University of Kansas Publications

Museum of Natural History

Volume 9, No. 8, pp. 337-346, 1 fig. in text, 1 table

August 15, 1956

Comments on the Taxonomic Status of

Apodemus peninsulae, with Description

of a New Subspecies from North China

BY

J. KNOX JONES, JR.

University of Kansas

Lawrence

1956

University of Kansas Publications, Museum of Natural History



Editors: E. Raymond Hall, Chairman, A. Byron Leonard,

Robert W. Wilson



Volume 9, No. 8, pp. 337-346, 1 fig. in text, 1 table

Published August 15, 1956

University of Kansas

Lawrence, Kansas

PRINTED BY

FERD VOILAND, JR., STATE PRINTER

TOPEKA, KANSAS

1956



26-3854

Comments on the Taxonomic Status of Apodemus peninsulae, with

Description of a New Subspecies from North China

BY

J. KNOX JONES, JR.

In the past several years the United States National Museum has received
a large number of mammals from central and southern Korea through the
auspices of the Commission on Hemorrhagic Fever of the Armed Forces
Epidemiological Board. Among these Korean collections are more than a
hundred specimens of a murine rodent originally described as "Micromys
speciosus peninsulae" by Oldfield Thomas but currently placed in the
genus Apodemus. In attempting to ascertain the specific relationships
of this mouse I have examined, through the generosity of Dr. David H.
Johnson, Acting Curator of Mammals, most of the other Oriental specimens
of the subgenus Sylvaemus in the U. S. National Museum and it is on
this combined material that the following comments and description are
based.

Three general groups of the genus Apodemus are presently known to
occur on the mainland of northeast Asia. One is the distinctive
Apodemus agrarius, lone representative of the subgenus Apodemus. The
others, both in the subgenus Sylvaemus and closely resembling each
other, are represented by a small animal that is currently regarded as
conspecific with Apodemus sylvaticus and a larger animal of which the
Korean mouse, peninsulae, is representative. The oldest trivial name
applied to the large Sylvaemus is major of Radde, 1862, in the
combination [Mus sylvaticus] vrt. major. This is, however, twice
preoccupied (see Ellerman and Morrison-Scott, 1951:566). The next
available name is peninsulae of Thomas, 1907, which was applied to
mice from central and southern Korea (type from Mun'gyong, 110 mi. SE
Seoul, Korea), and was originally proposed as a subspecies of the
insular Japanese species, Apodemus speciosus. G. M. Allen (1940:949),
who recognized peninsulae as a monotypic species, was the first
investigator to make the important distinction that it was not
conspecific with the Japanese speciosus, although Hollister (1913:1-2)
and Miller (1914:89) had previously used the combination Apodemus
peninsulae, evidently with the same thought in mind.[Pg 340]

Fig. 1. Ventral views of skulls and left maxillary
tooth-rows of two species of the genus Apodemus.


a. Apodemus flavicollis flavicollis (Melchior), Lolland, Denmark,
adult , No. 141691 USNM, ×2.


b. Apodemus flavicollis flavicollis (Melchior), Mauseklippe,
Germany, young , No. 112895 USNM, ×10.


c. Apodemus peninsulae peninsulae (Thomas), Central Nat'l Forest,
near Pup'yong-ni, 200 m., Korea, subadult ϓ2, No. 300650 USNM,
×10.


d. Apodemus peninsulae peninsulae (Thomas), 6 mi. S Yongdongp'o,
Korea, adult , No. 299554 USNM, ×2.

In comparing the ventral views of skulls note especially the size and
location of incisive foramina and posterior palatine foramina as well as
the breadth of mesopterygoid fossae. In comparing the left maxillary
tooth-rows note especially the size of M3 and the reduced
posterointernal cusp on Ml in A. peninsulae.]

More recently, Ellerman (1949:32) and Ellerman and Morrison-Scott
(1951:566) have arranged peninsulae as a subspecies of Apodemus
flavicollis under the assumption that all the members of the subgenus
Sylvaemus on the eastern Asiatic mainland are subspecies of one or
another of the species of western Europe, A. flavicollis or A.
sylvaticus. Ellerman (in Ellerman and Morrison-Scott, 1951:564) states:
"The majority of the forms I distribute in a somewhat arbitrary manner
between sylvaticus, average smaller skull, and flavicollis, average
larger skull; occurring together nearly throughout the Palaearctic. I
feel fairly sure that there are some errors of judgment in my
arrangement, and equally sure that there is no other way to define
species in this very large and difficult group." I have compared the
specimens of peninsulae available to me from central and southern
Korea with specimens of A. f. flavicollis from Denmark, Germany and
Sweden and find, although the[Pg 341] two are similar in many ways, that
peninsulae differs from flavicollis in several important characters:
Mammae 1-2=6 in flavicollis, and 2-2=8 in peninsulae; incisive
foramina reaching level of alveoli of M1, or nearly so, in
flavicollis, but ending conspicuously short of that level in
peninsulae; posterior palatine foramina large in flavicollis and
opposite a point where M1 and M2 meet, but small in peninsulae and
situated farther back on the palate, opposite M2. Moreover, peninsulae
lacks the characteristic buffy throat patch of flavicollis, has a much
reduced posterointernal cusp on the M1, a relatively (frequently
actually) larger M3 and, on the average, a broader mesopterygoid fossa.
In view of these differences, all of which appear to be constant, I
consider peninsulae specifically distinct from flavicollis.
Throughout its known geographic range (see below) peninsulae is
evidently confined to wooded terrain, either scrub or brush types or
forested areas, and the vernacular name wood mouse, therefore, seems
appropriate for this species.

