(crocidura) of southern china

Journal of Mammalogy, 82(4):1059–1079, 2001

A REVISION OF THE WHITE-TOOTHED SHREWS (CROCIDURA) OF SOUTHERN CHINA XUE-LONG JIANG

AND

ROBERT S. HOFFMANN*

Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan 650223, People’s Republic of China (XLJ) Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (RSH)

Many systematic relationships among Chinese white-toothed shrews of genus Crocidura are presently unresolved. In this paper, a taxonomic revision of Crocidura from Southern China is presented. We studied 338 specimens from Burma, China, Korea, Pakistan, Turkey, Middle and Central Asia, and Russia (Appendix I), 285 of which had complete skulls that were analyzed with principal component and discriminant analyses. Results indicated that 6 species of Crocidura can be recognized in South China. C. fuliginosa occurs in Southwestern and Eastern China, C. attenuata is broadly distributed throughout Southern China, and C. horsfieldii is restricted to the southern part of China. C. vorax and C. rapax, usually placed as synonyms of the European C. russula, are recognized as 2 valid species whose ranges overlap in Southwestern China. C. shantungensis of Eastern Asia extends to the northern part of Southern China and is distinct from C. suaveolens and C. gmelini of Middle and Central Asia, respectively. Key words:

Crocidura, distribution, multivariate analysis, Southern China, taxonomic revision

The genus of white-toothed shrews, Crocidura, with 164 recognized species (Wolsan and Hutterer 1998), is the most speciose of all mammalian genera. White-toothed shrews occur through much of the Palearctic and Ethiopian and extend to the Oriental faunal regions. At least 15 species and subspecies of Crocidura have been described or recorded from China, spanning 2 faunal regions (Palearctic and Oriental), but most of them have been relegated to synonymy; e.g., only 4 species were listed by Ellerman and Morrison-Scott (1951), 5 species were recognized by Allen (1938) and, with different composition, by Jenkins (1976), 6 by Hutterer (1993), and 7–9 by Wolsan and Hutterer (1998) and Zhang et al. (1997). Compared with studies of Crocidura in neighboring countries, such as Malaya and Indonesia (Jenkins 1982; Ruedi 1995,

1996), Vietnam (Heaney and Timm 1983), Philippines (Heaney and Ruedi 1994), Korea (Jones and Johnson 1960), and Taiwan (Fang et al. 1997; Jameson and Jones 1977), there have been few taxonomic studies of Chinese Crocidura other than checklists or short comments (Corbet and Hill 1992; Ellerman and Morrison-Scott 1951; Hutterer 1993; Lazell 1998) since Allen (1938). The taxonomy and distribution of Chinese Crocidura is confusing and inconclusive. Foremost among questions are the status of small Crocidura in Eastern China. C. shantungensis (Miller 1901) was described but reduced to a subspecies of C. ilensis (Allen 1938) or C. suaveolens (Ellerman and Morrison-Scott 1951; Jenkins 1976). Hoffmann (1996) raised it provisionally to species rank again. C. vorax and C. rapax were first described by Allen (1923) as spe-

* Correspondent: [email protected] 1059

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cies, but were assigned to C. russula as 2 subspecies (Ellerman and Morrison-Scott 1951), or treated as synonyms of C. gueldenstaedtii (Corbet and Hill 1992) or C. pullata (Hutterer 1993), when C. russula was restricted to Western Europe and Northwestern Africa, based on chromosomal, biochemical, and morphologic analyses (Catzeflis et al. 1985). However, C. gueldenstaedtii is most certainly conspecific with C. suaveolens (Catzeflis et al. 1985; Hutterer 1993), leading to the view that the small C. suaveolens of Western Europe occurs across Eurasia all the way to Korea and Southeastern Siberia (Jenkins 1976; Zhang et al. 1997). Geographic ranges of most Chinese Crocidura are unclear because of this taxonomic confusion. Our purpose was to conduct a morphology-based taxonomic revision of Southern Chinese white-toothed shrews, especially C. rapax and C. vorax, and their synonyms, and then define distributions of the revised species. First, however, we discuss the status of C. shantungensis from Northern China, and its relatives because they extend into the northern part of South China. MATERIALS

AND

METHODS

South China is defined in this study as the Chinese Oriental Region. Because of intermingling of Palearctic and Oriental faunas in China, especially in the east, it is hard to draw an accurate line between them; the arbitrary boundary of Corbet and Hill (1992) at 358N has been followed. All materials came from museum collections. External measurements were recorded from specimen tags. Because these measurements are likely to show considerable interobserver variation, multivariate analyses relied only on measurements of cranial characters. All 15 cranial characters were measured by X.-L. Jiang with a digital caliper graduated to 0.01 mm: condyloincisive length (CIL), condyles to anterior tip of incisors; interorbital breadth (IOB), measured at anterior end of orbit; braincase breadth (BB), greatest breadth of braincase; braincase height (BH) from the highest point of the sagittal crest to the basisphenoid–basioccip-

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ital suture; rostral breadth (RB), greatest width of rostrum; rostral length (RL), anterior tip of incisors to anteriormost portion of orbit on right side; postrostral length (PRL), anteriormost portion of orbit on right side to posterior end of sagittal crest; palato-incisor length (PIL), measured from posterior edge of palate to anterior tip of incisors; postpalatal length (PPL), posterior edge of palate to anteriormost point on foramen magnum; upper toothrow length (UTR), greatest length of upper toothrow from posterior alveolus of M3 to anterior tip of incisors; breadth across M2–M2 (M–M), greatest width measured at anterior labial margins of 2nd upper molars; mandibular length (ML), measured from posteriormost point of condyle to anterior bony tip of mandible; lower toothrow length (LTR), greatest length of lower toothrow at alveolus of m3 to anterior tip of incisors; length of lower incisor (ilL), greatest length of right lower incisor; breadth of coronoid process (BCP), least breadth of coronoid process. Overall similarities of skulls were assessed first through principal component analyses (PCA) based on the 15 variables described earlier. Groups of individuals sharing a comparable morphology were then discriminated through a 2nd multivariate approach, namely, discriminant analysis (DA). To clarify the status of C. shantungensis, our 1st objective, specimens of C. suaveolens from Turkey, Iran, Afghanistan, and Pakistan, C. coreae from North and South Korea and Southeastern Russia, C. shantungensis and C. phaeopus from Northern and Eastern China, and C. gmelini from Iran, Kazakhstan, and Western China, were included in the multivariate analyses. Our 2nd objective was to determine the status of C. rapax, C. vorax, and other specimens from Southern China referable to C. fuliginosa, C. attenuata, C. grisescens, C. grisea, C. horsfieldii, and C. phaeopus, and C. pullata from Pakistan, and specimens allocated to C. shantungensis in the 1st analyses, which were included in a 2nd set of multivariate analyses. Both PCA and DA programs were run using SYSTAT for WINDOWS (1998) version 8.0 programs separately. The different groups identified were then named according to current taxonomic descriptions of holotypes, or decisions were made from a comparison with the literature in cases where type specimens were not intact or not examined. Species ranges were then based

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JIANG AND HOFFMANN—WHITE-TOOTHED SHREWS

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TABLE 1.—Means (6SD) and ranges of external and cranial measurements (mm) of Crocidura shantungensis, C. gmelini and C. suaveolens from Turkey, Iran, Afghanistan, Kazakhstan, Pakistan, China, Korea, and southeastern Russia. Variablea HB TL CIL IOB BB BH RL PRL RB PIL PPL UTR M2–M2 ML LTR ilL BCP a

