Mammal Study 32: 155–168 (2007) © the Mammalogical Society of Japan

A survey of small from Huu Lien Nature Reserve, Lang Son Province, Vietnam

Darrin P. Lunde1,*, Nguyen Truong Son2 and Guy G. Musser1 1 Department of Mammalogy, American Museum of Natural History, Central Park West @ 79th Street, New York, NY 10024 2 Department of Vertebrate Zoology, Institute of Ecology and Biological Resources, Vietnamese Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam

Abstract. A survey of the small mammals of Huu Lien Nature Reserve, Lang Son Province, Vietnam (21°40'52''N, 106°20'28''E) resulted in the capture of 187 specimens representing 21 species of bat, insectivore, and : Suncus etruscus, Rhinolophus pearsonii, Rhinolophus pusillus, Rhinolophus stheno microglobosus, Aselliscus stoliczkanus, Hipposideros armiger, Hipposideros larvatus, Harpiocephalus harpia, Pipistrellus paterculus, Scotomanes ornatus, Callosciurus erythraeus, Dremomys rufigenis, fulvescens, Niviventer langbianis, Tonkinomys daovantieni, Rattus andamanensis, Rattus losea, Rattus tanezumi, Mus caroli, Rhizomys pruinosus, and Atherurus macrourus. In addition Ratufa bicolor and Tamiops sp. are reported from observations. The recently described genus and species Tonkinomys daovantieni is currently known only from material collected during the present survey and is apparently endemic to karst habitats in northeastern Vietnam.

Key words: biodiversity, Huu Lien, mammals, Tonkinomys, Vietnam.

Across the northeastern part of Vietnam extends a stun- resentative of the entire Viet Bac karst zone. The first ning landscape of forested limestone towers of the Viet efforts towards a comprehensive survey of the Huu Lien Bac karst zone (Tuyet 1998). Confined to Vietnam east fauna were begun by the staff of the Institute of of the Red River, and extending into southern China, it is Ecology and Biological Resources (Can et al. 2000), the largest of at least five distinct limestone regions in which reported fifty-seven species of mostly large ani- Vietnam (Fig. 1). Despite its close proximity to Hanoi; mals based on interviews of local hunters. Additional the entire Viet Bac karst zone remains very poorly sur- surveys of bats were conducted in 1998 and 2000 veyed for small mammals. Delacour and Lowe passed (Hendrichsen et al. 2001; Son et al. 2004) but a de- through Bac Can and Lang Son Provinces in the winter tailed survey of and shrews has been long out- of 1926–1927, but their expedition, focused as it was on standing. avian surveys, neglected the smaller terrestrial mammals In 2004 we surveyed small mammals from the tower so that little was added to our knowledge of these groups karst forests of Huu Lien Nature Reserve as part of an (Thomas 1928). It was only in the last years of the twen- effort to further document the fauna of this region. Our tieth century that comprehensive mammal surveys were specific goals were to: 1) fill a void in our understanding begun at the Huu Lien Nature Reserve. of the rodent and shrew fauna; 2) supplement the previ- Established in 1986 for the purpose of conserving ous surveys of bats; 3) specifically sample karst habitats limestone biodiversity and to protect the Musk deer, for endemic taxa; and, 4) establish an additional geo- Moschus berezovskii, the Huu Lien Nature Reserve is graphic reference point in a continuing effort by Lunde just 90 kilometers northeast of Hanoi and is accessible by towards eventually understanding the biogeography of a one-day car trip and a second day of hiking into the small mammals across all of northern Indochina. mountains. The reserve is an ideal site for implementing The location of our survey site is: Vietnam: Lang Son a mammal survey that could be regarded as broadly rep- Province; Huu Lung District; Huu Lien Commune; Huu

*To whom correspondence should be addressed. E-mail: [email protected]. 156 Mammal Study 32 (2007)

Fig. 2. Tower karst landscape typical of the Huu Lien Nature Reserve. Tropical moist deciduous forest covers the steep slopes, but the valley bottom has been cleared for agriculture. Photo by D. P. Lunde, April 2004.

eroded limestone towers. The topography of the forest habitat was marked by sheer cliffs and limestone rubble (Figs. 2 & 4). There was abundant evidence of forest degradation as the local Dao villagers depend heavily on forest resources. Imperata grass (300–400) — A dense stand of tall Imperata grass. Bamboo forest — A thick stand of tall, hollow- stemmed bamboo and scattered trees. Continuing down the far side of the massif that was the focus of our study site we discovered an entirely deforested hillside of Fig. 1. Map of Vietnam showing five limestone regions recognized Imperata grass and swidden agriculture. by Tuyet (1998). The location of our study site in the Huu Lien Nature Reserve is indicated at 21°40'52''N, 106°20'28''E. Materials and methods

Lien Nature Reserve; Lan Dat Village, 21°40'52''N, Fieldwork was continued for 20 consecutive days be- 106°20'28''E (Fig. 1). Trapping efforts were concen- ginning on April 6, 2004. Victor snap-traps, Sherman trated in several distinct habitats leading from our camp live traps, Tomahawk cage traps and Conibear traps were in a broad flat valley at 150 m elevation up to the top of used to trap small mammals (see Voss and Emmons an eroded karst tower at approximately 450 m elevation (1996) for descriptions of traps). Trapping efforts were (Fig. 2). Habitats sampled and their elevation range (in aimed at maximizing the number of species sampled, and parenthesis) are as follows: so in addition to setting traps on the ground, we tied Rice field (150 m) — A large expanse of fallow rice Victor and Conibear traps to woody vines to sample ar- fields occupying the relatively flat lowlands near the vil- boreal species. We also placed traps on elevated lime- lage (Fig. 2). At the time of our survey the fields were stone ledges and lowered them into caverns and cavities used for grazing small herds of buffalo. beneath limestone rubble. The type of trap employed Scrub (150–200 m) — A 30–70 m wide swath of and the length of time that a trap was allowed to remain scrub (grasses, shrubs, low secondary tree growth) defin- in one place depended on the experience of the trapper. ing the border between rice paddies in the valley and No attempts were made to standardize trapping effort evergreen forest on the karst slopes above (Fig. 3). but we did record the numbers, and kinds of traps set in Tropical moist deciduous forest (200–300) — moist different habitat types and these data are summarized in deciduous forest growing on the steep slopes of the Table 1. Traps were baited with a mixture of peanut Lunde et al., Small mammals in Vietnam 157

