Microhyla fanjingshanensis, Li & Zhang & Xu & Lv & Jiang & Liu & Wei & Wang, 2019

Li, Shize, Zhang, Meihua, Xu, Ning, Lv, Jingcai, Jiang, Jianping, Liu, Jing, Wei, Gang & Wang, Bin, 2019, A new species of the genus Microhyla (Amphibia: Anura: Microhylidae) from Guizhou Province, China, Zootaxa 4624 (4), pp. 551-575 : 556-569

Microhyla fanjingshanensis sp. nov.

Holotype. CIBFJS20180425013 ( Figs. 4 , 5 ), adult male, collected by S. Z. Li in Fanjing Mountain (27.9153°N, 108.61026°E; elevation 1139 m a.s.l.), Yinjiang County, Guizhou Province, China on 25 April 2018.

Paratypes. Eleven specimens including ten males and two females from the Fanjing Mountain of Yinjiang County, Guizhou Province, China. Five males: CIBFJS20180425002, CIBFJS20180425003, CIBFJS20180425004, CIB- FJS20180425005, CIBFJS20180425006 and two females: CIBFJS20180425011, CIBFJS20180425012 were collected by S.Z. Li on 25 April 2018 . Four males: CIBFJS20180425007, CIBFJS20180425008, CIBFJS20180425009 and CIBFJS20180425010 were collected by J.C. Lv from the same site on 25 April 2018.

Diagnosis. Microhyla fanjingshanensis sp. nov. is assigned to the genus Microhyla based on the molecular phylogenetic analyses and the following morphological characters: relatively small body size; maxillary and vomerine teeth absent; vomer divided into two parts and disappearing at the posterior edge of the choana; tongue posteriorly rounded; skin smooth or with tubercles; tympanum hidden; 1−2 row of horizontal skin ridges on the palate; fingers without webbing; toes webbed or free of webbing; 2 or 3 metacarpal tubercles; supernumerary tubercles absence below the base of fingers and toes; absence of skin ridge or skin projection between the subarticular tubercles of toes III and IV.

Microhyla fanjingshanensis sp. nov. is distinguished from its congeners by a combination of the following characters: (1) body of medium size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females); (2) disk and dorsal median longitudinal groove on finger tips absent; (3) toe with rudimentary webbing at base; (4) disk with dorsal median longitudinal groove present at toe tips except for the toe I; (5) two metatarsal tubercles on palm; (6) tibiotarsal articulation reaching the level between eye to nostril when leg stretched forward; (7) a distinct V-shaped white stripe on the upper midsection.

Description of holotype. SVL 22.7 mm; head wider than long (HDW/HDL = 1.15); snout short, rounded and projecting beyond the lower jaw in dorsal view; nostril rounded, distinct, and closer to the tip of the snout than the eye; internasal distance (2.6 mm) greater than interorbital distance (2.0 mm) and upper eyelid (1.8 mm); eye large and convex, eye diameter 29.5% of head length; tympanum hidden; canthus rostralis and supratympanic fold distinct; tongue posteriorly oval and not notched behind; maxillary and vomerine teeth absent.

Forelimbs slender, the length of lower arm and hand 43.6% of SVL; fingers short, relative finger lengths: I <II <V <III; tips of digits rounded, not webbed, no disks and dorsal longitudinal grooves; subarticular tubercles roughly circle-shaped and distinct; supernumerary tubercles below the base of finger absent; two metacarpal tubercles, inner palmar tubercle elliptical and bigger than outer palmar tubercle; nupital pads and spines absent.

Hindlimbs slender (TL/SVL = 0.58); heels overlapped when thighs are positioned at right angles to the body, tibiotarsal articulation reaching the level between eye to nostril when leg stretched forward; tibia length longer than thigh length; relative toe lengths I <II <V <III <IV; toes distally rounded, and equipped with disks and longitudinal grooves except for the toe I, toes with rudimentary webbing at bases; two metatarsal tubercles, inner one is long and thin, and the outer is relatively small and rounded; subarticular tubercles approximately circle-like and markedly prominent; supernumerary tubercles below the base of toes absent.

Dorsal skin surface roughish with tiny tubercles; ventral surface of posterior part of body, cloacal region, thigh granules. Throat, chest, and ventral part of thigh and tibia smooth.

