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Leptolalax puhoatensis sp. nov. Figs. 3–5. Holotype. VNMN 2016 A.22, adult male, calling in seep in evergreen forest at Pu Hoat Nature Reserve, Nghe An Province, Vietnam (19.6983° N, 104.7373° E, 1177 m a.s.l; Fig. 1). Collected on 5 June 2010 by Jodi J L Rowley, Trung Tien Cao, Vinh Quang Dau, Huong Thu Phung, Tuan Ngoc Le, Luong Thi Nguyen, Trung Danh Hoang, Thang Thanh Le. Paratypes. VNMN 2016 A.23, VNMN 2016 A.24, NCSM 79649, three adult males, same collection locality and date as holotype. VNMN 2016 A.26 and AMS R184850, two adult males collected in evergreen forest at Pu Hoat Nature Reserve, Nghe An Province, Vietnam (19.7044° N, 104.7401° E, 1128 m a.s.l on 6 June 2010). AMS R184851, adult male, collected in evergreen forest at Pu Hoat Nature Reserve, Nghe An Province, Vietnam (19.6986° N, 104.7317° E, 1352 m a.s.l on 7 June 2010). VNMN 2016 A.21, adult female, collected in evergreen forest at Pu Hoat Nature Reserve, Nghe An Province, Vietnam (19.7001° N, 104.7381° E, 1110 m a.s.l on 5 June 2010). AMS R184853, AMS R184854, two adult females collected in evergreen forest at Pu Hoat Nature Reserve, Nghe An Province, Vietnam (21.9156° N, 104.3608° E, ~ 1300 m a.s.l in May 2010). Specimens from June 2010 were collected by Jodi J L Rowley, Trung Tien Cao, Vinh Quang Dau, Huong Thu Phung, Tuan Ngoc Le, Luong Thi Nguyen, Trung Danh Hoang, Thang Thanh Le and specimens collected in May 2010 were collected by Vinh Quang Dau, Hoang Van Bui, Trung Quang Dau and Luong Thi Nguyen. Etymology. specific epithet is in reference to the type locality of Pu Hoat Nature Reserve. Diagnosis. Assigned to the genus Leptolalax on the basis of the following: small size, rounded finger tips, the presence of an elevated thenar tubercle not continuous to the thumb, presence of macroglands on body, vomerine teeth absent, tubercles on eyelids, anterior tip of snout with vertical white bar (Dubois 1980; Lathrop et al. 1998; Delorme et al. 2006). Leptolalax puhoatensis sp. nov. is distinguished from its congeners by a combination of (1) a body size range of 24.2–28.1 mm in eight adult males and 27.3–31.5 mm in three adult females, (2) males with a reddish-brown venter, often with faint white dusting and females with a pale pink venter, (3) toes with webbing basal and narrow lateral fringes, (4) iris copper in upper half and golden in lower half, and (5) a call consisting of a single note and a dominant frequency of 4.9–5.6 kHz (at 22.3–25.8° C). Description of holotype. Head wider than long; snout bluntly rounded in profile and obtusely pointed in dorsal view, projecting slightly over lower jaw; nostril closer to tip of snout than eye; canthus rostralis rounded, indistinct; lores sloping, slightly concave; vertical pupil; diameter of eye 78% length of snout; tympanum distinct, round, diameter approximately 62% that of the eye; vomerine teeth absent; pineal ocellus absent; large oval vocal sac openings present, located on either side of floor of mouth; tongue large, moderately broad, with wide notch at tip; distinct, raised supratympanic ridge running from corner of eye to axillary gland. Tips of fingers rounded, slightly wider than width of fingers; relative finger lengths I Colour of holotype in life (Figs. 3 A, 4A). Dorsum dark reddish brown with faint paler brown markings between eyes and W-shaped marking on axilla; black supratympanic fold and darker brown wash over upper half of tympanum, faint barring on upper lip, and transverse barring on dorsal surface of limbs including fingers and toes; ventral surface of elbow and upper arm without dark bars; deep coppery wash on elbows. Ventral surfaces deep reddish brown, faint white speckling on chest and belly, concentrated on chest; ventral surface of arms and legs with small whitish spots concentrated on edges. Supra-axillary gland copper, pectoral gland white, femoral glands white. Iris copper in upper half and golden in lower half with minute, black reticulations. Colour of holotype in preservative (Fig. 5 A): Dorsum medium brown with faint paler markings between eyes and on axilla, indistinct darker barring on surface of limbs including fingers and toes. Ventral surface, including throat, brown. Margins of throat and ventrolateral surfaces of arms and thighs, and entire tibiotarsus and upper arm pale brown with white speckling; macroglands cream. Measurements. Holotype: SVL 24.8, HDL 9.2, HDW 9.4, SNT 3.8, EYE 3.2, IOD 2.9, TMP 1.7, TEY 0.9, TIB 12.2, EN 2.1, IN 2.6, NS 2.0, ML 7.3, PL 12.3, F1 2.3, F2 3.2, F3 5.6. Weight in life 1.2 g. Measurements Males Females Range; Mean (N=8) Range; Mean (N=3) SVL 24.2–28.1; 25.8 27.3–31.5; 29.6 HDL 9.2–10.4; 9.9 10.9–12.3; 11.8 HDW 9.4–10.0; 9.4 10.5–12.1; 11.3 SNT 3.8–4.1; 3.9 4.1–4.5; 4.3 EYE 3.2–3.9; 3.4 3.6–4.1; 3.9 IOD 2.9–3.4; 3.1 3.1–3.3; 3.2 TMP 1.7–2.0; 1.9 2.2–2.4; 2.3 TEY 0.8–1.2; 1.0 1.0–1.7; 1.4 TIB 11.6–12.8; 12.3 12.8–15.6; 12.8 EN 1.9–2.4; 2.1 2.3–2.6; 2.5 IN 2.4–2.9; 2.6 2.7–3.1; 2.8 NS 1.6–2.0; 1.8 1.7–2.1; 2.0 ML 6.8–7.7; 7.2 6.9–9.0; 7.9 PL 11.8–13.1; 12.2 12.3–15.8; 13.9 Weight (g) 1.3–1.4; 1.3 (n=8) – Range; Median (N=8) Range; Median (N=3) HDL:HDW 0.97–1.05; 1.02 1.02–1.06; 1.04 HDL:SVL 0.36–0.42; 0.38 0.39–0.40; 0.40 TIB:SVL 0.45–0.52; 0.47 0.45–0.49; 0.47 TMP: SVL 0.07–0.08; 0.07 0.07–0.08; 0.07 Variation. Dorsal colour in life and in preservative is relatively uniform in males but the three females display brighter colours in life and more distinct patterning (Figs. 3, 5). In life, the ventral surfaces are deep reddish-brown in males, with varying amounts of faint white dusting in most individuals, and a pale pinkish colour in females (Fig. 4). In preservative, ventral surfaces are brown in males and very pale brownish in females. On the basis of the individuals collected, females also appear slightly larger than males (male SVL 24.2–28.1mm [N=8] and female SVL 27.3–31.5mm [N=3]). Measurements of the type series are shown in Table 3. Advertisement call. Call descriptions are based on the calls of the holotype, recorded at 22.3°C ambient temperature. Calls were an average of 9.8 ms in duration and invariably consisted of a single note. The dominant frequency was 5.2–5.6 kHz, and harmonics were present at approximately 10.9 and 16.5 kHz (Table 4, Fig. 6). A fundamental frequency was not evident. Calls were repeated at a rate of approximately ten calls per second, and had an average intercall interval of 86 ms. VNMN 2016 NCSM 79649 VNMN 2016 AMS AMS A.22* A.26 R184851 R184852 Primary Call duration (ms) 9.8 (6–14) 6.2 (6–7) 8.9 (8–10) 9.9 (7–11) 10.6 (8–14) call Intercall interval (ms) 86.4 (82–93) 106.7 (102–115) 102.6 (81–128) 78.8 (74–52) 85.6 (82–99) In one individual (VNMN 2016 A.26), a secondary call type was evident (Fig. 6 Biii), this call was 299 ms in duration and consisted of 41 notes of a similar frequency and lower amplitude than other calls. In the calls of the five individuals recorded, the number of notes in primary calls was invariably one. The dominant frequency varied only slightly among individuals, from 4.9–5.6 kHz (over 3.5 °C difference in ambient temperature). To the human ear, the advertisement call of L. puhoatensis sp. nov. is a very rapid, high-pitched ticking. Ecology. All specimens were found at night in evergreen forest between ~ 1123–1352 m a.s.l. Males were observed calling on rocks, stream banks and on vegetation within or adjacent to rocky streams (Fig. 7 A–B). The species was observed in smaller streams and seeps compared to the sympatric Leptolalax eos and Leptolalax ventripunctatus (Rowley et al. unpublished data). Conservation status. The species is known only from Pu Hoat Nature Reserve in Nghe An Province, Vietnam. Its true extent of occurrence is unknown but the species probably extends further into adjoining areas; suitable forested regions may also include adjacent forested areas in Nghe An Province as well as parts of eastern Huaphanh Province, Laos. The predicted Extent of Occurrence (EOO) for the species is 241 km 2.Given the small number ( Comparisons. Leptolalax puhoatensis sp. nov. differs from all known congeners by a combination of (1) a body size range of 24.2–28.1 mm in eight adult males and 27.3–31.5 mm in three adult females, (2) distinct dorsolateral markings including blackish spots on the flank and dark canthal and/or temporal streaks (3) males with a reddish-brown venter, often with faint white dusting, and females with a pale pink venter, (4) skin on dorsum with