C. shantungensis, n 5 33

C. gmelini, n 5 14

C. suaveolens, n 5 70

60.9 6 5.00 51.0–71.0 38.1 6 3.73 28–45 16.82 6 0.39 16.16–17.52 3.86 6 0.15 3.48–4.09 7.64 6 0.27 6.98–8.11 3.95 6 0.19 3.51–4.23 5.40 6 0.18 5.06–5.79 11.20 6 0.39 10.43–12.04 5.11 6 0.20 4.51–5.42 7.48 6 0.15 7.06–7.79 7.75 6 0.34 7.12–8.37 6.90 6 0.13 6.65–7.14 4.85 6 0.24 4.07–5.18 8.68 6 0.20 8.34–9.18 6.66 6 0.11 6.44–6.86 2.92 6 0.17 2.41–3.21 0.92 6 0.06 0.81–1.10

59.8 6 4.37 52.0–68.0 31.8 6 3.47 25.0–38.0 17.77 6 0.36 17.09–18.29 4.20 6 0.13 3.96–4.42 8.08 6 0.25 7.40–8.38 4.13 6 0.18 3.79–4.50 5.85 6 0.22 5.35–6.19 11.53 6 0.23 10.98–11.89 5.51 6 0.18 5.19–5.81 8.18 6 0.18 7.84–8.48 7.92 6 0.23 7.44–8.30 7.62 6 0.18 7.30–7.93 5.27 6 0.33 4.12–5.60 9.32 6 0.24 8.72–9.68 7.26 6 0.17 6.96–7.65 3.25 6 0.14 2.93–3.41 1.03 6 0.08 0.86–1.21

71.0 6 5.80 53.0–84.0 43.2 6 4.46 31.0–51.0 18.82 6 0.58 17.32–20.68 4.39 6 0.15 4.04–4.69 8.68 6 0.19 8.32–9.11 4.52 6 0.21 4.07–5.07 6.41 6 0.26 5.58–6.98 12.17 6 0.38 11.33–13.14 5.76 6 0.18 5.36–6.20 8.62 6 0.32 7.64–9.53 8.37 6 0.29 7.69–8.99 7.94 6 0.30 7.23–8.64 5.53 6 0.18 5.09–6.05 9.84 6 0.36 9.08–10.63 7.65 6 0.29 6.96–8.26 3.28 6 0.34 2.14–3.77 1.00 6 0.07 0.87–1.19

Acronyms defined in ‘‘Materials and Methods’’ section.

on those specimens examined, supplemented by the literature records.

RESULTS Status of Middle and East Asian whitetoothed shrews.—The PCA of C. suaveolens, C. gmelini, and C. shantungensis was based on 117 complete skulls; 17 from China together with 100 others (15 from Korea, 3 from the former USSR, 20 from Iran, 22 from Turkey, 15 from Afghanistan, 15 from Pakistan, and 10 from Kazakhstan) were in-

cluded in a PCA to assess the overall similarity. External and cranial measurements and sample sizes are given in Table 1. The 1st principal component (83% of variance) was a size factor; most variables were strongly correlated with that axis: e.g., CIL, PIL, ML, UTR, LTR, RL, and PPL. The 2nd (4.8%) was strongly correlated with CH, IOB, and i1L. The 3rd (3.3%) was correlated with BCP. The result (not shown) demonstrated that C. coreae and C. phaeopus greatly overlapped with C. shantungen-

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FIG. 1.—Plot of Crocidura shantungensis (s), C. gmelini (g), and C. suaveolens (u) on principal components factors 1 and 2.

FIG. 2.—Plot of Crocidura shantungensis (s), C. gmelini (g), and C. suaveolens (u) on discriminant canonical axes 1 and 2.

sis, as concluded by Hoffmann (1996), so they were combined with C. shantungensis for further analysis. The East Asian sample represented a distinct taxon, C. shantungensis (Fig. 1), which was easily distinguished from Middle and Central Asian samples of C. gmelini and C. suaveolens on the 1st principal component. A DA was designed to differentiate the 3 groups. Four variables (CIL, PIL, IOB, and BB) were used to achieve that discrimination (Fig. 2). This result showed that East Asian shrews formed a distinct group, as did C. gmelini; all specimens were classified correctly (including the neotype of C. gmelini) except for 3 (4%) specimens of C. suaveolens which were classified as C. gmelini. Although the holotype of C. shantungensis was damaged, available measurements were comparable to other East Asian shrews, leading us to assign small Crocidura from East Asia to C. shantungensis. That species was distinguished from C. gmelini, and especially C. suaveolens, by its smaller size, shorter palate (PIL), and shorter toothrows (UTR, LTR) (Table 1). The eastern end of the range of C. suaveolens was also clarified. In addition to its occurrence in Kazakhstan and Iran (Hoffmann 1996; Lay 1967), as C. russula, C. suaveolens also oc-

curs in Northeastern Pakistan and Northwestern Afghanistan, and is probably sympatric with C. gmelini there, as it is in Iran (Hoffmann 1996). Status of Southern Chinese white-toothed shrews.—All 156 intact skulls of Southern Chinese Crocidura, as well as 20 skulls of C. pullata from Pakistan and 8 skulls of C. horsfieldii from Burma, were included in the PCA. External and cranial measurements and sample sizes are given in Table 2. Results indicate that most of the variation (93%) was explained by the 1st principal component, the size factor. The 2nd (2.1%) was strongly correlated with BCP and i1L. The last component (1.5%) was most correlated with IOB. Most of the species included in the PCA fell into 2 groups readily defined by size (Figs. 3A and 3B). C. fuliginosa formed a group of large specimens well separated from all other species and had values .2 on the 1st factor, suggesting a high level of distinctiveness (not shown in subsequent figures). The 2nd group consisted of all other specimens, including holotypes of C. grisea, C. grisescens, C. pullata, C. vorax, and C. rapax. In that group, the larger C. attenuata–C. pullata subgroup was easily distinguished from the smaller C. rapax, C. shantungensis, and C. hors-

Variablea HB TL

CIL IOB BB BH RL PRL RB PIL PPL UTR M2–M2 ML LTR i1L

a

C. pullata, n 5 20

C. vorax, n 5 12

C. rapax, n 5 10

C. horsfieldii, n 5 13

C. shantungensis, n 5 16

86.3 6 6.7 72.0–100.0 74.1 6 5.4 62.0–89.0 16.5 6 1.1 15.0–19.0 23.26 6 0.64 22.04–25.00 5.18 6 0.24 4.64–5.71 10.33 6 0.35 9.40–11.19 5.47 6 0.22 4.42–5.95 8.33 6 0.34 7.61–9.14 14.61 6 0.39 13.81–15.46 7.39 6 0.29 6.84–8.11 10.76 6 0.38 9.91–11.91 10.44 6 0.36 9.30–11.18 10.06 6 0.35 9.21–10.85 6.95 6 0.24 6.23–7.46 12.61 6 0.41 11.74–13.45 9.51 6 0.28 8.74–10.33 4.21 6 0.28 3.09–4.75 1.37 6 0.10 1.11–1.58

76.0 6 6.60 60.0–89.0 50.5 6 5.29 43.0–60.0 13.4 6 1.12 11.0–16.0 20.13 6 0.70 18.51–21.49 4.68 6 0.21 4.20–5.01 9.02 6 0.32 8.32–9.82 4.87 6 0.20 4.17–5.23 6.87 6 0.32 6.36–7.99 12.96 6 0.48 11.95–14.04 6.43 6 0.23 5.92–6.89 9.21 6 0.31 8.63–9.77 9.14 6 0.37 8.51–9.95 8.50 6 0.28 8.07–9.10 6.09 6 0.23 5.58–6.55 10.77 6 0.44 9.68–11.55 8.11 6 0.28 7.36–8.70 3.52 6 0.19 2.95–3.86 1.16 6 0.12 0.82–1.44

