Leptobrachium waysepuntiense, Hamidy, Amir & Matsui, Masafumi, 2010

Hamidy, Amir & Matsui, Masafumi, 2010, A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia, Zootaxa 2395, pp. 34-44 : 36-40

Leptobrachium waysepuntiense sp. nov.

Holotype: MZB Amp 15862, an adult female collected among the leaf litter in primary forest in Way Sepunti trail (05°03’51’’S, 104°02’05’’E, 691 m a.s.l.; Figure 1 ), Kubu Perahu village, District of Liwa, Lampung Province, Sumatra, Indonesia, collected by A. Hamidy, S. Kirono, D. Susanto, Marji, Andi, and Habsi at 20:00 h on 14 February 2009 ( Figures 2 , 3 ).

Paratypes: MZB Amp 14592, an adult male collected by A. Ul Hasanah and W. Endarwin from near the type locality (05°03’51’’S, 104°02’07’’E, 852 m a.s.l.) in November 2004; KUHE 42805, a juvenile, collected on 13 February 2009, collectors and locality same as for holotype.

Etymology: The specific name refers to the locality where this species was collected.

Diagnosis: The new species is placed in Leptobrachium by having the combination of: femoral glands present; oval, flat axillary glands present; inner palmar tubercle circular, not extending along first metacarpal; vomerine teeth absent; snout and/or dermal palpebral projections absent; rictal glands and ventrolateral glandular ridges absent; spines on the upper lip absent. A medium-sized Leptobrachium (adult female 58.3 mm and male 50.0 mm in SVL; Table 1 ); iris in adult light blue with fine black reticulations, but is light grey in juvenile; dorsum uniform dark grey; dark grey laterally with white and orange dots; top of head from interorbital to parietal region, and dorsal side of fingers and toes faintly orange; greyish on belly and brownish on throat with white dots particularly on chest; no markings at groin; femoral glands very small.

Species Males Females

Description of Holotype (measurements in mm): Body tapering to the groin (SVL 58.3), head broad and depressed, slightly longer (HL 25.5: 43.7 % of SVL) than wide (HW 24.4: 41.9 % SVL); snout rounded from above, truncate in profile, very slightly projecting beyond lower jaw; eye large and obviously projecting from sides of head, smaller (EL 8.6: 14.8% SVL) than snout (SL 10.5: 18.0% SVL); canthi sharp, lores oblique, moderately concave; nostrils lateral, below canthus, distinctly closer to tip of snout (S-NL 4.8: 8.2% SVL) than to eye (N-EL 6.0: 10.3% SVL); internarial distance ( IND 4.2: 7.2% SVL) much shorter than interorbital distance, (IOD 8.1: 13.9% SVL), latter wider than upper eyelid (UEW 7.5: 12.9% SVL); no pineal spot; tympanum distinct, diameter (TD 5.0: 8.6% SVL) about three-fifth that of eye and separated from eye by half of its diameter (T-EL 2.5: 4.3%SVL); vomerine teeth absent; tongue heart-shaped, without papillae, notched posteriorly.

Forelimb slender and long (FLL 43.2: 74.1% SVL), about three-fifths of hindlimb; fingers moderately slender, unwebbed; first finger (FFL 6.6: 11.3% SVL) slightly longer than fourth and second, third much longer (TFL 10.3: 17.7% SVL); tips rounded, not swollen; inner palmar tubercle large (IPTL 2.8: 4.8% SVL), not extending onto first metacarpal and smaller outer palmar tubercle (OPTL 2.5: 4.3% SVL); subarticular tubercles indistinct, replaced by low callous tissue.