tiny, indistinct, low tubercles in preservative, more distinct and forming low dorsal ridges on dorsal surface in life (5) toes with webbing basal and narrow lateral fringes, (6) iris copper in upper half and golden in lower half, and (7) a call consisting of a single note of and a dominant frequency of 4.9–5.6 kHz (at 22.3–25.8° C). Leptolalax puhoatensis sp. nov. differs morphologically from all known Leptolalax species north of the Isthmus of Kra by having males with a reddish-brown venter, often with faint white dusting, and females with a pale pink venter (Table 5). A combination of male body size, presence of distinct blackish dorsolateral markings (canthal stripe and/or lateral blotches), and dorsal skin texture further distinguishes the new species from all other known species in the region (Table 5). In particular, the new species differs from the most closely related species, L. petrops, by having a reddish-brown venter in males and a pale pink venter in females (versus an immaculate white chest and belly), distinct blackish dorsolateral markings (versus no distinct black markings on the head) and dorsal skin with longitudinal skin ridges (versus highly tuberculate skin texture with no longitudinal ridges), having a call consisting of a single note and with a dominant frequency of 4.9–5.6 kHz at 22.3–25.8° C (versus a call with an average of four notes and a dominant frequency of 5.6–6.4 kHz at 24.5–25.3° C), and by molecular divergence (6.3% at the 16S gene fragment examined). puhoatensis sp. nov. Species Ventral Coloration Male SVL (mm) Distinct đorsolateral markings Dorsal skin texture In addition to morphological differences (Table 5), the advertisement call of the new species further differentiates Leptolalax puhoatensis sp. nov. from all other congeners north of the Isthmus of Kra with known calls (20 of 36 species). In having a call consisting invariably of a single note, Leptolalax puhoatensis sp. nov. differs from L. aereus, L. alpinus, L. applebyi, L. ardens, L. bidoupensis, L. botsfordi, L. croceus, L. firthi, L. fuliginosus, L. isos, L. kalonensis, L. liui, L. maculosus, L. melicus, L. oshanensis, L. pallidus, L. phyrrhops, and L. tadungensis. Of the Leptolalax species in the region with known calls, only L. tuberosus has an invariably single– note call, but this call is of a longer duration (54–78 ms at 22.4–22.5° C vs 6–14 ms at 22.3–25.8° C in the new species). In addition, the dominant frequency of 4.9–5.6 kHz (at 22.3–25.8° C) further distinguishes the call of the new species from that of the higher frequency calls of L. aereus (6.2–7.9 kHz at 22.4–25.7° C), L. alpinus (6.7 kHz at 16° C) and L. isos (5.8–6.2 kHz at 22.4–22.8° C), and the lower frequency calls of L. applebyi (4.0–4.3 kHz at 21.5° C), L. ardens (3.1–3.4 kHz at 21.4–24.7° C), L. bidoupensis (1.9–3.8 kHz at 19–21° C), L. botsfordi (2.6–3.4 kHz at 14° C), L. croceus (3.0–4.3 kHz at 21.6–25.1° C), L. fuliginosus (2.1–2.8 kHz at 19.3–19.6° C), L. kalonensis (2.8 kHz at 26.4° C), L. maculosus (2.7–2.8 kHz at 23.3–24.1° C), L. melicus (2.6–4.0 kHz at 26.1–26.2° C), L. oshanensis (4.4–4.6 kHz at 14° C), L. pallidus (2.4–2.7 kHz at 14.0–21.4° C), L. pyrrhops (1.91–2.23 kHz at 25° C), L. tadungensis (2.6–3.1 kHz at 12.9–22.3° C) and L. tuberosus (2.8–2.8 kHz at 22.5–24.5° C). Calls of the sympatric congeners L. eos and L. ventripunctatus remain unknown.

Leptolalax petrops sp. nov. Figs. 3���4. Holotype: VNMN 2016 A.06, adult male, 0.5 m from 5 m wide, rocky stream in evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2045 �� N, 105.0754 �� E, 715 m; Fig. 1). Collected on 25 May 2013 by Jodi J. L. Rowley, Dau Quang Vinh, Hoang Duc Huy, and Hoang Anh Tuan. Paratypes: AMS R 84828, adult female, and AMS R184828, VNMN 2016 A.08, VNMN 2016 A.09, VNMN 2016 A.10, VNMN 2016 A.11, AMS R184829, six adult males, collected at same location and date as holotype; AMS R184826 and AMS R1848287, two adult males, collected at same location as holotype on 24 May 2013. AMS R184830, AMS R184832, AMS R184833, three adult males collected at same location as holotype on 26 May 2013. AMS R184831, adult male, collected 2 m from 5 m wide rocky stream in evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2061 �� N, 105.0747 �� E, 587 m) on 26 May 2013. AMS R184834, adult male collected 0.5 m from 15 m wide, rocky stream in disturbed evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2105 �� N, 105.0619 �� E, 234 m) on 1 June 2013. VNMN 2016 A.12, adult female collected near 1���2 m wide creek in evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2118 �� N, 105.0824 �� E, 996 m) on 27 May 2013. VNMN 2016 A.13, AMS R184836, two adult females, and AMS R184835, adult male, collected in disturbed evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2067 �� N, 105.0783 �� E, 679 m) on 28 May 2013. AMS R184837, adult female, and AMS R184838, AMS R184839, AMS R184841, three adult males, collected in evergreen forest near 5���6 m wide rocky stream at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2062 N, 105.0811 E, 814 m) on 29 May 2013. AMS R184840, adult male, collected in evergreen forest at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2054 �� N, 105.0824 �� E, 835 m) on 29 May 2013. All specimens were collected by Jodi J. L. Rowley, Dau Quang Vinh, Hoang Duc Huy, and Hoang Anh Tuan. AMS R184842, AMS R184843, AMS R184845, three adult females, and AMS R184844, one adult male, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2758 �� N, 103.2637 �� E, 1257 m) on 14 September 2013. VNMN 2016 A.14, adult male, and VNMN 2016 A.15, VNMN 2016 A.16, two adult females, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2758 �� N, 103.2637 �� E, 1257 m) on 15 September 2013. VNMN 2016 A.17, VNMN 2016 A.18, VNMN 2016 A.19, three adult females, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2758 �� N, 103.2637 �� E, 1257 m) on 18 September 2013. VNMN 2016 A.20, one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2740 �� N, 103.2648 �� E, 1237 m) on 17 September 2013. AMS R184846, one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2773 �� N, 103.2626 �� E, 1260 m) on 19 September 2013. AMS R184847, one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2770 �� N, 103.2593 �� E, 1283 m) on 19 September 2013. AMS R184848, one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2687 �� N, 103.2589 �� E, 1171 m) on 21 September 2013. All specimens were collected in evergreen forest over limestone terrain by Jodi J. L. Rowley, Le Thi Thuy Duong, Dau Quang Vinh, Chad Minshew, and Hoang Anh Tuan. Etymology: The specific epithet derives from the Latin petra, meaning rock, and ���ops, meaning having the appearance of, in reference to both the stone-like appearance of the frog and the limestone karst in which the species was found in Lai Chau Province. Diagnosis: Assigned to the genus Leptolalax on the basis of the following: small size, rounded finger tips, the presence of an elevated thenar tubercle not continuous to the thumb, presence of macroglands on body, vomerine teeth absent, tubercles on eyelids, anterior tip of snout with vertical white bar (Dubois 1980; Lathrop et al. 1998; Delorme et al. 2006). The species differs from its congeners by having (1) a medium-sized body of 23.6���27.6 mm in 21 adult males and 30.3���47.0 mm in 17 adult females, (2) immaculate white chest and belly, (3) no distinct black markings on the head, (4) highly tuberculate skin texture, (5) toes lacking webbing and with narrow lateral fringes, and (6) a call consisting of an average of four notes and a dominant frequency of 5.6���6.4 kHz (at 24.5���25.3 ��C). Description of holotype: Head longer than wide; snout rounded in profile and obtusely pointed in dorsal view, projecting slightly over lower jaw; nostril approximately equidistant to tip of snout and eye; canthi rostralis rounded, indistinct; lores sloping, slightly concave; vertical pupil; diameter of eye 84% length of snout; tympanum distinct, round, diameter approximately half that of the eye; vomerine teeth absent; pineal ocellus absent; large oval vocal sac openings present, located on either side of floor of mouth; tongue large, moderately broad, with wide notch at tip; distinct, raised supratympanic ridge running from corner of eye to axillary gland. Tips of fingers rounded, slightly wider than width of fingers; relative finger lengths I Colour of holotype in life: Dorsum dull medium brown with slightly darker brown markings, the most distinct