Coloration of holotype in life. Dorsally grayish brown, prominent dark brown markings outlined in pale brown; a thick dark brown inverted triangle marking between the eyes and on the dorsum with three pairs of symmetrical figures or stripes, the anterior pair of figures shorter and extending laterally to the shouler; the middle pair relatively long and extending laterally; the anterior pair and the middle pair connect with each other; posterior pair of stripes shorter and disconnect with the middle pair stripe; a ^ stripe distinctly between the posterior dorsum to the anterior hip; dorsum of posterior part of body and legs brownish scattered with dense dark brown oval spots; supra- tympanic fold dark brown; dark stripe on flanks contiguous; limbs indefinitely barred with dark brown; throat dark brown, belly white mottled with brown; a white stripe from the lower jaw to the cloacae then from the cloacae to the upper jaw and another white stripe from the left wrist joint to the right forming a V-shape on the upper midsection, and the two stripes converges on the upper midsection.

Coloration of preserved holotype. The stripes on dorsum and limbs are still clear, but on dorsum color fades to grey, and belly color fades to white-grey.

Variation. All specimens were similar in morphology ( Table 1 ). However, the coloration and pattern of stripes were different between individuals ( Fig. 6 ). The stripes on dorsum were darker and not asymmetrical in the middle pair of stripes in some individuals; the inverted triangle marking between the eyes disconnect with the anterior pair of stripes in some individuals; in some individuals some brown spots were present on the venter; in some individuals the venter dark purple and on the midsection had dark brown mottling like throat; in some adult individuals the white stripe on dorsal was absent and the venter was darker mottled with white.

Sexual dimorphism. Adult males with a single external subgular vocal sac ( Fig. 5C ) and nupital pads and spinesabsent.

Morphological comparisons. Ten Microhyla species were not included in the molecular phylogenetic analyses because so far, no molecular data is available for them. So we provide a comparative note with these species as follows. Microhyla fanjingshanensis sp. nov. differs from M. darevskii by a smaller body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females) and toes with rudimentary webbing at bases (vs. SVL 27.0– 32.6 mm in males and webbing reaching to disks at most toes; Table 3 ). The new species differs from M. annamensis by toes with rudimentary webbing at bases, lacking disks and longitudinal grooves on fingers and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. webbing reaching to disks at most toes, disks and longitudinal grooves on fingers present and tibiotarsal articulation of adpressed hindlimb reaching the level to snout). The new species differs from M. arboricola by a bigger body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females), toes with rudimentary webbing at bases, lacking disks and longitudinal grooves on fingers and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. SVL 13.2–15.0 mm in males and 15.9–17.0 mm in females, toe webbing reaches the level of distal subarticular tubercles, disks and longitudinal grooves on fingers present and tibiotarsal articulation of adpressed hindlimb reaching the level beyond snout). The new species differs from M. fusca by having disks and longitudinal grooves on the toes tip except for the toe I (vs. both fingers and toes with disks and longitudinal grooves). The new species differs from M. maculifera by a bigger body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females) and having two metatarsal tubercles on palm and having dorsal median longi- tudinal grooves on the toe disks except for the toe I (vs. SVL 12.0– 13.3 mm in males, 11.8 mm in female, only one metatarsal tubercle on palm and dorsal median longitudinal grooves on the toe disks absent). The new species differs from M. minuta by longitudinal grooves on the toe disks except for the toe I (vs. both fingers and toes with disks and longitudinal grooves). The new species differs from M. picta by a smaller body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females), having disks on toes tip except for the toe I and throat dark brown, chest and belly white mottled with brown in life (vs. SVL 25.5–30.1 mm in males and 27.2–33.4 mm in females, both finger and toe disks entirely absent and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. pineticola by lacking disks and longitudinal grooves on fingers (vs. both fingers and toes with disks). The new species differs from M. pulchella and M. pulverata by toes with rudimentary webbing at bases, having disks only on the toes tip and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. toe webbing reaches the level of distal subarticular tubercles, both fingers and toes with disks and tibiotarsal articulation of adpressed hindlimb reaching the level beyond snout).