Hindlimb slender and relatively short (HLL 71.0: 121.8% SVL); heels not meeting when legs are held at right angles to body; tibia slightly longer (TL 20.3: 34.8% SVL) than foot (FL 19.8: 34.0% SVL); tibiotarsal articulation of adpressed limb reaching to the middle of tympanum ( Table 3 ); third toe longer than fifth; toe tips similar to those of fingers; toe webs poorly developed, webbing formula I 2 -- 2+ II 1 + -- 3 III 2 + -- 4- IV 4 - -- 2 V ( Figure 3 C); inner metatarsal tubercle low, oval, length (IMTL 2.6: 4.5% SVL) more than half distance between tip of first toe and tubercle (1TOEL 4.1:7.0%); outer metatarsal tubercle absent; subarticular tubercles obscure, but elongate, replaced by low callous tissue.

Skin above nearly smooth, with minute granules scattered posteriorly, especially around waist; granules denser laterally on body and on posterior thigh; ventrally slightly granular; a very low supratympanic ridge from eye to behind tympanum; indistinct low of dermal ridges on upper surface of forelimb; a flat pectoral gland at median border of axilla behind arm insertion; minute femoral gland on posterior surface of thigh.

Colour: In life, dorsally dark brownish grey with a faint V-shaped brownish orange interorbital and parietal marking ( Figure 3 A); grey colour fading laterally to light grey on ventral side ( Figure 3 B); laterally and ventrally dotted with white and orange, especially densely on sides; white dots denser on chest and throat than on abdomen; toes orange brown dorsolaterally; iris light blue with black reticulations; light blue orbital arc surrounding iris visible when eye opened maximally; no black bars around lips; supratympanic ridge bordered by a very thin brownish orange line; forelimb dorsally vaguely barred with dark brown; posterior thigh spotted with white and orange; no dark markings around groin from posterior flank to anterior thigh. In preservative, the aspects of the colour pattern remain, but the dorsal ground colour has been darkened and orange dots have faded to white.

Variation: The male paratype (MZB Amp 14592) is morphologically similar to holotype, but has smaller body, fewer dark reticulations on blue iris, more rugose dorsal skin, more distinct dark crossbars on the dorsal sides of limbs, and more distinct markings between interorbital and on upper half of tympanum. It has an internal vocal sac and a pair of vocal sac openings. The juvenile paratype (KUHE 42805; Figure 4 ) has a light grey iris with irregular black reticulations, and a distinct orange line from canthus through margin of upper eyelid and above tympanum to arm insertion. Narrow dark brown bars each with orange spots medially are distinct on dorsal half of limbs, and toes are dorsally light brown.

Comparisons: Small number of samples limited statistical comparisons of morphometric characters, but some dimensions did not overlap between L. waysepuntiense and other species ( Table 2 ). The new species tended to have longer LAL and FL than L. hasseltii , longer FL than L. hendricksoni , and longer LAL, FL, and IMTL than L. abbotti , all relative to SVL. In addition, the new species seemed to have larger FL/TL ratio than L. nigrops ( Table 2 ). In the new species, the tibiotarsal articulation reached the centre of the tympanum, but it barely reached the posterior end of the tympanum in many specimens of L. gunungense , and exceeded the anterior end of the tympanum in more than half specimens of L. montanum ( Table 3 ).

More discrete differences were found in qualitative characters ( Table 4). Leptobrachium waysepuntiense differs in eye colour from all the other congeneric species including the subgenus Vibrissaphora . In most species of Leptobrachium from China and Indochina like L. chapaense , L. hainanense and L. huansen ( Liu et al. 1973; Yang 1991; Dubois & Ohler 1998; Lathrop et al. 1998; Fei 1999; Fei et al. 2009), L. xanthospilum Lathrop, Murphy, Orlov & Ho and L. banae Lathrop, Orlov, Murphy & Ho , as well as species of Vibrissaphora , light blue or white colour is limited to the dorsal half of the iris, unlike L. waysepuntiense with totally light blue iris. Of the Indochinese species, L. buchardi has upper part of iris pale green ( Ohler et al. 2004), and L. mauhoti has black iris surrounded by an orange-red crescent dorsally ( Stuart et al. 2006). Leptobrachium hendricksoni from southernmost Thailand through Peninsular Malaysia, Sumatra to Borneo ( Berry 1975; our own observations), L. pullum from southern Vietnam ( Smith 1921) and L. smithi from Thailand to southern Malay Peninsula ( Matsui et al. 1999) share dorsal half of the iris scarlet or yellow in colour. Leptobrachium nigrops from Peninsular Malaysia, Sumatra and Borneo ( Berry 1975; our own observations) and all Bornean species, L. montanum , L. abbotti ( Inger et al. 1995; Malkmus et al. 2002; our own observation), and L. gunungense ( Malkmus et al. 2002; our own observations) have a totally black iris. The iris of L. hasseltii from Sumatra to Bali, once reported as scarlet ( Iskandar 1998), is actually totally black (our observations).

