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Russian Journal of Herpetology Vol. 5, No. 1, 1998, pp.

51 -60

TWO NEW SPECIES OF Leptobrachium (ANURA: MEGOPHRYIDAE)


FROM THE CENTRAL HIGHLANDS OF VIETNAM
WITH A REDESCRIPTION OF Leptobrachium chapaense

Amy ~ a t h r o p Robert
,~ W. ~ u r p h y Nikolai
,~ L. 0rlov: and Cuc Thu H O ~

Submitted 15 March 1998.

Two species of Leptobrachium are described from the Central Highlands of Vietnam in Gia Lai Province.
These descriptions increase the number of species of Leptobrachium in Indochina from four to six. We also
include a redescription of a northern species, Lepfobrachium chapaense, based on recent collections from
Tam Dao, Vinh Phu Province, Vietnam. A key to the species of Leptobrachium from Indochina is provided.

Key words: Amphibia, Anura, new species, Leptobrachium chapaense, Leptobrachium xanthospilum,
Leptobrachium banae, taxonomic key.

Our field work in Vietnam for the past five years with L. hasseltii because the differences between the
has led to the discovery of several new species of am- two species can be observed across several popula-
phibians (e.g., Lathrop et al., in press), as well as tions of L. hasseltii alone (Inger, 1966: p. 3 1). A
known species not previously recorded from Viet- smaller species, L. nigrops, is found on the Malay
nam. These discoveries include localities collected Peninsula as well as Singapore and Borneo (Berry and
heavily by pourret. Indeed, at every locality visited Hendrickson, 1963). It is very similar to L. hasseltii,
thus far, we have found either new species, or new but differs by having a more acurninate snout and
records of species occurrences in Vietnam. Among broader head. The detailed description of one species,
these discoveries, in the forests of Vietnam we have L. pullum, is wanting. All that is known for the spe-
found two new sympatric species of frogs belonging cies can be traced back to a meager description by
to the rare megophryid genus Leptobrachium. These Smith (1921: p. 440) in which he describes it as a
terrestrial amphibians may be found in high mountain "variant" of L. hasseltii and comparisons of the for-
regions either near streams or on the forest floor. mer made with L. nigrops (Berry and Hendrickson,
Their broad heads, and often dual-colored iris, easily
1963). Dubois (1983) elevated two Vietnamese
distinguishes them from other genera of anurans.
forms, L. hasseltii pullus (Smith, 1921) and L.
Until now only four species of Leptobrachium hasseltii chapaensis, (Bourret, 1937) to full species
were known from Indochina. One species, L. status without accompanying justification. In light of
hasseltii, has a broad distribution (Myanmar, Thai- this taxonomic change, and because we have col-
land, Borneo, Java, and the Philippines) and the vari-
lected L, chapaense, we believed it desirable to rede-
ation seen throughout its distribution has been care-
scribe L. chapaense. Notwithstanding, confirmation
fully detailed by Inger (1954, 1966). Leptobrachium
of the validity ofL. pullum as a distinct species must
hendrichoni (Taylor, 1962) was described from three
specimens from Malaya, Malaysia and Thailand. wait until type material can be examined. In the in-
However, this species was subsequently synonomized terim, the comparisons of our new species with L. pul-
lum are based on the brief description of Smith
Centre for Biodiversity and Conservation Biology, Royal On- '(1921).
tario Museum, 100 Queen's Park, Toronto, Ontario, M5S 2C6.
email: AL -AmyL@rom.on.ca; RWM -drbob@rom.on.ca.
MATERIAL AND METHODS
Zoological Institute, Russian Academy of Sciences, 119034 St.
Petersburg, Russia.
email: nbanoms@glas. apc. org.
All measurements were taken with digital calipers
Institute of Ecology and Biological Resources, Nghia Do, Tu to the nearest 0.01 mm and rounded to 0.1 rnm. Meas-
Liem, Hanoi, Vietnam. urements are summarized in Table 1. Below, snout

