated; Ml with three roots; M2 with two or three

roots; M3 usually with one or two enamel islands; posterior

loop broad and ShOI1, where the posterior enamel island can be

accommodated; M, usually with the Mimomys angle, but no

enamel island. . Villanyia

Borsodia

Fig. 1 Schematic diagram showing the buccal views of M 1 of Mimomys and its allies. The ontogeneticchange from a juvenile individual (j) to an aged individual (a) is also shown.

Yoshinari KAWAMURA and Yingqi ZHANG 3

AL

BorsodiaeiAL

~-""""'-

VillanyiaMimomysei

Promimomys

LRA2

,,[ "\ "[",,\,~ positive

negative negative typetype type

AL

Fig. 2 Schematic diagram of M3 of Mimomys and its allies showing theiJ' occlusal patterns with terminology anddifferentiation types in enamel thickness. M3 of Promimomys is unknown in China. AL: anterior loop,BRA: buccal reentrant angle, ei: enamel island, LRA: lingual reentrant angle, PL: posterior loop, T:triangle. BRA, LRA and T are numbered for showing the homology of each part.

Promimomys Mimomys Villanyia Borsodia

T3T4

T4T2

T3 T3T3

/ ~ I T1

PL <: ~ PL <: PLP~

negative negative positive

type type type

Fig. 3 Schematic diagram of M1 of Mimomys and its allies showing their occlusal patterns with terminology anddifferentiation types in enamel thickness. ACe: anteroconid complex, AL: anterior loop, BRA: buccalreentrant angle, ei: enamel island, LRA: lingual reentrant angle, ma: Mimomys angle, PL: posterior loop,T: triangle. BRA, LRA and T are numbered for showing the homology of each part.

-Cementum absent; enamel positive type (Microtus-type) in

differentiation; M1 and M2 with two roots; M3 with no enamel

island; posterior loop of M3 slenderer, where its dentine field

does not have enough space for the enamel island; M1 with

neither Mimomys angle nor enamel island (except in youngindividuals) Borsodia

3. Taxonomic notes on Promimomys

following European and North American species are

considered to be included in the genus: P. cor Kretzoi,

1955; P. insuliferus Kowalski, 1958; P. microdon Janossy,

1974; P. mimus (Shotwell, 1956); P. moldavicus (Kormos,

1932). On the other hand, Zazhigin (1980) listed antiquus,

baschkirica, cor, gracilis, moldavicus, occitanus and

stehlini as the members of Promimomys, but all of them

except cor and moldavicus are assignable to Mimomys inthe present treatment.

The genus Promimomys was described by Kretzoi

(1955) on the basis of a material with uncertain geological

age from Hungary. In China, only one mandible with I, M 1

and M 2 is referred to this genus. The mandible was

collected from the Lower Pliocene of Xindong Cave,

Huainan (Fig. 4), and was named P. asiaticus as a new

species by Jin and Zhang (2005). Besides this, the

4. Taxonomic notes on Mimomys

The genus Mimomys is an extensive genus comprising

many species and was established by Forsyth Major (1902)on the basis of remains from the Lower Pleistocene of

England. Aratomys described by Zazhigin in Gromov and

4 Revision of the Extinct Voles of Mimomys and its Allies

Fig. 4 Representative localities yielding Mimomys and its allies in China andthe adjacent area. Detailed information on each locality is given inTable 1. "* Promimomys,. Mil1'lomys, eVillanyia, 0 Borsodia.

Polyakov (1977) and Cromeromys by Zazhigin (1980) are

considered to be synonymous with Mimomys. Remains

assigned to Mimomys have been recorded from many

localities in China (Fig. 4). We examined the literatures

de cribing the remains as well as the remains stored in the

Institute of Vertebrate Paleontology and Paleoanthro­

pology, Chinese Academy of Sciences. As the result of the

examinations, we recognize the following species as themembers of this genus:

Mimomys bilikeensis (Qiu and Storch, 2000)

M. orientalis You ng, 1935

M. youhenicus Xue, j 981

M. banchiaonicus Zheng et al., 1975

M. gansunicus Zheng, 1976

M. peii Zheng and Li, 1986

Besides these, Zheng and Li (1986) described the

remains assigned to M. cf. intermedius and Mimomys sp.