Molecular phylogenetic analyses revealed that Microhyla fanjingshanensis sp. nov. was genetically closer to M. beilunensis , M. mixtura and M. okinavensis . The new species can be distinguished from these species by a series of morphological characters as follows. Microhyla fanjingshanensis sp. nov. differs from M. mixtura in the following set of characters: SVL 22.5–23.0 mm in females (vs. SVL 23.8–26.6 mm in females; Table 3 ); having a V-shaped stripe on the upper midsection (vs. the stripe absent); tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. tibiotarsal articulation of adpressed hindlimb reaching the eyes or the posterior margin of the eyes); belly white mottled with brown in life (vs. white to whitish); additionally the new species differs from M. mixtura by having significantly higher ratios of IOD, ED, LAL, TWand FL to SVL and lower ratios of HDL, HDW, SL, IND, UEW and LW to SVL in males (all p -values <0.05; Table 1 ). The new species differs from M. okinavensis in the following set of characters: SVL 22.5–23.0 mm in females (vs. SVL 26.5–30.8 mm in females); having a V-shaped stripe on the upper midsection (vs. the stripe absent); belly white mottled with brown in life (vs. white to whitish); having disks and dorsal median longitudinal grooves on the toes tip except for the toe I (vs. all toes with disks and dorsal median longitudinal grooves). The new species differs from M. beilunensis in the following set of characters: SVL 22.5–23.0 mm in females (vs. SVL 26.4–28.3 mm in females); having a V-shaped stripe on the upper midsection (vs. the stripe absent); tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. tibiotarsal articulation of adpressed hindlimb reaching the level to eyes or the posterior margin of eyes); having disks and dorsal median longitudinal grooves on the toes tip except for the toe I (vs. all toes with disks and dorsal median longitudinal grooves); additionally the new species differs from M. beilunwensis by having significantly higher ratios of HDL, HDW, SL, IND, LAL, LW, HAL, HLL, TL, TW, TFL and FL to SVL and lower ratios of IOD, and ED to SVL in males (all p -values <0.05; Table 1 ).