Fig. 3. A pitfall trapline in the dense scrub at the base of a karst Fig. 4. Tropical moist deciduous forest covering the steep slopes of tower. The only specimen captured in this line was a single Suncus a karst tower. Musser is seated near the entrance to a small cave-like etruscus, although several specimens of Niviventer fulvescens, Rattus structure where we captured specimens of Tonkinomys daovantieni, andamanensis, and Mus caroli were captured here using conventional Niviventer fulvescens, and Rattus andamanensis. Photo by D. P. traps. Photo by D. P. Lunde, April 2004. Lunde, April 2004. butter, oatmeal, raisins and bacon mixed in a 6 : 2 : 2 : 1 Liver samples were preserved in lysis buffer prior to ratio. In addition to conventional traps we also employed fixation. Skulls were extracted and cleaned from most pitfall-trap lines after a design depicted in Voss et al. specimens. Limits of standard external body measure- (2001); except our lines differed in the numbers of pit- ments (in mm) taken in the field are abbreviated: TL falls per line with one in secondary scrub consisting of (total length); T (tail length); HF (hind foot length); E 7 buckets along 40 meters of continuous drift fence (ear length); and FA (forearm length, bats only). Weight (Fig. 3), and a second in bamboo forest consisting of 19 was recorded in grams. Cranial measurements (in mm) buckets along 100 meters of continuous drift fence. Bats follow Bates and Harrison (1997) for bats, and Musser formed a secondary component of our survey and were (1979) for murid rodents. Specimens were cataloged in collected by providing mist nets to survey helpers from the American Museum of Natural History (AMNH) and Lan Dat village. will be eventually split with the Department of Zoology Specimens were fixed in formalin solution (3.75% at the Institute for Ecology and Biological Resources formaldehyde) and are now preserved in 70% ethanol. (IEBR). This paper is the sixth in a series on the small

Table 1. Summary of trapping effort based on numbers of trap nights (a trap night is defined as one trap set for one night) per trap type and habitat type

Victor Traps Sherman Traps Tomahawk Traps Conibear Traps Total trap nights Proportion of effort Scrub/Rice field 716 349 34 0 1099 26% Forest 2065 352 205 124 2746 65% Bamboo 200 0 47 0 247 6% Imerata grass 0 110 0 0 110 3% Total trap nights 2981 811 286 124 4202 100% Proportion of effort 71% 19% 7% 3% 100% – When expressed as a percentage of total trap nights it is clear that most of our sampling efforts were directed toward the forest habitat using Victor traps. Overall trap success is calculated at 2.9% based on 120 captures in 4202 trap-nights total. 158 Mammal Study 32 (2007) mammal fauna of Vietnam by Lunde and colleagues 4.4, anterior width of upper M2 1.3, length of M3 0.4. (Lunde and Son 2001; Lunde and Musser 2002; Lunde et al. 2003a, b; Lunde et al. 2004; Musser et al. 2006). Order Chiroptera Family Rhinolophidae Results Rhinolophus pearsonii Horsfield, 1851 We report 21 species of bat, insectivore and rodent This species was first reported from the Huu Lien from Huu Lien Nature Reserve based on 187 voucher Nature Reserve by Hendrichsen et al. (2001) and to their specimens. A complete listing of specimens collected, single specimen we add eighteen more. Most of our along with catalog numbers, is provided in the Appendix. specimens were netted in secondary forest just outside The following accounts provide descriptions of species the village but two were captured deep within a lime- and relevant taxonomic notes along with natural history stone cave and one was brought in by a villager without information for each species collected. habitat data. R. pearsonii is a medium-sized Rhinolo- phus (Tables 2 & 3) with long brown wooly fur. The Order Soricomorpha noseleaf is characterized by a broad horseshoe that com- Family Soricidae pletely covers the muzzle. There is a deep notch at the anterior end of the horseshoe, and an absence of lateral Suncus etruscus (Savi, 1822) lappets. The sella is broad at the base but then narrows Among the smallest of terrestrial mammals, Suncus abruptly and is roughly parallel sided through the mid- etruscus was only recently reported from Vietnam dle, ending in a rounded tip. The connecting process is (Feiler and Nadler 1997), and to these we add two more. broadly rounded, joining the sella at its apex such that it Our first specimen was captured in a pitfall on April 9 in does not form a notch. The posterior margins of the bamboo forest at the top of a limestone massif at 450 m horseshoe join the base of the lancet, which is slightly elevation, while our second was taken four days later in a convex sided and with a tall and pointed tip. The nose- pitfall in scrub at 100 m elevation. Both specimens are leaf of R. pearsonii is similar to that of R. luctus, but R. similar in external appearance. The pelage is silvery luctus is much larger and has basal lappets lateral to the brown over both the upperparts and underparts of the horseshoe. head and body; the tail is brown along its dorsal surface, There are two species with which R. pearsonii is likely paler below, and with long pilose hairs extending along to be confused: R. affinis and R. yunanensis. Rhinolo- the entire length of the tail. Feiler and Nadler (1997) phus affinis is usually smaller in body size (Hendrichsen reported an astonishing black upper lip in one of their et al. 2001) with shorter fur that is not wooly; the sella is specimens, but we found no trace of such pigmentation narrow and concave in the middle and the connecting in our two specimens although this character was noted process joins the sella below its tip thus forming a dis- in Crocidura kegoensis (Lunde et al. 2004), another tinct notch. small-bodied shrew from Vietnam. Suncus etruscus is Rhinolophus yunanensis is a larger species that is distinguished by the combination of its very small size reported to be morphologically indistinguishable from R. and the presence of four unicuspids in each cranial tooth- pearsonii (Hill 1986; Bates and Harrison 1997; Csorba et row (the genus Crocidura has only three unicuspids in al. 2003). Measurements of our series of R. pearsonii the cranial toothrow). External measurements of our two from Huu Lien (Tables 2 & 3) average greater than those specimens are as follows: TL 74–84; Head and body reported for the species and just overlap R. yunanensis. length 44–53 (obtained by subtracting tail length from Hill (1986) reviewed R. pearsonii and R. yunanensis in total length); T 30–31; HF 8–9; E 5. Weight: 2 grams. detail and recognized the two as separate species differ- We extracted and measured the skull from the smaller of ing only in size; however, in light of the fact that our Huu the two specimens collected (275574); cranio-dental Lien specimens seem to bridge these two size categories, measurements: condylo-incisive length 14.1, condylo- the limits between these two species warrants further basal length 13.4, upper toothrow length 5.8, incisor to investigation. 3rd unicuspid length 2.8, upper M2-M2 breadth 3.7, least interorbital breadth 3.1, braincase breadth 6.2, braincase Rhinolophus pusillus Temminck, 1834 height 3.0, length of braincase 5.9, postglenoid breadth A single specimen was collected from Huu Lien; an Lunde et al., Small mammals in Vietnam 159