The representative localities yielding these forms are

mapped in Fig. 4, and detailed information on each locality

is given in Table 1. This figure and table also show

Transbaikal ian localities.

In regard to European species, we follow a

comprehensive review by Kowalski (2001) who listed

many species of Mimomys, although he did not treat pre­

Villanyian (early Pliocene) species. He included

Pitymimomys described by Tesakov (1998) in Mimomys.

The species listed by him are considered to have thediagnositic characters of Mimomys mentioned above. On

the other hand, Zazhigin (1980) showed a different scheme

on the taxonomy of Mimomys, in which European andSiberian species were treated. We include all the species

of Promimomys except P. cor and P. moldavicus, and allthe species of Cromeromys except "c. newtoni" listed by

him, in Mimomys.

Yoshinari KAWAMURA and Yingqi ZHANG

Table I Detailed information on the representative localities yielding Mimomys and its allies in China

and Transbaikalia. For location of each locality see Fig. 4.

5

Locality

ChinaI Xiyingzi, Linxi

2 Bilike

3 Xiashagou, Majuangou

ill, Xiaochangliang and

Danangou in the Nihe­wan Basin

4 Chaojiayan, Lishi

5 Haiyan and Jizigou in the

Yushe Basin

6 Xicun, Tunliu

7 Dachai, Xiangfeng

8 Dongyan, Pinglu

9 Youhe, Weinan

10 Jingou and Langgou,

J:-Ieshui

I J Localities 77076 and 77078in the Gonghe Basin

12 Longdan, Dongxiang

13 Xindong Cave, Huainan

14 Renzidong Cave,

Fanchang

15 Longgupo, Wushan

16 Nixi, Diqing

Transbaikalia17 Zasukh ino

18 Klochnevo 1, 11

19 Dodogol

20 Tologoi I.

21 Udunga

22 Beregovaya

Taxon (partly revised)

Borsodia chinensis

Mimomys bilikeensis

Mimomys orientalis , M cf.

youhenicus, M. gansunicus,Borsodia chinensis

Mimomys cf. intermedius

Mimomys orientalis

Mimomys sp.

Mimomys pei i

Mimomys orientalis

Mimomys youhenicus, Morientalis, M sp.

Mimomys banchiaonicus,Borsodia chinensis

Borsodia chinensis

Mimomys cf. gansunicus

Promimomys asiaticus

Alimomys cE gansunicus, Mcf peii, Villanyiafanchangensis

Mimomys peii

Villanyia hengduanshanensis

Mimomys cf. reidi, M.pseudintermedius, Borsodiaklochnevi

Mimomys cf. reidi, M. cf.

pusillus, M. psedintermedius,Borsodia klochnevi

Mimomys cf. pusillus,Borsodia chinensis

Mimomys gracilis, M aff.

stehlini, M cf. minor,Villanyia cf. eleonorae,"Pitymimomys koenigswaldi"

Mimomys cf. gracilis, M cf

stehlini, M cf. minor,Villanyia ex. gr. eleonorae

Mimomys minor, M cf reidi,M pseudintermedius,Villanyia eleonorae

Geological age

? Early Pleistocene

Early Pliocene

Late Pliocene to

Early Pleistocene

? Early Pleistocene

? Middle Pliocene

"Early Pleistocene"

? Late Pliocene(="Early Pleistocene")

? Middle Pliocene

Middle Pliocene

Middle Pliocene and

"Early Pleistocene"

? "Early Pleistocene"

Late Pliocene

(="Early Pleistocene")

Early Pliocene

Late Pliocene(="Early Pleistocene")

"Early Pleistocene"

"Early Pleistocene"

Late Pliocene

Late Pliocene

Early Pleistocene

Middle Pliocene

Middle Pliocene

Middle Pliocene

Reference

Zheng and Li (1986)

Qiu and Storch (2000)

Kormos (1934), Zheng and Li (1986), Zheng

and Cai (1991), Zhang, Kawamura and Cai

(2008), Cai et al. (2008)