In addition to the iris colour, L. waysepuntiense differs from five of the seven congeners from Sundaland as follows; Leptobrachium waysepuntiense , without distinct dorsal markings, is differentiated from L. hasseltii and L. nigrops with distinct blotches on back, and L. smithi with confluent dark markings ( Matsui et al. 1999). Furthermore, L. waysepuntiense clearly differs in ventral marking from L. hendricksoni with many black spots and L. abbotti with large dark lead-grey to black spots. The remaining two species, L. montanum and L. gunungense , with totally dark iris can be easily differentiated from L. waysepuntiense with light blue iris in life, but are otherwise very similar, although femoral glands are larger and more distinct in L. montanum than L. waysepuntiense ( Figure 4 B).

Species N Sex Beyond anterior edge Between anterior and Behind posterior edge

of tympanum posterior edge of tympanum of tympanum Species Iris colour Marking at groin Femoral gland

L. waysepuntiense all parts light blue with fine black reticulations no very small white spot L. hasseltii all parts black yes very large white blotch L. hendricksoni upper part orange yes large white blotch L. nigrops all parts black yes large white blotch L. montanum all parts black no small white spot L. abbotti all parts black no large white blotch L. gunungense all parts black no very small white spot L. smithi upper part yellow, orange, or scarlet yes medium-sized white spot Range: Southwestern Sumatra, Indonesia. Known from the type locality at Way Sepunti trail, Kubu Perahu, Liwa, West Lampung, but probably also occurs in western of Jambi Province, central Sumatra (Matsui et al., unpublished data, see below) ( Fig. 1 ).

Natural history: Larval, acoustic and other ecological data are unknown. Way Sepunti trail, where the type series of L. waysepuntiense was collected, is in primary forest and located approximately 250 m from a rocky stream. On this trail, the new species was found at a higher elevation (691–852 m a.s.l.) than where L. hasseltii occurred (300 m a.s.l.). Other species found on this trail are: Leptophryne borbonica Tschudi , Megophrys nasuta (Schlegel) , Kalophrynus pleurostigma Tschudi , Limnonectes kuhlii (Tschudi) , Odorrana hosii (Boulenger) , Hylarana crassiovis (Boulenger) , Huia sumatrana Yan g, Microhyla palmipes Boulenger , M. berdmorei (Blyth) , and Rhacophorus sp. Neither eggs nor larvae were found and calls were not heard in mid February.

Figures

FIGURE 1. Map of southern Sumatra, showing the type locality (closed circle) of Leptobrachium waysepuntiense sp. nov.

FIGURE 2. The holotype of Leptobrachium waysepuntiense sp. nov. (SVL = 58.3 mm) in life (MZB Amp 15862).

FIGURE 3. Dorsal (A) and ventral (B) views, and ventral view of the right foot (C) of the holotype of Leptobrachium waysepuntiense sp. nov. (MZB Amp 15862). Scale indicates 10 mm in A and B, and 5 mm in C.

FIGURE 4. The juvenile paratype (KUHE 42805) of Leptobrachium waysepuntiense sp. nov. (SVL = 33.5 mm) in life (A) and its posteroventral view (B: TL = 13.8 mm).

Tables

TABLE 1. Morphometric variation of SVL (in mm) in Leptobrachium from Sundaland. For L. smithi, data for the topotypic population (Kao Chong) adopted from Matsui et al. (1999).