83.2 6 4.00 73.0–89.0 44.2 6 4.38 39.0–53.0 15.06 6 0.66 14.0–16.0 20.26 6 0.46 19.45–21.21 4.76 6 0.13 4.51–5.03 9.27 6 0.20 8.92–9.63 4.63 6 0.18 4.37–4.99 7.01 6 0.21 6.62–7.43 12.90 6 0.28 12.50–13.44 6.25 6 0.17 5.90–6.57 9.30 6 0.27 8.93–9.93 9.03 6 0.25 8.51–9.57 8.61 6 0.22 8.22–9.00 6.03 6 0.14 5.75–6.31 10.59 6 0.26 10.12–11.09 8.94 6 0.22 7.77–8.70 3.73 6 0.25 3.08–4.13 1.19 6 0.06 1.07–0.32

67.3 6 9.77 54.0–90 46.7 6 3.34 41.0–51.0 12.3 6 0.78 11.0–14.0 19.09 6 0.77 17.74–20.11 4.02 6 0.20 3.70–4.36 8.40 6 0.48 7.57–9.06 4.75 6 0.22 4.37–5.03 6.30 6 0.49 5.59–7.08 12.43 6 0.45 11.65–13.26 5.85 6 0.26 5.37–6.13 8.38 6 0.35 7.84–8.94 8.88 6 0.39 8.37–9.56 7.77 6 0.39 7.16–8.23 5.53 6 0.22 5.10–5.78 9.60 6 0.37 8.77–10.12 7.37 6 0.32 6.79–7.86 3.28 6 0.28 2.83–3.88 1.14 6 0.11 1.01–1.30

64.1 6 4.51 56.0–70.0 42.4 6 2.80 38.0–47.0 11.9 6 0.60 11.0–13.0 18.14 6 0.27 17.45–18.33 4.06 6 0.13 3.78–4.21 8.32 6 0.35 7.64–8.78 4.51 6 0.19 4.12–4.73 6.20 6 0.30 5.75–6.58 11.76 6 0.37 11.25–12.23 5.72 6 0.13 5.48–5.86 8.13 6 0.18 7.80–8.36 8.55 6 0.34 7.88–9.03 7.57 6 0.30 7.11–8.07 5.45 6 0.14 5.25–5.71 9.58 6 0.39 8.70–10.13 7.25 6 0.27 6.87–7.79 3.04 6 0.22 2.60–3.34 1.08 6 0.09 0.95–1.23

61.1 6 6.18 49.0–71.0 40.8 6 5.97 30.0–48.0 11.0 6 1.04 10.0–13.0 17.18 6 0.40 16.14–17.82 4.06 6 0.16 3.81–4.40 8.17 6 0.25 7.69–8.61 4.39 6 0.21 3.88–4.75 5.77 6 0.26 5.09–6.13 11.38 6 0.28 10.84–11.78 5.34 6 0.21 5.02–5.67 7.54 6 0.29 7.01–7.92 8.04 6 0.18 7.77–8.40 7.07 6 0.33 6.46–7.72 5.10 6 0.20 4.76–5.43 9.18 6 0.26 8.78–9.85 6.83 6 0.18 6.58–7.26 2.79 6 0.20 2.46–3.23 0.96 6 0.12 0.83–1.16

58.7 6 3.57 51.0–64.0 36.0 6 4.63 28.0–45.0 11.2 6 0.79 10.0–12.5 16.63 6 0.28 16.16–17.22 3.79 6 0.16 3.48–4.09 7.49 6 0.17 7.14–7.77 3.94 6 0.20 3.67–4.35 5.34 6 0.17 5.06–5.67 10.99 6 0.30 10.43–11.59 5.07 6 0.23 4.51–5.47 7.44 6 0.16 7.13–7.83 7.57 6 0.33 7.12–8.37 6.91 6 0.14 6.65–7.20 4.86 6 0.21 4.39–5.34 8.63 6 0.27 8.00–9.24 6.66 6 0.11 6.49–6.86 2.94 6 0.17 2.70–3.25 0.91 6 0.06 0.81–1.03

Acronym defined in ‘‘Materials and Methods’’ section.

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BCP

C. attenuata, n 5 40


HF

C. fuliginosa, n 5 73

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TABLE 2.—Means (6SD) and ranges of external and cranial measurements (mm) of South Chinese Crocidura and Pakistan C. pullata.

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FIG. 3.—Plots of Southern Chinese Crocidura and Pakistan C. pullata (p) on principal components, A) factors 1 and 2, and B) factors 1 and 3: C. fuliginosus (f), C. attenuata (a), C. vorax (v), C. rapax (r), C. horsfieldii (h), and C. shantungensis (s).

fieldii, but C. vorax overlapped the 2 subgroups on each side. Neither of the other two principal components separated them easily, although C. shantungensis was slightly separated from the others (Figs. 3A and 3B). A DA was then designed to distinguish groups using 5 variables (CIL, PIL, ML, IOB, and BB). The Chinese taxa were all easily differentiated, but 5 specimens labeled C. attenuata were misclassified as C. pullata from Western Asia (see below). Four groups can be recognized readily (Fig. 4A): C. attenuata–C. pullata, with all neg-

FIG. 4.—Plots of Southern Chinese Crocidura and Pakistan C. pullata (p) on discriminant canonical axes, A) 1 and 2, B) 1 and 3, and C) 1 and 4: C. attenuata (a), C. vorax (v), C. rapax (r), C. horsfieldii (h), and C. shantungensis (s).

ative values on the 1st axis; C. shantungensis–C. horsfieldii, with higher positive values on the 1st axis; C. vorax with lower positive values on the 1st axis but higher

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positive values on the 2nd axis; and C. rapax with lower positive values on the 1st axis and lower values on the 2nd axis. The combined Chinese group may be differentiated with the help of the 3rd and 4th discriminant functions. C. shantungensis had higher scores on the 3rd axis and lower scores on the 4th than C. horsfieldii (Figs. 4B and 4C). Chinese C. attenuata and Pakistan C. pullata, however, are not readily differentiated even with the 3rd and 4th discriminant functions because 12% of C. attenuata and 20% of C. pullata specimens are misclassified with each other. Their taxonomic status is discussed below. In conclusion, the Southern Chinese Crocidura can be resolved as follows: a very large shrew, C. fuliginosa; a somewhat smaller shrew, C. attenuata; 2 intermediate-sized species, C. rapax and C. vorax; and 2 small shrews, C. shantungensis and C. horsfieldii. TAXONOMIC ACCOUNTS

AND

DISCUSSION

Crocidura fuliginosa (Blyth 1856) Sorex fuliginosa Blyth 1856, Journal of the Asiatic Society of Bengal, 24:362, Schwegyin, Pegu, Burma. Crocidura dracula Thomas 1912, Annals and Magazine of Natural History, 9:686. Mongtze (5Mengzi), Southern Yunnan, China; Allen 1938:125; Ellerman and Morrison-Scott 1951:84; Zhang et al. 1997:14. Crocidura praedax Thomas 1923, Annals and Magazine of Natural History, 11: 656. Likiang (5Lijiang) Valley, Yunnan, China. Crocidura fuliginosa Jenkins 1976, Bulletin of the British Museum (Natural History) Zoology, 25:282; Corbet and Hill 1992: 39; Hutterer 1993:85. This is the largest white-toothed shrew in China and clearly distinct from all other species. CIL . 22.0 mm (Fig. 5A). Tail proportionately (.80% of head–body length) and absolutely ($62 mm) longer. Only a few vibrissae-like hairs on basal part