Zheng and Li (1986)

Zheng and Li (1986)

Zong et al . (1982), Zheng and Li (1986)

Zheng and Li (1986)

Young (1935), Zheng and Li (1986)

Xue (1981), Zheng and Li (1986)

Research Group for the Huanghe Elephant

(1975), Zheng (1976), Zheng and Li (1986)

Zheng et al. (1985), Zheng and Li (J 986)

Qiu et al. (2004)

Jin and Zhang (2005)

Jin et al . (2000), Zhang, Kawamura and

Jin (2008)

Zheng (1993)

Zong (1987)

Vangengejm et al. (1990), Erbajeva (1998)

Erbajeva (1998), Alexeeva et af. (200 I)

Erbajeva (1973), Erbajeva and AJexeeva

(2000)

Alexeeva et al. (200 I), Erbajeva et al . (2006)

Alexeeva et al. (2001), Erbajeva et al. (2003)

Bazarov et al . (1976), Erbajeva (1976)Erbajeva et af . (2003)

6 Revision of the Extinct Voles of Mimomys and its Allies

S. Taxonomic notes on Villanyia

The genus Villanyia was described by Kretzoi (1956)

from Early Pleistocene remains collected from Hungary.

Its distinction from Mimomys and from Borsodia have

been controversial. The diagnostic characters given in the

key, however, distinguish the following two species

described in China as Villanyia:

Villanyiajanchangensis Zhang, Kawamura and Jin, 2008

V. hengduanshanensis (Zong, 1987)

These two species occurred from one locality each in

southern China (Fig. 4, Table 1). Moreover V.

hengduanshanensis is known from only one specimen,

although V. fanchangensis is represented by abundant

specimens.

Using the diagnostic characters, we include the

following European and Siberian species in Villanyia:

Villanyia exilis Kretzoi, 1956; Kowalski (200 I) included V.

veterior Kretzoi 1969, V. kowalskii Terzea, 1991 and V.

paraexilis Terzea 1991 in this species.

V. petenyii (Mehely, 1914); possibly including "Mimomys

praeh.ungaricus Schevtschenko, 1965" and "M.

tanaitica Schevtschenko, 1965."

V. eleonorae Erbajeva, 1976

V. novoasovica (Topachevsky and Scorik, 1977)

V. steklovi Zazhigin, 1980

V. betekensis Zazhigin, 1980

6. Taxonomic notes on Borsodia

Janossy and van der Meulen (1975) first described

Borsodia as a new subgenus of Mimomys, which was

characterized by the molars without cementum and with

positive-type enamel differentiation. Subsequently,

Borsodia was often treated as a full genus (Tesakov, 1993;

Kowalski, 2001 and others). We adopt the generic

treatment of Borsodia and consider that it is dis­

tinguishable from Mimomys and Villanyia by Llsing the

diagnostic characters given in the key. In China, only one

species, B. chinensis, is allocated to this genus. "Mimomys

heshuinicus Zheng, 1976" is considered to be a synonym

of this species (Zheng and Li, 1986). In Transbaikalia,Erbajeva (1973) described "Mimomys (Villanyia)

laguriformes", which was subsequently synonymized with

"M. (V.) chinensis" (= B. chinensis) by Zheng and Li

(1986). Additionally, Villanyia gromovi Erbajeva, 1976 is

probably a subspecies of "M. (V.) laguriformes" (Gromov

and Polyakov, 1977). Thus V. gromovi is considered to be

included in B. chinensis. As shown in Fig. 4 and Table 1,

B. chinensis is known from several localities in northern

China and Transbaikalia.

We regard the positive-type enamel differentiation in

the molars as the most important character to distinguish

Borsodia from the other genera. The following European

and Siberian species are considered to be included in

Borsodia because they have the diagnostic characters in the

key:

Borsodia newtoni (Forsyth Major, 1902); Mayhew and

Stuart (1986) included "Mimomys hungaricus Kormos,

1938" in this species. "Mimomys lagurodontoides

Schevtschenko, 1965" seems not to belong Borsodia,

because its M, usually has the Mimomys angle judging

from the figures of Schevtschenko (1965).

