Limnonectes quangninhensis Pham, Le, Nguyen, Ziegler, Wu & Nguyen, 2017, sp. nov.
Creators: name: Pham, Cuong The
name: Le, Minh Duc
name: Nguyen, Tao Thien
name: Ziegler, Thomas
name: Wu, Zheng Jun
name: Nguyen, Truong Quang
Publisher: name: Zenodo
Year: 2017
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Dicroglossidae
Limnonectes
Limnonectes quangninhensis

description: Limnonectes quangninhensis sp. nov.
(Figs. 2–5)
Holotype: IEBR 3907 (Field number QN 2015.7), adult male, collected by C. T. Pham and H. N. Ngo on 26 May 2015 in the evergreen forest near Tai Chi Village (21o31.785’N, 107o38.965’E, at an elevation of 195 m), Quang Son Commune, Hai Ha District, Quang Ninh Province, Vietnam.
Paratypes: Two adult females, VNMN A.2016.17, A.2016.18 (Field numbers TAO 936, 937), collected by T. T. Nguyen and C. T. Pham on 21 September 2009 (21o06.498’N, 107o34.708’E, at an elevation of 15 m) from the evergreen forest of Cai Lim (or Ba Mun) Island, Bai Tu Long National Park, Quang Ninh Province. One adult male, ZFMK 95213, and one adult female, ZFMK 95210, collected by C. T. Pham, A. Gawor, and A. Dogra on 20 May 2011 (21o04.333’N, 107o35.297’E, at an elevation of 5 m) from Cai Lim Island, Bai Tu Long National Park, Quang Ninh Province. Eight specimens collected from the evergreen forest of Quang Son Commune, Hai Ha District, Quang Ninh Province, Vietnam: Two adult females, IEBR 3908, 3909 (Field numbers QN 2015.8, 2015.9), collected by C. T. Pham and H. N. Ngo on 26 May 2015 (21o31.785’N, 107o38.965’E, at an elevation of 195 m); one adult male, IEBR 3913 (Field number QN 2016. 36), and three females, IEBR 3910, 3911, 3912 (Field numbers QN 2016.24, 2016.25, 2016.37); one adult male, IEBR 3914 (Field number QN 2016.44), collected by C. T. Pham and T. V. Nguyen on 6 May 2016 (21o30.751’N, 107o38.550’E, at an elevation of 119 m); one adult male, IEBR 3915 (Field number QN 2016.60), collected by C. T. Pham and T. V. Nguyen on 7 May 2016 (21o32.296’N, 107o39.376’E, at an elevation of 229 m).
Diagnosis. The new species from Quang Ninh Province is assigned to the genus Limnonectes on the basis of the following characters: the presence of fanglike odontoid processes on the lower jaw and sexual dimorphism (with males larger than females and hypertrophy of the head in males) (Emerson et al. 2000). The new species is also strongly supported as a member of Limnonectes based on molecular analyses. Limnonectes quangninhensis sp. nov. is distinguishable from its congeners by a combination of the following morphological characters: (1) size large (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in femsles); (2) males with moderately enlarged head (HL/SVL 0.48); (3) head longer than wide; (4) vomerine teeth present; (5) external vocal sacs absent; (6) rostral length short (RL/SVL 0.16 in males, 0.15 in females); (7) tympanum distinct (TD/ED 0.63 in males, 0.60 in females); (8) dorsal surface of head, body and flanks with flattened tubercles; (9) dorsal surface of tibia with small tubercles; (10) supratympanic fold present; (11) dorsolateral fold absent; (12) webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V; (13) in life, dorsum yellowish brown with a dark brown marking; (14) throat and chest white with dark brown marking, ventral surface of fore and hind limbs, as well as belly white.
Description of holotype. Adult male; SVL 55.9 mm; habitus robust with enlarged head (HL/SVL 0.47, HW/ SVL 0.42); head longer than wide (HL 26.5 mm, HW 23.6 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to the snout tip than to eye (NS 4.8 mm, EN 5.1 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 9.5 mm, ED 7.3 mm); internarial distance slightly wider than interorbital distance and upper eyelid (IND 4.8 mm, IOD 4.4 mm, UEW 3.8 mm); tympanum distinct, round, 66% of eye diameter (TD 4.8 mm); vomerine teeth in two oblique ridges; tongue cordiform, deeply notched posteriorly; lower jaw with two tooth-like; external vocal sac absent.
Forelimbs: Arms short; forelimb length (FLL) 10.6 mm, hand length (HAL) 24.5 mm; relative finger lengths: II<I<IV<III; fingers free of webbing; tips of fingers blunt, not expanded; subarticular tubercles prominent, round, formula 1, 1, 2, 2; inner metatarsal tubercle oval, elongate; outer metatarsal tubercle small; finger I with nuptial pad, composed of minute spines.