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of tail. Pelage brownish-gray dorsally and slightly more grayish ventrally. Before Jenkins (1976), populations from China were referred to as C. dracula (Allen 1938; Ellerman and Morrison-Scott 1951), which Jenkins assigned to C. fuliginosa as a subspecies and was followed by others (Corbet and Hill 1992; Hutterer 1993). In the current literature, Chinese C. fuliginosa includes C. dracula, C. praedax, and C. grisescens. As early as 1938, Allen placed C. praedax as a synonym of C. dracula, citing no distinguishable characters. We also found no differences between C. praedax and other Yunnan representatives based on 10 specimens from Lijiang (type locality of C. praedax) and 63 from other parts of Yunnan. The assignment of C. grisescens, however, was problematic. A year after Howell (1928) first described the taxon as a species, he reduced it to a subspecies of C. dracula (Howell 1929). Others followed, including Ellerman and Morrison-Scott (1951), who suspected that this assignment was invalid because of the small size of the subspecies. We compared the holotype of C. grisescens with our other specimens, and it shared no similarities with large Chinese C. fuliginosa, but instead was affiliated with the smaller C. attenuata in PCA (Fig. 3), and was classified with C. attenuata in DA (Fig. 4). On the other hand, a large shrew was recorded in Zhejiang, the province adjacent to Fujian where the holotype of C. grisescens was obtained. Although those specimens were identified as C. dracula (5fuliginosa) grisescens (Zhuge 1993), their size was even greater than that of specimens of C. fuliginosa from Southwestern China (Yunnan, Guizhou, and Guangxi), and their tails were shorter. CIL and UTR in the Zhejiang population were 25.3 mm (n 5 13, range 24.2–26.6) and 11.5 mm (n 5 13, range 11.0–12.1), respectively. Average tail length was 58.7 mm in males and 53 mm in females, (range 45–65 mm), and relative tail length was only about 58% of head– body length (Zhuge 1993:75). In contrast,

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FIG. 5.—Skulls of 6 species of Chinese Crocidura: A) C. fuliginosa (NMNH 260769, CIL 5 23.0 mm); B) C. attenuata (NMNH 279336, CIL 5 20.8 mm); C) C. vorax (NMNH 239534, CIL 5 18.01 mm, occiput broken); D) C. rapax (NMNH 277598, CIL 5 17.31 mm, occiput broken); E) C. horsfieldii (NMNH 357862, CIL 5 17.6 mm); and F) C. shantungensis (NMNH 298933, CIL 5 16.4 mm). Top row, left lateral skull and mandible; middle row, dorsal skull; bottom row, ventral skull.

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FIG. 6.—Distribution of Crocidura fuliginosa and C. gmelini in China.

in the southwestern population of C. fuliginosa we studied, CIL and UTR were only 23.3 mm (22.0–25.0) and 10.1 mm (9.2– 10.8), respectively; tail length was 74.1 mm (62–89), and average relative tail length was 86% of head–body length (Table 2). The Zhejiang population (Fig. 6, far right on map) may be affiliated with C. fuliginosa, but it probably represents an undescribed taxon; unfortunately, no specimens were available to us. Distribution.—On the basis of specimens examined and records in the literature, C. fuliginosa occurs in Southwestern China from Northwestern Yunnan (Lijiang) and adjacent Xizang (Medog, Zayu, not mapped; Zhang et al. 1997) southward to the China–Burma border, in Southwestern Guizhou (not mapped) and Western Guangxi (Fig. 6), and possibly in Zhejiang,

Eastern China. The distribution seems to be disjunct from west (Guangxi) to east (Fujian, Zhejiang; Fig. 6). The published record from Fujian was based on the type of C. grisescens, which we consider a synonym of C. attenuata (see later), and there is only 1 other published record of C. fuliginosa in Fujian, from Wuyishan (Zhang et al. 1997; not mapped). Crocidura attenuata Milne-Edwards 1872 Crocidura attenuata Milne-Edwards 1872, Recherches pour servir a l’Histoire Naturelle des Mammiferes, p. 263. Moupin, Szechuan (5Baoxing, Sichuan), China; Jenkins 1976:297; Corbet and Hill 1992: 43; Hutterer 1993:82; Zhang et al. 1997: 15.

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Crocidura grisea Howell 1926, Proceedings of the Biological Society of Washington, 39:137. Yenpingfu, Fujian. Crocidura grisescens Howell 1928, Journal of Mammalogy, 9:60. Kuatun, Fujian, China. Crocidura tanakae Kuroda 1938, A list of the Japanese mammals. Shohosha (5She-hou-hsia) Horigai, Taichusiu, Taiwan (258049N, 1218369E). This is a smaller shrew (CIL 18.5–21.5 mm; Fig. 5B) with proportionately (67% of head and body length) and absolutely (#60 mm) shorter tail compared with C. fuliginosa. Pelage grayish-brown dorsally and slightly more grayish ventrally, with summer pelage much darker. Tail sparsely haired or naked, with only a few vibrissae on the basal part. Traditionally, C. attenuata in continental Southern China includes only 1 synonym, C. grisea, which was first described as a species by Howell (1926) but then relegated to subspecific status in C. attenuata (Howell 1929). Allen (1938) went further, regarding it as a synonym of C. attenuata; we have now added C. grisescens (see above). Although C. attenuata is widely distributed in Southern China, our results do not support the recognition of subspecies for either eastern or western populations, based on external or cranial characters, except for the insular C. tanakae in Taiwan. Crocidura pullata specimens from Pakistan were also compared with Chinese shrews. The PCA and DA results showed similarity between C. pullata and C. attenuata (Figs. 3 and 4); 20% of C. pullata and 12% of C. attenuata were misclassified with each other. However, the 2 taxa are not conspecific; all specimens that we examined exhibited consistent differences in external characters. C. pullata from Pakistan have a shorter, densely haired tail (53% of head and body length), with bristle hairs present on the proximal four-fifths of the tail, and more brownish dorsal pelage. Specimens of Chinese C. attenuata have a

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longer, sparsely haired or naked tail (67% of head and body length), with fewer bristle hairs scattered on the basal part, and dorsal pelage is more grayish, tipped with silver. Distribution.—Crocidura attenuata is widely distributed in Southern China, including the islands of Hainan and Taiwan (Zhang et al. 1997). The northwestern limit of the range of this species was reported in Lanzhou, Gansu (Zheng and Li 1991; Fig. 7). It is thus widely separated from the morphologically similar C. pullata, whose easternmost occurrence is in Northeastern Pakistan (Fig. 8). Crocidura vorax G. Allen 1923 Crocidura vorax G. Allen 1923, American Museum Novitates, 100:8. Li-kiang (5Lijiang) Valley, Yunnan, China. Crocidura russula vorax Ellerman and Morrison-Scott 1951, Checklist of Palaearctic and Indian Mammals, 1758– 1946, p. 81; Zhang et al. 1997:13. Crocidura gueldenstaedtii Corbet and Hill 1992, The Mammals of the Indomalayan Region, p. 45; as a synonym. Crocidura pullata Hutterer 1993, Mammal Species of the World, p. 94; as a synonym. This is a medium-sized Chinese whitetoothed shrew (CIL 17.7–20.1 mm; Fig. 5C) with relative tail length similar to C. attenuata (69%). Pelage is pale grayishbrown with a narrow gray band below brownish tips of the dorsal guard hairs. Tail bicolor, covered with short hairs, except for bristles on basal two-thirds of the tail. Crocidura vorax has been one of the most problematic of Chinese shrews. It was originally described by Allen (1923), based on 3 specimens caught in the Lijiang valley, Yunnan, China. Later, he decided it was related to C. russula of Europe, and perhaps should have been regarded as a subspecies, but he hesitated to make a definite assignment (Allen 1938). Ellerman and MorrisonScott (1951) subsequently listed vorax as a subspecies of C. russula, based only on

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FIG. 7.—Distribution of Crocidura attenuata and C. dsinezumi in China.

