šM“®ŠwŽ•(J. Mamm. Soc. Japan) 10 (2): 69-80 (1984)

A Review of the -moles (Genus ) of China and Burma

Robert S. HOFFMANN

The University of Kansas, Museum of Natural History and Department of Systematics and Ecology, Lawrence, Kansas 66045, U.S.A.

The genus Uropsilus was first described by MILNE-EDWARDS (1871) from material sent back from Muping (Moupin, now Baoxing), Sichuan, by Pere Armand DAVID;

the type species is U. soricipes. These small insectivores are shrew-like in external

appearance, but exhibit a -like skull and dentition. The tail is long and forefeet are not enlarged, while the zygomatic arch is complete, and the tympanic bones form an auditory bulla (Fig. 1). Thus, this•gshrew-mole•hlacks skeletal specializations for digging found in more derived moles, and the derived characters of skull and

dentition found in . However, Uropsilus soricipes exhibits a derived dental formula compared to the primitive eutherian condition seen in some genera of moles (Ta/pa, Parascalops, , Condylura; i 3/3, c 1/1, p 4/4, m 3/3=44), possessing, i 2/1, c 1/1, p 3/3, m 3/3=34 according to Mime-Edwards (Fig. 1a). Subsequent to the initial record, other specimens from Yunnan (POUSARGUES,

1896) and southern Sichuan (THOMAS, 1911) came to light in which the dental formula differed from that of the specimen described by Mime-Edwards; additional upper and lower incisors seemed to be present (i 3/2, c 1/1, p 3/3, m 3/3=38). This was in- terpreted as individual variation (Fig. 1b). Soon, however, THOMAS (1912) changed his mind. Receipt of new specimens from western and southeastern Sichuan revealed still a third dental formula, and prompted THOMAS to erect two new monotypic genera to encompass the variation. These were: Rhynchonax andersoni THOMAS, 1911, type locality Omi-san (Emei Shan), Sichuan; dental formula i 2/2, c 1/1, p 4/3, m 3/3=38

(corrected from THOMAS, 1911, see above); and Nasillus gracilis Thomas, 1911, type locality Mt. Chin-fu-san (Jingfu Shan) near Nan-chwan (Nanchuan), Sichuan (Fig. 2); dental formula i 2/1, c 1/1, p 4/4, m3/3=38 (Fig. 1c).

This arrangement was not greeted with universal acceptance. ALLEN (1912a) continued to regard the several dental formulae as individual variation within a single species, U. soricipes, as did HOWELL (1929) and OsGooD (1937). Later, however,

ALLEN (1938) changed his views somewhat, stating:•gThe fact that... these three groups [genera] seem to have fairly distinct areas of geographic distribution, may be interpreted as favoring their distinctness.•h

That Allen may have begun to change his mind earlier is evidenced by his having described two new subspecies, Rhynchonax andersoni atronates from the Salween drainage in southwestern Yunnan, and R. a. nivatus from the Likiang (Lichiang)

Range, western Yunnan (Fig. 2) (ALLEN, 1923). These new subspecies were described 70 哺 乳 動 物 学 雑 誌 Vol. 10, No. 2 1984”N8ŒŽ Shrew-moles of China and Burma 71

Fig. 2. Localities of occurrence of the genus Uropsilus. Open circles, U. gracilis; squares, U. andersoni; solid circles, U. soricipes.

as smaller than typical R. andersoni, and having different tooth proportions. The reason for these differences became apparent when OSGOOD (1937) pointed out that the topotypical series of these two taxa, including both holotypes, possessed the dentitions described by THOMAS (1912) for Nasillus rather than Rhynchonax. I have subsequently examined these specimens and confirm OsGooD's conclusion. Meanwhile, THOMAS (1922) had described a new species, Nasillus investigator, from the Kiu-kiang•\Salween divide, Yunnan. This type locality is north, up the

Salween River from the type locality of•gR. a.•hatronates, and not far west of the type locality of•gR. a.•hnivatus (Fig. 2). THOMAS (op. cit.) described N. investigator as larger than N. gracilis, but ALLEN (1938) observed that •gundoubtedly it will eventual- ly prove to be at most a slightly larger subspecies of N. gracilis ...•h 72 哺乳動 物 学雑誌 Vol. 10, No. 2

It is curious that even though ALLEN (op. cit.) listed OscooD's 1937 paper (wherein

atronates and nivatus were said to have Nasillus dentition) in his bibliography, he did

not cite the paper in the pertinent text passages. He instead retained these taxa as subspecies of Rhynchonax andersoni, even though recognizing the occurrence of