Table 2. Summary statistics (mean ± one standard deviation, observed range, number of specimens) for external measurements, and weights of nine species of bat collected at Huu Lien Nature Reserve

Species FA HB E T HF TIB Weight Rhinolophus pearsonii 54 ± 1.4 63 ± 3.7 27 ± 1.7 24 ± 1.4 13 ± 0.6 28 ± 0.7 16 ± 1.7 52–57 54–68 24–29 21–26 12–14 26–29 13–20 18 18 18 18 18 18 16 Rhinolophus pusillus 38 37 29 21 7 15 5 Rhinolophus stheno 44 42 17 23 9 20 8 Aselliscus stoliczkanus 42 ± 1.2 44 ± 1.7 9 ± 0.9 38 ± 3.1 7.3 ± 0.5 18 ± 0.3 6 ± 1.7 40–43 42–46 8–12 34–41 7–8 18.1–18.7 4–6 3333323 Hipposideros armiger 931023458183850 Hipposideros larvatus 58 ± 1.3 68 ± 4.8 23.2 ± 2.1 38.1 ± 3.0 11.4 ± 1.0 23.4 ± 0.6 16.8 ± 1.6 56–61 59–79 20–27 31–43 10–13 22–25 14–21 26 26 26 26 26 25 18 Harpiocephalus harpia 49 ± 1.9 61 ± 6.3 16 ± 2.9 48 ± 3.1 12 ± 0.5 23 ± 1.2 16 ± 3.3 46–50 55–70 13–20 45–52 11–12 21–24 14–21 3333333 Pipistrellus paterculus 31 ± 0.8 39 ± 1.2 11 ± 0.8 32 ± 1.4 7 ± 0.5 12 ± 0.5 5.0 ± 0.5 30–32 38–41 10–12 31–34 6–7 11–12 4–5 3333333 Scotomanes ornatus 54 ± 2 67 ± 1.5 20 ± 0 55 ± 3 11.5 ± 0.5 23.5 ± 0.5 20 ± 1 52–56 65–68 20–20 52–58 11–12 23–24 19–20 2222222

Table 3. Summary statistics (mean ± one standard deviation, observed range, number of specimens) for craniodental measurements of nine species of bat collected at Huu Lien Nature Reserve

