Leptolalax tadungensis Rowley &amp; Tran &amp; Le &amp; Dau &amp; Peloso &amp; Nguyen &amp; Hoang &amp; Nguyen &amp; Ziegler 2016, sp. nov.
Creators: name: Rowley, Jodi J. L.
name: Tran, Dao T. A.
name: Le, Duong T. T.
name: Dau, Vinh Q.
name: Peloso, Pedro L. V.
name: Nguyen, Truong Q.
name: Hoang, Huy D.
name: Nguyen, Tao T.
name: Ziegler, Thomas
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Megophryidae
Leptolalax
Leptolalax tadungensis

description: <i>Leptolalax tadungensis</i> sp. nov. Figs. 8D, 14. <b>Holotype.</b> UNS 00516, adult male, in dry stream in evergreen forest in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8350º N, 108.0081º E, 1262 m elevation, Fig. 1, 9G). Collected on 13 July 2010 by Dao T. A. Tran, Vinh Q. Dau, and Huy D. Hoang. <b>Paratypes.</b> AMS R 177666 and ZFMK 96601, two adult males, collected on same date and locality as holotype. UNS00515, AMS R 177661, AMS R 177662, ZFMK 96602 and ZFMK 96603, five adult males collected on 17 July 2010 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8388º N, 107.8945º E, 720 m elevation). UNS 0 0 526 and UNS 0 0 527, two adult males, collected on 17 July 2011 in Ta Dung Nature Reserve, Dak Nong Province, Vietnam (11.8561º N, 108.0371º E, 1932 m elevation); UNS 0 0 517 adult female, collected at same site on 16 July 2011. Specimens were collected by Dao T. A. Tran and Huy D. Hoang. <b>Etymology.</b> Specific epithet “ <i>tadungensis</i> ” is in reference to the type locality of the species, Ta Dung Nature Reserve. <b>Diagnosis.</b> Assigned to the genus <i>Leptolalax</i> on the basis of the following characters: small size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra–axillary, pectoral, femoral and ventrolateral glands), the absence of vomerine teeth, the presence of tubercles on eyelids, and anterior tip of snout with pale vertical bar (Dubois 1983; Lathrop <i>et al</i>. 1998; Delorme <i>et al</i>. 2006). <i>Leptolalax tadungensis</i> <b>sp. nov.</b> is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) dark brownish red ventral surface with white speckling; (3) small/medium SVL for the genus (23.5–26.0 mm in eight adult males, 32.1 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 43–48% of SVL in males; (6) pectoral gland 3.3–5.4% of SVL in males, (7) smooth dorsum, (8) iris dark gold, (9) distinct black supratympanic line present, (10) an advertisement call with 4–7 notes with the first, introductory note encompassing less than a tenth of each call. <b>Description of holotype</b>. Head width less than head length; snout rounded in dorsal view and slightly truncate in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores straight; vertical pupil; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim slightly elevated relative to skin of temporal region; vomerine teeth absent; pineal ocellus absent; vocal sac openings slit–like, located posteriolaterally on floor of mouth; tongue wide with broad, shallow notch at posterior tip; supratympanic ridge distinct, running from eye to axillary gland. Tips of fingers rounded, very slightly swollen; relative finger lengths I &lt;IV = II &lt;III; nuptial pad absent; subarticular tubercles absent; a large, round inner palmar tubercle separated from a much smaller, laterally compressed outer palmar tubercle; finger webbing and dermal fringes absent. Tips of toes like fingers; relative toe length I &lt;II &lt;V &lt;III &lt;IV; subarticular tubercles absent; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toe webbing rudimentary, slight lateral fringes present. Tibia 47% of snout–vent length; tibiotarsal articulation reaches snout. Skin on dorsum smooth; ventral skin smooth; pectoral gland oval, 1.0 mm diameter; femoral gland small, oval, approximately 0.6 mm diameter, on posteroventral surface of thigh, closer to knee than to vent; supra–axillary gland oval, raised, 1.0 mm diameter. Ventrolateral glands indistinct. <b>Colour of holotype in life</b>. Dorsal surface dark brown; very faint transverse darker brown bars on the dorsal surface of the thighs, tibia, tarsus, lower arms, fingers and toes. Black line along supratympanic ridge, terminating above axilla, encompassing posterodorsal half of tympanum; anteroventral