1026-2296/98/0501-0051 O 1998 Folium Publishing Company


52 Amy Lathrop et al.

vent length has been abbreviated SVL. The formula curs on the throat and darker mottling on the legs
for toe webbing follows Myers and Duellman (1982). (Fig. 3) distinguishing it from L. hasseltii, L. hen-
Abbreviations used: ROM) Royal Ontario Museum, dricksoni, L. pullum, L. nigrops, and the little known
Toronto, Canada; MVZ) Museum of Vertebrate Zool- L. hainanensis (Ye and Fe) (Ye et al., 1993) that are
ogy, University of California, Berkeley, USA; KIZ) said to have predominantly light venters (either cream
Kunming Institute of Zoology, Yunnan Province, or white) with some dark spotting. The absence of
China; IEBR) Institute of Ecology and Biological Re- distinct, dark spots on the dorsum distinguishes L.
sources, Hanoi, Vietnam. chapaense from most other species of Lepto-
brachium, including L. pullum, L. nigrops, L. hai-
Leptobrachium chapaense (Bourret) nanensis, many variants of L. hasseltii, and L. mon-
Figs. 1,2 tanum.
Megophlys hasseltii chapaensis Bourret, 1937 Description of species (based on 4 males, ROM
Leptobrachium (Leptobrachium) chapaense Dubois, 31199 -200; 32175 - 176 and 3 females, ROM
1980: p. 476 3 1196, 3 1198, IEBR specimen bearing ROM field
Diagnosis. A medium-sized Leptobrachium (fe- number 30006). Body short, head 42% in males, 36%
male 58.7 - 79.9 mm; male 53.5 - 65.5 mrn) with females of SVL; head broad, width slightly greater
short legs; heels do not overlap when legs are at right than length, greater than width of body; head dor-
angles to body; webbing basal on toes. Dorsal color soventrally compressed; canthus rostralis rounded,
pattern dark brown with small, diffuse orange loreal region oblique, slightly concave; interorbital
blotches on the sacral region, flanks, and limbs; ven- and frontal areas flat and sloping ventrally near snout;
tral coloration of limbs black with white reticulations nostril to eye distance almost twice that of nostril to
and banding on the thigh and knee; groin and flanks tip of snout; snout in dorsal view rounded, in lateral
black with white pustules that change to dark mottling view sloped. Eyes large and protuberant; eye length
on white near the chest and throat. Axillary gland lo- equal to snout length; tympanum small, half diameter
cated in armpit, behind insertion of arm. Dorsal half of eye, and separated from eye by a distance equal to
of iris white. or 75% tympanum diameter; supratympanic fold ex-
Leptobrachium chapaense has a relatively dark tends from rear comer of eye to articulation of jaw;
belly with distinct white punctuations; mottling oc- eyes broadly separated, interorbital distance 50% of

TABLE 1. Measurement for Leptobrachiurn from Indochina

Measurements,
L. chapaense L. pullum L. hasseltii L, nigrops L. xanthospilum ' L. banae '
mm 0" 9 0" 0 0" 0" 0 0" 9 0" 9
n=4 n=3 n=5 n=3 n=6 n=l n=7 n=8 n=2 n=14 n=2
SVL 53.5 - 65.5 58.7 -79.9
59.0 f 6.0 71.6 f 11.3
Head length/SVL 0.39 - 0.43 0.36 - 0.38
0.41 f 0.02 0.37 f 0.01
Head width/SVL 0.41 - 0.45 0.38 - 0.40
0.43 f 0.02 0.39 f 0.01
Arm/SVL 0.26-0.30 0.26 -0.28
0.28 f 0.02 0.27 f 0.01
Tibia/SVL 0.35 -0.36 0.35 -0.37
0.36 f 0.01 0.36 f 0.01

Note. Except SVL, where available, each measurement is given as a ratio with the mean k one standard deviation above the range (in
parentheses). Numbers in bold are measurements of the holotype. NA) Not available.
This study.
From Smith (1921: p. 440).
From Taylor (1962) for Thailand material. For further data on this species across its distribution see Inger (1954, 1966).
From Beny and Hendrickson (1963).
Vietnamese Leptobrachium
54 Amy Lathrop et al.
Vietnamese Leptobrachium
56 , Amy Lathrop et al.