Hindlimbs: Tibia and thigh short (FeL 27.8 mm, TbL 28.2 mm), approximately three times longer than wide (TbW 8.5 mm); tips of toes blunt, slightly rounded; relative length of toes: I<II<V<III<IV; webbing well developed, formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V; subarticular tubercles prominent, elongate, formula 1, 1, 2, 3, 2; inner metatarsal tubercle elongate; outer metatarsal tubercle absent; tibio-tarsal articulation reaching between eye and tip of snout.
Skin texture in life: Dorsal surface of head, body and flanks with flattened tubercles; small tubercles on upper eyelid; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; dorsal surface of limbs and thighs smooth; dorsal surface of tibia with fine tubercles; throat, chest, belly and ventral surface of thighs smooth.
Coloration in life: Iris black; head with a narrow light brown bar in interorbital region; dorsum yellowishbrown with a light yellow vertebral stripe and a dark brown marking; upper part of flanks yellowish brown, lower part of flanks yellowish white; lips with dark bars; tympanum dark brown; dorsal surface of fore and hind limbs yellowish brown with dark crossbars; throat and chest white with dark brown marking; ventral surface of fore and hind limbs, as well as belly white; toe webbing brown.
Coloration in preservative: Dorsum greyish brown with a light vertebral stripe and a dark brown marking; upper part of flanks light brown, lower part cream; lips with dark bars; tympanum dark brown; dorsal surface of fore and hind limbs light brown with dark crossbars; throat and chest white with dark brown marking; ventral surface of fore and hind limbs, as well as belly white; toe webbing brown.
Variation and sexual dimorphism. Measurements and morphological characters of the type series are provided in Table 3. Two specimens (IEBR 3908, 3914) have a yellowish vertebral stripe on the dorsum. The male specimens have nuptial pads on finger I. The males also have a larger head than the females (HL/SVL 0.48 vs. 0.42, HW/SVL 0.43 vs. 0.40, respectively).
TABLE ³. Measurements (in mm) and proportions of the type series of Limnonectes quangninhensis sp. nov. (H = holotype, P = paratype, SD = standard deviation, for other
abbreviations see Material and methods).
Specimen IEBR IEBR IEBR IEBR ZFMK Mean±SD IEBR IEBR IEBR IEBR IEBR VNMN VNMN ZFMK Mean±SD 3907 3913 3914 3915 95213 (n=5) 3908 3909 3910 3911 3912 A.2016.17 A.2016.18 95210 (n=8) Ƌ Ƌ Ƌ Ƌ Ƌ ♀ ♀ ♀ ♀ ♀ ♀ ♀ ♀
status H P P P P P P P P P P P P
55.9 67.6 50.1 62.2 68.9 60.9±7.9 56.9 57.4 62.9 58.1 63.0 49.3 45.5 62.6 57.0±6.5 26.5 33.2 23.3 31.1 34.4 29.7±4.7 24.6 24.8 25.8 24.2 24.3 20.5 19.0 26.2 23.7±2.5 23.6 30.3 20.1 29.5 32.7 27.4±4.9 23.4 22.9 23.4 23.1 23.0 20.2 18.4 25.3 22.5±2.0 23.1 29.8 20.7 28.1 30.4 26.4±4.3 21.4 21.9 22.3 21.5 20.4 18.0 16.6 22.3 20.3±2.2 18.8 23.9 16.7 23.4 24.9 21.5±3.6 17.6 18.0 17.6 17.5 16.1 14.3 13.0 17.1 16.3±1.9
MBE 12.0 17.2 9.9 16.3 16.0 14.3±3.1 10.4 10.5 10.7 10.0 8.9 8.5 7.4 9.8 9.5±1.2 9.5 11.5 8.2 10.1 11.4 10.1±1.4 9.2 9.3 9.4 9.2 9.7 7.6 6.0 9.4 82±1.3 7.3 8.7 6.8 7.9 9.4 8.2±1.1 8.3 8.7 8.4 7.9 7.8 6.1 5.9 8.8 7.6±1.1
UEW 3.8 4.4 3.2 4.0 4,5 4.0±0.5 3.4 3.6 4.0 4.0 4.6 3.7 3.7 4.8 3.9±0.4 4.8 5.7 4.4 5.6 5.8 5.3±0.6 4.4 4.4 4.7 4.2 4.7 3.9 3.9 4.9 4.4±0.4 4.4 5.5 3.9 5.4 5.6 5.0±0.7 4.7 4.9 5.0 4.5 5.1 4.2 4.1 5.4 4.7±0.4 9.6 12.1 8.3 10.8 13.6 10.9±2.1 9.6 9.4 10.3 9.1 9.8 8.9 8.8 10.4 9.5±0.6 15.5 17.6 13.8 17.0 18.9 16.6±1.9 15.7 14.9 16.7 14.7 16.6 12.6 11.0 15.5 14.7±2.0 4.8 5.2 4.4 4.9 5.3 4.9±0.4 3.8 3.7 4.6 4.4 5.0 3.3 3.2 4.8 4.1±0.7 5.1 5.9 4.6 5.6 6.2 5.5±0.7 4.2 4.2 4.9 5.2 5.1 4.0 3.9 5.1 4.6±0.5 4.8 5.5 4.0 5.3 5.7 5.1±0.7 5.0 5.0 4.8 4.5 4.7 4.1 4.3 4.4 4.6±0.3 3.3 5.0 2.6 4.7 5.5 4.2±1.2 2.2 2.3 2.5 2.1 2.4 2.4 2.2 2.6 2.3±0.2 10.6 13.4 8.6 11.2 12.9 11.3±1.9 10.4 9.0 9.8 9.7 10.9 9.0 8.5 11.4 9.6±0.6 24.5 28.4 22.6 27.8 30.2 26.7±3.1 25.7 25.5 25.6 24.8 26.2 21.7 20.4 27.0 24.6±2.3 3.0 2.9 3.0 3.0 3.6 3.2±0.4 2.9 3.3 2.6 2.6 2.8 2.5 2.2 3.2 2.7±0.4 27.8 31.5 25.3 30.0 30.3 28.9±2.5 27.8 27.2 27.9 27.7 29.0 23.8 21.3 29.3 26.7±2.7 Etymology. The specific epithet “ quangninhensis ” refers to the type locality of the new species, Quang Ninh Province. For the common names we suggest Quangninh Wart Frog (English) and Ếch nhẽo quảng ninh (Vietnamese).