The type specimens of Apodemus praetor Miller (type from Sungari
River, 60 mi. SW Kirin, Manchuria) and Apodemus nigritalus Hollister
(type from Tapucha, Altai Mountains, Siberia) agree with peninsulae as
concerns the above characters and differ from it only in minor external
and cranial features. They are, therefore, here considered as subspecies
of the latter.

Ellerman (1949:32) and Ellerman and Morrison-Scott (1951:567) regarded
nigritalus, like peninsulae, as a subspecies of flavicollis. The
subspecies praetor, on the other hand, has generally been regarded as
a synonym of peninsulae by recent authors. Howell (1929:58) noted that
the holotype was, "... a phenomenally large specimen such as is
encountered occasionally in almost all groups of rodents." He ascribed
the color differences noted by Miller to "seasonal" variation. The
holotype of praetor is undeniably larger than the other adult
specimens listed in the original description. These paratypes and other
specimens of praetor available to me are approximately the same size
externally and average only slightly larger cranially than specimens of
peninsulae from central and southern Korea. However, the dorsal
coloration of praetor is somewhat darker and duller than that of
peninsulae, especially in summer pelage when praetor lacks the
conspicuous bright ochraceous tinge of the Korean specimens. In
addition, praetor has broader zygomatic plates with correspondingly
deeper zygomatic notches and the color on the face of the upper incisors
averages much more orange than in peninsulae.[Pg 342]

In the north then, wood mice range from Korea and Manchuria westward at
least as far as the Altai Mountains. For mice from the intervening
Siberian areas Russian workers have used the name major which, as
noted above, is unavailable. The exact relationships of the mice of
these areas to previously named subspecies is unknown to me and I have
not seen specimens of "Mus (Alsomys) major rufulus" of Dukelsky, 1928,
the type locality of which is 75 versts (approximately 50 miles) SE
Vladivostok, Siberia. It appears to be of the same species as
peninsulae and judging from the original description it closely
resembles praetor. Neither have I seen specimens of the Sakhalin
Island mouse, giliacus, which Ellerman (1949:32) regards as a
subspecies of Apodemus sylvaticus. I feel reasonably sure, however,
that it will prove to be a subspecies of peninsulae. In the original
description giliacus was referred to as, "Most closely allied to the
Korean subspecies..." (Thomas, 1907:411).

In China the extent of the distribution of Apodemus peninsulae is also
uncertain. Allen (1940:949-50) reported its occurrence from Jehol and
Hopeh in the northeast, southwestward through Shansi, Shensi and eastern
Kansu to Szechuan and northwestern Yunnan. Throughout most of this
region it occurs with another mouse, currently regarded as conspecific
with Apodemus sylvaticus, and the two kinds have been confused by some
previous authors. Howell (1929:58), for instance, reported twelve
specimens of peninsulae from 65-75 mi. NE Peking but my examination of
these mice indicates that only four are peninsulae while the others
are referrable to what is currently regarded as Apodemus sylvaticus
draco. Another subspecies of sylvaticus, A. s. orestes, occurs in
Szechuan and Yunnan and it is certain that some records of distribution
ascribed to peninsulae from those provinces actually represent
orestes (see Allen, 1940:949-50). A. sylvaticus is distinguishable
from peninsulae by darker ears, blackish preauricular patches, dark
eye rings, a noticeably smaller skull, incisive foramina that reach the
level of Ml (or nearly so), much larger auditory bullae, and a more
fully developed posterointernal cusp on M1. Too, sylvaticus typically
has 1-2=6 mammae although Allen reports finding a 2-2=8 formula in some
specimens. Apodemus latronum, regarded as a full species by Osgood
(1932:318) and G. M. Allen (1940:950) but as a subspecies of
flavicollis by Ellerman (1949:32) and Ellerman and Morrison-Scott
(1951:567), also occurs in Szechuan and Yunnan. Its relatively dark
color, large feet and large ears, flavi[Pg 343]collis-like skull and large
molar teeth immediately separate it from peninsulae although the two
possibly have been confused in the earlier literature. Until a complete
revisionary study of the Asiatic members of the subgenus Sylvaemus can
be undertaken the presence of peninsulae in southwestern China must
remain in question.

The western limits of the geographic range of Apodemus peninsulae are
unknown. Apodemus gurkha Thomas, 1924, from Nepal is said to have
2-2=8 mammae but the description is not otherwise suggestive of close
relationship to peninsulae. Farther to the west, Apodemus flavicollis
rusiges Miller, 1913, from Kashmir seems to have been properly assigned
as a subspecies of flavicollis (cotypes and large series in USNM).