head width; posterior 112 of tongue free, distinctly lighter tan, with light-brown mottling. Banding on the
notched; vomerine teeth absent; choanae separated by legs dark brown.
a distance slightly less than that between nares. Distribution and ecology. The type material is
Fingers long (27% SVL), tips not wider than dig- known from Sa Pa (Chapa), Lao Cai Province (22'22'
its; first finger slightly longer than second, latter N, 103'52' E), which is about 30 km south of the Chi-
slightly shorter than fourth; webbing absent, lateral nese border. Seven specimens, 4 males, ROM 3 1199
fringes weak; inner metacarpal tubercle, round, not -200; 32175 - 176 and 3 females, ROM 31 196,
protuberant, same size or slightly larger than outer 3 1198, IEBR specimen bearing ROM field number
metacarpal tubercle; midventral ridge incorporating 30006 and three specimens, 2 adult females and 1
subarticular tubercles of various sizes on each finger, adult male (MVZ 23976 - 77; 2398 I), were collected
no distinct tubercle at base of digits; webbing on toes in Tam Dao, Vinh Phu Province (21°27'N, 105'38'
basal; inner metatarsal tubercle large, slightly more
E), approximately 250 km SE of Sa Pa at an elevation
than half of length of first toe, outer metatarsal tuber-
approximately 400 m. Yang (1991) has records for
cle absent; subarticular tubercles on toes 3, 4 and 5;
ten individuals collected from five localities in central
one subarticular tubercle on second toe, none on first;
heels do not meet when legs are at right angles to and southern Yunnan: Tengchong (25'0' N, 98'5' E);
body. Menglian (24'5' N, 98'8' E); Mengla (21'4' E,
Skin on dorsum smooth with microscopic net- 101'5' E); Luchun (23'0' N, 102'4' E); Jingdong
work of ridges that are evident around the sacral re- (24'4' N, 100'8' E) from elevations between 1000-
gion and less evident on head and limbs (in preserva- 1900 m. Five adult frogs (two males and three' fe-
tion); flanks, belly, chest, and throat, with small white males) collected in Tam Dao on 4 May 1996 were
pustules; ventral surface of legs smooth with similar found in reproductive period in the rapid mountain
pustules adjacent to vent. Femoral gland distinct, its stream, and one couple was observed in amplexus
distance to vent is twice that to knee; chest gland im- (Fig. 2). This stream full down the steep mountain
mediately behind insertion of arm; area between jaw slope covered by the rain polydominant forest
articulation and eye with minute tubercles. (Fig. 4).
Coloration in life (based on specimens from Tam Remarks. Bourret (1937, 1942) considered this
Dao). Dorsum and limbs chocolate-brown. Irregu- species to be a subspecies of L. hasseltii. He distin-
larly shaped orange spots on back and limbs; legs guished it from L. hasseltii hasseltii by relative length
with alternating orange and black bands on the infe- of the 3rd finger, greater interorbital space, and the
rior surface of the tibiotarsus; feet, ventral side of condition and color of the metatarsal tubercle. It was
arms, and posterior thighs with black and white mot- distinguished from L. pullum by the size of the meta-
tling; groin black with small, white spots that are tarsal tubercle and color pattern. In addition, the white
more concentrated on the chest and throat, chest and color of the dorsal part of the eye distinguishes it from
throat predominantly white with some black mottling.
these two species, both of which have a scarlet-col-
Hands and feet gray, including tubercles; tips of digits
ored dorsal part of the eye. Although there appears to
white. Dark patch on canthus, just behind nares, and
one at posterior corner of eye; lower edge of su- be some variation in the eye color for populations of
pratyrnpanic fold bordered by dark line; margin L. chapaense, the eye color is never scarlet.
around eyelid black; nictatating membrane dark; dor- Yang (199 1: p. 69) reports that the dorsal portion
sal half of iris white. of the iris is blue. This may be similar to the very dark
Coloration in preservative. Dorsum dark brown or black iris described by Bourret (1937,1942). In the
with tan spots on back and limbs; ventral color pattern live specimens from Tam Dao, the dorsaI half of the
does not change in preservative. An older specimen iris is strikingly white, and very distinguishable from
(KIZ 8409186) in the collections of the Kunming In- the rest of the eye. This is evident even in our pre-
stitute of Zoology, Yunnan Province, China has re- served specimens, as they match very well in all other
tained a similar color pattern, but is slightly faded. respects to the description and illustration of L. cha-
The dorsurn is tan with some dark spotting; venter paense by Bourret (1942).
Vietnamese Leptobrachium 57