of these being a interorbital bar, a W-shaped mark on axilla, indistinct darker wash over upper tympanum, distinct dark brown barring on upper lip, and transverse barring on dorsal surface of limbs including fingers and toes; ventral surface of elbow and upper arm without dark bars; pale yellowish brown on elbows. Throat and ventral surface of arms pale pink and slightly transparent, particularly at either side of throat; edges of throat pale brown with white speckling, concentrated towards snout; ventral surface of chest and abdomen immaculate white; ventral surface of legs pinkish grey with small whitish spots concentrated on edges. Supra-axillary gland pale yellowish brown, pectoral gland cream, femoral glands cream. Iris gold in lower half and copper in upper half, with minute, black reticulations. Colour of holotype in preservative: Nearly uniform dark brown dorsal surface, with darker barring on surface of limbs including fingers and toes. Ventral surface creamy white. Ventrolateral surfaces of lower arms and tibiotarsus pale brown; Supra-axillary, pectoral and femoral glands creamy white. Measurements: Holotype (mm): SVL 24.3, HDL 9.4, HDW 9.0, SNT 43.7, EYE 2.9, IOD 3.4, TMP 1.8, TEY 0.9, TIB 12.2, EN 2.1, IN 2.4, NS 1.7, ML 6.3, PL 11.3, F1 2.3, F2 3.0, F3 4.8. Weight in life 1.8 g. Measurements Tuyen Quang Province Lai Chau Province Range; Mean (N=19) Range; Mean (N=2) SVL 24.3���27.6; 26.0 23.6���27.0; 25.3 HDL 9.4���10.8; 10.1 9.1���10.9; 10.0 HDW 8.9���10.1; 9.5 8.5���10.2; 9.4 SNT 2.3���4.3; 3.8 3.7���4.0; 3.9 EYE 2.9���3.7; 3.4 3.3���3.4; 3.3 IOD 2.6���3.6; 3.2 3.0���3.2; 3.1 TMP 1.7���2.3; 2.0 1.5���2.0; 1.7 TEY 0.7���1.2; 1.0 0.9���1.2; 1.1 TIB 12.0���13.5; 12.6 11.6���13.8; 12.7 EN 1.5���2.6; 2.2 2.3���2.6; 2.5 IN 2.3���2.9; 2.6 2.5���2.7; 2.6 NS 1.3���1.8; 1.6 1.4���1.7; 1.6 ML 6.3���7.8; 7.0 6.2���7.0; 6.6 PL 11.1���12.9; 11.9 10.8���12.1; 11.5 Weight (g) 1.8���2.2; 1.9 1.6���2.2; 1.9 Range; Median (N=19) Range; Median (N=2) HDL:HDW 1.01���1.09; 1.06 1.07���1.07 HDL:SVL 0.36���0.41; 0.39 0.39���0.40 TIB:SVL 0.46���0.51; 0.48 0.49���0.51 TMP: SVL 0.06���0.09; 0.08 0.06���0.07 Variation: Male L. petrops sp. nov. are smaller than females (23.6���27.6 mm in 21 adult males versus 30.3��� 47.0 mm in 17 adult females). The 12 females from Lai Chau Province were larger than the three from Tuyen Quang Province (34.0���47.0 versus 30.3���33.5 mm; Table 4). Dorsal colour in life varies from pale to medium brown, and vary in their degree of dorsal patterning. AMS R184841, AMS R184840, AMS R184831, AMS R184827, have a nearly completely unpatterned dorsum, others have weak patterning. Individuals vary in the number and size of dark spots along the flanks. Females are more tuberculate than males. In both sexes, skin texture more tuberculate in life compared to in preservative. Tubercles form ridges in some individuals. Measurements of the type series are shown in Tables 3���4. Advertisement call: Call descriptions are based on the calls of the holotype, recorded at 24.7��C ambient temperature. Most calls were an average of 55 ms in duration and consisted of an average of 3.9 notes (Table 5). Notes contained a single pulse. The dominant frequency was 6.0 kHz, and harmonics were present at approximately 12.2 and 18.2 kHz (Fig. 5). A fundamental frequency was not evident. Calls were repeated at a rate of 3.4 calls per second, and had an average intercall interval of 237 ms. In six of the seven males recorded an additional call type was evident (Fig. 5 Bi, iii). In the holotype, these calls were 347 and 351 ms in duration and consisted of 24 and 23 notes of a similar frequency and amplitude to other calls. In the calls of the seven individuals recorded, the number of notes in the most common calls varied from one to six but four was the typical number of notes. The dominant frequency varied only slightly among individuals, from 5.6���6.4 kHz (over 0.8 ��C difference in ambient temperature). To the human ear, the advertisement call of L. petrops sp. nov. is a rapid, high-pitched rasping, similar to an orthopteran. Ecology: All specimens were found at night in forest between ~ 234���1283 m asl. In Tuyen Quang Province, males were observed calling from on rocks, stream banks and on vegetation within or adjacent to rocky streams in May. During surveys in Lai Chau Province in September, no males were heard calling and most frogs encountered were females, mostly located away from water on limestone karst. Conservation status: The species is currently known from Lai Chau and Tuyen Quang Provinces, Vietnam. The true extent of occurrence is unknown but the species probably extends further into adjoining areas of northwestern Vietnam and southern China. Suitable forested regions may include parts of Hoang Lien Son Nature Reserve, Muong Nhe Nature Reserve and Nam Don Nature Reserve in Vietnam. We recommend that the species is listed as Vulnerable on the Red List of Threatened Species as its predicted Extent of Occurrence is 19,874km 2, it likely occurs in only two threat-defined locations, and there is an observed continuing decline in the quality of its habitat, satisfying Red List Criteria B1ab(iii). AMS VNMN 2016 VNMN VNMN AMS AMS AMS R184826 A.06* 2016 A.09 2016 A.11 R184831 R184832 R184833 Primary Call duration 42.6 (10��� 55.0 52.1 55.5 (51��� 53.2 (50��� 54.8 (52��� 59.9 (57��� call (ms) 57) (44���57) (5���56) 60) 55) 58) 62) Secondar Duration none 347, 351 379 228, 266 215 378, 354 217, 196 y call Notes/call 24, 23 28 16, 18 18, 21, 21 14, 13 Comparisons: Leptolalax petrops sp. nov. differs morphologically and/or bioacoustically to all known Leptolalax species. From species in the subgenus Lalos (Dubois et al. 2010), distributed south of the Isthmus of Kra, Leptolalax petrops sp. nov. differs by lacking ventrolateral glands (present in the subgenus Lalos; Delorme et al. 2006) and by molecular divergence (Table 2, Fig. 2). From the 35 Leptolalax species considered to be in subgenus Leptolalax (Delorme et al. 2006) or in the Leptolalax applebyi group (subgeneric placement remains unresolved; Rowley et al. 2016), Leptolalax petrops sp. nov. differs by a combination of morphological, molecular and/or bioacoustic data. From the 23 species with molecular data available, the new species differs in a high divergence in the 16S gene fragment examined (Table 2, Fig. 2), and by a combination of morphological and bioacoustic characters. From L. aureus, the new species differs by having highly tuberculate dorsal skin (versus finely tuberculate), having black lateral spots (versus no lateral spots), and a call consisting of 1���6 notes (average of four) (versus a call consisting of 2���16 notes). From L. applebyi, the new species differs by having a body size of 23.6���27.6 mm in 21 adult males and 30.3���47.0 mm in 17 adult females (versus 19.6���22.3 mm in males and 21.7���25.9 mm in females), having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black markings present), and a call with a dominant frequency of 5.6��� 6.4 kHz at 24.5���25.3 ��C (versus 4.0���4.3 kHz at 21.5 ��C). From L. ardens, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black markings present), and a call with a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 3.1���4.3 kHz at 21.5 ��C). From L. bidoupensis, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black markings present), and a call consisting of an average of four notes (versus 6���9 notes) and a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 1.9���3.8 kHz at 19���21 ��C). From L. bourreti, the new species differs by having a body size of 23.6���27.6 mm in adult males (versus 36.2 mm), highly tuberculate dorsal skin (versus mostly smooth), and lacking distinct black markings on the head (versus black markings present). From L. eos, the new species differs by having a male body size of 23.6���27.6 mm (versus 33.1���34.7 mm), and having highly tuberculate dorsal skin (versus shagreened). From L. firthi, the new species differs by having black lateral spots (versus no lateral spots), feet with narrow but distinct lateral fringes (versus feet with wide lateral fringes in males), and a call repetition rate of 1.7���3.1 calls/s at 24.5���25.3 ��C (versus L. isos, the new species differs by having black lateral spots (versus no lateral spots), feet with narrow but distinct lateral fringes in both sexes (versus feet with wide lateral fringes in males), and a call consisting of an average of four notes (versus three notes). From