head–body and tail length, but they did not describe any other characters uniting them. However, after Catzeflis et al. (1985) restricted C. russula to Western Europe and Northwestern North Africa, Corbet and Hill (1992) assigned vorax to C. gueldenstaedtii, which they believed to be distributed from Europe to Asia, including also pullata. Hutterer (1993), however, thought C. gueldenstaedtii was most certainly conspecific with suaveolens, and referred to the larger, darker, montane shrew of Afghanistan and Pakistan as C. pullata, whose distribution he extended to Yunnan, China, to include C. vorax. Although C. vorax was not readily distinguished from others in our comparisons of C. pullata with Chinese Crocidura (Figs. 3A and 3B), all specimens of C. vorax were classified correctly (100%) and dif-

ferentiated from C. pullata and other Chinese white-toothed shrews in DA on the basis of a high score on the 2nd axis (.2; Fig. 4A). It is distinguished from C. fuliginosa and C. attenuata by its smaller size, wellhaired tail, and more brownish pelage, and from C. pullata by its smaller size and relatively long tail. Distribution.—Crocidura vorax is known to occur in Yunnan and Hunan (Fig. 9). It is reported (as C. russula) from Guizhou and Guangxi and Southern Sichuan, and elsewhere by Zhang et al. 1997; not mapped, but this must be confirmed. Crocidura rapax G. Allen 1923 Crocidura rapax G. Allen 1923, American Museum Novitates, 100:9. Yinpankai, Mekong River, Yunnan, China.

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FIG. 8.—Distribution of Crocidura gmelini, C. pullata, and C. suaveolens in Middle and Central Asia.

Crocidura russula rapax Ellerman and Morrison-Scott, 1951, Checklist of Palaearctic and Indian Mammals, 1758– 1946, p. 81; Zhang et al. 1997:13. Crocidura horsfieldii kurodai Jameson and Jones 1977, Proceedings of the Biological Society of Washington, 90:461. Linkou (5Ling-kou), Taipei Hsien (228469N, 1208269E). Comparison of holotype of C. kurodai (USNM 358115) with specimens of C. rapax, apparently not done by Jameson and Jones, clearly place C. kurodai closer to C. rapax morphologically than it is to C. horsfieldii. Crocidura gueldenstaedtii Corbet and Hill 1992, The Mammals of the Indomalayan Region, p. 45; as a synonym. Crocidura pullata Hutterer 1993, Mammal Species of the World, p. 94; as a synonym. Like C. vorax, this is another mediumsized Chinese shrew (CIL 17.5–18.3 mm; Fig. 5D) with relative tail length 66%. Dorsal pelage is similar to C. vorax, although

more brownish. Tail is less well haired and indistinctly bicolor, the bristle hairs scattered on two-thirds of the tail. Allen (1923) originally described both C. rapax and C. vorax in the same paper, and their taxonomic destinies followed much the same course (see above). Results of PCA and DA showed that C. rapax was not only different from other well-defined species of Chinese Crocidura and Pakistan C. pullata, but also different from C. vorax (Figs. 3 and 4A); all specimens were correctly classified. C. rapax differs from C. fuliginosa and C. attenuata in smaller size and more brownish pelage. It differs from C. vorax in lacking a pronounced sagittal crest and in its indistinctly bicolor tail. The average CIL is shorter (Table 2; Fig. 5D). Distribution.—Crocidura rapax is distributed in Yunnan, Western and Southern Sichuan, Taiwan (see above), and reported (as C. russula) in Guizhou and on Hainan Island (not mapped; Zhang et al.1997; Fig. 10). A specimen from Burma was also identified as C. rapax (see Appendix I).

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FIG. 9.—Distribution of Crocidura vorax and C. shantungensis in China.

Crocidura horsfieldii Tomes (1856) Sorex horsfieldii Tomes 1856, Annals and Magazine of Natural History, 17:23. Sri Lanka. Crocidura horsfieldii indochinensis Robinson and Kloss 1922, Annals and Magazine of Natural History, 9:88. South of Annam, Viet Nam; Jenkins 1976:285; Zhang et al. 1997:12. Crocidura horsfieldii wuchihensis Shaw et al. 1966, Acta Zootaxonomica Sinica, p. 261. Mount Wuchih, Hainan Island, China; Jenkins 1976:285; Zhang et al. 1997: 12. This is a small white-toothed shrew distributed in Southern China (mean CIL 17.2 mm; Fig. 5E) Pelage brownish-gray dorsally and more grayish ventrally. Tail (relative

length, 67%), thinly covered or naked, with few bristles on basal part. Because of the small sample available from China, we added 8 specimens from Northeastern Burma (Appendix I) to the statistical analysis. The DA results showed that the few specimens from Yunnan, Guizhou, Southern Sichuan, and Fujian were grouped with Burmese specimens and were distinguished correctly from all other Chinese Crocidura (Fig. 4). The species is easily distinguished from large or mediumsized Crocidura by its small size, grayish color, and almost naked tail. Jenkins’ report of C. suaveolens in Yunnan was probably based on C. horsfieldii because of the similar size and proportions of their skulls (Jenkins 1976). Distribution.—According to Zhang et al.

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FIG. 10.—Distribution of Crocidura rapax and C. sibirica in China.

(1997), C. horsfieldii occurs in Yunnan, Guizhou, and Hainan and Taiwan islands; we examined specimens from Guangxi, Fujian, and Yunnan (Fig. 11). It was also reported from Shaanxi (Baihe), but this is far north of its main range and needs confirmation. Crocidura shantungensis Miller 1901 Crocidura shantungensis Miller 1901, Proceedings of the Biological Society of Washington, 14:158. Chimeh, Shantung (5Shandong), China; Hoffmann 1996: 162. Crocidura coreae Thomas 1907, Proceedings of the Zoological Society of London, 1906:860. Mingyong, Korea. Crocidura ilensis phaeopus G. Allen 1923, American Museum Novitates, 100:7.

Wanhsien, Szechuan (5Wanxian, Sichuan), China. Crocidura ilensis shantungensis G. Allen 1938, Mammals of China and Mongolia, p. 131. Crocidura suaveolens shantungensis Ellerman and Morrison-Scott, 1951, Checklist of Palaearctic and Indian Mammals, 1758–1946, p. 77; Jenkins 1976:288; Zhang et al. 1997:12. Crocidura russula hosletti Jameson and Jones 1977, Proceedings of the Biological Society of Washington, 90:465. 10 miles W Taichung, Taichung Hsien, Taiwan (248159N, 1208439E). Comparison of holotype of C. hosletti (USNM 398640) with other Chinese shrews reveal it to be closest morphologically to C. shantungensis. This conclusion was also reached

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FIG. 11.—Distribution of Crocidura horsfieldii and C. lasiura in China.

by Fang et al. (1997), although they used the name C. suaveolens. This is the smallest white-toothed shrew (CIL 16.2–17.2 mm; Fig. 5F) in Eurasia, and it extends its distribution into the northern part of Southern China (Fig. 9). Dorsal pelage grayish-brown with a velvety gray sheen. Tail is densely covered with hairs, and bristle hairs occur throughout the length of the tail. Since Miller (1901) first described this species, it has been surrounded by taxonomic confusion. Allen (1938) thought it was the eastern representative of the common small reddish-brown shrew found across Central Asia and reduced it to a subspecies of C. ilensis, which was subsequently allocated to C. suaveolens as a subspecies (Ellerman and Morrison-Scott 1951). However, after a detailed review of

C. suaveolens and C. gmelini, Hoffmann (1996) provisionally raised the small East Asian shrews, C. shantungensis, C. coreae, and C. phaeopus, to species rank as C. shantungensis based on pelage darker than C. gmelini, noting that it occupied more mesic habitats than the latter. Specimens of C. shantungensis were widely separated from the Middle Asian shrews, C. gmelini and C. suaveolens, in PCA analysis (Fig. 1). DA analysis also showed clear differentiation among the 3 taxa (Fig. 2). C. shantungensis is also easily differentiated from other Southern Chinese shrews (Figs. 3 and 4) by its small size (Table 2), darker pelage, and short densely haired tail. Specimens referred to C. ilensis phaeopus from Wanxian, Sichuan, were not differentiated from C. shantungensis from East China based on PCA and DA. How-