Leptobrachium xanthospilum (sp. nov.) Description of holotype. Body long (SVL 84.8
Figs. 5 , 6 rnm) head 39% SVL; head large, width equal to
Holotype. ROM 32 181, an adult female from the length, as broad as body; interorbital and frontal areas
forested hills about 2 krn east a farming village flat and sloping ventrally near snout; head dorsoven-
(14O20'29" N, 108O28'46" E elevation 850 m), Gia trally compressed; canthus rostralis rounded, loreal
Lia Province, Vietnam, collected on 25 September region oblique, slightly concave; nostrils to tip of
1997 by local Ba Na hunters. snout 1/3 distance of nostril to eye; snout in dorsal
Paratypes. ROM 32180 (female) collected at the view rounded, in lateral view sloped. Eyes large and
same time and in the vicinity of the holotype; ROM protuberant, diameter greater than snout length; tym-
32 182 - 88 (males) from Tram Lap (14O26'24" N, panum large, 63% of eye diameter, and separated
108O32'58" E elevation 900 m), Gia Lia Province, from eye by a distance 84% of tympanum diameter;
Vietnam (approximately 35 km NE [by road] of the supratympanic fold extends from rear comer of eye to
type locality) collected by N. L. Orlov, R. W. Mur- articulation of jaw; interorbital distance 55% of head
phy, and R. Bain on 16 - 27 June 1996. width; posterior 1/2 of tongue free, distinctly
Diagnosis. A large-sized Leptobrachium (two fe- notched; vomerine teeth absent; choanae separated by
males 83.2 - 84.8 mm SVL, eight males 62.8 - 73.4 a distance nearly 1.5 times internasal distance.
rnrn), with long legs, heels overlap when legs are at Fingers long (23% SVL), tips not wider than dig-
right angles to body; toes 1/3 webbed. Dorsal color its; first finger longer than second, latter equal to
pattern light brown to dark chocolate brown; flanks, fourth; webbing absent, lateral fringes weak; inner
posteroventral surface of legs; and sometimes chin metacarpal tubercle, oval, protuberant, outer metacar-
with distinct large yellow spots; limbs with or without pal tubercle similar shape as inner, but slightly
distinct dark bands, venter uniformly dark with min- smaller; midventral ridge of subarticular tubercles of
ute white spots on chest, throat and chin. Chest gland various sizes on each finger, no distinct tubercle at
located in armpit, behind insertion of arm. Dorsal half base of digits; toes about 1/3 webbed (11-1.5111-3-
of iris white. 1112-3.5IV3-5-1.5V); inner metatarsal tubercle large,
This large Leptobrachium can be distinguished length equal to first toe, outer metatarsal tubercle ab-
from all other species of Leptobrachium by the dis- sent; subarticular tubercles like those of hand on toes
tinct, large yellow (creamy-white in preservation) 3, 4, and 5, one on second toe, none on first; heels
spots on the flanks, posterior legs, and sometimes on broadly overlap when legs are at right angles to body.
the chin. Furthermore, the uniformly dark venter with Skin on dorsurn, smooth with microscopic net-
scattered white punctuation (Fig. 3) distinguishes it work of ridges that extend onto dorsal surface of
from L. nigrops, L. chapaense, L. hasseltii, and L. limbs and ventral surface of arms; belly, chest, throat,
hendrichoni (syn. of hasseltii; Inger, 1966) that are and area adjacent to vent with small pustules. Femoral
not uniformly dark, but predominantly either white gland, indistinct, midway between knee and vent;
marbled or speckled with darker pigmentation. In life chest gland immediately behind insertion of arm.
(sometimes visible in preserved specimens) the white Color in life. Dorsum and limbs dark brown; legs
dorsal part of the iris easily distinguishes it from L. with darker bands on the posterodorsal surface of
hasseltii and L. pullum that are said to be scarlet thigh and tibiotarsus; flanks with 7 - 8 large, well-de-
(Beny and Hendrickson, 1963: p. 618; Smith, 1921: fined yellow spots; venter uniformly dark brown-lav-
p. 440) and L. nigrops that has a uniformly black iris. ender with evenly dispersed light-brown pustules on
The long legs of L. xanthospilum, where the heels belly, chest, and adjacent to vent; more concentrated
broadly overlap when at right angles to the body, dis- pustules on throat and chin. Tips of digits and tuber-
tinguishes it from all species ofLeptobrachium except cles on hands and feet white. Margin of eyelid par-
Leptobrachium nigrops. Although the latter species tially yellow. Dorsal half of iris white.
has been reported to have relatively short legs (Berry Color in preservative. Dorsum dark brown with
and Hendrickson, 1963: p. 644), it is unclear if this microscopic network of ridges lighter brown; large
means the heels do not contact when drawn in toward spots on flanks, chest and chin, creamy-white. Limbs
the body. light brown with dark brown bands. Venter brown
58 Amy Lathrop dt al.