Nasillus in that part of Yunnan. Subsequently ANTHONY (1941) compounded this

confusion by referring a series of Burmese shrew-moles with Nasillus dentition to Rhynchonax a. andersoni, after comparing them to•g... a series of over eighty a.

andersoni, collected in western Szechwan by the Sage West China Expedition and nine a. atronates Allen, including the type ...•hANTHONY (op. cit.) concluded that atronates was a synonym of andersoni•g... not only on the basis of the characters shown but on geographical grounds, since the range of atronates appears to be midway of the range of andersoni (as I am now extending it) [i.e., from western Sichuan to northern Burma] and not isolated from it by virtue of special conditions such as altitude.•h

There the matter rested, until ELLERMAN and MoRRISON-ScoTT (1951) suggested that there was insufficient evidence to retain the taxa as distinct, monotypic genera.

They recognized one species, Uropsilus soricipes, with five subspecies : U. s, soricipes; U. s. gracilis; U. s. andersoni (incl. atronates); U. s. investigator and U. s. nivatus.

Since then this arrangement has been adopted by other workers (CRANBROOK 1960-

61; CORSET and HILL, 1980; HONACKI et al., 1982), although GUREEV (1979) con- tinued to recognize three genera and four species. I have examined all but one of the holotypes, as well as the majority of extant specimens in North America and Europe, and have concluded that the present taxonomic arrangement is incorrect.

For reasons detailed below, I recognize three species in a single genus, Uropsilus.

Materials and Methods

A total of 522 specimens was examined. Many of these had broken skulls and the number of intact skulls studied was much lower (see Specimens Examined; Tables). Three external measurements were taken, where available, from collector's original labels. Seven cranial measurements were made to the nearest 0.05 mm with dial calipers. These included: 1) condyloincisor length (CIL); occipital condyles to anterior tips of first incisors; 2) cranial breadth (CB), greatest breadth of braincase, approximately across the mastoids; 3) maxillary breadth (MB), greatest distance between tips of maxillary processes; 4) M2 to M2 breadth (M2-M2), greatest distance between anterior labial margins of second upper molars ; 5) interorbital breadth

(IOB), least lateral diameter of skull at anterior end of orbits, just posterior to maxil- lary processes; 6) palatoincisor length (PIL), posterior margin of palate in midline to anterior tips of first incisors; 7) upper tooth row length (UTRL), greatest length of tooth row from posterior margin of alveolus of M3 to anterior tip of I1. These meas- urements were selected for the speed and accuracy with which they could be made, and in some cases differ from those of earlier investigators. For example, condy- lobasal length is a traditional measure of skull length, but it is usually difficult and time-consuming to measure the anterior margin of the premaxillary bone. Employ- ing the anterior tips of the first incisors for the anterior terminator results in a more accurate and repeatable measurement. 1984年8月 Shrew-moles of China and Burma 73

Results The basis for placing all Chinese shrew-moles in a single species, Uropsilus soricipes, is the view that the different dental patterns, formerly the basis for recogniz- ing other genera and species, are in fact attributable to individual variation. There- fore, particular attention was paid to the occurrence of dental anomalies. The largest series of specimens were those from the Chen Lliang Shan (Qionlai Shan), western Sichuan, collected by the Sage Expedition (see SHELDON,1975; LAWRENCE,1982); from northern Burma by the Vernay-Cutting Expedition (see ANTHONY,1941); and from several adjacent localities in west-central Sichuan by F. T. SMITHfor the Field

Table 1. Individual variability in dentition among Uropsilus pop- ulations.

Table 2. Altitudinal distribution of Uropsilus soricipes and U. gracilis in the Qionlai Shan, Sichuan, China. 74 哺 乳 動 物 学 雑 誌 Vol. 10, No. 2 1984年8月 Shrew-moles of China and Burma 75 76 哺 乳 動 物 学 雑 誌 Vol. 10, No. 2

Museum (see OsGooD, 1937). In these series, a few dental anomalies were found, but overall, dentition was not highly variable (Table 1). Note that the three basic dental patterns formed the basis for identification of three taxa. If all specimens from Sichuan in Table 1 were to be regarded as belonging to a single species, then out of 386 individuals, 99 (25.6 %) would possess dental anomalies, and individual variation might indeed be considered frequent in U. soricipes.