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ever, specimens of phaeopus are distinguished from the nominate form by their hind-foot color (light in C. shantungensis, dark in C. phaeopus), so we provisionally retained them as subspecies. Distribution.—The range of C. shantungensis includes the Korean peninsula, Southeastern Siberia, and Eastern China. It also extended westward to Shaanxi, Qinghai, and Northern Sichuan, and south to Jiangxi and Zhejiang (Fig. 9), and Taiwan Island (Fang et al. 1997). Zhang et al. (1997) also reported the species from Guizhou and Guangxi, but this is far south of its described range and needs confirmation. The results of PCA and DA also show that the 2 specimens from Qinghai (USNM 576004, -5) and 3 from Shanxi (USNM 172539, -40, -41), referred to C. gmelini by Hoffmann (1996), are assigned to C. shantungensis in the present study (Fig. 8). DISCUSSION Although 6 Crocidura species can be recognized in Southern China, they are unevenly distributed. In the southwest, especially in Yunnan and Western Guizhou, 5 species occur (all except C. shantungensis), while there are only 2–3 in the north and east. The widespread C. attenuata is distributed throughout the whole of Southern and much of Central China and is sympatric with several other species with partial distributions in Southern China. The largest shrew, C. fuliginosa, widely distributed in Southeastern Asia, reaches its northern limit in Southern China. The smallest whitetoothed shrew, C. shantungensis, extends from Southeastern Siberia to Northern South China, where its range overlaps with C. fuliginosa and C. attenuata in Zhejiang and C. attenuata in Anhui and Sichuan. Conversely, C. shantungensis is separated geographically on the mainland from the medium-sized shrews, C. vorax and C. rapax, whose ranges overlap in Yunnan, Guizhou, and Sichuan, and another small southern shrew, C. horsfieldii, whose northern distributional limit is in Southern China.

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Sympatry in Crocidura is common throughout much of China. In Yunnan, 4 of the 5 species have been caught at Jingdong (C. horsfieldii, C. vorax, C. attenuata, C. fuliginosa). Elsewhere in Yunnan, at Lijiang, C. rapax, C. vorax, and C. fuliginosa occur together. At other localities in the southern provinces, we recorded various combinations of species pairs; the most common pairing was C. attenuata–C. fuliginosa (7), while others included C. shantungensis–C. attenuata (2), C. rapax–C. vorax (2), C. horsfieldii–C. fuliginosa (2), and single instances of C. horsfieldii–C. rapax, C. vorax–C. attenuata, C. vorax–C. fuliginosa, and C. horsfieldii–C. attenuata. A combined analysis of our records and those extracted from Zhang et al. (1997) revealed species records from 249 localities, 24 (9.6%) of which indicated sympatry. However, sympatry is not evenly spread; there is a cline of increasing frequency from north to south. The northeastern range limit of C. attenuata occurs at about 308309N, and there are 9 localities of sympatry south of this line between C. shantungensis and C. attenuata, but no other combinations involving shantungensis. Northward, however, C. shantungensis overlaps with the large C. lasiura, a very distinctive species (Figs. 9 and 11). We did not study this taxon, but Zhang et al. (1997) recorded it at 17 localities from Heilongjiang southward to Jiangsu, and C. shantungensis from another 22 different localities within their joint range. The 2 species were sympatric at only 2 (5%) of those 39 collection localities: Mishan (Heilongjiang) and Antu (Jilin). These 2 provinces are the northernmost in China (.408N) and support only these 2 species of Crocidura. Temperate Central China from Beijing south to about Shanghai (;308N) has an intermediate number of species (2–4), and has an intermediate level of sympatry (12.5%). The greatest species diversity of Crocidura (and other shrews—Hoffmann 1986, 1987) is found in subtropical and tropical Southern China, as well as the

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highest level of sympatry (18%). Taiwan Island, although fairly far south (;23–248N), has high relief, and supports a mammalian fauna fairly evenly divided between Holarctic and Oriental species (Hoffmann 2001). Below 1,100 m, 1–2 species of Crocidura are found (C. attenuata, C. shantungensis), while above 1,200 m, only one species occurs, either C. attenuata or C. rapax. Thus, the small Crocidura fauna in Taiwan mirrors the larger mainland fauna, with 1 widespread species (C. attenuata), 1 northern species (C. shantungensis), and 1 southern montane species (C. rapax). The Japanese C. dsinezumi has also been reported from Taiwan (Fig. 7), but its status is uncertain (Hutterer 1993). The overall pattern, both on the mainland and on the continental shelf island of Taiwan, is probably correlated with several north–south gradients: insolation and growing season, primary productivity, greater structural diversity of habitats, and greater temporal stability. However, details of the ways in which sympatric congeneric shrews share or compete for resources in sympatry need to be studied. KEY 1.

2.

3.

TO

SOUTHERN CHINESE CROCIDURA

Large; CIL . 22.0 mm; tail long ($62 mm), hind foot $ 15 mm, pelage brownish-gray, tail dark brown above and slightly paler below . . . C. fuliginosa Smaller; CIL , 21.5 mm; tail shorter (#60 mm), hind foot # 16 mm . . . . 2 Pelage darker grayish-brown, especially in summer; base and tip of dorsal hairs not, or only indistinctly bicolored; tail thinly covered with short hairs or naked, few bristles scattered on basal part of tail . . . . . . . . . . . . . . 3 Pelage lighter, pale grayish brown; base and tip of dorsal hairs distinctly bicolored; tail well covered with hairs, bristles scattered on more than one half of tail . . . . . . . . . . . . . . . . . . . . . 4 Smaller; CIL , 17.8 mm, RB , 5.7 mm, UTR , 7.7 mm; average tail length 40.8 mm . . . . . . . . C. horsfieldii Larger; CIL . 18.5 mm, RB . 5.9 mm,

4.

5.

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UTR . 8.0 mm; average tail length 51.0 mm . . . . . . . . . . . . . . C. attenuata Smaller; CIL 16.2–17.5 mm, PIL , 7.8 mm; tail short, bicolored, densely covered with hair, bristles on most of tail; velvety gray sheen on dorsal surface . . . . . . . . . . . . . . C. shantungensis Larger; CIL . 17.5–20.1 mm, PIL . 7.8 mm; tail longer, less densely haired, more or less bicolored, long bristles only on basal two-thirds of tail . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Larger; CIL 17.7–20.1 mm; sagittal crest distinct; dorsal pelage paler, grayish; tail bicolored . . . . . . . C. vorax Smaller; CIL 17.5–18.3 mm; sagittal crest indistinct; dorsal pelage darker, brownish; tail indistinctly bicolored . . . . . . . . . . . . . . . . . . . . . . . . . C. rapax

ACKNOWLEDGMENTS This work was completed at National Museum of Natural History, Smithsonian Institution, Washington, D.C., where X.-L. Jiang studied on a fellowship from Chinese Academy of Sciences. We are indebted to Y.-X. Wang, Kunming Institute of Zoology, Chinese Academy of Sciences, Z.-J. Feng, Institute of Zoology, Chinese Academy of Sciences, R. S. Voss and G. G. Musser, American Museum of Natural History, L. R. Heaney, The Field Museum of Natural History, M. Rutzmoser, Museum of Comparative Zoology, Harvard University, and P. F. Whitehead, Peabody Museum of Natural History, Yale University, for access to or loan of specimens. Distribution maps were produced by Y. Xei, Institute of Zoology, Academia Sinica, Beijing, and by GIS Laboratory, World Wildlife Fund, Washington, D.C., courtesy of J. Morrison and J. Lamoreax. X.-L. Jiang’s special thanks are due to S. A. Hoffmann for her kind help when he was in Washington, D.C. The work was funded by Key Project of the Chinese Academy of Sciences, Special Support for Youth of the Chinese Academy of Sciences, and National Science Foundation of China (39730080).