L. kalonensis, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus dark brownish with white speckling), and a call with a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 2.8 kHz at 26.4 ��C). From L. laui, the new species differs by having highly tuberculate dorsal skin (versus shagreened), and feet with narrow but distinct lateral fringes in both sexes (versus feet with wide lateral fringes). From L. liui, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), lacking distinct black markings on the head (versus black markings present), and feet with narrow but distinct lateral fringes in both sexes (versus feet with wide lateral fringes). From L. maculosus, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black markings present), and a call with a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 2.7���2.8 kHz at 23.3���24.1 ��C). From L. melicus, the new species differs by having a male body size of 23.6��� 27.6 mm (versus 19.5���22.7 mm in males), highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black markings present), and a call with a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 2.9��� 3.8 kHz at 26.1���26.2 ��C). From L. minimus, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth). From L. nyx, the new species differs by lacking dark, regularly set rounded spots on the dorsum (versus dark, regularly set rounded spots present), and highly tuberculate dorsal skin (versus mostly smooth). From L. oshanensis, the new species differs by having highly tuberculate dorsal skin (versus smooth with fine glandular ridges), no distinct black markings on the head (versus black markings present), and a call with an average of four short repeated notes (versus three) and a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 4.4���4.6 kHz at 14 ��C). From L. pallidus, the new species differs by having an immaculate white chest and belly (versus brown with white speckling), and a call with a dominant frequency of 5.6���6.4 kHz at 24.5���25.3 ��C (versus 2.4���2.7 kHz at 14.0���21.4 ��C). From L. pyrrhops, the new species differs by having a male body size of 23.6���27.6 mm (versus 30.8���34.3 mm), highly tuberculate dorsal skin (versus finely shagreened), an immaculate white chest and belly (versus brownish with white speckling), no distinct black markings on the head (versus black markings present), and a call with an average of four notes (versus 5���6 notes) and a, Published as part of Rowley, Jodi J. L., Dau, Vinh Q., Hoang, Huy D., Le, Duong T. T., Cutajar, Timothy P. & Nguyen, Tao T., 2017, A new species of Leptolalax (Anura: Megophryidae) from northern Vietnam, pp. 544-564 in Zootaxa 4243 (3) on pages 551-558, DOI: 10.11646/zootaxa.4243.3.7, http://zenodo.org/record/400196, {"references":["Lathrop, A., Murphy, R. W., Orlov, N. & Ho, C. T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibia-Reptilia, 19, 253 - 267.","Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24, 6 - 21.","Dubois, A., Grosjean, S., Ohler, A., Adler, K. A. & Zhao, E. - M. (2010) The nomenclatural status of some generic nomina of Megophryidae (Amphibia: Anura). Zootaxa, 2493, 66 - 68.","Rowley, J. J. L., Tran, D. T. A., Le, D. T. T., Dau, V. Q., Peloso, P. L. V., Nguyen, T. Q., Hoang, H. D., Nguyen, T. T. & Ziegler, T. (2016) Five new, microendemic Asian Leaf-Litter Frogs (Leptolalax) from the southern Annamite Mountains, Vietnam. Zootaxa, 4085 (1), 63 - 102."]}

Leptolalax pallidus sp. nov. Figs. 8A, 11. Holotype. UNS 0 0 510, adult male, calling on plant leaf 0.1 m from ground, 1 m from 2–3 m wide rocky stream in evergreen forest, Bidoup Nui Ba National Park, Lam Dong Province, Vietnam. (12.1520 º N, 108.695º E; 1644 m elevation, Fig. 1, 9E). Collected at night on 27 June 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. UNS 0 0 512, AMS R 177657, ZFMK 96599, three adult males collected on 26 July 2011 in in evergreen forest in Bidoup Nui Ba National Park, Lam Dong Province, Vietnam (12.1519º N, 108.6940º E; 1681 m elevation). UNS 0 0 511, AMS R 177659, ZFMK 96598, three adult males collected on same date and location as holotype. AMS R 177658, collected on 27 June 2011 at same location as holotype. All specimens were collected by Dao T. A. Tran and Duong T. T. Le Etymology. The specific name “ pallidus ”, meaning pale, is used as an adjective in reference to the paler coloration and less distinct markings of this species, particularly in preservative, compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax 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 et al. 1998; Delorme et al. 2006). Leptolalax pallidus sp. nov. is distinguished from its congeners by a combination of (1) supra–axillary and ventrolateral glands present; (2) dark brownish red ventral surface with faint white speckling; (3) small SVL for the genus (24.5.1– 27.7 mm in 8 adult males), (4) toes lacking webbing and lateral fringes; (5) tibia length 45–51% of SVL in males; (6) pectoral gland 3.4–6.7% of SVL in males, (7) tuberculate skin texture on dorsum, (8) iris copper in upper half, gold in lower half; (9) distinct black supratympanic line absent, (10) an advertisement call with 4–7 notes with the short introductory note encompassing a quarter of each call and only one note in non-introductory notes. Description of holotype. Head width slightly subequal to head length; snout truncate in dorsal view and in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to 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 small, 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 Colour of holotype in life. Dorsal surface brown with slightly darker interorbital bar with pale brown patches anteriorly and posteriorly; faint dark W-shaped marking between axilla (Fig. 11A). 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; numerous, 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. Colour of holotype in preservative. Dorsum relatively uniform medium brown with darker interorbital bar and slightly paler posterior to this; dorsal surface of brachium, fingers and toes paler brown, dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8A). Elbows and upper arms pale brown. Ventral surface medium brown with faint white speckles. Macroglands creamy white. Measurements (mm). Holotype: SVL 25.2, HDL 9.4, HDW 9.2, SNT 3.9, EYE 3.4, IOD 3.2, TMP 1.7, TEY 1.4, TIB 13.0, EN 2.5, IN 2.6, NS 1.2, ML 6.8, PL 12.6. Variation. Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Fig. 11. All specimens have a degree of darker brown patterning (mostly marbling but also some spots) on the dorsum and dark banding on the tibiotarsus and antebrachium, hands and feet, but both are more pronounced in UNS00512, AMS R 177657 and UNS00511. ZFMK 96599 has faint dark brown spots evenly distributed around tubercles on dorsum. UNS00510–00512 have more distinctly pale elbows in preservative. Size and distribution of white speckles on the venter varies a little between specimens, more clumped on chest in AMS R 177659 and ZFMK 96598. Skin texture less tuberculate in preservative than in life. Advertisement call. Call descriptions are based on the calls of three individuals, recorded at 14.0–21.4 ºC ambient temperature. Calls were an average of 673 ms in duration and consisted of 4–7 notes (Table 7, Fig. 6A). Short introductory notes of low relative amplitude were present in all calls. Introductory notes were 22–26% of the call duration, contained up to 63 pulses. Non-introductory notes invariably contained a single pulse. Amplitude peaked towards the middle of the call. Notes were repeated an average of 6.8 notes/s. The average dominant frequency was 2.5 kHz. Harmonics were weakly present at ~7.3 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. pallidus sp. nov. consists of a squelch followed by clicking. Ecology. All specimens of Leptolalax pallidus sp. nov. were found in evergreen forest in Bidoup Nui Ba National Park between 1644–1681 m elevation. Males were located on the forest floor, on leaf-litter or rocky crevices and clay holes in the banks of a dry stream. Fig 9E. Distribution. Leptolalax pallidus sp. nov. is only known from Gia Rich mountain in Bidoup Nui Ba National Park, only about 5 km from L. bidoupensis. The two species occur in continuous forest, with no obvious current biogeographic barrier between them. Comparisons. Leptolalax pallidus sp. nov. differs from all other Leptolalax 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 L. aereus by ventral coloration and iris coloration, from