Ecological notes. Specimens were found at night between 19:00 and 23:00 h in small rocky streams (Fig. 6).
Most of specimens were found in the water. The surrounding habitat was secondary evergreen forest of medium and small hardwoods mixed with bamboo, shrubs and vines. Air temperature was 25.4–29.1o C and relative humidity 70–82%. Advertisement calls and tadpoles of the species are not known. The females collected in May 2011 and in May 2016 contained yellow eggs suggesting that the breeding season of this species is in summer. Other amphibian species found at the sites included: in the mainland: Amolops ricketii (Boulenger), Hylarana maosonensis Bourret, and Rhacophorus dennysi Blanford and on Cai Lim Island: Microhyla heymonsi Vogt, Occidozyga martensii (Peters), Hylarana guentheri (Boulenger), Hylarana sp., Polypedates mutus (Smith), and Theloderma albopunctatum (Liu & Hu) (see Gawor et al. 2016).
Distribution. Limnonectes quangninhensis sp. nov. is currently known only from Quang Ninh Province, northeastern Vietnam, where it can be found both on the mainland and on islands in the Gulf of Tonkin (Fig. 7). The record of the new species on Cai Lim Island is approximately 50 km far from the type locality in Hai Ha District.
Comparisons. We compared the new species with its congeners from Vietnam and neighbouring countries in mainland Indochina including Laos, Cambodia, Thailand and China.
Limnonectes quangninhensis sp. nov. differs from L. fujianensis Ye & Fei by having a slightly larger size (average SVL 60.9 mm in males, n=5 and 57.0 mm in females, n=8 vs. 53.9 mm in males, n=20 and 47.5 mm in females, n=20), distinct tympana (vs. indistinct in L. fujianensis), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. fujianensis), and males with nuptial pad on finger I only (vs. males with nuptial pad on fingers I and II in L. fujianensis); from L. namiyei Stejneger by having a smaller size (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in females vs. 83.4 mm in males, 75.1 mm in females of L. namiyei), distinct tympana (vs. indistinct in L. namiyei), different webbing between toes (webbing formula I0– 0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. namiyei), and males without vocal slits (present in L. namiyei); from L. bannaensis Ye, Fei & Jiang by having distinct tympana (vs. indistinct in L. bannaensis), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. bannaensis), different digit tips of toes (tips of toes blunt, slightly round vs. tips of toes expanded into discs, round), and dorsal skin with tubercles (vs. smooth in L. bannaensis); from L. dabanus (Smith) by having a smaller ratio of TD/ED (0.63 in males, 0.60 in females vs. 0.98 in males, 1.0 in females of L. dabanus) and the absence of caruncle on head (vs. present in L. dabanus); from L. fragilis (Liu & Hu) by having distinct tympana (vs. indistinct in L. fragilis), different size of subarticular tubercles (small vs. large in L. fragilis), and males with nuptial pads on finger I (vs. nuptial pads absent in males of L. fragilis); from L. gyldenstolpei (Andersson) by having a smaller ratio of TD/ED (0.63 in males vs. 1.0 in males of L. gyldenstolpei) and the absence of caruncle on head in males (vs. present in L. gyldenstolpei); from L. hascheanus (Stoliczka) by having a larger size (SVL 50.1–68.9 mm in males, 45.5–63.0mm in females vs. 18.8–25.4 mm in males, 20.5–25.0 mm in females of L. hascheanus) and the absence of dorsolateral folds (vs. present in L. hascheanus); from L. isanensis McLeod, Kelly & Barley by having distinct tympana (vs. indistinct in L. isanensis), dorsal skin with tubercles (vs. feebly crenulated in L. isanensis), and different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. isanensis); from