half paler brown; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms slightly paler, brownish copper; numerous, small darker brown spots on sides from groin to axilla. Dark brown ventral surface with fine white speckling on ventral surfaces of throat, chest, belly, legs and arms; most dense on chest. Supraaxillary gland copper; femoral glands white; pectoral glands white. Iris copper, fine black reticulations throughout. <b>Colour of holotype in preservative</b>. Dorsum uniform dark brown with scattered small black blotches on flanks and larger black spot on groin; fine black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8D). Ventral surface medium brown with only very faintly visible, fine white speckling. Macroglands white. <b>Measurements (mm)</b>. Holotype: SVL 24.9, HDL 9.4, HDW 8.9, SNT 3.8, EYE 2.9, IOD 3.4, TMP 1.6, TEY 0.9, TIB 11.7, EN 2.2, IN 2.2, NS 1.7, ML 6.0, PL 10.4. <b>Variation</b>. Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Fig. 14. Little variation in dorsal colouration. Single female specimen UNS 0 0 527 is paler in colour with a more truncate snout. The white speckling on venter is more distinct in UNS 0 0 527, UNS 00517, and AMS R 177661, forming small blotches in UNS 0 0 527, and UNS 00517; the dorsolateral line is more distinct and less broken in these specimens. UNS 0 0 526 and UNS 00517 have a slightly shagreened skin texture in preservative, possibly an artefact of preservation (with these specimens being better preserved). <b>Advertisement call.</b> Call descriptions are based on the calls of four individuals, recorded at 12.9–22.3ºC ambient temperature. Calls were an average of 291 ms in duration and consisted of 4–7 notes (Table 7, Figure 6D). Introductory notes were very short and quite similar to non-introductory notes, comprising only 3–9% of the call duration, and were indistinctly pulsed. Non-introductory notes contained 1–4 pulses. Notes were repeated an average of 14.3 notes/s. The average dominant frequency was 2.8 kHz. Harmonics were weakly present at ~6.3, 13.2, 16.6 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of <i>L. pallidus</i> <b>sp. nov.</b> is a short raspy chirp, similar to an orthopteran. <b>Ecology.</b> All specimens of <i>Leptolalax tadungensis</i> <b>sp. nov.</b> were found in evergreen forest in Ta Dung Nature Reserve between 720–1932 m elevation (Fig. 9H). All frogs were found calling during rainy conditions. Frogs collected at the type locality were in underground rocky crevices. Individuals collected on Ta Dung Mountain (~ 1930 m elevation) were calling from the ground or under leaf litter. <b>Distribution.</b> <i>Leptolalax tadungensis</i> <b>sp. nov.</b> is only known from Ta Dung Nature Reserve, Dak Nong Province, Vietnam. The greatest distance between known localities for the species is 15.7 km. <b>Comparisons.</b> <i>Leptolalax tadungensis</i> <b>sp. nov.</b> differs from all other <i>Leptolalax</i> species in mainland Southeast Asia on the basis of morphology, and from all congeners in the region for which comparable data is available on the basis of molecular and acoustic data. The new species differs from <i>L. aereus</i> by ventral coloration and presence of distinct black supratympanic line, from <i>L. bourreti</i> by ventral coloration, from <i>L. botsfordi</i> by male body size, from <i>L. croceus</i> by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from <i>L. eos</i> by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from <i>L. firthi</i> by ventral coloration, iris coloration, skin texture and lateral fringes on toes; from <i>L. fuliginosus</i> by ventral coloration and iris coloration; from <i>L. heteropus</i> by ventral coloration and iris coloration; from <i>L. kecil</i> by ventral coloration, male body size and iris coloration; from <i>L. melanoleucus</i> by ventral coloration and iris coloration; from <i>L. minimus</i> by ventral coloration; from <i>L. nahangensis</i> by ventral coloration and male body size; from <i>L. nyx</i> by ventral coloration; from <i>L pelodytoides</i> by ventral coloration; from <i>L. platycephalus</i> by ventral coloration and male body size; from <i>L. pluvialis</i> by ventral coloration and male body size; from <i>L. solus</i> by ventral coloration and iris coloration; from <i>L. sungi</i> by ventral coloration, male body size and iris coloration; from <i>L. tuberosus</i> by ventral coloration, presence of black supratympanic line and skin texture; from <i>L. ventripunctatus</i> by ventral coloration and iris coloration; and from <i>L. zhangyangpingi</i> by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the <i>L. applebyi</i> group, <i>L. tadungensis</i> <b>sp. nov</b>. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from <i>L. applebyi</i> by having a significantly greater body size (Wilcoxon post-hoc Z=3.6, p=0.001), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.5, p=0.013), and greater relative pectoral gland size (Wilcoxon posthoc Z=2.9, p=0.004). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. bidoupensis</i> by having a significantly smaller relative head width (Wilcoxon post-hoc Z=-2.1, p=0.032), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-2.1, p=0.032), and gold iris (versus reddish upper and silver below in <i>L. bidoupensis</i>). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. melicus</i> by having a significantly larger body size (Wilcoxon post-hoc Z=3.5, p=0.001), and smaller relative tibia length (Wilcoxon post-hoc Z= -2.3, p=0.019). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. pyrrhops</i> by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.4, p=0.001), smaller relative tibia length (Wilcoxon post-hoc Z= -3.2, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=-3.1, p=0.002) and smaller relative diameter of the eye (Wilcoxon post-hoc Z= -3.3, p=0.001). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. ardens</i> <b>sp. nov.</b> by having a significantly larger body size (Wilcoxon post-hoc Z= 3.4, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.0, p=0.003). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. pallidus</i> <b>sp. nov.</b> by having smaller relative tibia length (Wilcoxon post-hoc Z= -3.1, p=0.002), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.019), a black supratympanic line (versus no black supratypmanic line in <i>L. pallidus</i> <b>sp. nov</b>) and mostly smooth skin texture (versus tuberculate in <i>L. pallidus</i> <b>sp. nov</b>). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. kalonensis</i> <b>sp. nov.</b> by having a significantly smaller body size (Wilcoxon post-hoc Z= -3.3, p=0.001) and smaller relative tibia length (Wilcoxon post-hoc Z= -3.4, p=0.001). <i>L. tadungensis</i> <b>sp. nov</b>. differs from <i>L. maculosus</i> <b>sp. nov.</b> by having a significantly smaller relative tibia length (Wilcoxon post-hoc Z= -2.5, p=0.013). See Table 4 and Fig. 4. The male advertisement call of <i>L. tadungensis</i> <b>sp. nov.</b> differs from all species in the <i>L. applebyi</i> group. The call of the new species differs from <i>L. applebyi</i> in dominant frequency; from <i>L. bidoupensis</i> in having an introductory note present and in dominant frequency; from <i>L. melicus</i> in relative length of introductory note and dominant frequency; from <i>L. pyrrhops</i> by dominant frequency; from <i>L. ardens</i> <b>sp. nov.</b> by relative length of introductory note; from from <i>L. pallidus</i> <b>sp. nov.</b> by call duration and relative length of introductory note; from <i>L. kalonensis</i> <b>sp. nov.</b> by relative length of introductory note; from <i>L. maculosus</i> <b>sp. nov.</b> by call duration, number of notes per call, distinctness of introductory note and different relative duration of introductory note. See Table 7 and Fig. 5–6. <i>Leptolalax tadungensis</i> <b>sp. nov.</b> differs from all species within the <i>L. applebyi</i> group by&gt;6.3% divergence at the 16S gene fragment examined. Interspecific variation in four <i>L. tadungensis</i> <b>sp. nov.</b> collected from up to ~ 15 km apart was 0.0–0.2%.
description: Published as part of <i>Rowley, Jodi J. L., Tran, Dao T. A., Le, Duong T. T., Dau, Vinh Q., Peloso, Pedro L. V., Nguyen, Truong Q., Hoang, Huy D., Nguyen, Tao T. &amp; Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1)</i> on pages 96-99, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459
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