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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.
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