B.jejervaryi (Kormos, 1934)

B. arankoides (Alexandrova, 1976)

B. prolaguroides (Zazhigin, 1980)

B. klochnevi (Erbajeva, 1998)

The specific compositions of Borsodia and Villanyia

in this paper are considerably different from those in

Tesakov (1993) and Kowalski (2001). Among the species

listed above, B. klochnevi is known from Transbaikalia,

and was originally allocated to Villanyia (Erbajeva, 1998).

Its generic allocation to Borsodia was discussed in Zhang,

Kawamura and Jin (2008).

7. Chronospatial distribution of Villanyia and Borsodia

In China and Transbaikalia, the chronospatial

distributions of Villanyia and Borsodia are more limited

than that of Mimomys (Fig. 4, Table 1). Before discussing

their distributions, we emend the chronological confusion

having occurred during the publication of Zhang,

Kawamura and Jin (2008). In China, the boundary

between the Pliocene and Pleistocene is generally placed at

ca 2.6 Ma, which is different from the geochronological

scheme accepted in other countries (adopting ca 1.8 Ma as

the boundary; Fig. 5). In the draft of Zhang, Kawamura

and Jin (2008), the usage in the Chinese sense was

distinguished as "Early (Lower) Pleistocene" and

"Pleistocene" from that in the other scheme. These

quotation-marked words mean the Late Pliocene plus Early

Pleis tocene and the Late Pli ocene pI us Pleis tocene

respectively, in the other scheme (Fig. 5). Thus the words

Early Pleistocene, Lower Pleistocene and Pleistocene in

the following sentences in Zhang, Kawamura and Jin

(2008) should be emended as follows:

·'In this paper, "Early" or "Lower Pleistocene" means the

usage in Chinese sense, ... ' in page 163.

·'a cooling event at the beginning of the "Pleistocene"

caused southward migration of the Palaearctic elements.' in

page 164.

Yoshinari KAWAMURA and Yingqi ZHANG 7

TransbaikaliaNorthern ChinaSouthern China(3)

Q)c

Borsodia chinensis Borsodia chinensisQ)u (= B. laguriformes).8

M~ "ien c

M~ i'w co0:: -Q)

.~

>- c .c~ Q) i:!iuw 0

1ii- 'w0::

t?>- - 1.95Q) ~ Villanyiafanchangensis Borsodia klochnevic

~Q)

ei~~ "~ "~"iuQ

MN0::2 17 "0

co--J

Cco

2.6 '>, 1'?c~ Villanyia eleonorae'>

~:iQ)

Q)cQ) c MNu Q)

.Q u 16.Q0:: 0::Q)

Q)u rou~

--Jf--3.4

MN I·~.~15 ~ cca 3.6

ca 1.8

ca 2.6

caG8 (1) (2)

Fig. 5 Chronospatial distribution of Villanyia and Borsodia in China and the adjacent area. Note that two schemes ongeological age are shown for the difference in the Pliocene / Pleistocene boundary. In China, the boundary isgenerally placed at ca. 2.6 Ma. In this sense, we use "Early Pleistocene" for the late Pliocene plus the EarlyPleistocene in another scheme. (1): chronological division adopted here (boundary ages mainly by Gradstein et ai.,2004), (2): chronological division generally used in China, (3): mammal biozones and units in Europe (boundaryages by Steininger et ai., 1996). Transbaikalian species redrawn from Bazorov et ai. (1976) and Erbajeva (1998).ei: enamel island, ma: Mimomys angle.

.'which seems affirmative for the assignment of the "Early

Pleistocene" age of V.janchangensis.' in page 169.