Species GTL CCL ZB BB PC C-M3 M3-M3 C1-C1 Rhinolophus pearsonii 25.2 ± 0.5 21.8 ± 0.3 11.9 ± 0.4 9.5 ± 0.3 2.6 ± 0.2 9.7 ± 0.2 8.9 ± 0.2 6.6 ± 0.2 23.8–25.9 21.2–22.4 11.4–12.8 9.2–10.1 2.2–2.8 9.4–10.2 8.5–9.2 6.3–7.0 16 17 17 17 17 17 17 17 Rhinolophus pusillus 15.2 13.3 7.5 6.5 2.3 5.7 5.5 3.7 Rhinolophus stheno 19.5 16.3 9.0 7.9 1.7 7.2 7 4.9 Aselliscus stoliczkanus 15.3 ± 0.1 13.1 ± 0.0 7.7 ± 0.1 6.2 ± 0.0 1.8 ± 0.1 5.3 ± 0.1 5.4 ± 0.1 3.5 ± 0.1 15.3–15.4 13.1–13.2 7.6–7.7 6.2–6.2 1.8–1.9 5.2–5.4 5.6–5.8 3.4–3.5 33333333 Hipposideros armiger 31.9 28.3 18.1 11.4 4.1 12.6 12.7 8.7 Hipposideros larvatus 23.0 ± 0.3 19.9 ± 0.3 12.7 ± 0.1 11.3 ± 0.1 3.3 ± 0.1 8.6 ± 0.1 8.6 ± 0.2 5.5 ± 0.1 22.5–23.5 19.5–20.5 12.5–12.9 11.1–11.5 3.2–3.6 8.4–8.8 8.3–8.8 5.3–5.9 22 22 22 22 22 22 22 22 Harpiocephalus harpia 22.1 18.9 14.2 9.5 5.4 6.3 7.6 6.7 Pipistrellus paterculus 12.3 ± 0.0 11.2 ± 0.2 7.8 ± 0.3 6.1 ± 0.0 3.4 ± 0.1 4.2 ± 0.0 5.1 ± 0.1 3.9 ± 0.0 12.3–12.4 11.0–11.4 7.6–8.1 6.0–6.1 3.3–3.5 5.1–5.3 5.1–5.3 3.9–3.9 22222222 Scotomanes ornatus 19.9 18.6 15.3 10.0 4.5 7.8 8.9 6.7 adult female netted in disturbed forest near our camp. (Tables 2 & 3). The dorsal pelage is pale brownish white Son et al. (2004) reported this species from Huu Lien for with darker brown tips while the ventral pelage is simi- the first time, and our capture confirms its presence in larly colored but paler overall. The noseleaf is simple the reserve. This is a very small species of Rhinolophus with a small and slightly hourglass-shaped sella; pointed 160 Mammal Study 32 (2007) connecting process; and a short pointed lancet. The mor- caves, in forest along streams, in agricultural scrub, and phology of the noseleaf of our specimen is similar to the in open clearings near houses. This is a medium sized description and illustration in Csorba et al. (2003) and it Hipposideros; external and cranial measurements are is an example of an individual with a slightly more elon- provided in Tables 2 and 3. The dorsal pelage is varying gated and pointed connecting process as described by shades of brown with the lighter brown bases of the hairs these authors. showing through; the ventral pelage is similar in color but slightly paler. Rhinolophus stheno K. Andersen, 1905 A single specimen netted in secondary growth just Harpiocephalus harpia (Temminck, 1840) outside Lan Dat village represents the first record of the Both Hendrichsen et al. (2001) and Son et al. (2004) species from the Huu Lien Nature Reserve. This is a reported H. harpia from the reserve and we document medium-sized horseshoe bat (Tables 2 & 3). The fur of three more; two were netted over a small pool in the for- the dorsal pelage is pale brownish white with darker red- est near camp and a third was collected without habitat dish brown tips; the ventral pelage is similarly colored information. External and cranial measurements are pro- but paler. The noseleaf has a rather broad horseshoe vided in Tables 2 and 3. The dorsal pelage is reddish- covering much of the muzzle; the sella is parallel sided; brown; ventral pelage similarly colored but somewhat the connecting process well rounded; and the lancet as grayer. The species is most readily identified in the field rather long and pointed with straight sides. Our speci- by its long tubular nostrils. The only other bats occur- men is referred to the subspecies R. s. microglobosus ring in the region with similarly tubular nostrils belong to (Csorba and Jenkins 1998). the genus Murina but only one of these, Murina leuco- gaster, approaches the large size of Hariopcephalus. Aselliscus stoliczkanus (Dobson, 1871) These two genera can be distinguished by the size of the First recorded from the reserve by Son et al. (2004); third upper molar: in Murina it fairly large, wheras in we collected three additional specimens. One was netted Harpiocephalus it is minute (0.30 mm in the one Huu inside a small cave at 17:30 hrs; the remaining two were Lien specimen from which the skull was removed). netted in secondary growth just outside the village. This is a small bat with short pointy ears; external and cranial Pipistrellus paterculus Thomas, 1915 measurements are provided in Tables 2 and 3. The dor- Three specimens were collected; two females netted sal pelage is dark brown, but with the white bases of the over a small pool in forest near camp, and one male net- hairs showing through; and the ventral pelage is pale ted over a small stream feeding the same pool. This is a whitish brown. This species is most readily identified by small pipistrelle; external and cranial measurements are its posterior noseleaf, which is divided into three upright provided in Tables 2 and 3. The dorsal pelage is uniform points. dark brown; the ventral pelage is similarly colored but slightly paler. The morphology of P. paterculus is very Hipposideros armiger (Hodgson, 1835) similar to that of P. abramus with the two differing pri- Both Hendrichsen et al. (2001) and Son et al. (2004) marily in the relative sizes of the anterior dentition: in P. reported this species from the reserve and we document paterculus the second upper incisor attains the height of an additional individual netted in the secondary growth the posterior cusp of the first upper incisor whereas in near our camp. This is a large species of Hipposideros; P. abramus they are subequal; the posterior cutting external and cranial measurements are provided in edge of the canine lacks a secondary cusp in P. paterculus Tables 2 and 3. The pelage is a rather uniform brown, whereas in P. abramus it is variably developed; and being only slightly paler on the ventral side. The nose- while the first upper premolar of P. paterculus is small it leaf is small, with a smoothly rounded horseshoe flanked is neither as minute nor so displaced out of the toothrow by four lateral lappets. as in many individuals of P. abramus. We agree with Bates et al. (1997) that these distinguishing dental char- Hipposideros larvatus (Horsfield, 1823) acters are not always consistent and therefore of only Twenty-six specimens were collected, making this the limited value for identification. Males of the two species single most commonly collected bat during our survey. are unequivocally distinguished based on the morphol- This species was caught wherever we set nets: inside ogy of the baculum: in P. paterculus the baculum is long Lunde et al., Small mammals in Vietnam 161 and nearly straight whereas in P. abramus the baculum is ventral pelage is mahogany red, but the very tip of the doubly curved (Thomas 1915; Hill and Harrison 1987; chin is colored like the sides of the face and the fur of the Bates et al. 1997). In all three of our specimens from scrotum and penis are grizzled with buff against dark Huu Lien the morphology of the anterior dentition fits brown. The tail hairs have alternating bands of black and the description of P. paterculus, and the cleaned baculum buff, with the terminal band of each hair being buff. We of our one male has the long and straight configuration regularly observed individuals of this species in forested characteristic of this species. Our specimens constitute areas throughout our survey area. The only other species the first records of P. paterculus from the Huu Lien of Callosciurus occurring in the region with which this Natural Reserve. species might be confused is Callosciurus inornatus, but this species is readily distinguished by its grey ventral Scotomanes ornatus (Blyth, 1851) pelage. Two specimens were collected; both were netted over a forest stream. The species was also reported by Son et Dremomys rufigenis (Blanford, 1878) al. (2004). This is a large and very distinctive species; We collected three specimens in forest: one on the external and cranial measurements are provided in ground and two others captured two days apart on the Tables 2 and 3. The dorsal pelage of this species is same 30 cm diameter woody liana. Ranges of external orange-brown with white patches on top of the head, measurements: TL 390–410, T 165–175, HF 52–55, E shoulders, flanks, and down the middle of the back. The 24–26. Range of weights: 270–310. The dorsal pelage is distribution of white splotches on the dorsal pelage var- dark blackish brown grizzled with pale orange. The ies between individuals. One of our specimens had a cheeks, sides of the face, and the top of the head are white splotch on top of the head, behind the ears, on the bright orange. The underside of the chin and throat are shoulders, between the shoulders, and on just one flank. pale orange but the fur of the remainder of the ventral A second specimen was similarly patterned but differed body pelage is gray with buff orange tips. The ears are in having splotches on both flanks, and in having an brown at the tip but with pale orange hairs at the base. irregularly defined white stripe running from the center The dorsal surfaces of the hands and feet are colored like of the back between the shoulders to a point midway the dorsal body pelage but with a little more orange between the flanks. The ventral pelage is darker brown color, and the sides of the arms and legs have a faint with somewhat less visually striking grayish white marks wash of buffy orange fur. The tail is bright orange along under the jaw, and along the chest to a grayish yellow the entire underside but the fur on the dorsal surface of patch on the abdomen. the tail is black with grayish white highlights. The spec- imens agree with descriptions of typical D. rufigenis Order Rodentia (Lunde and Son 2001). Dremomys gularis is one other Family Sciuridae species occurring in Vietnam but this species has a darker grey ventral pelage and extensive reddish-orange Callosciurus erythraeus (Pallas, 1779) fur on the throat and neck, but not on the cheeks. A third A single adult male was shot at dusk at the edge of for- species, Dremomys pernyi, may extend as far south as est just outside our camp. External measurements and northernmost Vietnam but this species is readily distin- weight taken from the fresh specimen: TL 440, T 190, guished from D. rufigenis in having the reddish-orange HF 57, E 23, Wt. 370 g. The dorsal pelage is finely griz- on the underside of the tail limited to the anal region, and zled and appears dark olivaceous in overall effect, the in having a complete absence of red markings on the individual hairs being dark brown with light buff-colored cheeks and throat (Moore and Tate 1965; Lunde and Son bands. The sides of the body are somewhat richer buff 2001). compared to the middorsal pelage. The dorsal surfaces of the hands and feet are very dark brown with just a fine Ratufa bicolor (Sparrman, 1778) speckling of buff, and the fingers and toes appear black An adult female shot by a local hunter was measured to the tip. The outer sides of the face, chin, arms and legs and photographed but not preserved as a voucher speci- are darker gray like the midback but are more finely men. speckled with buff. The ears are pale buff and contrast conspicuously with the darker dorsal pelage. Most of the 162 Mammal Study 32 (2007)