with very small light pustules. Toe tips, metatarsal, mental stages and three couples in amplexus (Fig. 6)
and metacarpal tubercles creamy white. were noted on 6 - 10 May, i.e., in the beginning of the
Variation. Dorsal color pattern is either dark pur- rainy season. Stony streams with Leptobrachium xan-
ple-black (specimens from Krong Pa and the majority thospilum filled by the deciduous leaves, flow in the
of those from Tram Lap) or brown (Tram Lap ROM aerial layered polydominant rain forest; the canopy
32187) with darker diffuse mottling. Venter in all above the streams is closed (Fig. 8).
specimens uniformly one color, either very dark Etymology. The specific name is derived from
(Krong Pa) or lighter brown (Tram Lap). In the lighter the Greekxanthos meaning yellow and the Greek spi-
colored specimens, the granular nature of the throat 10smeaning spot. The name refers to the conspicuous
and chin can be more easily detected because the yellow spots this frogs bears.
granules show up as creamy-white. Two Tram Lap
specimens have yellow dots on the chin and chest. Leptobmchium banae (sp. nov.)
Axillary gland more distinct in male Tram Lap speci- Fig. 9
mens. However, all males are from Tram lap and all Holotype. ROM 32200, an adult female from the
. females are from Krong Pa and, thus, this may reflect forested hills about 2 km east of a farming village,
an artifact of collecting. Arms of males are thicker Gia Lia Province, Vietnam (14°20'29" N,
and more stout; females have longer, weaker arms. 108'28'46" E, elevation 850 m) collected on 2 1 Sep-
Banding on arms distinctly obvious in the light brown tember 1997 by local Ba Na hunters.
specimen from Tram Lap, and less distinct in other Paratypes. ROM 32189 - 32198, 32201 - 207
Tram Lap specimens, and very difficult to see in the (males) and ROM 32 199 (female) from vicinity of the
dark females from Krong Pa. Interorbital bar on the type locality, collected by N. Orlov, R. Murphy, L.
one light brown specimen from Tram Lap. Some yel- Lowcock, A. Lathrop, and local hunters between 21 -
low spotting may occur on the head and around the 26 June 1998.
edge of the upper eyelid. Diagnosis. A large-sized Leptobrachium (two fe-
Remarks. Two gravid females were collected in males, 72.1 - 84.2 and 17 males, 57.2 - 70.0 mrn)
late September. As well, two metamorphs (ROM with short legs, heels do not overlap when legs are at
32178 - 32179) (Fig. 7), Gosner (1960) Stage 46, right angles to body; basal webbing on toes. Dorsal
were collected in late June from the same stream color pattern black with small red-orange diffuse
where the adults were collected. We are unsure blotches near groin, flanks and on back; forelimbs and
whether they are L. xanthospilum or the other sympa- hind limbs with alternating red (sometimes red-or-
tric species, described below. Nonetheless, if these ange) and black bands; ventral surface of limbs and
are L. xanthospilum and gravid females are found in flanks black, groin and belly brown with white pus-
September, this suggests that either the L. xanthospi- tules that extend up to chin. Dorsal half of iris white
lum have an extended larval period lasting nearly nine with a strikingly white nictitating membrane around
months, or that the breeding season is long. A similar the eye.
finding was reported by Pope (1 93 1) for Chinese L. Leptobrachium banae has distinct red (gray in
hasseltii in which larvae were found in January, April preservative) bands on the limbs that distinguish it
and July. No conclusions can be drawn until addi- from L. hendricboni which has no bands on the
tional collections can be made at different times of the limbs. The strikingly white porti.on of the iris distin-
year. guishes it from all species ofLeptobrachium except L.
Besides the localities noted above, we have ob- xanthospilum, and some forms of L. chapaense. From
served this species in the vicinity of Buoenloy village these two, it can be identified by its ventral color pat-
(14"20f38" N, 108O36'18" E, elevation 700 m), Gia tern, as illustrated in Fig. 3. The venter of L. banae is
Lai Province. One adult male was found in October dark with some white mottling and spotting, while L.
1993 during the rainy night on the leaf litter in 100 m chapaense is dark with distinct mottling and L. xan-
out from the stream; 26 metamorphs were collected in thospilum is entirely dark with indistinct spotting.
March- beginning of May 1995 in small mountain Description of holotype. Body long, head 37%
stream flowing into Cha River (tributary of Ba River), SVL; head large, width slightly greater than length,
In the latter locality the tadpoles on the latest develop- nearly as broad as body; interorbital and frontal areas
Vietnamese Leptobrachium