In China and Transbaikalia, Villanyia occurred from

Late Pliocene sediments of Renzidong Cave (V.

janchangensis; Zhang, Kawamura and Jin, 2008), "Early

Pleistocene" (?) sediments of Nixi (V. hengduanshanensis;

Zong, 1987), the Middle Pliocene (MNI6b) of Beregovaya

(V. eleonorae; Erbajeva and Alexeeva, 2000) and the

Middle Pliocene (MNI6a) of Tologoi 1.1 and Udunga (V.

cf. eleonorae or V. ex. gr. eleonorae; Alexeeva et al., 2001,

Erbajeva et al., 2003). Among the localities, Tologoi 1.1

and Udunga are the earliest in geological age; Beregovaya

is next; and Renzidong Cave is the latest, although Nixi is

uncertain in age. Thus V. eleonorae is earlier than V.

janchangensis (Fig. 5). The descriptions of V. eleonorae

by Erbajeva (1976) and Bazarov et al. (1976) permit us its

morphological comparison with V. janchangensis. M2 of

V. eleonorae sometimes has three- roots, while the same

tooth of V. janchangensis always has two roots. M t of V.

eleonorae often has the Mimomys angle, but the same tooth

of V. janchangensis usually lacks the angle (Fig. 5). These

differences suggest that V. eleonorae is more primitive

than V. janchangensis. In M3, however, V. eleonorae has

the posterior enamel island easily disappeared by wear and

lacks an anterior one. In V. janchangensis, both of the

islands appear more frequently (Fig. 5). This character

suggests that V. eleonorae is more advanced than V.

janchangensis. V. eleonorae is considered to be a sister

species of V. janchangensis, although the phylogenetic

relationship between them is still unclear, owing to the

scarcity of the information on Villanyia in China.

As shown in Fig. 5, it is inferred that Villanyia

inhabited Transbaikalia in the Middle Pliocene, but its

distribution to China in this period is still unknown. In the

Late Pliocene, it was already replaced by Borsodia in

Transbaikalia, but still survived in southern China. Even

in sou them China, Villanyia was absent in the Early

Pleistocene fauna.

In regard to Borsodia, we recognize the two species,B. klochnevi and B. chinensis, in Transbaikalia and China

(Fig. 4, Table 1). B. klochnevi occurs from the Late

Pliocene sediments (MN17) of Klochnevo (1-1 and II-I)

and Zasukhino I in Transbaikalia (Erbajeva, 1998). B.

chinensis is known from many localities in China and

Transbaikalia, which are dated to the Early Pleistocene

(Table 1). Thus B. klochnevi is earlier than B. chinensis

8 Revision of the Extinct Voles of Mimomys and its Allies

(Fig. 5). Erbajeva (1998) pointed out that B. klochnevi had

several morphological characters intermediate between V.

eleonorae and B. chinensis, and stated "Villanyia klochnevi

(= B. klochnevi) n. sp. is considered to be transition form

between the archaic species Villanyia eleonorae and

advanced species Borsodia laguriformes (= B. chinensis)."

This statement is coincident with the chronological order

of the three species (Fig. 5). Thus it is likely that the tlu'ee

species represent a single evolutionary lineage, in which

Borsodia originated from Villanyia.

In Europe, Borsodia (B. newtoni, B. fejervari and B.

arankoides as its components) ranges from early

Villanyian to early Biharian (Kowalski, 2001). In westernSiberia, B. prolaguroides was recorded from the

Eopleistocene (=Villanyian) by Zazhigin (1980). In

Transbaikalia, B. klochnevi is dated to MN17 (late

Villanyian) as mentioned above. If the early Villanyian

occurrences of Borsodia in Europe are reliable, Borsodia

first appeared in Europe, and then spread eastward to

Transbaikalia and China. If Villanyia eleonorae, B.

klochnevi and B. chinensis form a single evolutionary

lineage as mentioned above, Villanyia gave rise to

Borsodia in Transbaikalia then spread vice versa. At any

rate, more chronological and taxonomical data are required

to clarify the origin and evolution of Borsodia.

As shown in Fig. 5, Borsodia appeared in Trans­

baikalia in the Late Pliocene, then its distribution expanded

to northern China in the Early Pleistocene. Borsodia

survived at least at 1.36 Ma in northern China (Zhang,

Kawamura and Cai, 2008).

8. Conclusion

Mimomys and its allies from China and Transbaikalia

comprise four genera such as Promimomys, Mimomys,

Villanyia and Borsodia. These genera are distinguishable

from each other by the molar characters given in the key.