Table 4. Summary statistics (mean ± one standard deviation, observed range, number of specimens) for external and craniodental measure- ments, and weights of six species of murine rodent collected at Huu Lien Nature Reserve (See Musser et al. 2006, for summary statistics derived from the sample of Tonkinomys daovantieni, the seventh murine species encountered at the Reserve)

Rattus Niviventer Niviventer Rattus tanezumi Rattus losea Mus caroli andamanensis langbianis fulvescens Total length 391 ± 25 364 300 340 ± 10.0 332 ± 26.9 152 ± 8.5 353–435 325–358 212–372 135–161 22 9 60 10 Tail length 208 ± 16 185 155 181 ± 8.0 181 ± 15.9 77 ± 4.7 183–250 169–196 111–203 70–84 22 9 58 10 Hindfoot length 35.7 ± 1.6 34 29 29.4 ± 1.0 30 ± 1.3 17 ± 2.3 32–39 28–31 26–32 12–21 22 9 62 10 Ear length 24.1 ± 1.1 21 20 21.7 ± 0.8 23 ± 1.3 13.8 ± 1.7 22–26 20–23 20–25 12–18 22 9 58 10 Occipital-nasal length 41.8 ± 1.9 41.2 38.7 37.4 ± 0.6 37.6 ± 1.4 20.1 ± 1.4 39.3–45.8 36.7–38.5 34.8–40.0 18.9–22.1 18 8 51 10 Zygomatic breadth 20.6 ± 0.9 20.0 18.7 17.6 ± 0.4 16.8 ± 0.6 9.9 ± 0.7 19.4–22.3 17.0–18.0 15.1–18.0 9.4–11.0 20 8579 Interorbital breadth 5.8 ± 0.2 6.2 5.7 6.0 ± 0.2 5.7 ± 0.3 3.6 ± 0.1 5.3–6.2 5.7–6.4 4.9–6.3 3.5–3.7 21 8 58 10 Length of rostrum 12.9 ± 0.8 12.5 11.0 11.4 ± 0.3 12.0 ± 0.6 5.1 ± 0.4 11.7–14.4 11.0–12.1 10.8–13.5 4.7–5.7 22 8 57 10 Breadth of rostrum 7.3 ± 0.4 6.9 6.9 6.2 ± 0.3 6.4 ± 0.3 3.6 ± 0.4 6.7–8.2 5.6–6.7 5.8–7.0 3.2–4.1 22 8 57 10 Breadth of braincase 16.2 ± 0.4 15.7 15.5 15.0 ± 0.3 14.9 ± 0.4 9.0 ± 0.3 15.5–16.9 14.6–15.5 13.2–16.1 8.8–9.4 18 9 57 10 Height of braincase 11.2 ± 0.3 10.6 11.0 9.7 ± 0.2 10.0 ± 0.3 6.6 ± 0.3 10.5–11.7 9.4–10.0 8.6–10.7 6.3–7.0 19 9 57 10

Tamiops sp. Family Although we often heard or caught glimpses of Tamiops while tending our traplines, we did not pro- Niviventer fulvescens (Gray, 1847) cure any specimens. One of our camp helpers showed Sixty-four specimens were collected: 14 in scrub, 39 us a Tamiops that was trapped locally and that he kept in forest (including three under piles of limestone rub- caged in his home as a pet. This individual appeared to ble), and 11 in bamboo. Nearly all of these were trapped be much darker than Tamiops we had seen as museum on the ground or under piles of rocks but nine were specimens, and we were unable to identify it to species. trapped on woody lianas and one was trapped in a tree branch within arms reach of the forest floor. External and cranial measurements and weight are summarized in Table 4. Niviventer fulvescens is somewhat variable in Lunde et al., Small mammals in Vietnam 163

Table 4. (continued)