Wood mice almost certainly do not occur in the Gobi Desert. They are
known as far west as the Altai Mountains to the north of the Gobi and at
least as far west as Kansu (see below) to the south of it. Whether the
geographic range of the species skirts the western edge of the arid
regions of northern China is at present unknown; perhaps it does not. At
any rate, mice available to me from the North Chinese provinces of
Jehol, Shansi, Shensi and Kansu are notably different in certain
external and cranial features from other known races of Apodemus
peninsulae and are here given subspecific recognition. All measurements
are in millimeters. Capitalized color terms are from Ridgway (1912).

Apodemus peninsulae sowerbyi, new subspecies

Table Key:

A: Occipitonasal length
B: Zygomatic breadth

C: Mastoid breadth

D: Interorbital length

E: Frontonasal length

F: Nasal length

G: Depth of skull

H: Alveolar length of maxiary tooth-row

TABLE 1.—Cranial Measurements of Adults of Several Subspecies of Apodemus peninsulae

Comparisons.—From Apodemus peninsulae peninsulae (specimens from
various localities in central Korea), A. p. sowerbyi differs in:
External size smaller throughout, especially hind foot; upper parts,
especially in summer pelage, and dorsal aspect of tail paler;[Pg 345] skull
smaller and less massive; braincase proportionally more inflated;
rostrum shorter and noticeably down-curved. From Apodemus peninsulae
praetor of Manchuria (holotype and paratypes), A. p. sowerbyi differs
in most of the same ways in which it does from peninsulae as well as
in having more shallow zygomatic notches, narrower zygomatic plates and
smaller, more slender, upper incisors. From Apodemus peninsulae
nigritalus of the Altai Mountains of Siberia (holotype and paratypes),
A. p. sowerbyi differs in: Smaller size, both external and cranial;
paler dorsal coloration; less convex cranial outline in lateral view;
smaller auditory bullae.

Remarks.—Apodemus peninsulae sowerbyi is named in honor of the late
Arthur de Carle Sowerby whose collections of mammals from North China
and Manchuria have added so much to our meager knowledge of that part of
the world.

Four specimens from Hsin-lung-shan, 65 mi. NE Peking, here assigned to
sowerbyi, are darker dorsally than mice from farther to the west and
in this respect may show approach to A. p. praetor. In all other
features, however, they closely resemble the new subspecies.

All of the specimens of sowerbyi available to me are from altitudes of
3000 feet or higher. At lower elevations in North China, destruction of
wooded habitats owing to intense land-use practices has probably
restricted the distribution of sowerbyi primarily to hilly and
mountainous areas where brushy, scrub and forest habitats still prevail.

Apodemus peninsulae, then, is known or suspected to occur over much of
southeastern Siberia, Manchuria, Korea and North China. The western
limits of its geographic range are unknown. Over this vast area only
four subspecies, one newly named, can be ascribed with certainty to
peninsulae whereas only two other kinds, giliacus of Thomas from
Sakhalin and rufulus of Dukelsky from extreme southeastern Siberia are
probably conspecific with it, the latter possibly a synonym of
praetor. These considerations underscore the preliminary nature of the
present paper. The mammalian fauna of northeastern Asia is scarcely
better known today than was[Pg 346] that of North America in 1885 when Dr. C.
Hart Merriam organized what was later to become the U. S. Biological
Survey.

It seems to me that the correct names of four kinds of wood mice
discussed above are as follows:

Apodemus peninsulae peninsulae (Thomas, 1907)

Apodemus peninsulae nigritalus Hollister, 1913

Apodemus peninsulae praetor Miller, 1914

Apodemus peninsulae sowerbyi Jones, 1956

LITERATURE CITED

Allen, G. M.

1940. The mammals of China and Mongolia. Amer. Mus. Nat. Hist., New
York, 2:XXVI + 621-1350, September 3.

Ellerman, J. R.
1949. The families and genera of living rodents. British Mus.,
London, 3:V + 1-210, March.

Ellerman, J. R., and T. C. S. Morrison-Scott.
1951. Checklist of Palaearctic and Indian mammals, 1758 to 1946.
British Mus., London, 810 p., November 19.

Hollister, N.
1913. Two new mammals from the Siberian Altai. Smith. Misc. Coll.,
60:1-3, March 13.

Howell, A. B.
1929. Mammals from China in the collections of the United States
National Museum. Proc. U. S. Nat. Mus., 75:1-82, June 7.

Miller, G. S., Jr.
1914. Two new murine rodents from eastern Asia. Proc. Biol. Soc.
Washington, 27:89-92, May 11.

Osgood, W. H.
1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic
expeditions. Field Columb. Mus. Nat. Hist., Zool. Series,
18:193-339, August 19.

Ridgway, R.
1912. Color standards and color nomenclature. Washington, D. C.,
published by the author.

Thomas, O.
1907. The Duke of Bedford's zoological explorations in eastern
Asia.—IV. A list of small mammals from the islands of Saghalien
and Hokkaido. Proc. Zool. Soc. London, 1907:404-414, August 1.

Transmitted May 12, 1956.

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The proportions (x2 and x10) in the figure caption are taken from the original text;
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