Among the genera, Promimomys is known by the only

species P. asiaticus from the Lower Pliocene of China.

Mimomys is more diversified and has a longer cl1fono­

logical range from the Early Pliocene to Early Pleistocene.

It consists of M. bilikeensis, M. orientalis, M. youhenicus,

M. banchiaonicus, M. gansunicus, M. peii and severalother forms in this area. The taxonomic positions of

Villanyia and Borsodia have been controversial not only in

China but also in other countries. We consider that thefollowing species belong to Villanyia: V. exilis, V. petenyii,

V. eleonorae, V. novoasovica, V. steklovi, V. betekensis, V.

fanchangensis and V. hengduanshanensis. We also referthe following species to Borsodia: B. newtoni, B.

fejervaryi, B. arankoides, B. prolaguroides, B. klochnevi

and B. chinensis.

Among the species of Villanyia, V. eleonorae and V.

fanchangensis are known from the Middle Pliocene of

Transbaikalia and from the Upper Pliocene of southern

China, respectively (Fig. 5). Among the species of

Borsodia, B. klochnevi and B. chinensis are recorded from

the Upper Pliocene of Transbaikalia, and from the Lower

Pleistocene of Transbaikalia and northern China,

respectively.These records illustrate the chronospatial distributions

of Villanyia and Borsodia in China and Transbaikalia. In

the Middle Pliocene, Villanyia was distributed in Trans­

baikalia, where it was replaced by Borsodia in the Late

Pliocene. In southern China, however, Villanyia still

survived in the Late Pliocene. In the Early Pleistocene,

Villanyia was already extinguished from China, but

Borsodia was distributed from Transbaikalia to northern

China. These changes can be interpreted as the

replacement of the more archaic genus (Villanyia) by the

more advanced one (Borsodia) during the later part of the

Pliocene.

Acknowledgments

We wish to thank Professors Jin Changzhu and Zheng

Shaohua of the Institute of Vertebrate Paleontology and

Paleoanthropology, Chinese Academy of Sciences, and

Professor Shusaku Yoshikawa of Osaka City University

for giving us valuable discussion and facilities for this

study. Thanks are also due to Mr. Hiroyuki Taruno of the

Osaka Museum of Natural History for reviewing and

improving our draft.

References

Alexeeva, N. V., Erbajeva, M. A. and Sen, S. (2001)Geology and fauna, and preliminary correlation ofsediments of the main Late Cenozoic sites of theTransbaikal area. Quat.lnternat., 80-81, 93-100.

Bazarov, D. B., Erbajeva, M. A. and Rezanov, I. N. (1976)Geology and Fauna of the Anthropogene Key Sectionsof West Transbaikalia. Moscow, Nauka, 147pp. (inRussian)

Cai, B. Q., Li, Q. and Zheng, S. H. (2008) Fossil mammalsfrom Majuangou section of Nihewan Basin, Chinaand theil' age. Acta Anthrop. Sinica, 27, 129-142. (inChinese with English abstract)

Erbajeva, M. A. (1973) Early Anthropogene vole(Microtinae, Rodentia) with the features of Mimomysand Lagurodon from Transbaikalia. QuaternaryCommission of the USSR Bulletin, noAO, 134-138. (inRussian)

Erbajeva, M. A. (1976) The Early Eopleistocene rootteethvoles from the Transbaikal area. Vestnik Zoologii,

Yoshinari KAWAMURA and Yingqi ZHA G 9

no.5, 13-18. (in Russian with English summary)Erbajeva, M. A. (1998) Late Pliocene Itantsinian faunas in

Western Transbaikalia. Meded. Nederl. Inst.Toegepaste Geowetensch. TNO, no.60, 417-429.

Erbajeva, M. A. and Alexeeva, N. V. (2000) Pliocene andPleistocene biostratigraphic succession of Trans­baikalia with emphasis on small mammals. Quat.Internat., 68-71, 67-75.

Erbajeva, M. A., Alexeeva, . V. and Khenzykhenova, F.1. (2003) Pliocene small mammals from the Udungasite of the Transbaikal area. Coloquios de Pale­ontologia, vol. ext. I, 133-145.