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of Gansu (X. T. Wang, ed.). Gansu Science and Technology Publishing House, Lanzhou, China. ZHUGE, Y. 1993. Crocidura (Insectivora: Soricidae). Pp. 19–23 in Fauna of Zhejiang: Mammalia (Y. Zhuge and H.-Q. Gu, eds.). Zhejiang Science and Technology Publishing House, Hangzhou, China. Submitted 28 April 2000. Accepted 13 February 2001. Associate Editor was Brett R. Riddle.

APPENDIX I Specimens examined.—Specimens were examined from the following museums: American Museum of Natural History, New York (AMNH), Institute of Zoology, Beijing, China (BIZ), The Field Museum of Natural History, Chicago, Illinois (FMNH), Institute of Zoology, Kunming, China (KIZ), Peabody Museum of Natural History, Yale University, New Haven, Connecticut (PMNH), and National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Crocidura suaveolens Turkey.—Icel (5Mersin) Prov., Cilician Toros, 8 km S Namrun (5Camliyayla), 378039N, 348469E, 3 (USNM 327230, -32, -34); Icel District, Tarsus, 368529N, 348529E, 1 (USNM 327227). Kayseri Prov., 15 km NE Keyseri, 388439N, 358309E, 2 (USNM 3327235, -36). Istanbul Prov., Yalova, Taskopru, 408419N, 298239E, 1 (USNM 327238); Istanbul, Derbent Army T.B. Sanitarium, 418029N, 288579E, 3 (USNM 327243, -44, -45); Kemerburgaz, Istanbul, 418099N, 288549E, 1 (USNM 327247). Izmir Prov., S Izmir, Yalikahve, 388259N, 278109E, 3 (USNM 327240, -41, -42). Sinop Prov., Bektasaga, 418569N, 358019E, 4 (USNM 327222, -23, -24, -25). Amasya Prov., Amasya, 408379N, 358509E, 1 (USNM 327218); Tasova, Borabey Lake, 408469N, 368209E, 2 (USNM 327219, -20). Iran.—Kerman Prov., 42 km W Kermanshahan, 308179N, 578059E, 5 (USNM 354501, -02, -07, -08, -11). E Azarbaijan Prov., 5 km SE Meyaneh, 378269N, 478429E, 5 (USNM 354503, -05, -14, -17, -19); 11 km N Reziyeh, 378339N, 458049E, 1 (USNM 354509). Teheran Prov., 11 km ENE Fasham, 358579N, 508349E, 1 (USNM 369535). Qazin Prov., Karaj, 358489N, 508599E, 1 (AMNH 171173). Luristan Prov., Khurramabad Dist., Bakhitiari, 338309N, 488209E, 3 (AMNH 171174, -76, -205). Fars Prov., 25 mi

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NE Yasuj, 308409N, 518369E, 3 (FMNH 110990, -91, -95). Gorgan Prov., Dar Kaleh, 368549N, 548529E, 1 (AMNH 88735). Pakistan.—Hazara Dist., Dungagali, 348039N, 738229E, 1 (USNM 3266281); Lalazar (5Lulusar), 358059N, 738569E, 2 (USNM 326285, -86). Northwest Frontier Prov., Naran, 348539N, 738399E, 11 (USNM 326287, -88, -89, -90, -91, -94, -95, -96, -300, -302, -303); Saif-ul-Maluk Lake, 348529N, 738419E, 1 (USNM 326299). USSR (former).—Georgia, Abkhazia, ;438N, 408459E, 1 (AMNH 245352). Afghanistan.—Badakhshan Prov., 19.3 km E Ishkashim (5Eshkashem), 368429N, 718349E, 3 (FMNH 102101, -02, -04); S Iskashim, 32 km SW Zebak, 368349N, 718229E, 11 (FMNH 102105, -06, -08, -09, -10, -11, -12, -13, -15, -16, -17); 25 km W Faizabad, 378069N, 708349E, 1 (FMNH 102120). Crocidura shantungensis North Korea.—3 mi SW Yanggu, 388069N, 1288009E, 1 (USNM 298934); Unson-ni, 388039N, 1278129E, 2 (USNM 299474, -77); Taegwang-ni, 388119N, 1278069E, 1 (USNM 298933); Chipo-ri, 388089N, 1278199E, 2 (USNM 299478, -79). South Korea.—½ mi N Tongjin, 378429N, 1268339E, 2 (USNM 298938, -39); 6 mi E Seoul, 378329N, 1278009E, 1 (USNM 299476); Seoul, 1 (USNM 299596); 5 mi ENE Pusan, 358089N, 1298049E, 1 (USNM 299046); Pupyang-ni, 378449N, 1278129E, 1 (USNM 299045); near Pupyang-ni, Central National Forest, 2 (USNM 299045, 300640); 2 mi SW Sochon-ni, 378599N, 1278049E, 1 (USNM 298937); 5 mi W Kwangju, 358099N, 1268559E, 1 (USNM 299047); Yang Ju Kung, SE Seoul, 378459N, 1278069E, 1 (USNM 283674). Russia.—Primor’sk, Pogranichnii Krai, Nesterovka, 448289N, 1318469E, 2 (E. Shvarts 606, 847). China.—Shandong Prov., Chimeh, 378429N, 1208129E, 1 (USNM 86151, holotype); Weifang (5Weixian), 368429N, 1198129E, 1 (AMNH 38368). Shanxi Prov., 5 mi S Taiyuan, 378499N, 1128349E, 3 (USNM 172539, -40, -42). Shaanxi Prov., Qingling, 338549N, 1078549E, 1 (AMNH 56086); Feng Xian, 348309N, 1078309E, 2 (FMNH 18935; AMNH 32298). Zhejiang Prov., Tunglu, 298509N, 1198359E, 3 (USNM 268760; MCZ 24320, -24). Jiangsu Prov., Huai Yuan, 328549N, 1178069E, 1 (USNM 256688); Sichuan,