L. bourreti by ventral coloration, male body size, absence of black supratympanic line, skin texture and iris coloration, from L. botsfordi by male body size, absence of black supratympanic line, iris coloration and relative male body weight (0.42–0.50 versus 0.75–0.94 g /mm), from L. croceus by ventral coloration and iris coloration; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, absence of black supratympanic line, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size, skin texture and iris coloration; from L. heteropus by ventral coloration, male body size, skin texture and absence of black supratympanic line; from L. kecil by ventral coloration, male body size, absence of black supratympanic line and iris coloration; from L. melanoleucus by ventral coloration, body size, absence of black supratympanic line, skin texture and iris coloration; from L. minimus by ventral coloration, skin texture and male body size; from L. nahangensis by ventral coloration, male body size, skin texture, absence of black supratympanic line and iris coloration; from L. nyx by ventral coloration, skin texture and absence of black supratympanic line, from L pelodytoides by ventral coloration and absence of black supratympanic line; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration, body size, skin texture and absence of black supratympanic line; from L. solus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, absence of black supratympanic line, skin texture and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size, skin texture and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. pallidus sp. nov. differs by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=3.4, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. applebyi). L. pallidus sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=2.7, p=0.006), larger relative tibia length (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum size (Wilcoxon post-hoc Z=2.0, p=0.041), dark brown iris (versus reddish upper and silver below), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. bidoupensis) and tuberculate skin texture (versus mostly smooth in L. bidoupensis). L. pallidus sp. nov. differs from L. melicus by having a significantly larger and non-overlapping male body size (Wilcoxon post-hoc Z=-3.3, pL. applebyi) and tuberculate skin texture (versus mostly smooth in L. melicus). L. pallidus sp. nov. differs from L. pyrrhops by having a significantly smaller and non-overlapping male body size (Wilcoxon post-hoc Z=3.2, p=0.002), smaller relative diameter of the eye (Wilcoxon post-hoc Z=3.2, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. pyrrhops) and tuberculate skin texture (versus only slightly shagreened in L. pyrrhops). L. pallidus sp. nov. differs from L. ardens sp. nov. by having a significantly larger body size (Wilcoxon post-hoc Z=-3.8, pL. ardens sp. nov) and tuberculate skin texture (versus mostly smooth in L. ardens sp. nov). L. pallidus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=3.0, p=0.002), larger relative tympanum diameter (Wilcoxon post-hoc Z=-3.1, p=0.002), indistinct, brown supratympanic line (versus distinct black supratypmanic line in most L. kalonensis sp. nov) and tuberculate skin texture (versus mostly smooth in L. kalonensis sp. nov). L. pallidus sp. nov. differs from L. maculosus sp. nov. by having indistinct, brown supratympanic line (versus distinct black supratypmanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). L. pallidus sp. nov. differs from L. tadungensis sp. nov. by having a significantly larger relative tibia length (Wilcoxon post-hoc Z=-3.1, p=0.002), larger relative distance between the eye and tympanum (Wilcoxon post-hoc Z=-2.4, p=0.019), indistinct, brown supratympanic line (versus distinct black supratympanic line in L. maculosus sp. nov) and tuberculate skin texture (versus mostly smooth in L. maculosus sp. nov). See Table 4 and Fig. 4. The male advertisement call of L. pallidus sp. nov. differs from all members of the L. applebyi group. The call of L. pallidus sp. nov. differs from L. applebyi in call duration, having a relatively long, pulsed introductory note, having one pulse in non-introductory notes, and by average note repetition rate and average dominant frequency. L. pallidus sp. nov. differs from L. bidoupensis in call duration, in having a distinct, pulsed introductory note, and by average note repetition rate and perhaps average dominant frequency (non-overlapping but similar values); from L. melicus in call duration, number of pulses in the introductory note, average note repetition rate and average dominant frequency; from L. pyrrhops in call duration, average number of pulses in introductory note, average note repetition rate and potentially by dominant frequency; from L. ardens sp. nov. differs by call duration, number of pulses in the introductory note, note repetition rate and dominant frequency; from L. kalonensis sp. nov. in average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. in average call duration number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. in average call duration, average duration of introductory note, and note repetition rate. See Table 7 and Figs. 5–6. Leptolalax pallidus sp. nov. differs from all species within the L. applebyi group by>4.7% divergence at the 16S gene fragment examined. Interspecific variation in three L. pallidus sp. nov. collected from

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., Ziegler, Thomas (2016): Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa 4085 (1): 63-102, DOI: http://doi.org/10.11646/zootaxa.4085.1.3

Leptolalax maculosus sp. nov. Figs. 8C, 9C, 13. Holotype. UNS 00513, adult male, in dry stream bed in evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0152º N, 108.7271º E, 900 m elevation, Fig. 1). Collected on 24 September 2011 by Dao T. A. Tran and Duong T. T. Le. Paratypes. ZFMK 96600, adult male from same date and location as holotype. AMS R 177660, adult male, and UNS 00514, adult female, collected on 27 September 2011 from evergreen forest in Phuoc Binh National Park, Ninh Thuan Province, Vietnam (12.0413º N, 108.7385º E, 1166 m elevation). All specimens were collected by Dao T. A. Tran and Duong T. T. Le. Etymology. The specific name “ maculosus ”, meaning spotted, speckled, mottled or variegated, is used as an adjective in reference to the more mottled dorsal pattern of this species. Diagnosis. Assigned to the genus Leptolalax 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 et al. 1998; Delorme et al. 2006). Leptolalax maculosus sp. nov. 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 (24.2–26.6 mm in three adult males, 27.0 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 48–50% of SVL in males; (6) pectoral gland 3.5–4.2% of SVL in males, (7) mostly smooth dorsum, (8) iris copper in upper half and gold in lower half (9) distinct black supratympanic line present, (10) an advertisement call with 7–38 notes, lacking a distinct introductory note. Description of holotype. Head width equal to 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 to eye; canthus rostralis indistinct, gently rounded; lores straight; pupil vertical; eye diameter slightly larger 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 Colour of holotype in life. Dorsal surface dark brown with irregular, darker brown blotch down back; small, pale brown blotches scattered over dorsum; dark interorbital bar with pale copper edging anteriorly; two small, pale copper V-markings between axilla (Fig. 13A, B). Black line along supratympanic ridge, terminating above axilla, encompassing most of tympanum; irregular copper band under black supratympanic line; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms copper; numerous, small darker brown spots on sides from groin to axilla. Brownish pink ventral surface with white speckling on ventral surfaces of throat, chest, belly, 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. Colour of holotype in preservative. Dorsum medium brown with small, pale brown blotches on top of head, barely visible paler intraorbital blotch; dark brown/black banding on dorsal surface of tibiotarsus, antebrachium, hands and feet (Fig. 8C). Black blotch on groin. Ventral surface medium brown with white speckling. Macroglands creamy white. Measurements (mm). Holotype: SVL 24.2, HDL 8.9, HDW 9.0, SNT 3.3, EYE 3.4, IOD 3.5, TMP 1.5, TEY 0.9, TIB 11.6, EN 2.3, IN 2.3, NS 1.1, ML 6.0, PL 10.1. Variation: Measurements of the type series are shown in Tables 5–6 and representative photographs of paratypes are shown in Figs. 9C and 15. AMS R 177660 and to a lesser extent ZFMK 96600 have more distinct patterning, with paler blotches on the dorsum Advertisement call. Call descriptions are based on the calls of two individuals, recorded at 23.3–24.1 ºC ambient temperature. Calls were an average of 907 ms in duration and consisted of 7–38 notes (Table 7, Fig. 6C). Calls were highly amplitude modulated, with a slow rise in amplitude, and amplitude peaking towards the middle of each note. Notes near-invariably contained a single pulse. Notes were repeated an average of 17 notes/s, but note repetition rate was higher at the start of each call. The average dominant frequency was 2.7 kHz, with peak frequency spread over ~2.7–3.5 kHz. A fundamental frequency was not visible. To the human ear, the advertisement call of L. maculosus sp. nov. is a squealch grading gently into a chirp. Ecology. All specimens of Leptolalax maculosus sp. nov. were found in evergreen forest between 900–1166 m elevation at Phuoc Binh National Park. Specimens were found within leaf-litter and in clay holes in a dry stream bed of 1.5–2.0 m width, connecting to a larger stream. Distribution. Leptolalax maculosus sp. nov. is only known from Phuoc Binh National Park, Ninh Thuan Province, Vietnam. The maximum distance between known localities is only 3.2 km. Comparisons. Leptolalax maculosus sp. nov. differs from all other Leptolalax 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 L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration and male body size, from L. botsfordi by male body size and relative body weight (0.41–0.47 versus 0.75–0.94 g /mm in L. botsfordi), from L. croceus by ventral coloration, presence of black supratympanic line and iris coloration; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. maculosus sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly greater body size (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative head width (Wilcoxon post-hoc Z=2.7, p=0.007), greater relative diameter of the eye (Wilcoxon post-hoc Z=2.2, p=0.025), and greater relative pectoral gland size (Wilcoxon post-hoc Z=2.4, p=0.017). L. maculosus sp. nov. differs from L. bidoupensis by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=2.4, p=0.018), and a bronze iris (versus reddish upper and silver below in L. bidoupensis). L. maculosus sp. nov. differs from L. melicus by having a significantly greater body size (Wilcoxon post-hoc Z=2.6, p=0.009), and greater length between eye and tympanum (Wilcoxon post-hoc Z=2.1, p=0.034). L. maculosus sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z= -2.6, p=0.010), smaller relative length between eye and tympanum (Wilcoxon post-hoc Z= -2.4, p=0.018) and smaller relative diameter of the eye (Wilcoxon posthoc Z=-2.4, p=0.018). L. maculosus sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. maculosus sp. nov. differs from L. pallidus sp. nov. by having a black supratympanic line (versus no black supratypmanic line in L. pallidus sp. nov) and smooth- finely shagreened skin texture (versus tuberculate in L. pallidus sp. nov). L. maculosus sp. nov. differs from L. kalonensis sp. nov. by having a significantly smaller body size (Wilcoxon post-hoc Z=-2.7, p=0.007). L. maculosus sp. nov. differs from L. tadungensis sp. nov. by having a significantly greater relative tibia length (Wilcoxon post-hoc Z=-2.5, p=0.013). See Table 4 and Fig. 5. The male advertisement call of L. maculosus sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in call duration, number of notes per call, an introductory note, and dominant frequency; from L. bidoupensis in call duration, average number of notes per call, presence of an introductory note, and average dominant frequency; from L. melicus in call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. pyrrhops by call duration, number of notes per call, having an indistinct introductory note, and dominant frequency; from L. ardens sp. nov. by call duration, number of notes per call and dominant frequency; from L. pallidus sp. nov. by call duration and number of notes per call; from L. kalonensis sp. nov. by average call duration, and number of notes per call; from L. tadungensis sp. nov. by call duration, number of notes per call, having a less distinct introductory note and different relative duration of introductory note. See Table 7 and Figs. 5–6. Leptolalax maculosus sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. maculosus sp. nov. collected from up to ~ 3 km apart was 0.0–0.2%.

Leptolalax tadungensis sp. nov. Figs. 8D, 14. Holotype. 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. Paratypes. 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. Etymology. Specific epithet “ tadungensis ” is in reference to the type locality of the species, Ta Dung Nature Reserve. Diagnosis. Assigned to the genus Leptolalax 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 et al. 1998; Delorme et al. 2006). Leptolalax tadungensis sp. nov. 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. Description of holotype. 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 Colour of holotype in life. 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. Colour of holotype in preservative. 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. Measurements (mm). 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. Variation. 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). Advertisement call. 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 L. pallidus sp. nov. is a short raspy chirp, similar to an orthopteran. Ecology. All specimens of Leptolalax tadungensis sp. nov. 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. Distribution. Leptolalax tadungensis sp. nov. 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. Comparisons. Leptolalax tadungensis sp. nov. differs from all other Leptolalax 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 L. aereus by ventral coloration and presence of distinct black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by male body size, from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line and lateral fringes on toes; from L. firthi by ventral coloration, iris coloration, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and iris coloration; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration; from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, presence of black supratympanic line and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. tadungensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi 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). L. tadungensis sp. nov. differs from L. bidoupensis 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 L. bidoupensis). L. tadungensis sp. nov. differs from L. melicus 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). L. tadungensis sp. nov. differs from L. pyrrhops 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). L. tadungensis sp. nov. differs from L. ardens sp. nov. 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). L. tadungensis sp. nov. differs from L. pallidus sp. nov. 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 L. pallidus sp. nov) and mostly smooth skin texture (versus tuberculate in L. pallidus sp. nov). L. tadungensis sp. nov. differs from L. kalonensis sp. nov. 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). L. tadungensis sp. nov. differs from L. maculosus sp. nov. 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 L. tadungensis sp. nov. differs from all species in the L. applebyi group. The call of the new species differs from L. applebyi in dominant frequency; from L. bidoupensis in having an introductory note present and in dominant frequency; from L. melicus in relative length of introductory note and dominant frequency; from L. pyrrhops by dominant frequency; from L. ardens sp. nov. by relative length of introductory note; from from L. pallidus sp. nov. by call duration and relative length of introductory note; from L. kalonensis sp. nov. by relative length of introductory note; from L. maculosus sp. nov. 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. Leptolalax tadungensis sp. nov. differs from all species within the L. applebyi group by>6.3% divergence at the 16S gene fragment examined. Interspecific variation in four L. tadungensis sp. nov. collected from up to ~ 15 km apart was 0.0–0.2%.