Erbajeva, M. A., Alexeeva, N. V. and Khenzykhenova, F.1. (2006) Review of the Pliocene-Pleistocenearvicolids of the Bajkalian region. Palaeontographica,Abt. A, 278,113-123.

Forsyth Major, C. 1. (1902) Exhibition of, and remarksupon some jaws and teeth of Pliocene voles(Mimomys gen. nov.). Proc. Zool. Soc. London, 1902(1),102-107.

Grad tein, F. M., Ogg, 1. G., Srillth, A. G., Bleeker, W. andLourens, L. 1. (2004) A new Geologic Time Scale,with special reference to Precambrian and Neogene.Episodes, 27, 83-100.

Gromov, 1. M. and Polyakov, 1. Ya. (1977) Fauna oj theUSSR, Mammals Vol.3, No.8, Voles (Microtinae).Leningrad, auka. (English translation in 1992,Leiden, E. 1. Brill, 725pp.)

Janos y, D. and van der Meulen, A. 1. (1975) On Mimomys(Rodentia) from Osztramos-3, North Hungary. Proc.Kon. Nederl. Acad. Wetensch., Ser. B, 78, 381-39l.

Jin, C. Z. and Zhang Y. Q. (2005) First discovery ofPromimomys (Arvicolidae) in East Asia. Chinese Sci.Bull., 50, 327-332.

Jin, C. Z., Zheng, L. T., Dong, W., Liu, J. Y., XU, Q. Q.,Han, L. G., Zheng, 1. 1., Wei, G. B. and Wang, F. Z.(2000) The Early Pleistocene deposits andmammalian fauna from Renzidong, Fanchang, AnhuiProvince, China. Acta Anthrop. Sinica, 19, 184-198.(in Chinese with English abstract)

Kormos, T. (1934) Premiere preuve de I'existence du genreMimomys en Asie orientale. Trav. Lab. Geol. Fac. Sci.Lyon, fasc. 24, memo 20, 3-8.

Kowalski, K. (2001) Pleistocene rodents of Europe. FoliaQuaternaria, 72, 3-389.

Kretzoi, M. (1955) Promimomys cor n. g. n. sp., einaltertUmlicher Arvicolide au dem UngarischenUnterpleistozan. Acta Geol. Acad. Sci. Hung., 3, 89­94.

Kretzoi, M. (1956) Die altpleistozanen Wirbertierfaunendes Villanyer Gebirges. Geologica Hungarica, Ser.Palaeont., fasc. 27,1-264.

Mayhew, D. F. and Stuart, A. 1. (1986) Stratigraphic andtaxonomic revision of the fossil vole remains(Rodentia, Microtinae) from the Lower Pleistocenedeposits of eastern England. Phil. Trans. Royal Soc.London, Ser. B, 312, 431-485.

Qiu, Z. D. and Storch, G. (2000) The early Pliocenemicromammalian fauna of Bilike, Inner Mongolia,

China (Mammalia: Lipotyphla, Chiroptera, Rodentia,Lagomorpha). Senckenbergiana lethaea, 80, 173-229.

Qiu, Z. X., Deng, T. and Wang, B. Y. (2004) EarlyPleistocene mammalian fauna from Longdan,Dongxiang, Gansu, China. Palaeont. Sinica, New Ser.C, no. 27, 1-198, pls.I-34. (in Chinese with Englishummary)

Research Group for the Huanghe Elephant (1975) TheHuanghe Elephant. Beijing, Science Press, 46pp.,20pls. (in Chinese)

Schevtschel1ko, A. 1. (1965) Faunistic complexes of smallmammals from Upper Cenozoic deposits in the south­western part of the Russian Plain. In: StratigraphicImportance oj Small Mammalian Anthropogen Fauna(Nikiforova, K. V. ed.), Moscow, Nauka, 7-59. (inRussian with English summary)

Steininger, F. F., Berggren, W. A., Kent, D. V., Bernor, R.L., Sen, S. and Agusti, 1. (1996) Circum­Mediterranean Neogene (Miocene and Pliocene)marine-continental chronologie correlations ofEuropean mammal units. In: The Evolution ojWestern Eurasian Neogene Mammal Faunas (Bernor,R. L., Fahlbusch, V. and Mittmann, H.-W. eds.), ewYork, Columbia University Press, 7-46.