Several considerations militate against this view. First, of the 99•ganomalies•h, all but five are bilateral in occurrence. This preponderance of bilateral consistency in presence or absence of teeth is not what would be expected if individual variation were the cause of the anomalies. Second, in the Quonlai Shan, 25-30 mi W of

Wenchuan, Sichuan, both U. soricipes and U. gracilis dental patterns are represented by large samples (Table 1). If only a single species with a high frequency of individual tooth variation was present here, one would expect that dental anomalies might be randomly distributed through the local populations of the Qionlai Shan. This is not the case; the U. gracilis pattern is more frequent at higher elevations, while the U. soricipes pattern is more frequent lower down (Table 2). Moreover, U. gracilis is a larger , with a longer tail, than is U. soricipes, at least in this area of geographic sympatry. While head-body lengths overlap considerably, there is little overlap in tail length, condyloincisor length of skull, palatoincisor length, or length of upper tooth row (Table 3). Based on these data, I regard U. soricipes and U. gracilis as distinct species. While U. soricipes has a very limited distribution in western Sichuan (Fig. 2), U. gracilis extends far to the south and west, to northern Burma and western Yunnan, and eastward to the type locality in southeastern Sichuan. Considerable variation in size is exhibited by U. gracilis throughout its range. The holotype, from the eastern- most edge of the species range, is also from a low elevation (4,000 ft.), and is a small individual (Table 3). The populations named atronates and nivatus are also small, while investigator from northwestern Yunnan and northern Burma are larger. In none of these populations, however, does the tail average as long as in U. gracilis from Sichuan (Table 3) (although comparisons of measurements taken by different collectors are suspect). The monotypic Uropsilus andersoni (since atronates and nivatus are properly assigned to U. gracilis) is known only from the type locality, Emei Shan, and that vicinity (Wa Shan, Liang-ho-kio, Ta-chiao), and from two other localities in western Sichuan, Lu-ting Shan and Luan-shih-gow. Uropsilus andersoni possesses a distinc- tive dental pattern and more robust mandible (Fig, lb), but also has a longer average tail length (93 % of head-body length) than soricipes (82-85 %), approaching the proportions of the long-tailed 97-98 %) Sichuan population of U. gracilis, though averaging smaller in size (Table 3). Where all three forms occur in central Sichuan, U, gracilis is the largest, with the longest tail, followed by U. andersoni, while the smallest, shortest tailed specimens are U. soricipes.

Discussion

Given that dental patterns are the chief characters used in distinguishing the 1984”N8ŒŽ Shrew-moles of China and Burma 77 taxa of Chinese shrew-males, it is necessary that the homologies of the various teeth be understood. Fortunately, ZIEGLER (1971) has examined in detail not only Uro- psilus, but most talpids. His evidence indicates that while•gNasillus-like•h U. gracilis and•gRhynchonax-like•hU. andersoni each possesses a total of 38 teeth, there are differences in their respective dentnl fnrmulae, as follows

U. gracilis I1, 12,•\, C, P1, P2, P3, P4, Ml, M2, M3•\

, i2,•\, c, p1, p2, p3, p4, ml, m2, m3

U. andersoni I1, 12,•\, C, P1, P2, P3, P4, M1, M2, M3•\

, i2, i3, c, p1, p2,•\, p4, ml, m2, m3 Thus, U. andersoni possesses an additional lower incisor pair (i3), but lacks a lower premolar pair (p3) compared to U. gracilis (Fig. lb, c). Uropsilus soricipes has a total of 34 teeth, and ZIEGLER(op. cit.) gives its dental formula as :

U. soricipes I1, 12,•\, C, P1, P2,•\, P4, Ml, M2, M3•\

, i2,•\, c, p1, p2,•\, p4, ml, m2, m3

This formula can be derived either by 1) loss of a lower incisor pair (i3) and upper premolar pair (P3) from the•gRhynchonax-like•hdentition of U. andersoni, or by 2) loss of both upper and lower premolar pairs (P3, p3) from the •gNasillus-like' dentition of U. gracilis (Fig. 1). OsGoon (1937) suggested that the •eRhynchonax- like•hdentition was •g... only a gradient between Nasillus and Uropsilus.•hFor this to be the case,•gNasillus•hwould first have to lose a pair of lower premolars (p3) and gain a pair of incisors (i3) to form•gRhynchonax•h, and then lose the incisor pair again, and lose an upper premolar pair (P3) as well, to form Uropsilus. This complex process of sequential gain and loss of i3, and non-synchronous loss of upper and lower pre- molars seems improbable, since 13, P3, and p3 are small teeth, and are hardly likely to be regained if once lost. This evidence that•gRhynchonax•his not intermediate between•gNasillus•hand •gUropsilus•hdentitions further strengthens my contention that the variable dentitions of these taxa are not due to individual variation. The small number of individual variants that do occur (Table 1) all involve the small i3,

