Leptolalax kalonensis 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 kalonensis

description: <i>Leptolalax kalonensis</i> sp. nov. Figs. 8B, 9B, 11. <b>Holotype.</b> IEBR A.2014.15, adult male, collected from rocky crevice, 0.5 m from 4–5 m wide, rocky stream in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748º N, 108.2109º E; 200 m elevation; Figs 1, 9F). Collected 5 August 2011 by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. <b>Paratypes.</b> AMNH A191768–9, 2 adult males, and IEBR A.2014.22, one adult female, collected on 6 August 2011; AMNH A191770, adult male collected on 11 August 2011; AMNH A191761 adult male collected on 12 August 2011; IEBR A.2014.18, adult male collected on 13 August 2011; all from disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2748º N, 108.2109º E; 200 m elevation). AMNH A191762, AMNH A191765, AMNH A191763, IEBR A.2014.19, AMNH A191764, IEBR A.2014.21, AMNH A191767, seven adult males, and AMNH A191766, IEBR A.2014.20, two adult females, collected on 5 August 2011; IEBR A.2014.15, adult male, collected on 12 August 2011; IEBR A.2014.16 and IEBR A.2014.17, two adult males, collected on 13 August 2011 in disturbed evergreen forest in Song Luy Watershed Forest, Binh Thuan Province, Vietnam (11.2919º N, 108.2150º E, 571 m elevation). Paratypes were collected by Dao T. A. Tran, David Kizirian, Truong Q. Nguyen, Pedro L. V. Peloso, and Cuong T. Pham. <b>Etymology.</b> Specific epithet “ <i>kalonensis</i> ” is in reference to the type locality of the species, which is in close proximity to the village previously known as Kalon. <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 kalonensis</i> <b>sp. nov.</b> is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) pale brownish pink ventral surface with white speckling; (3) small/medium SVL for the genus (25.8–30.6 mm in 16 adult males, 28.9–30.6 mm in three females); (4) toes lacking webbing and lateral fringes; (5) tibia length 45–52% of SVL in males; (6) pectoral gland 2.5–7.0% of SVL in males, (7) mostly smooth dorsum, (8) iris gold, coppery orange in upper third in some specimens (9) distinct black supratympanic line present in most specimens, (10) an advertisement call with 4–5 notes including introductory note, comprising almost one-third of each call duration. <b>Description of holotype</b>. Head width slightly subequal to head length; snout slightly truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum indistinct, 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 distinctly separated from small, 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, replaced by dermal ridges, distinct on base of toes II and III; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toe webbing rudimentary; lateral fringes absent. Tibia 46% of snout–vent length; tibiotarsal articulation reaching to anterior edge of eye. Skin on dorsum smooth; ventral skin smooth; pectoral gland oval, 1.0 mm diameter; femoral gland oval, 1.1 mm diameter, on posteroventral surface of thigh, closer to knee than to vent; supra–axillary gland oval, raised, 1.2 mm diameter. Ventrolateral glands indistinctly present as broken lines. <b>Colour of holotype in life</b>. Dorsal surface brown with slightly darker interorbital bar; faint dark W-shaped marking between axilla (Fig. 12A). Darker brown line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms paler brown; several small dark brown spots on sides from groin to axilla. Dark brownish ventral surface with white speckling on ventral surfaces of throat, chest and belly, more sparse on legs and arms. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris copper in upper half, gold in lower half, black reticulations throughout. <b>Colour of holotype in preservative</b>. Dorsum greyish brown with darker brown, elongated, oval blotches and intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet. Groin on blotch (Fig. 8B). Ventral surface cream. Macroglands creamy white. <b>Measurements (mm)</b>. Holotype: SVL 27.5, HDL 10.2, HDW 10.1, SNT 4.3, EYE 3.6, IOD 2.9, TMP 1.7, TEY 1.3, TIB 12.6, EN 2.5, IN 2.8, NS 1.6, ML 6.7, PL 11.5. <b>Variation</b>. Measurements of the type series are shown in Tables 2–3 and representative photographs of paratypes are shown in Figs. 9B and 12. AMNH A191768, AMNH A191769, IEBR A.2014.16, IEBR A.2014.17, and IEBR A.2014.22 are darker in dorsal colouration and have less distinct patterning. IEBR A.2014.18 and AMNH A191763 are pale and have only dark brown, rather than black markings. Specimens vary in degree of patterning on dorsum. Specimens vary in degree of medium brown speckling on venter; AMNH A191768, IEBR A.2014.16, AMNH A191762, IEBR A.2014.20, and AMNH A191766 have variably medium brown throats with white speckling and medium brown speckling on chest and belly (especially IEBR A.2014.16). The ventrolateral line varies in distinctness and completeness; the most distinct and complete in AMNH A191769. <b>Advertisement call.</b> Call descriptions are based on the call of AMNH A191770, recorded at 26.4 ºC ambient temperature. Calls were an average of 201 ms in duration and consisted of 4–5 notes (Table 7, Fig. 6B). Introductory notes of low relative amplitude were present in all calls, were an average of 29% of the call duration, and contained up to an average of 15 pulses. Non-introductory notes near-invariably contained a single pulse. Notes were repeated an average of 16 notes/s. The average dominant frequency was 2.8 kHz, spread more widely (2.6–2.9 kHz) in non-introductory notes. Harmonics and a fundamental frequency were not visible due to poor recording quality. To the human ear, the advertisement call of <i>L. kalonensis</i> <b>sp. nov.