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Wanxian, Yen-Ching-Kao, 308379N, 1088179E, 9 (AMNH 56010, -11, -14, -21, -24, -26, -38; MCZ 20713, -14). Hubei Prov., I-Chang, 308439N, 1118229E, 1 (MCZ 7230). Qinghai Prov., Haixi, N Wutomeiren, 368599N, 938119E, 2 (USNM 576004, -05). Crocidura gmelini China.—Xinjiang Prov., Shache (5Yarkand), 388279N, 778169E, 1 (AMNH 30551). Kazakhstan.—150–160 mi N Kzyl Orda, Tasbulak, 468139N, 668579E, 5 (AMNH 85312, -13, -14, -16, -17); 75 mi NE Kyzl Orda, 508129N, 658119E, 2 (AMNH 85318, -19). Taldy-Kurgan Prov., Panfilov, 448109N, 808019E, 1 (AMNH 30549). Iran.—Gorgan Prov., Dar Kaleh, 368549N, 548529E, 2 (AMNH 88734, -36). Khorasan Prov., between Turbat-i-Haidari and Sangan, 1 km S Ayyubi, 358129N, 598169E, 1 (USNM 327916). Isfahan Prov., Isfahan, 328409N, 518389E, 1 (AMNH 110989). Khorassan Prov., 6 km E Bujnurd, 378289N, 578199E, 1 (USNM 340947); Dasht, 378199N, 568019E, 3 (AMNH 88745, neotype, -46, 88906). Crocidura pullata India/Pakistan.—Kashmir, Kotihar Dist., Kotihar Valley, 338509N, 738309E, 5 (USNM 173213, holotype, -16, -17, -19, -21); Nowboog Valley, 338409N, 738309E, 2 (USNM 173224, -25); Khistwar Dist., Dachin, 338309N, 758459E, 3 (USNM 173909, -12, -13); Lolab Valley, 348309N, 748309E, 5 (USNM 175905, -06, -10, -15, -16); Dandwar, 338309N, 758109E, 4 (USNM 201134, -35, -37, -38); Ladak, Sobu, 348099N, ;778E, 1 (USNM 198451). Crocidura fuliginosa China.—Yunnan Prov., Li-kiang (5Li-jiang, 5Li-chiang), 268469N, 1008009E, 9 (USNM 260769; AMNH 44418, -19, -26, -29, -32, -33, -34, -37); Ho-Mu-Shu (Pass), ;258009N, 988499E, 1 (AMNH 44465); Chung Pa (Mekong R.), ;268449N, 998499E, 1 (AMNH 44449); Namting River (Burmese border), 238369N, 998269E, 8 (AMNH 44478, -79, -80, -81, -83, -85, -90, -93); Yang-pi (5Shangjie) River, Yangpi, 258409N, 1008009E, 8 (AMNH 44461, -62; BIZ 17476, -77, -78, -79, -80, -81); Yung-Chang Fu, 258099N, 998049E, 1 (AMNH 44452); Jingdong, 248289N, 1008549E, 23 (KIZ 640373, -480, -495; 98458, -59, -61, -62, -66, -67, -68,

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-74, -75, -78, -79, -80, -89, -90, -99; 96355; 97511; 98500, -08; 98655); Liuku, 258529N, 988529E, 2 (KIZ 74385, -91); Bijiang (5Zhiziluo), 268339N, 988569E, 5 (KIZ 780246, -83, -99; 780350, -51); Jianchuan, 268289N, 998529E, 2 (KIZ 650364, -91); Jizushan, Binchuan, 258589N, 1008219E, 13 (KIZ 810627, -28, -29, -30, -57, -60, -61, -99; 810700, -01, -02, -03, -04); Yuanmu, 248519N, 1018549E, 1 (KIZ 690022); Kunming, 258049N, 1028419E, 8 (KIZ 631135; 640207; 310308, -33; 73396; 75612; 840045; 85022); Menglian (5Menglien), 228199N, 998239E, 1 (KIZ 55); Ximeng, 228459N, 998299E, 1 (KIZ 0016); Menglun, 218569N, 1018159E, 1 (KIZ 78567); Jiangcheng, 248269N, 1028489E, 1 (KIZ 72180); Luchun (5Pingshan), 258359N, 1028289 E, 1 (KIZ 72058). Guizhou Prov., Xinyi, 258009N, 1048009E, 1 (KIZ 631093). Guangxi Prov., Dibao (5Debao, Tepao), 238209N, 1068379E, 1 (BIZ 21425). Crocidura attenuata China.—Fujian Prov., Kuatun, 278399N, 1178229E, 2 (USNM 252187, holotype of grisescens; KIZ 113); Chongan, 278429N, 1188009E, 1 (AMNH 84801); Yanping (5Yengping), 268369N, 1188069E, 2 (USNM 238229, holotype of grisea; BIZ 0037); Fuqing, 258429N, 1198249E, 1 (AMNH 44631). Guangdong Prov., Nan Ao island, 238269N, 1178039E, 4 (PMNH 5091, -92, -93, -94). Hongkong, Lantau, 228159N, 1138569E, 1 (PMNH 5090). Hainan Prov., Dongfang, 198049N, 1088519E, 1 (BIZ 70650); Nodda (5Nodoa), 198299N, 1098269E, 1 (AMNH 597780). Hunan Prov., Yochow (5Yueyang), 298239N, 1138069E, 1 (USNM 239533); Changsha, 288129N, 1128589E, 1 (BIZ 21522). Guizhou Prov., Kweiyang (5Guiyang), 268349N, 1068409E, 6 (USNM 279334, -35, -36, -37, -38; 282624); Jiangkou, 278419N, 1088449E, 1 (KIZ 650006). Sichuan Prov., Tacho, 298129N, 1038189E, 1 (USNM 260746); Wenchuan, 318249N, 1038369E, 3 (USNM 259367, -69, -70); Kuan Shien (5Guanxian), 308598N, 1038399E, 1 (USNM 258176); Baoxing, 308189N, 1038489E, 2 (BIZ 19690, -91); Omei, 298359N, 1038119E, 1 (KIZ 17404); Kangding 308009N, 1018549E, 2 (KIZ 820290, -344); Wanxian, 308379N, 1088179E, 9 (AMNH 56006, -16, -27, -28, -40, -61; 57192; 58301, -02). Yunnan Prov., Namting River (near Burmese border), 238369N, 998269E, 1 (AMNH 44489); Jingdong, 248289N, 1008549E,

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1 (KIZ 808); Pingbian (Miaozu Zizhixian), 228549N, 1038409E, 1 (KIZ 84101); Ninglang, 278129N, 1008599E, 1 (KIZ 810754). Crocidura vorax China.—Yunnan Prov., Li-jiang, Ssu Shan, 268469N, 1008009E, 1 (AMNH 44383, holotype); Yinjiang (5Pingyuan), 248439N, 978569E, 1 (KIZ 24023); Jingdong, 248289N, 1008549E, 2 (KIZ 640067, -476); Kunming, 258049N, 1028419E, 6 (KIZ 310335, -56, -57; 630095, -97; 631134); Mile (5Milo, Miyang), 248249N, 1038279E, 1 (KIZ 84297). Sichuan Prov., Luzhou, 288539N, 1058239E, 2 (KIZ 147, -48). Hunan Prov., Yochow (5Yueyang), 298239N, 1138069E, 3 (USNM 239530, -31, -34). Crocidura rapax China.—Yunnan Prov., Ying-Pan-Kai (5Yingpan), 268279N, 998099E, 1 (AMNH 44321, holotype); Deqin (5Deqen), 288489N, 988489E, 1 (BIZ 79743); Li-jiang, 268469N, 1008009E, 2 (AMNH 44381, -84); Luxi (5Mangshi), 248279N, 988369E, or (5Zhongshu), 248319N,

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1038469E, 2 (BIZ 24025, -27); Mile (5Milo, Miyang), 248249N, 1038279E, 1 (KIZ 84298). Sichuan Prov., Xichang, 278539N, 1028189E, 2 (BIZ 17392, -93); Miyi (5Miyi-kai), 268509N, 1028049E, 1 (BIZ 17391); Luzhou, 288539N, 1058239E, 1 (KIZ 149). Burma.—12 mi N Myitkyina (W side of Irawaddy R.), 258249N, 978259E, 1 (USNM 277598). Crocidura horsfieldii China.—Yunnan Prov., Ho-Mu-Shu (Pass), ;258N, 988499E, 1 (AMNH 44356); Deqin (5Deqen), 288489N, 988489E, 4 (KIZ 79741, -42; BIZ 17419, 24098); Bijiang, 268339N, 988569E, 1 (KIZ 780235); Yongde (5Dedang), 248049N, 998189E, 1 (BIZ 24098); Jingdong, 248289N, 1008549E, 5 (KIZ 64148, -79, -433, -461, -462). Fujian Prov., Chongan, 278429N, 1188009E, 1 (AMNH 84797). Burma.—Rawngaw, 268159N, 988209E, 1 (AMNH 114724); Hpare, 258509N, 988259E, 1 (AMNH 114749); Gangfang, 268009N, 988389E, 5 (AMNH 114659, -711, -12, -13, -14); Tang Tung, 268049N, 988359E, 1 (AMNH 114724).