Rattus Niviventer Niviventer Rattus tanezumi Rattus losea Mus caroli andamanensis langbianis fulvescens Breadth of zygomatic plate 4.5 ± 0.3 4.6 4.3 3.8 ± 0.2 3.5 ± 0.2 2.0 ± 0.2 4.0–5.2 3.6–4.1 2.9–4.0 1.7–2.2 22 8 58 10 Length of diastema 11.4 ± 0.6 10.6 10.4 9.7 ± 0.3 9.7 ± 0.6 5.6 ± 0.7 10.3–12.7 9.4–10.1 8.4–10.8 5.1–6.6 22 8 57 10 Postpalatal length 14.1 ± 0.8 14.9 13.5 13.4 ± 0.5 12.9 ± 0.7 7.1 ± 0.7 13–15.8 12.7–14.4 11.6–14.2 6.5–8.2 19 8 53 10 Palatal length 9.0 ± 0.4 8.2 7.4 6.3 ± 0.3 6.7 ± 0.3 3.5 ± 0.1 8.3–9.9 5.8–6.8 6.0–7.4 3.4–3.7 22 8 58 10 Palatal breadth 4.3 ± 0.3 3.8 3.5 3.6 ± 0.2 3.6 ± 0.2 1.9 ± 0.2 3.8–5.0 3.4–3.9 3.0–4.1 1.8–2.1 22 8 58 10 Breadth mesopterygoid 3.1 ± 0.2 2.5 2.4 3.1 ± 0.3 2.9 ± 0.2 1.1 ± 0.1 2.7–3.4 2.7–3.6 2.4–3.3 0.9–1.2 22 8 58 10 Length incisive foramina 7.7 ± 0.3 6.9 7.7 6.9 ± 0.5 6.7 ± 0.4 4.3 ± 0.5 7.1–8.2 6.1–7.8 5.8–7.6 4.0–5.0 22 8 57 10 Breadth incisive foramina 2.6 ± 0.2 2.2 2.2 2.7 ± 0.2 2.6 ± 0.2 1.5 ± 0.1 2.2–3.1 2.4–3.0 2.3–3.1 1.4–1.6 22 8 57 10 Length of bulla 6.8 ± 0.3 7.4 6.7 5.4 ± 0.2 5.0 ± 0.3 3.5 ± 0.2 6.3–7.4 5.1–5.7 4.1–5.6 3.1–3.6 22 9 57 10 Crown length of molars 7.3 ± 0.2 6.7 6.2 6.0 ± 0.1 5.8 ± 0.2 3.4 ± 0.1 6.8–7.7 5.8–6.1 5.3–6.3 3.3–3.5 22 9 58 10 Breadth M1 2.1 ± 0.1 2.0 2.1 1.8 ± 0.1 1.7 ± 0.1 1.2 ± 0.0 2.0–2.3 1.7–1.9 1.5–1.9 1.2–1.2 22 9 58 10 Body Weight 129.4 ± 28.5 130 120 88.1 ± 8.0 79.0 ± 17.0 12.4 ± 2.8 85–180 70–100 20–120 9–19 22 9 62 10 external appearance; the dorsal pelage is spiny and of brown on the ventral side of the tip of the tail, while ranges in color from deep fulvous orange to a rather the more muted yellow-brown individuals tended to have muted yellowish brown. Among the specimens in our a tail that was more whitish toward the tip of the dorsal large sample there was a tendency for specimens col- surface. Like N. langbianis, the ventral pelage is creamy lected in bamboo and agricultural scrub to have a more white and sharply demarcated from the dorsal pelage; but orange dorsal pelage, while specimens from forest unlike N. langbianis, N. fulvescens does not have so dis- tended to be more yellowish brown. The tail of this spe- tinct a fringe of buffy orange fur where the dorsal and cies is brown on the dorsal surface and creamy white on ventral pelages meet. The dorsal surfaces of the hands the ventral surface. As with the dorsal pelage, the color- and feet of N. fulvescens are usually pale brownish white ation of the tail varied between individuals: the more but they sometimes have a darker brown patch in the orange-colored specimens tended to have a tail with a middle of the hand or foot. darker brown dorsal surface and sometimes even a trace 164 Mammal Study 32 (2007)

Niviventer langbianis (Robinson and Kloss, 1922) scrub, one in tall Imperata grass near the top of a lime- Nine specimens were collected: 8 in forest and one in stone massif, and one at the entrance to a cave-like struc- bamboo. Of these, three were caught a few meters above ture in a pile of limestone rubble. External and cranial ground on woody lianas, three were on the ground in measurements are provided in Table 4. The dorsal pel- dense vegetation, three were on the ground under lime- age is orange-brown and distinctly shaggy. Long black stone blocks. All are fairly uniform in external morphol- hairs run down the center of the back but there are no ogy. The dorsal pelage is spiny and dull brown in overall spines. The ventral pelage is creamy white and fairly color; the ventral pelage is creamy white and sharply well demarcated against the dorsal pelage. The dorsal demarcated from the dorsal pelage. The margin of the surfaces of the hands and feet are pale brownish white dorsal and ventral pelages is marked by patches of bright and often with pale brown patches in the center. The tail buffy orange fur. The tail is monocolored brown and is monocolored brown all around. This species is distin- only slightly paler on the underside. The dorsal surfaces guished from other species of Rattus occurring in the of the hands and feet are brownish white and sometimes area by its larger body size, the long coarse guard hairs have a darker brown patch in the center. External and running along the length of the back, and by its broader cranial measurements and weights are summarized in rostrum as reflected in measurements of breadth of ros- Table 4. Niviventer langbianis is distinguished from N. trum (Table 4). This is the species usually recorded in fulvescens by its somewhat duller dorsal pelage and char- the literature as koratensis, sikkimensis, or remotus (see acteristic buffy fur along the margins of the dorsal and the explanation in Musser and Carleton, 2005: 1464). ventral pelage, and by a monocolored brown tail and shorter hind feet with larger more bulbous plantar pads Rattus losea (Swinhoe, 1871) (see Musser et al. 2006; Fig. 9). Based on our trapping Our record is a specimen brought to us by a local vil- experience in Vietnam, N. langbianis is primarily arbo- lager claiming to have caught it near his home. External real and most often captured on woody lianas looping and cranial measurements are provided in Table 4. The down from the forest canopy; we were therefore sur- dorsal pelage is soft and orange-brown. The ventral pel- prised to have captured such a high proportion of our age is gray-based orange. The tail is monocolored brown Huu Lien on the ground. on both the dorsal and ventral side, with the ventral side being almost imperceptivity paler and appearing more Tonkinomys daovantieni Musser, Lunde and Son 2006 finely scaled compared to the tails other species of Rattus Our 14 specimens include the holotype and type series collected from Huu Lien. The dorsal surfaces of the of this recently described genus and species. This is a hands and feet are brownish white with darker brown large spiny rat with a grayish black dorsal pelage that is patches marking the center. This species is distinguished typically marked with a small white blaze on the top of from the two other species of Rattus recorded from the the head and a larger irregular blaze on the center of the region by its smaller body size and relatively short and chest. The basal two-thirds of the dorsal side of the tail narrow rostrum (Table 4). Although expected to occur in is dark brown but the underside and distal one-third of the region, our record from northeastern Vietnam fills in the tail is pale whitish brown. See Musser et al. (2006) a conspicuous gap in the documented distribution of the for a full description and comparisons. Restricted to species (Musser and Newcomb 1985). A revision of this areas of karst limestone, T. daovantieni were captured species is currently being prepared by Angela Frost, Ken only in the vicinity of rocky habitats in forest. Other spe- Aplin, and Guy Musser; Frost and Aplin examined our cies captured in association with this species include specimen and confirmed our identification. three specimens of Niviventer fulvescens and one speci- men of Rattus andamanensis. The species is most likely Rattus tanezumi Temminck, 1844 rather patchily distributed in suitably rocky habitats One example of this species was brought to us by a throughout the Viet Bac karst region. See discussion and local villager claiming to have caught it near his home. Musser et al. (2006) for further details. External and cranial measurements are provided in Table 4. Our one specimen is very similar to R. losea in exter- Rattus andamanensis (Blyth, 1860) nal appearance but differs in being larger and in having a We collected 22 specimens of this species, all on the somewhat more coarsely scaled tail. This species is dis- ground in highly modified non-forest habitats: 20 in tinguished from Rattus andamanensis by its shorter Lunde et al., Small mammals in Vietnam 165 molar toothrow length and from R. losea by its longer tion of its dark grey-brown pelage scattered with white- rostrum (Table 4). tipped hairs; and by having the two posterior pads of the foot separate (Lunde and Son 2001). Mus caroli Bonhote, 1902 We collected ten specimens, all on the ground in non- Family Hystricidae forest habitats: two in a field; four in scrub; and four in Imperata grass. On several occasions we observed what Atherurus macrourus (Linnaeus, 1758) we believe to be this species resting just inside small bur- A single specimen was presented to us by villagers rows dug into the sides of rice paddy dykes and on one who had trapped it in the scrub near the village. External occasion Lunde observed a 0.5 meter long green snake measurements: TL 700; T 240; HF 75; E 43. Weight: slowly enter one of these burrows just moments after the 3700 g. mouse had turned and retreated into its burrow. After waiting several minutes neither the mouse nor the snake Discussion were seen coming out of the burrow. External and cranial measurements are summarized in Table 4. Our The results of our small mammal survey at Huu Lien specimens have a soft brownish-grey dorsal pelage; a are just one more step in a continuing effort by the grayish buff ventral pelage; brownish white hands and authors to document and describe the diversity and distri- feet; and a long and sharply bicolored tail that is brown butions of small mammals across northern Vietnam. Of above and whitish below. This species is sometimes the rodents collected from the site, most are typical of the confused with Mus cervicolor but M. caroli is distin- Indochina fauna, with Callosciurus erythraeus, Dremo- guished by its longer tail, which is about the same length mys rufigenis, Niviventer fulvescens, and Niviventer of the head and body. langbianis being readily captured wherever forested hab- itats persist (Table 5). Likewise Rattus andamanensis, Rhizomys pruinosus Blyth, 1851 R. tanezumi, R. losea, Mus caroli, Atherurus macrourus A single specimen was obtained from local villagers and Rhizomys pruinosus are all commonly captured in and was said to have been trapped in a small stand of non-forest or disturbed habitats throughout Indochina. bamboo near our camp in the lowland valley. External One species that did stand apart is Tonkinomys daovan- measurements and weight: TL 463, T 130, HF 33, E 30, tieni, which, as discussed in Musser et al. (2006), is cur- Wt 205. This species is distinguished by the combina- rently known only from the presently reported type series