Teilhard de Chardin, P. and Piveteau, 1. (1930) Lesmammiferes fossiles de Nihowan (Chine). Ann.Paleont., 19,1-134, pls.I-23.

Tesakov, A. S. (1993) Evolution of Borsodia (Arvicolidae,Mammalia) in the Villanyian and in the EarlyBiharian. Quat. Internat., 19,41-45.

Tesakov, A. S. (1998) Voles of the Tegelen fauna. Meded.Nederl. inst. Toegepaste Geowetensch. TNO, no.60,71-134.

Vangengejm, E. A., Erbajeva, M. A. and Sotnikova, M. V.(1990) Pleistocene mammals from Zasuhino, westernTransbaikalia. Quartarpalaontologie, 8, 257-264.

Xue, X. X. (1981) An Early Pleistocene mammalian faunaand its stratigraphy of the river You, Weinan, Shenxi.Vertebrata PalAsiatica, 19, 35-44, pls.I-2. (in Chi­nese with English abstract)

Young, C. C. (1935) Miscellaneous mammalian fossilsfrom Shansi and Honan. Palaeont. Sinica, Ser. C, 9(fasc. 2), 1-57.

Zazhigin, V. S. (1980) Late Pliocene and Anthropogenerodents of the south of western Siberia. Acad. Sci.USSR Trans., 339, 1-155. (in Russian)

Zhang, Y. Q., Kawamura, Y. and Caj, B. Q. (2008) Smallmammal fauna of Early Pleistocene age from theXiaochangliang site in the Nihewan Basin, Hebei,northern China. The Quat. Res. (Daiyonki-Kenkyu),47,81-92.

Zhang, Y. Q., Kawamura, Y. and Jin, C. Z. (2008) A newspecies of the extinct vole Villanyia from RenzidongCave, Anhui, East China, with discussion on relatedspecies from China and Transbaikalia. Quat. Internat.,179,163-170.

Zheng, S. H. (1976) Small mammals of Middle Pleistocenein Heshui, Gansu. Vertebrata PalAsiatica, 14, 112­119, pl.l. (in Chinese)

10 Revision of the Extinct Voles of Mimomys and its Allies

Zheng, S. H. (1993) Quaternary Rodents of Sichuan­Guizlwu Area, China. Beijing, Science Press, 270pp.(in Chinese with English summary)

Zheng, S. H. and Cai, B. Q. (1991) Micromammalianfossils from Danangou of Yuxian, Hebei. In:Contributions to the XIII INQUA, Institute ofVertebrate Paleontology and Paleoanthropology,Academia Sinica, Beijing, Beijing Scientific andTechnological Publishing House, 100-131. (inChinese with English summary)

Zheng, S. H. and Li, C. K. (1986) A review of ChineseMimomys (Arvicolidae, Rodentia). VertebrataPalAsiatica, 24,81-109, pl.1. (in Chinese withEnglish summary)

Manuscript received September 1, 2008.

Revised manuscript accepted October 29, 2008.

Zheng, S. H., Wu, W. Y., Li, Y. and Wang, G. D. (1985)Late Cenozoic mammalian faunas of Guide andGonghe Basins, Qinghai Province. VertebrataPalAsiatica, 23, 89-134, pls.l-lO. (in Chinese withEnglish summary)

Zong, G. F. (1987) Note on some mammalian fossils fromthe Early Pleistocene of Di-qing county, Yunnan.Vertebrata PalAsiatica, 25, 69-76. (in Chinese withEnglish abstract)

Zong, G. F., Tang, Y. 1., XU, Q. Q. and Yu, Z. Q. (1982)The Early Pleistocene in Tunliu, Shanxi. VertebrataPalAsiatica, 20, 236-247, pl.1. (in Chinese withEnglish abstract)