</b> is a short rasp, similar to an orthopteran. <b>Ecology.</b> All specimens of <i>Leptolalax kalonensis</i> <b>sp. nov.</b> were found in evergreen forest between 200–791 m elevation in Song Luy Watershed Forest. Specimens were located on the ground, 0.5–4.0 m from 2–5 m wide, shallow, rocky streams (Fig. 9F). Forest in the area was disturbed secondary forest, consisting of discontinuous patches of small hardwoods, bamboo and shrubs. Forest on mountain slopes was replaced by plantations and cultivated fields. <b>Distribution.</b> <i>Leptolalax kalonensis</i> <b>sp. nov.</b> is only known from Song Luy Watershed Forest in Binh Thuan Province, Vietnam. The greatest distance between known localities for the species is 43 km. Given the relatively large distance between localities, low elevation, and relatively larger streams, this species is likely to be more widely distributed than other species in the <i>L. applebyi</i> group. <b>Comparisons.</b> <i>Leptolalax kalonensis</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 black supratympanic line, from <i>L. bourreti</i> by ventral coloration, from <i>L. botsfordi</i> by ventral coloration, from <i>L. croceus</i> by ventral coloration, presence of black supratympanic line, 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, presence of black supratympanic line, 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 male body size; 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, iris coloration 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. kalonensis</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 larger body size (Wilcoxon post-hoc Z=4.0, p&lt;0.001), larger relative tibia length (Wilcoxon post-hoc Z=2.8, p=0.005), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.9, p=0.004), and larger relative pectoral gland size (Wilcoxon post-hoc Z=2.6, p=0.009). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. bidoupensis</i> by having a significantly larger body size (Wilcoxon post-hoc Z=4.4, p&lt;0.001), larger relative tibia length (Wilcoxon post-hoc Z=3.2, p=0.001), larger relative eye size (Wilcoxon post-hoc Z=2.3, p=0.019), and gold iris (versus reddish upper and silver below). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. melicus</i> by having a significantly larger body size (Wilcoxon post-hoc Z= -3.9, p&lt;0.001), and greater length between eye and tympanum (Wilcoxon post-hoc Z= -3.0, p=0.003). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. pyrrhops</i> by having a significantly smaller body size (Wilcoxon post-hoc Z=3.7, p&lt;0.001), smaller relative tympanum size (Wilcoxon post-hoc Z=2.7, p=0.006), smaller relative distance between eye and tympanum (Wilcoxon post-hoc Z=2.2, p=0.025) and smaller relative eye width (Wilcoxon post-hoc Z=3.5, p=0.001). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. ardens</i> <b>sp. nov.</b> by having a significantly greater body size (Wilcoxon post-hoc Z=4.7, p&lt;0.001). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. pallidus</i> <b>sp. nov.</b> by having a significantly greater body size (Wilcoxon post-hoc Z=3.0, p=0.002), smaller relative tympanum diameter (Wilcoxon post-hoc Z= -3.1, p=0.002), black supratympanic line in most specimens (versus no black supratypmanic line in <i>L. pallidus</i> <b>sp. nov</b>) and smooth skin texture (versus tuberculate in <i>L. kalonensis</i> <b>sp. nov</b>). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. maculosus</i> <b>sp. nov.</b> by having a significantly greater body size (Wilcoxon post-hoc Z= -2.7, p=0.007). <i>L. kalonensis</i> <b>sp. nov</b>. differs from <i>L. tadungensis</i> <b>sp. nov.</b> by having a significantly greater body size (Wilcoxon post-hoc Z= -3.3, p=0.001), and greater relative tibia length (Wilcoxon post-hoc Z=-3.5, p=0.001). See Table 4 and Fig. 4. Although based on a single call recording, the male advertisement call of <i>L. kalonensis</i> <b>sp. nov.</b> differs from all members of the <i>L. applebyi</i> group with the exception of <i>L. ardens</i> <b>sp. nov.</b> In particular, the species differs from <i>L. applebyi</i> in length of introductory note and dominant frequency; from <i>L. bidoupensis</i> in call duration, presence of a distinct introductory note and dominant frequency (although similar); from <i>L. melicus</i> in dominant frequency; from <i>L. pyrrhops</i> by dominant frequency; from <i>L. pallidus</i> <b>sp. nov.</b> by average call duration, average number of pulses in the introductory note and average note repetition rate; from <i>L. maculosus</i> <b>sp. nov.</b> by average call duration, number of notes per call, and having a distinct introductory note; from <i>L. tadungensis</i> <b>sp. nov.</b> by average call duration, and average duration of introductory note. See Table 7 and Figs. 5–6. <i>Leptolalax kalonensis</i> <b>sp. nov.</b> differs from all species within the <i>L. applebyi</i> group by&gt;5.2% divergence at the 16S gene fragment examined. Interspecific variation in four <i>L. kalonensis</i> <b>sp. nov.</b> collected from up to 43 km apart was 0.0–0.6%.
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 90-93, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459
description: {"references": ["Dubois, A. (1983) Note preliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espece novelle du Vietnam. Alytes, 2, 147 - 153.", "Lathrop, A., Murphy, R. W., Orlov, N. &amp; Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267. http: // dx. doi. org / 10.1163 / 156853898 X 00160"]}