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Title: Comments on the Taxonomic Status of Apodemus peninsulae, with Description of a New Subspecies from North China
Author: Jones, J. Knox, 1929-1992
Language: English
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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.

[Illustration: 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 [Male], No. 141691 USNM, ×2.

_b._ _Apodemus flavicollis flavicollis_ (Melchior), Mauseklippe,
Germany, young [Male], No. 112895 USNM, ×10.

_c._ _Apodemus peninsulae peninsulae_ (Thomas), Central Nat'l Forest,
near Pup'yong-ni, 200 m., Korea, subadult [Female], No. 300650 USNM,
×10.

_d._ _Apodemus peninsulae peninsulae_ (Thomas), 6 mi. S Yongdongp'o,
Korea, adult [Male], 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 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_.

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, _flavicollis_-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

     _Type._--Adult female molting from winter to summer pelage, skin
     and skull, U. S. National Museum no. 175523, from 30 miles west of
     Kuei-hua-cheng, 7000 ft., northern Shansi, China; obtained on 23
     May 1912 by Arthur de Carle Sowerby, original no. 456.

     _Distribution._--Known presently from eastern Kansu eastward
     through Shensi, Shansi and Hopeh to southern Jehol, probably also
     in northeastern Szechuan, exact limits of range unknown.

     _Diagnosis._--Size small for species (see measurements). Color:
     Upper parts (fresh summer pelage) averaging near (15'_a_)
     Ochraceous-Buff, suffused with blackish (especially mid-dorsally);
     winter pelage much paler; underparts grayish-white, individual
     hairs plumbeous at base, tipped with white; ears pale brownish;
     feet whitish above, darker below; tail bicolor, pale brownish
     above, whitish below. Skull: Small (see measurements); rostrum
     somewhat shortened and conspicuously down-curved; zygomatic notches
     relatively shallow; zygomatic plates narrow; braincase
     proportionally more inflated than in other subspecies of the
     species; auditory bullae moderately inflated; upper incisors
     slender, their faces averaging bright yellowish-orange.

     _Measurements._--External measurements of the holotype, followed by
     those of an adult male and female from the type locality, are,
     respectively: Length of head and body, 101, 102, 100; length of
     tail, 93, ----, 102; length of hind foot (_su_), 21, 21.5, 23;
     length of ear from notch, 14, 16, 15.5. Corresponding measurements
     for an adult female from 20 mi. E Taiyuan, Shansi, are: 91, 99, 23,
     16. For cranial measurements see Table 1.


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
============================================================================
        Sex           |      |      |      |     |      |      |      |     |
    and catalogue     |   A  |   B  |   C  |  D  |   E  |   F  |  G   |  H  |
number or number of   |      |      |      |     |      |      |      |     |
individuals averaged  |      |      |      |     |      |      |      |     |
----------------------------------------------------------------------------

  _Apodemus peninsulae peninsulae_, various localities in central Korea
Average 10 (4M, 6F)   | 29.2 | 14.2 | 11.8 | 4.7 | 20.1 | 11.4 | 10.2 | 4.3 |
Minimum               | 28.3 | 13.8 | 11.5 | 4.6 | 19.2 | 10.8 |  9.9 | 4.1 |
Maximum               | 29.8 | 14.6 | 12.2 | 5.1 | 20.7 | 12.0 | 10.5 | 4.4 |

  _Apodemus peninsulae nigritalus_, Tapucha, Altai Mts., Siberia
USNM 175164, M (type) | 28.8 | 14.8 | 12.4 | 4.5 | 20.8 | 11.7 | 11.0 | 4.4 |
USNM 175171, F        | 28.2 | 13.7 | 11.8 | 4.5 | 19.8 | 11.2 | 10.3 | 4.5 |

  _Apodemus peninsulae praetor_, Sungari River, 60 mi. SW Kirin, Manchuria
USNM 197792, M (type) | 30.5 | .... | 12.5 | 4.7 | 21.5 | 12.5 | 10.3 | 4.6 |
USNM 197798, F        | 30.2 | 14.4 | 11.8 | 4.6 | 21.6 | 12.7 | 10.6 | 4.6 |

  Mukden, Manchuria
USNM 197782, M        | 29.5 | 14.8 | 12.4 | 4.8 | 20.6 | 12.2 | 10.5 | 4.2 |

  _Apodemus peninsulae sowerbyi_, Kuei-hau-cheng, Shansi
USNM 175523, F (type) | 27.9 | 13.3 | 11.7 | 4.5 | 19.6 | 11.4 |  9.9 | 4.0 |
USNM 175521, M        | 27.6 | .... | 11.5 | 4.6 | 18.9 | 11.4 |  9.7 | 4.1 |
USNM 175522, F        | 27.9 | .... | 11.8 | 4.6 | 19.4 | 11.3 |  9.8 | 4.2 |

  20 mi. E Taiyuan, Shansi
USNM 172558, F        | 27.4 | 13.8 | 11.5 | 4.6 | 19.4 | 11.6 | 10.1 | 4.4 |

  12 mi. S Yenan, Shensi
USNM 155072, M        | 27.8 | 14.1 | .... | 4.4 | 19.5 | 11.0 | .... | 4.3 |
USNM 155073, F        | 27.7 | 13.3 | 11.5 | 4.5 | 19.4 | 11.0 | 10.0 | 4.2 |
USNM 155075, M        | 27.9 | 13.5 | 11.4 | 4.5 | 19.2 | 11.0 | 10.0 | 4.3 |

  Hsin-lung-shan, 65 mi. NE Peking, Jehol
USNM 219229, M        | 27.7 | 13.8 | 11.4 | 4.5 | 19.0 | 10.9 | 10.4 | 4.4 |

15 mi. S Lanchow, Kansu
USNM 155171, M        | 27.7 | 13.6 | 11.7 | 4.6 | 19.0 | 11.3 |  9.9 | 4.5 |

_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; 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.

     _Specimens examined._--Thirty-three, all from North China, as
     follows: JEHOL: Hsin-lung-shan, 65 mi. NE Peking, 3000 ft., 4.
     KANSU: 15 mi. S Lanchow, 7400 ft., 1. SHANSI: Chiao-cheng-shan, 90
     mi. W Taiyuan, 7000-8000 ft., 4; 30 mi. W Kuei-hau-cheng, 7000 ft.,
     5; Lung-wang-shan, 20 mi. E Taiyuan, 4000 ft., 10; 18 mi. W
     Taiyuan, 5000 ft., 1; 50 mi. NW Taiyuan, 5500 ft., 4. SHENSI: 12
     mi. S Yenan, 4000 ft., 4.

_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 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._


26-3854





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