L. waysepuntiense	N Average (Range) 1 50.0	SD –	N Average (Range) 1 58.3	SD –
L. hasseltii	9 43.9 (40.0–49.7)	2.83	1 59.0	–
L. hendricksoni L. nigrops	5 42.7 (38.0–45.1) 8 31.3 (27.4–40.1)	2.91 4.18	5 55.0 (46.1–69.0) 8 40.5 (38.3–44.8)	9.62 2.32
L. montanum	10 57.9 (51.2–65.1)	4.79	2 65.1 (58.0–72.2)	–
L. abbotti L. gunungense	– – 4 43.9 (39.2–52.6)	– 5.94	2 68.7 (64.8–72.6) 4 59.4 (55.4–63.8)	– 3.47
L. smithi	17 41.4 (36.2–46.1)	2.49	2 55.6 (50.2–60.9)	–

TABLE 3. Variation in the point reached by the tibio-tarsal joint when hindlimb is bent forwards along body (percentage frequency in parentheses). For L. smithi, data for the topotypic population (Kao Chong) adopted from Matsui et al. (1999).

L. waysepuntiense 1 F –	1 (100%)	–
1 M –	1 (100%)	–
L. hasseltii 1 F –	–	1 (100%)
9 M –	6 (66.7 %)	3 (33.3%)
L. hendricksoni 5 F 5 (100%)	–	–
5 M 3 (60%)	2 (40%)	–
L. nigrops 8 F 7 (87.5%)	1 (12.5%)	–
8 M –	–	8 (100%)
L. montanum 2 F 1 (50%)	1 (50%)	–
10 M 7 (70%)	3 (30%)	–
L. abbotti 2 F –	1 (50%)	1 (50%)
L. gunungense 4 F –	–	4 (100%)
4 M –	2 (50%)	2 (50%)
L. smithi 2 F –	2 (100%)	–
17 M –	16 (94.1%)	1 (5.9%)

TABLE 2. Morphometric variation in Leptobrachium from Sundaland. Relative values (R) of each character to SVL (in %), and ratios of foot-tibia length and head width-head length are given in medians with ranges in parentheses. For L. smithi, data for the topotypic population (Kao Chong) adopted from Matsui et al. (1999). Juveniles and adults of both sexes combined.

Species	N	RHL	RHW	RLAL	RHLL	RTL
L. waysepuntiense	3	45.7 (43.7–46.0)	41.9 (41.6–43.6)	55.6 (55.0–58.2)	121.9 (114.2–134.3)	38.2 (34.8–41.2)
L. hasseltii	10	43.0 (40.8–44.2)	42.3 (40.5–43.8)	52.4 (49.3–54.6)	118.6 (110.7–126.0)	37.0 (34.7–39.3)
L. hendricksoni	10	44.8 (37.4–46.9)	43.9 (39.1–45.1)	56.8 (47.7–60.0)	123.9 (102.8–132.1)	38.4 (34.9–40.6)
L. nigrops	16	43.3 (41.6–45.8)	40.0 (38.1–42.3)	54.1 (48.3–58.7)	127.3 (117.7–135.0)	39.5 (35.5–42.3)
L. montanum	12	44.4 (42.9–46.7)	42.9 (42.3–50.4)	54.7 (50.0–61.5)	124.6 (115.5–137.7)	38.8 (34.6–43.2)
L. abbotti	2	44.3 (43.2–45.4)	42.3 (41.8–42.8)	53.3 (52.1–54.4)	117.6 (112.3–122.8)	35.5 (34.5–36.5)
L. gunungense	8	44.1 (43.0–47.6)	44.6 (42.0–45.3)	54.6 (41.4–57.1)	119.2 (115.9–124.7)	38.0 (35.9–39.7)
L. smithi	19	44.9 (43.1–45.8)	44.3 (41.2–46.0)	56.0 (52.2–57.8)	123.9 (115.5–132.8)	36.5 (34.0–38.6)
continued.