Leptolalax ardens sp. nov. Figs. 7D, 9A, 10. Holotype. VNMN 04707/AMS R 176454, adult male, calling on clay bank 1.5 m from headwaters of 1���2m wide rocky, medium gradient, high���flow stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2258�� N, 108.3230�� E; 1162 m elevation, Fig. 1, Fig 9E). Collected at 19:30 h on 21 August 2011 Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Paratypes. AMS R 176452���176453, two adult males (both calling), and VNMN 04708/AMS R 176455��� VNMN 04710/AMS R 176457, three adult males (not observed calling), collected on same date and locality as holotype. VNMN 04711 /AMS R 176458, adult female (gravid) and VNMN 04712 /AMS R 176459 adult male (calling), collected at 19:55 h on 23 August 2011 in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2263�� N, 108.3345�� E; 1450 m elevation). UNS 00530/AMS R 176460���UNS 00532/ 176462, AMS R 176463, AMS R 176464, UNS 00533/176465, UNS 00534 /AMS R 176466, 176467, and NCSM 79650, 9 adult males, collected at night on 24 August 2011 in swampy area with small rocky stream in evergreen forest in Kon Ka Kinh National Park, Gia Lai Province, Vietnam (14.2197 �� N, 108.3207 �� E; 1041 m elevation). All specimens were collected by Jodi J. L. Rowley, Vinh Quang Dau, Huy Duc Hoang, Luong Thi Nguyen, Tan Thanh Le, Dinh Djung, and Y H'Jun. Etymology. The specific name ��� ardens ���, meaning burning, fiery, shining, or brilliant, is used as an adjective in reference to the bold and conspicuous coloration of this species compared to other species in the group. Diagnosis. Assigned to the genus Leptolalax 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 et al. 1998; Delorme et al. 2006). Leptolalax ardens sp. nov. 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 SVL for the genus (21.6.1��� 24.7 mm in 15 adult males, 24.5 mm in one female); (4) toes lacking webbing and lateral fringes; (5) tibia length 44���49% of SVL in males; (6) pectoral gland 3.5���5.6% of SVL in males, (7) mostly smooth dorsum, (8) iris dark coppery brown, (9) distinct black supratympanic line present, (10) an advertisement call with 3���10 notes with the first, introductory note encompassing one���third of each call. Description of holotype. Head width equal to head length; snout slightly truncate in dorsal view and rounded in profile, projecting slightly beyond margin of the lower jaw; nostril closer to snout than to eye; canthus rostralis indistinct, gently rounded; lores sloping; pupil vertical; eye diameter smaller than snout length; tympanum distinct, round, diameter smaller than that of the eye; tympanic rim 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 Colour of holotype in life. Dorsal surface brown with irregular, darker brown blotch down back; dark interorbital bar with pale cream edging anteriorly; two small, pale markings between axilla. Blackish brown line along canthus rostralis through eye, and continuing below supratympanic ridge, terminating above axilla, encompassing nare and most of tympanum; transverse dark brown bars on dorsal surface of thighs, tibia, tarsus, lower arms, fingers and toes; elbows and upper arms orange; large, black blotch on posterior flank and numerous, smaller black spots on sides from groin to axilla (Fig. 10A,C). Upper arms pale coppery orange. Dark brownish ventral surface with white speckling on entire ventral surface including throat, arms and legs; ventral margin of throat bearing broken row of slightly larger white spots. Supra-axillary gland copper; femoral glands white; pectoral glands white, ventrolateral glands white. Iris brown; minute, black reticulations throughout. Colour of holotype in preservative. Dorsum relatively uniformly dark brown; dorsal surface of brachium, fingers and toes paler brown (Fig. 7D). Ventral surface of chest and thighs slightly medium brown with white speckles; throat dark brown speckled with white; and belly white speckled with brown. Macroglands creamy white. Measurements (mm). Holotype: SVL 22.0, HDL 9.0, HDW 8.9, SNT 3.4, EYE 2.9, IOD 2.4, TMP 1.6, TEY 0.8, TIB 10.5, EN 2.0, IN 2.5, NS 1.3, ML 5.7, PL 10.8, weight in life 1.1g Variation. Measurements of the type series are shown in Tables 2���3 and representative photographs of paratypes are shown in Figure 10. Some specimens (VNMN 04711/AMS R 176458��� VNMN 04712/1764589) have more clumped white speckling on the venter. UNS 00531/AMS R 176461 has a paler brown venter with barely distinct brown mottling. AMS R 176453, VNMN 04709/AMS R 176456, VNMN 04710/AMS R 176457, UNS 00532/AMS R 176462, AMS R 176463, UNS 00533/AMS R 176465, UNS 00534/AMS R 176466, have a distinctly pale line anterior to dark brown interorbital bar. The degree of black dorsolateral spots/flecks varies, pale edged marbling also visible on dorsum of some specimens, especially AMS R 176457, UNS 00532/AMS R 176462, UNS 00533/AMS R 176465. Advertisement call. Call descriptions are based on the calls of seven individuals, recorded at 21.4���24.7 ��C ambient temperature. Calls were an average of 239 ms in duration and consisted of 3���10 notes (Table 4, Fig. 5D). Calls were highly amplitude modulated, with amplitude peaking towards the end of each call. Calls contained 3���10 notes and introductory notes were present in most calls. Introductory notes were 25���41% of the call duration, contained up to 54 pulses. Non-introductory notes contained 1���3 pulses. Notes were repeated an average of 20 notes/s. The average dominant frequency was 3.3 kHz, spread more widely (~2.0���7.6 kHz) in non-introductory notes. Harmonics were weakly present at ~ 6, 9, and 12 kHz, and a fundamental frequency was not visible. To the human ear, the advertisement call of L. ardens sp. nov. is a weak squeak followed by a rasping, similar to an orthopteran. Ecology. All specimens of Leptolalax ardens sp. nov. were found in evergreen forest between 1,041���1,450 m elevation in Kon Ka Kinh National Park. Males were heard faintly calling on the forest floor, adjacent to small streams or seeps (Figure 9D). Distribution. Leptolalax ardens sp. nov. is only known from Kon Ka Kinh National Park, in Gia Lai Province, Vietnam. Comparisons. Leptolalax ardens sp. nov. differs from all other Leptolalax 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 L. aereus by ventral coloration, body size and the presence of a black supratympanic line, from L. bourreti by ventral coloration and body size, from L. botsfordi by male body size and relative body weight (0.41���0.47 versus 0.75���0.94 g /mm), from L. croceus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. eos by ventral coloration, male body size, iris coloration and lateral fringes on toes; from L. firthi by ventral coloration, male body size, presence of black supratympanic line, iris coloration, male skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration, male body size and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size, and iris coloration; from L. melanoleucus by ventral coloration, male body size, and iris coloration; from L. minimus by ventral coloration and male body size; from L. nahangensis by ventral coloration, male body size and iris coloration; from L. nyx by ventral coloration and male body size, from L pelodytoides by ventral coloration and male body size; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration; from L. solus by ventral coloration, male body size and iris coloration; from L. sungi by ventral coloration, male body size, presence of black supratympanic line, iris coloration and skin texture; from L. tuberosus by ventral coloration, presence of black supratympanic line, iris coloration and skin texture; from L. ventripunctatus by ventral coloration, male body size, iris coloration and skin texture; and from L. zhangyangpingi by ventral coloration, male body size, presence of black supratympanic line, and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. ardens sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=3.7, pL. ardens sp. nov. differs from L. bidoupensis by having a smaller SVL (Wilcoxon post-hoc Z= -2.0, p=0.048), larger relative tibia length (Wilcoxon post-hoc Z=3.2, p=0.001), larger relative eye diameter (Wilcoxon post-hoc Z=2.1, p=0.038) and having a relatively uniform iris coloration (versus reddish in upper and silvery gold below). L. ardens sp. nov. differs from L. melicus by having a larger SVL (Wilcoxon post-hoc Z= -3.3, pL. ardens sp. nov. differs from L. pyrrhops, by having a smaller and non-overlapping body size (Wilcoxon post-hoc Z= 3.7, pL. ardens sp. nov. differs from L. pallidus sp. nov. by having a smaller SVL (Wilcoxon post-hoc Z= -3.8, pL. ardens sp. nov. differs from L. kalonensis sp. nov. by having a non-overlapping, smaller body size (Wilcoxon post-hoc Z= 4.7, pL. ardens sp. nov. differs from L. maculosus sp. nov. by having a smaller body size (Wilcoxon post-hoc Z=2.8, p=0.006). L. ardens sp. nov. differs from L. tadungensis sp. nov. by having a smaller body size (Wilcoxon post-hoc Z= 3.4, pL. ardens sp. nov. differs from all members of the L. applebyi group with the exception