L. jarujini Matsui, Panha, Khonsue & Kuraishi by having a smaller size in males (SVL 50.1–68.9 mm vs. 72.9 in L. jarujini), distinct tympana (vs. indistinct in L. jarujini), length of fingers II<I (vs. fingers II=I in L. jarujini), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. jarujini), and dorsal skin with tubercles (vs. smooth in L. jarujini); from L. khammonensis Smith by having a larger size (SVL 45.5– 63.0 mm in females vs. 37 mm in female in L. khammonensis) and distinct tympana (vs. indistinct in L. khammonensis); from L. kohchangae Smith by having a larger size (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in females vs. 31.0– 42.2 mm in males, 32.2–41.5 mm in females of L. kohchangae), head longer than wide (vs. broader than long in L. kohchangae), and different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. kohchangae); from L. limborgi (Sclater) by having a larger size (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in females vs. 29.8–37.6 mm in males, 28.0– 36.2 mm in females of L. limborgi) and the absence of dorsolateral folds (vs. present in L. limborgi); from L. liui (Yang) by having a larger size (SVL 50.1– 68.9 mm in males, 45.5–63.0 mm in females vs. 32–38.5 in males, 32.7 in female of L. liui) and different color on belly (white vs. pale yellow); from L. longchuanensis by having smaller size in males (SVL 50.1–68.9 mm vs. 75.5 in L. longchuanensis), distinct tympana (vs. indistinct in L. longchuanensis), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. longchuanensis), and dorsal surface of legs with small tubercles (vs. with large tubercles on dorsal surface of legs in L. longchuanensis); from L. lauhachindai by having a larger size (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in females vs. 30.5–42.0 mm in males, 32.9– 37.9 mm in females in L. lauhachindai), the absence of caruncle on head (vs. present in L. lauhachindai), a smaller ratio of TD/ED (0.63 in males, 0.60 in females vs. 0.82 in L. lauhachindai), and males with nuptial pad on finger I (vs. absent in L. lauhachindai); from L. macrognathus (Boulenger) by having larger size (SVL 50.1–68.9 mm in males vs. 43.2–44.8 mm in males of L. macrognathus) and the absence of caruncle on head (vs. present in L. macrognathus); from L. megastomias McLeod by having distinct tympana (vs. indistinct in L. megastomias), males with nuptial pad on finger I (vs. males with nuptial pad on fingers I and II in L. megastomias), different webbing between toes (I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. megastomias), and dorsal skin with tubercles (vs. smooth in L. megastomias); from L. nguyenorum by having a larger size (SVL 50.1–68.9 mm in male, 51.6– 63.0 mm in females vs. 43.5–43.8 mm in males, 36.5–43.6 mm in females in L. nguyenorum), distinct tympana (vs. indistinct in L. nguyenorum), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. nguyenorum), and dorsal skin with tubercles (vs. smooth in L. nguyenorum); from L. poilani (Bourret) by having a smaller size (SVL 50.1–68.9 mm in males, 45.5–63.0 mm in females vs. 81.1–99.5 mm in males, 69.5–95.3 mm in females of L. poilani), rostral length greater than eye diameter (vs. eye diameter greater than rostral length of L. poilani), and different color on ventral surface of fore and hind limbs and belly (white vs. yellow in L. poilani); from L. taylori Matsui, Panha, Khonsue & Kuraishi by having distinct tympana (vs. slightly visible through skin in L. taylori), different webbing between toes (webbing formula I0–0II0–1/ 3III 0–1/ 3IV 1/2–0V vs. fully webbed in L. taylori), and dorsal skin with tubercles (vs. smooth in L. taylori).
description: Published as part of Pham, Cuong The, Le, Minh Duc, Nguyen, Tao Thien, Ziegler, Thomas, Wu, Zheng Jun & Nguyen, Truong Quang, 2017, A new species of Limnonectes (Amphibia: Anura: Dicroglossidae) from Vietnam, pp. 545-558 in Zootaxa 4269 (4) on pages 549-554, DOI: 10.11646/zootaxa.4269.4.8, http://zenodo.org/record/582778