flat and sloping ventrally near the snout; head dor- pattern. Almost half the specimens have a uniformly
soventrally compressed; canthus rostralis rounded, colored dorsum, brown (black in preservative) with
loreal region oblique, slightly concave; nostrils half only a few spots of red (light gray in preservative) and
the distance to tip of snout than to eyes; snout in dor- mottling dorsolaterally. Three specimens are much
sal view rounded, in lateral view sloped. Eyes large lighter brown (gray in preservative) with dark mot-
and protuberant, eye equal to snout length; tympanum tling, including an interorbital triangle. The remaining
large, 57% of eye diameter, and separated from eye by eight specimens have a moderate amount of dark mot-
a distance 81% of tympanum diameter; supratym- tling on the dorsum with no interorbital triangle.
panic fold extends from rear corner of eye to articula- Banding pattern on the limbs is present in all speci-
tion of jaw; interorbital distance 47% of head width; mens, but more obvious in some, and not consistent
posterior 3/4 of tongue free, distinctly notched; with dorsal color patterns. Ventral color pattern is
vomerine teeth absent; choanae separated by a dis- consistent in all specimens.
tance slightly more than internasal distance. Remarks. Our field work in Krong Pa coincided
Fingers long (2 1% SVL), tips not wider than dig- with the beginning of the rainy season when it rained
its; first finger shorter than second, latter equal to everyday between 13 - 28 September, 1997. Follow-
fourth; webbing absent, lateral fringes weak; inner ing a few days of the heavy rain, choruses of about 5
metacarpal tubercle, oval, protuberant, outer metacar- - 10 male L. banae could be heard in the distance.
pal tubercle similar shape as inner, but slightly Several adults were collected from a forest in which
smaller; midventral ridge of subarticular tubercles of all of the understory had been cut and largely re-
various sizes on each finger, no distinct tubercle at moved by local Vietnamese farmers. Individual frogs
base of digits; toes 1/3 webbed (11-2111.5-31112- were collected near a foot path up to 2 krn from the
4IV4-2V); inner metatarsal tubercle large, length nearest stream. One male was located by its call,
equal to or slightly larger than first toe, outer metatar- where it was in a hole, buried in mud and leaf litter
sal tubercle absent; subarticular tubercles like those of along with a Bufo galeatus. The call, repeated about
hand on toes 3 , 4 and 5, two tubercles on second toe, every minute, consisted of two loud "crock ... crock"
one on first; heels do not overlap when legs at right followed by 5 to 8 more staccato-like crocks.
angles to body. Etymology. The specific epitaph is a patronym
Skin on dorsum smooth with microscopic net- from the word Ba Na which is the name of the ethnic
work of ridges that extend to a lesser extent onto dor- minority people in the area where this species was
sal surface of limbs; flanks, belly, chest and throat collected. Their invaluable assistance allowed us to
with small pustules; ventral surface of legs smooth, document the herpetofauna of Krong Pa.
sparse pustules on the thighs that extend upward to KEY TO THE SPECIES
vent. Femoral gland distinct, more than 50% closer to OF LEPTOBRACHIUM
knee than to vent; chest gland immediately behind in- FROM INDOCHINA
sertion of arm; area between jaw articulation and eye
I. Large conspicuous yellow spots along sides L. xan/hospilum
with minute tubercles. No large yellow spots on flanks 2
Color in life. Dorsum and limbs dark brown with 2. Red or orange spots on legs and or flanks 3
red-orange spots on the back and limbs; legs and arms Red spots absent on legs and or flanks 4
with bright red and black bands; ventral side of arms, 3. Ventral surface of tibiotarsus dark with white L. chapaense
legs and belly black, the latter with small pustules that marbling; iris white, blue or black
Ventral surface of tibiotarsus entirely black with no L. banae
are more or less evenly distributed throughout. Hands marbling; iris and palpebral membrane white
and feet gray, including tubercles; tips of digits white. 4. Snout acuminate in dorsal view; iris black L. nigmps
Nictatating membrane white; dorsal half of iris white. Snout rounded in dorsal view, spots on dorsum L. pullurn
Color in preservative: Dorsurn dark brown-black. Or- present or absent; iris scarlet or L. hassel~ii
ange spots turn cloudy gray in preservative, red
stripes on limbs white or gray in preservative; venter ACKNOWLEDGMENTS
dark brown, pustules on belly white.
Variation. Among the 19 specimens in the type Collecting and export permits were made avail-
series, some variation is present in the dorsal color able through Vietnam's Institute of Ecology and Bio-
60 Amy Lathrop et al.