P3, or p3 pairs, with one exception, in which a small supernumary incisor (I3) was found in a specimen of U. gracilis. This suggests the possibility that both•gNasillus•hand•gRhynchonax•hwere derived independently from a common ancestor which possessed a dental formula closer to the primitive eutherian formula of 3I/3i, 1C/lc, 4P/4p, 3M/3m exhibited by a number of talpid genera. For example, a common ancestor with the following dental formula may be postulated:

U. ancestor I1, 12,•\, C, P1, P2, P3, P4, Ml, M2, M3•\

, i2, i3, c, pl, p2, p3, p4, ml, m2, m3

ZIEGLER (1971) noted that i 1 is•g.. . reduced or variable in many [lines of Insectivoral•h, and this, plus the evidence of the supernumary 13 mentioned above, suggest that this postulated ancestor is plausible.•gNasillus•hcan then be derived by loss of the small i3 pair, whereas•gRhynchonax•hcan be derived by loss of p3.•gUropsilus" in turn can be derived from•gNasillus•hby loss of both upper and lower premolars, or from•gRhynchonax•hby loss of the i3 pair and the P3 pair. The latter seems more probable to me but in either case•gUropsilus•h represents the derived dental pattern. 78 哺 乳 動 物 学 雑 誌 Vol. 10, No. 2

Conclusions Evidence presented here supports the view that more than one species of Uropsilus exists, and that variation in dental pattern, which has been interpreted as individual variation, is instead due to species differences. Uropsilus gracilis has a large range in western China and adjacent northern Burma. While several subspecies have been described (atronates, investigator, nivatus) based mainly on size and color (parhaps) due to undescribed seasonal variation in pelage), populations in close geographic proximity vary considerably in these respects (i.e., investigator and nivatus in northwest Yunnan (Fig. 3 ; Table 3), and in the absence of any clear pattern of geographic varia- tion, I regard the species as monotypic. Uropsilus gracilis is geographically sympatric with a smaller, shorter-tailed species, U. soricipes, in central Sichuan. The two are both known from the Qionlai Shan west and southwest of Wenchuan, and also at Luan-shih-gow; U. soricipes is presently known only from a very restricted area. While the two preceding species are known from a failry large number of speci- mens, U. andersoni is represented by few, from several separate ranges in central Sichuan (Emei Shan, Lu-ting Shan). Given the paucity of specimens, its status is less certain than that of the other two species. Nonetheless, reasons have been pre- sented above for not regarding it as an individual variant (or "intermediate"). Uropsilus andersoni is the only shrew-mole known from the vicinity of Emei Shan, but in the Lu-ting Shan one U. andersoni was captured with five U. grad/is, and at Luan-shih-gow all three forms are represented.

Acknowledgements I thank those curators who assisted me in using collections in their care; these include S. ANDERSON,K. KOOPMAN,M. LAWRENCE,G. MUSSER(American Museum of Natural History); I. BISHOP,G. CORBET,J. HILL, J. INGLES,P. JENKINS(British Museum (Natural History)); R. IzoR, B. PATTERSON,R. TIMM (Field Museum of Natural History); J. KIRSCH, E. RUTZMOSER(Museum of Comparative Zoology, Harvard University); M. CARLETON,R. FISHER,A. GARDNER,L. CORDON,C. HANDLEY, R. THORINGTON,D. WILSON (National Museum of Natural History); C. SMART (Academy of Natural Science of Philadelphia). I also thank the University of Kansas for the sabbatical leave during which this research was carried out; JAN ELDERand COLETTASPENCER, who typed the manuscript; and Deb BENNETT,who prepared the figures. G. B. CORBET,K. KOOPMAN,M. A. LAWRENCE,and G. G. MUSSER,read early drafts and contributed valuable suggestions.

Specimens Examined Uropsilussoricipes (330). CHINA, Sichuan, Mupin (Baoxing),1 (USNM 256119,topotype); Weichou(Wenchuan), 1 (BMNH 11.9.8.12);Cheng Wei, (Cheng Gou) ChengouCreek, 25 mi W Wenchuan,7000 ft, 181 (AMNH 110511through -37, -41, -42, -46 through-50, -52 through -55, -57 through-78, -80, -81, -83 through-85, -87, -88, -90 through-95, -97 through-99, 110601,-03, -04, -06 through-09, -11 through-25, -28 through-62, -64, 110704through -16, -18 through-40, 1984年8月 Shrew-moles of China and Burma 79