Table 5. Summaries of the numbers of specimens collected for 15 non-volant species according to elevation and habitat type

Rice field Scrub Forest Imperata Bamboo Species Totals 150 m 150–200 m 200–300 m 300–400 m 400–450 m Suncus etruscus 1–––12 Callosciurus erythraeus ––1–obs1 Dremomys rufigenis ––3––3 Ratufa bicolor ––obs––0 Tamiops sp – obs obs – obs 0 Niviventer fulvescens –1439–1164 Niviventer langbianis ––8–19 Tonkinomys daovantieni ––14––14 Rattus andamanensis –2011 –22 Rattus losea 1––––1 Rattus tanezumi 1––––1 Mus caroli 24–4–10 Rhizomys pruinosus –1––obs1 Atherurus macrourus –1–––1 Totals 5 40 66 5 13 128 Specimens were collected along a transect extending from our base camp in the rice field at 150 m to the summit of a limestone massif at 450 m. See text for further descriptions of habitats. Observations of animals from habitats where specimens were not collected are indicated with the abbreviation “obs” but these are not included in tallies of specimens. 166 Mammal Study 32 (2007) and is likely restricted to the Viet Bac karst region this list we add 4 more new species records: Rhinolo- although the full extent to which the species occurs phus pusillus, Rhinolophus stheno, Scotomanes ornatus within this region remains unknown. The fact that and Pipistrellus paterculus. Thus, combining the Tonkinomys daovantienti remained undiscovered and results of the present study with those of Hendrichson undescribed for so long points to the need for further et al. (2001) and Son et al. (2004), our best assessment small mammal surveys throughout Vietnam, and also of the number of bats present in Huu Lien stands at 18 highlights the value of applying novel and creative species — a number clearly under representing true bat methods to small mammal sampling efforts. Prior to test- species diversity in the reserve. ing the limestone rubble microhabitat we ran extensive One of the surprising outcomes of our survey was trap lines just a few meters away without ever capturing the paucity of shrews captured. The pitfall method Tonkinomys daovantieni, and had we not considered employed at Huu Lien was of essentially the same design moving traps into the adjacent microhabitat, we might and construction that Lunde has used with great success have easily missed this important and distinctive new throughout South America, Africa and southeast Asia genus and species. but, despite 443 bucket nights of pitfall trapping (331 Tonkinomys daovantieni is just one of two recently bucket nights in bamboo, and 112 in scrub) our only cap- described genera of rodents endemic to specific lime- tures were the two Suncus etruscus, one from each of the stone areas within Indochina, the other is Saxatilomys two lines. Certainly some specimens of the ubiquitous, paulinae (Musser et al. 2005). Although recorded from and typically abundant C. attenuata would have fallen only 14 specimens on the Laos side of a limestone habi- into the traps if they were present in any numbers. tat spanning the Annamite mountains, the species may Whether this paucity of shrews has something to do with occur in contiguous areas just a few kilometers to the trap design, some seasonal anomaly in shrew popula- east in Vietnam’s Central limestone region (Fig. 1). tions, or low shrew diversity in the Huu Lien limestone Together with two other recently described taxa, Lao- area remains a mystery. nastes aenigmamus (Jenkins et al. 2005) and Hylomys In conclusion, our survey of the small mammal fauna megalotis (Jenkins and Robinson 2002), it is part of a of the Huu Lien Nature Reserve resulted in the discovery recently discovered limestone endemic fauna. Does the of a new genus and species of petricolous rodent, and Viet Bac limestone region contain a similarly diverse documented the presence of many additional species endemic small mammal fauna? And what about the previously unrecorded from the reserve, however much remaining karst forming regions of Vietnam? Both Viet- more work remains to be done, and continued surveys nam’s Ha Long Bay and the Southern limestone regions are likely to yield more surprises. are virtually unexplored for small mammal species, and further collecting may uncover still more undescribed Acknowledgments: We extend our thanks to Dr. limestone endemic taxa. Nguyen Xuan Dang, Head of the Department of Zoology The first survey of the Huu Lien bat fauna was con- at IEBR, and Nancy Simmons, Chairperson of the Divi- ducted in 1998 (Son et al. 2004) but many of the species sion of Vertebrate Zoology, AMNH, for supporting our listed in this report cannot be confirmed with voucher field work in Vietnam. For assistance in the field we are specimens. Continuing surveys in the years to follow grateful to the following IEBR staff members: Nguyen made possible more firm results on the numbers of Xuan Nghia (mammalogy student), Le Dinh Thanh species present (Hendrichson et al. 2001). Just prior to (camp cook), and Doan Van Kien (herpetologist), and to our survey 11 species of bat were known to occur with our two local guides, Tran Van Chuyen and Trieu Sinh certainty in the Huu Lien Reserve: Cynopterus sphinx, Duong. Megaderma lyra, Rhinolophus marshalli, R. pearsonii, Hipposideros pomona, H. armiger, H. larvatus, Asellis- References cus stoliczkanus, Myotis chinensis, M. ricketti, and Harpiocephalus harpia (Hendrichson et al. 2001). Bates, P. J. J. and Harrison, D. L. 1997. Bats of the Indian Subconti- Hendrichson et al. (2001) also reported somewhat nent. Harrison Zoological Museum, Kent, 258 pp. less firm identifications of Hipposideros pratti and Bates, P. J. J., Harrison, D. L., Jenkins, P. D. and Walston, J. L. 1997. Three rare species of Pipistrellus (Chiroptera: Vespertilionidae) Miniopterus schreiber from the reserve and Son et al. new to Vietnam. Acta Zoologica Academiae Scientiarum (2004) documented Rhinolophus paradoxolophus. To Hungaricae 43: 359–374. Lunde et al., Small mammals in Vietnam 167