logical Resources (IEBR), Hanoi. Import permits for des Sciences naturelles de 1'UniversitC au cours de l'an-
frozen tissues and preserved specimens were issued n6e 1937. Descriptions de deux especes et de deux
vari6tes nouvelles," Annexe Bull. G6n. Instl: Publique,
by Agriculture Canada and all collecting and euthana- 1937,l- 80.
sia of specimens were performed under approved ani- Bourret R. (1942), "Les Batraciens de l'Indochine," Mkm.
mal welfare protocols. This study was supported by Inst. Ockanogr: I %rdochine, 6.
the Natural Sciences and Research Council (NSERC) Dubois A. (1980), "Notes sur la Systematique et la Repar-
of Canada grant A3 148, by the generous assistance of tition des Amphibiens Anoures de Chine et des Regions
the ROM Sciences Fieldwork Fund, the ROM Future Avoisinantes. IV. Classification Generique et Subgen-
erique des Pelobatidae Megophryinae," Bull. Mens.
Fund, the ROM Foundation, and the Department of
Soc. Linn. Lyon, 49,469 - 482.
ROM Volunteers. We are extremely grateful to Prof.
Dubois A. (1983), "Note Preliminaire sur le Genre Lep-
Cao Van Sung, Director of IEBR, for assist in many topalax Dubois, 1980 (Amphibiens, Anoures), avec Di-
ways with on-site arrangements. Critical translations agnose d'une Esp6ce Nouvclle du Vietnam," Alytes, 2,
of Chinese descriptions were provided by J. Fu. Dr. 147- 153.
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