of L. kalonensis sp. nov. From L. applebyi the call of the new species differs by having a longer introductory note and note repetition rate. From L. bidoupensis, the call of L. ardens sp. nov. differs by having an introductory note in most calls and having a higher dominant frequency. From L. melicus, the call of L. ardens sp. nov. differs by having a lower dominant frequency. From L. pyrrhops the call of L. ardens sp. nov. appears to differ by having a higher dominant frequency. From L. pallidus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration,a higher average number of pulses in the introductory note, a higher note repetition rate and a higher dominant frequency. From L. maculosus sp. nov. the call of L. ardens sp. nov. differs by having a shorter call duration, a higher number of notes per call, a distinct introductory note, and higher dominant frequency. From L. tadungensis sp. nov. the call of L. ardens sp. nov. differs by having a relatively long introductory note. No clear differences between the call of L. ardens sp. nov. and L. kalonensis sp. nov. could be determined, perhaps in part due to the limited sample size (n=1) for L. kalonensis sp. nov. See Table 7 and Figs. 5���6. Leptolalax ardens sp. nov. differs from all species within the L. applebyi group by>4.5% divergence at the 16S gene fragment examined. Interspecific variation in three L. ardens sp. nov. collected from ~ 2 km apart was 0%. 1 Leptolalax applebyi $. % $: $; L. applebyi L. L. melicus L. pyrrhops L. ardens L. pallidus L. kalonensis L. maculosus L. tadungensis bidoupensis : $;: $ +;: $ (;: $ ��;: $ 3;). (�� +. 3 (/)* 3)/ 3* ;: * K.).;:)��K.));: *(K ��;:)��K 3/;:*..K /;:(/K/ 3;:����3K3)/;:.��K +; ; * + *3 3 (+ (*�� 3 (. / *�� :*3)K).;: (*K (;: *.K. /;:*.)K *.;: K. *;:* 3K; * / *. ! Leptolalax 0 2 !> 1 2!:; 9 5 C H $ aereus H 0 0 $ $ + *K �� 3 6 A F bourreti H ��)K (B> " $ 2 K $ 0 $ botsfordi & 0 0 0 $ 3 *K (B 6 0 $ 2 $ croceus $ $ K / " 0 ��� $ eos > 0 0 0 $ *K. / 7 $ " $ $ 2 $ B $ 0 firthi > 0 (.K 3 $ $ 2 $ " B fuliginosis �� K)) B & $ " " 0 0 $ 0 heteropus 0: B = 2. (K (3; kecil 0 0 *3 K) + B 2 $ :; melanoleucus $ $ 0 $ ((K �� �� B 7 $ " " 0 0 0 minimus H + (K *. B = 2 nahangensis H 0 0 $ $.) �� B 6 $ " 0 0 nyx H 0 0 (/K (B = " 0 0 pelodytoides H " 0 / += B = " 0 0 platycephalus $ C + * B P 7 $ = $ 0 pluvialis 0 0 0 $ * K B $ 2 solus H 0 0 / (B " H $ 0 0 ������continued on the next page !:>; !> 1 2!:; 9 5 C H $ sungi H " $ $.�� K+ / $ 2 $ tuberosus H $ 0..K 3 + ��� $ $ ventripunctatus > 0 + +K ��) B> " $ $ $ 0 0 zhangyangpingi H 0 0 $ $.+ ��K+ + " $ " 0 0 0:; applebyi & 0 0 0 $ *3 (K:) ��; B 2 bidoupensis & 0 0 0 $ *�� +K +.: (; B & " 2 0 melicus & 0 0 0 $ *3 +K /:) /; B 2 pyrrhops & 0 0 0 $) ��K.:; B 7 $ 2 $ $ ardens & 0 0 0 $ * K. /: ��; B 0 2 K $ pallidus & 0 0 0 $. +K / /: + (; 0 5 kalonensis % " 0 + ��K) (: /.; B " 0 A 2 maculosus 0 " 0 $. K ((: + +; B A 2 K $ tadungensis & 0 0 0 $ K ��: +); B 2, Published as part of 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. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 75-86, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"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. & 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"]}

Leptolalax kalonensis sp. nov. Figs. 8B, 9B, 11. Holotype. 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 (11.2748�� N, 108.2109�� 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. Paratypes. 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 (11.2748�� N, 108.2109�� 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 (11.2919�� N, 108.2150�� 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. Etymology. Specific epithet ��� kalonensis ��� is in reference to the type locality of the species, which is in close proximity to the village previously known as Kalon. Diagnosis. Assigned to the genus Leptolalax 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 et al. 1998; Delorme et al. 2006). Leptolalax kalonensis sp. nov. 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. Description of holotype. 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 Colour of holotype in life. 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. Colour of holotype in preservative. 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. Measurements (mm). 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. Variation. 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. Advertisement call. Call descriptions are based on the call of AMNH A191770, recorded at 26.4 ��C 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 L. kalonensis sp. nov. is a short rasp, similar to an orthopteran. Ecology. All specimens of Leptolalax kalonensis sp. nov. 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. Distribution. Leptolalax kalonensis sp. nov. 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 L. applebyi group. Comparisons. Leptolalax kalonensis sp. nov. differs from all other Leptolalax 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 L. aereus by ventral coloration and presence of black supratympanic line, from L. bourreti by ventral coloration, from L. botsfordi by ventral coloration, from L. croceus by ventral coloration, presence of black supratympanic line, and skin texture; from L. eos by ventral coloration, male body size, presence of black supratympanic line, and lateral fringes on toes; from L. firthi by ventral coloration, presence of black supratympanic line, skin texture and lateral fringes on toes; from L. fuliginosus by ventral coloration and iris coloration; from L. heteropus by ventral coloration and male body size; from L. kecil by ventral coloration, male body size and iris coloration; from L. melanoleucus by ventral coloration and iris coloration; from L. minimus by ventral coloration; from L. nahangensis by ventral coloration and male body size; from L. nyx by ventral coloration, from L pelodytoides by ventral coloration; from L. platycephalus by ventral coloration and male body size; from L. pluvialis by ventral coloration and male body size; from L. solus by ventral coloration and iris coloration; from L. sungi by ventral coloration, male body size and iris coloration; from L. tuberosus by ventral coloration, iris coloration and skin texture; from L. ventripunctatus by ventral coloration and iris coloration; and from L. zhangyangpingi by ventral coloration, male body size and iris coloration. See Table 8 for details. From members of the L. applebyi group, L. kalonensis sp. nov. differs from all species by at least one morphological character (all following morphometric differences refer to males only). The new species differs from L. applebyi by having a significantly larger body size (Wilcoxon post-hoc Z=4.0, pL. kalonensis sp. nov. differs from L. bidoupensis by having a significantly larger body size (Wilcoxon post-hoc Z=4.4, pL. kalonensis sp. nov. differs from L. melicus by having a significantly larger body size (Wilcoxon post-hoc Z= -3.9, pL. kalonensis sp. nov. differs from L. pyrrhops by having a significantly smaller body size (Wilcoxon post-hoc Z=3.7, pL. kalonensis sp. nov. differs from L. ardens sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z=4.7, pL. kalonensis sp. nov. differs from L. pallidus sp. nov. 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 L. pallidus sp. nov) and smooth skin texture (versus tuberculate in L. kalonensis sp. nov). L. kalonensis sp. nov. differs from L. maculosus sp. nov. by having a significantly greater body size (Wilcoxon post-hoc Z= -2.7, p=0.007). L. kalonensis sp. nov. differs from L. tadungensis sp. nov. 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 L. kalonensis sp. nov. differs from all members of the L. applebyi group with the exception of L. ardens sp. nov. In particular, the species differs from L. applebyi in length of introductory note and dominant frequency; from L. bidoupensis in call duration, presence of a distinct introductory note and dominant frequency (although similar); from L. melicus in dominant frequency; from L. pyrrhops by dominant frequency; from L. pallidus sp. nov. by average call duration, average number of pulses in the introductory note and average note repetition rate; from L. maculosus sp. nov. by average call duration, number of notes per call, and having a distinct introductory note; from L. tadungensis sp. nov. by average call duration, and average duration of introductory note. See Table 7 and Figs. 5���6. Leptolalax kalonensis sp. nov. differs from all species within the L. applebyi group by>5.2% divergence at the 16S gene fragment examined. Interspecific variation in four L. kalonensis sp. nov. collected from up to 43 km apart was 0.0���0.6%., Published as part of 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. & Ziegler, Thomas, 2016, Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam, pp. 63-102 in Zootaxa 4085 (1) on pages 90-93, DOI: 10.11646/zootaxa.4085.1.3, http://zenodo.org/record/1052459, {"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. & 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"]}