Can, D. N., Dang, N. X. and Son, N. T. 2000. Biodiversity of mam- Moore, J. C. and Tate, G. H. H. 1965. A study of the diurnal squirrels, malian fauna in Huu Lien, Lang Son. Tap Chi Sinh Hoc [Journal Sciurinae, of the Indian and Indochinese subregions. Fieldiana: of Biology] 22: 117–121 (in Vietnamese with English summary). Zoology 48: 1–351. Csorba, G. and Jenkins, P. D. 1998. First records of a new subspecies Musser, G. G. 1979. Results of the Archbold expeditions. No. 102. of Rhinolophus stheno (Chiroptera, Rhinolophidae) from Viet- The species of Chiropodomys, Arboreal mice of Indochina and nam. Bulletin of the Natural History Museum, London 64: 207– the Malay Archipelago. Bulletin of the American Museum of 211. Natural History 162: 377–445. Csorba, G., Ujhelyi, P. and Thomas, N. 2003. Horseshoe Bats of the Musser, G. G. and Newcomb, C. 1985. Definitions of Indochinese World. Alana Books, United Kingdom, 160 pp. Rattus losea and a new species from Vietnam. 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Appendix

Specimens collected (indicated by AMNH catalog number).

Suncus etruscus — 275544, 275574 Rhinolophus pearsonii — 275547, 275557, 275567, 275584, 275585, 275609, 275620, 275621, 275697, 275698, 275699, 275701, 275703, 275704, 275705, 275706, 275707, 275722 Rhinolophus pusillus — 275592 Rhinolophus stheno microglobosus — 275720 Aselliscus stoliczkanus — 275554, 275718, 275719 Hipposideros armiger — 275702 Hipposideros larvatus — 275539, 275548, 275558, 275559, 275560, 275561, 275562, 275563, 275564, 275565, 275666, 275667, 275668, 275669, 275678, 275679, 275680, 275681, 275682, 275700, 275723, 275724, 275725, 275726, 275727, 275728 Harpiocephalus harpia — 275540, 275541, 275721 Pipistrellus paterculus — 275542, 275543, 275556 Scotomanes ornatus — 275555, 275573 Callosciurus erythraeus — 275612 Dremomys rufigenis — 275664, 275665, 275690 Niviventer fulvescens — 275538, 275545, 275549, 275550, 275551, 275552, 275568, 275569, 275570, 275571, 275572, 275578, 275581, 275582, 275583, 275590, 275595, 275596, 275597, 275598, 275599, 275600, 275601, 275603, 275604, 275605, 275606, 275610, 275611, 275613, 275616, 275617, 275622, 275626, 275628, 275629, 275632, 275633, 275638, 275639, 275640, 275642, 275645, 275647, 275651, 275654, 275655, 275658, 275661, 275662, 275663, 275670, 275673, 275674, 275675, 275676, 275677, 275689, 275694, 275708, 275713, 275714, 275716, 275717 Niviventer langbianis — 275579, 275580, 275594, 275625, 275641, 275648, 275709, 275710, 275715 Tonkinomys daovantieni — 275575, 275576, 275577, 275586, 275593, 275602, 275618, 275627, 275644, 275688, 275692, 275693, 275711, 275712 Rattus andamanensis — 275546, 275608, 275615, 275624, 275630, 275631, 275634, 275635, 275646, 275652, 275656, 275657, 275659, 275660, 275671, 275672, 275684, 275685, 275686, 275687, 275695, 275696 Rattus losea — 275553 Rattus tanezumi — 275566 Mus caroli — 275589, 275591, 275607, 275619, 275623, 275636, 275637, 275643, 275649, 275650 Atherurus macrourus — 275683 Rhizomys pruinosus — 275691