-42 through -50 , -53, -59; Chengou (Cheng Gou) Forks, 30 mi W Wenchuan, 7600 ft, 23 (AMNH 110760, -61, -64 through -66, -69, -71, -75, -77, -82, -84, -86 through -89, -92 through -95, -97, -98 , 110806, -17, -39, 110943); Tsao Po, 15 mi SW Wenchuan, 5000 ft, 7 (AMNH 110694 through -700); S slope , Tsao Po Ridge, 20 mi SW Wenchuan, 8700 ft, 2 (AMNH 110702, -03); Chou Tsen

(Chouhsien) Goh, 6 (BMNH 1938. 4.1. 10; FMNH 45930, -31, -43 through -45); Dan (Dun) Shi Go, 40 (BMNH 1938.4.1.1.11; FMNH 37039 through -47, -49 through -61, -63 through -72, -74 through -78; USNM 260743 , -51; Gan Yang Go, 10 (FMNH 36141 through -50); Goan Shih (Kuan Shui?) Dwe, 11 (FMNH 45932 through -42); Heh (Hei?) Yen Wo, 1 (BMNH 55.443): Hsiao Yang Chi, 1

(FMNH 39620); Luan Shih Gow, 5 (FMNH 36158 through -59, -61 through -63); Lu Erh Cheh, 1 (FMNH 39619); Pin Yang Goh, 40 (BMNH 1938.4.1.8., -.9; FMNH 45905, -06, -08 through -29, -47 through -53 , -55 through -61). Uropsilus gracilis (173). CHINA, Sichuan, Chinfu (Jinfo) Shan, 4000 ft, near Nan-chwan

(Nancfiuan), 1 (BMNH 11.9.8.13, holotype); Cheng Wei, Chengou (Cheng Gou) Creek, 23 mi W Wenchuan, 7000 ft, 5 (AMNH 110539, -40, -45, -51, -56); Chengou (Cheng Gou) Forks, 30 mi W

Wenchuan, 7600 ft, 38 (AMNH 110762, -63, -67, -68, -70, -72, -76, -77, -80, -81, -83, -85, -90, -96, -99 through 110805 , -07 through -16, -30, -31, -33, -37, -38, -40 through 42); Chen Lliang Shan, 9000 ft, 30 mi W Wenchuan, 29 (AMNH 110665 through -93); Mao Mo Gou, 8600 ft, 30 mi

W Wenchuan, 12 (AMNH 110818 through -29); S slope, Tsao Po Ridge, 8700 ft., 20 mi SW Wenchuan, 1 (AMNH 110701); Luan Shih Gow, 1 (FMNH 36160); Lu-ting (Lulling, Jagsamka)

Shan, 5 (FMNH 36152 through -56). Yunnan, Kui-chiang-Salween Divide, 11-14,000 ft., 7

(BMNH 22.9.1.16, (holotype, investigator) - .13 through -.15, -.17 through -.19); Gomba La, Kui- kiang (chiang)-Salween Divide, 13-14,000 ft, 1 (BMNH 23.3.7.7); Mu-cheng, Salween drainage, 7000 ft, 14 (AMNH 44343 (holotype, atronates), 44335 through -37, -41, -42, -47, -49; MCZ 20702, -03; FMNH 35805 through -08; Pei-ti-ping , Mekong drainage, 9000 ft, 2 (AMNH 44368; MCZ 56853); Lichiang (Likiang, Lijiang) Range, Ssu-Shan (Xue Shan) Chang, 12,000 ft, 9 (AMNH 44352

(holotype, nivatus), -60, -65, -70, -71; FMNH 35089, 10; MCZ 20704, -05). BURMA, Kachin Prov., Adung Valley, 28•‹20'[N], 97•‹40'[E], 8000 ft, 1 (FMNH 40923); road to Chimeli Pass, 10,000 ft, 17

(AMNH 114582 through -96, 115548, -49); Hpare-Saulang road, 8200 ft, 11 (AMNH 114597 through -607); Imaw Bum , 9000 ft, 9 (AMNH 114563 through -71); Nyetmaw River, 8600 ft, 8 (AMNH 114572 through -77, -80, -81); above Nyetmaw River, 9200-9700 ft, 2 (AMNH 114578, -79).

Uropsilus andersoni (19). CHINA, Sichuan, Omi-san (Emei Shan), 9500 ft, 7 (BMNH 11.2.1.25

(holotype), -.26 through -.31); Liang Ho Kao (Kow) [near Emei Shan], 1 (BMNH 13.9.13.1); Luan Shin Gow, 2 (FMNH 36157, 36164); Lu-ting (Lulling, Jagsamka) Shan, 1 (FMNH 36151); Tachiao

(Ta-chi'iao), 1 USNM 175142); Wa Shan, 7 (ANSP 16594; MCZ 7520 through -24, -27).

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