Except M. beilunensis , M. mixtura , M. okinavensis and the new species, 34 Microhyla species was used in molecular phylogenetic analyses ( Table S2 ). Of them, four species ( M. butleri , M. heymonsi , M. fissipes and M. pulchra ) were reported to be distributed in Guizhou Province, China ( Fei et al., 2012). We make a comparative note between them and the new species as follows. The new species differs from M. achatina by having disks only on the toes tip, lacking disks and longitudinal grooves on fingers and tibiotarsal articulation reaching the level between eye to nostril when leg stretched forward (vs. both fingers and toes with disks, disks on fingers present and tibiotarsal articulation of adpressed hindlimb reaching the level beyond snout). The new species differs from M. aurantiventris by a smaller body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females) and disks only on the toes tip (vs. SVL 25.2–27.0 mm in males, 30 mm in female and both fingers and toes with disks). The new species differs from M. berdmorei by a smaller body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females), toes with rudimentary webbing at bases and lacking disks and longitudinal grooves on fingers (vs. SVL 23.8–28.9 mm in males, 26.2–45.6 mm in female, webbing reaching to disks at most toes and disks and longitudinal grooves on fingers present in the latter). The new species differs from M. borneensis by a bigger body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females) and lacking disks and longitudinal grooves on fingers (vs. SVL 10.6–12.8 mm in males, 17.9–18.8 mm in female and disks and longitudinal grooves on fingers present). The new species differs from M. chakrapanii by having dorsal median longitudinal grooves on the toe disks except for the toe I (vs. median longitudinal grooves on toe disks absent). The new species differs from M. darreli and M. kodial by a bigger body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females of the new species vs. SVL 15.0– 15.7 mm in males of M. darreli , SVL 16.9–17.4 mm in males and 18.0– 20.4 mm in females of M. kodial ). The new species differs from M. gadjahmadai and M. karunaratnei by lacking disks and longitudinal grooves on fingers but having disks and on the toes tip with longitudinal grooves except for the toe I and throat dark brown, belly white mottled with brown in life (vs. both fingers and toes having disks and on the toes tip with longitudinal grooves and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. laterite by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females) and throat dark brown, belly white mottled with brown in life (vs. SVL 15.3–16.3 mm in males, 18.4 mm in female and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. malang by toes with rudimentary webbing at bases and lacking disks and longitudinal grooves on fingers (vs. webbing reaching to disks at most toes and disks on fingers present). The new species differs from M. mantheyi and M. marmorata by toes with rudimentary webbing at bases, lacking disks and longitudinal grooves on fingers, and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. webbing reaching to disks at most toes, disks on fingers present and tibiotarsal articulation of adpressed hindlimb reaching the level beyond snout). The new species differs from M. mihintalei by having two metatarsal tubercles on palm and having dorsal median longitudinal grooves on the toe disks except for the toe I (vs. only one metatarsal tubercle on palm and both finger and toe disks entirely absent). The new species differs from M. mukhlesuri by longitudinal grooves on the toe disks except for the toe I and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (both fingers and toes with disks and longitudinal grooves and tibiotarsal articulation of adpressed hindlimb reaching the level to snout). The new species differs from M. mymensinghensis by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), having disks only on toes tip except for the toe I and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. SVL 14.2–17.6 mm in males, 15.2–21.3 mm in female, both finger and toe disks entirely absent and tibiotarsal articulation reaching the level between the eyes to the tip of the snout). The new species differs from M. nanapollexa by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), lacking disks and longitudinal grooves on fingers, tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. SVL 16.6 mm in females, disks on fingers present, tibiotarsal articulation of adpressed hindlimb reaching the level beyond snout). The new species differs from M. nilphamariensis by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), having dorsal median longitudinal grooves on the toe disks except for the toe I and throat dark brown, belly white mottled with brown in life (vs. SVL 17.4 mm in males, median longitudinal grooves on toe disks absent and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. orientalis by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), having two metatarsal tubercles on palm, lacking disks and longitudinal grooves on fingers and throat dark brown, belly white mottled with brown in life (vs. SVL 15.8–17.4 mm in males, 15.8–17.4 mm in females, having one metatarsal tubercle on palm, disks on fingers present and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. ornata by having dorsal median longitudinal grooves on the toe disks except for the toe I, having a V- shape stripe on the upper midsection, tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward and chest and belly white mottled with brown in life (vs. dorsal median longitudinal grooves and disks absent on toes tip, absent the V-shaped stripe, tibiotarsal articulation of adpressed hindlimb reaching the level to shoulder and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. palmipes by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), lacking disks and longitudinal grooves on fingers, having dorsal median longitudinal grooves on the toe disks except for the toe I and throat dark brown, belly white mottled with brown in life (vs. SVL 16 mm in males, 21.8 mm in female, disks on fingers present, median longitudinal grooves on toe disks absent and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. perparva and M. petrigena by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females), toes with rudimentary webbing at bases, having two metatarsal tubercles on palm, lacking disks and longitudinal grooves on fingers and tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward (vs. SVL <18 mm, webbing reaching to disks at most toes, one metatarsal tubercle on palm, disks on fingers present and tibiotarsal articulation reaching reaching the level to snout). The new species differs from M. rubra by having disks only on toes tip except for the toe I and throat dark brown, belly white mottled with brown in life (vs. both finger and toe disks entirely absent and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. sholigari and M. superciliaris by a bigger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females in the new species vs. SVL 11.0–15.0 mm in females of M. sholigari and SVL 12 mm in female of M. super- ciliaris). The new species differs from M. taraiensis and M. zeylanica by having disks only on toes tip except for the toe I and throat dark brown, belly white mottled with brown in life (vs. both finger and toe disks entirely absent and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. butleri by lacking disks and longitudinal grooves on fingers, having a V-shaped stripe on the upper midsection, having two metatarsal tubercles on palm, tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward and chest and belly white mottled with brown in life (vs. disks and longitudinal grooves on fingers present, the V-shaped stripe absent, having three metatarsal tubercles on palm, tibiotarsal articulation of adpressed hindlimb reaching the level to eyes and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. fissipes by having disks on toes tip except for the toe I, tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward and throat dark brown, belly white mottled with brown in life (vs. both finger and toe disks entirely absent, tibiotarsal articulation of adpressed hindlimb reaching the level to snout and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. heymonsi by lacking disks and longitudinal grooves on fingers, having a V-shaped stripe on the upper midsection, tibiotarsal articulation of adpressed hindlimb reaching the level between eye to nostril when leg stretched forward and chest and belly white mottled with brown in life (vs. disks and longitudinal grooves on fingers present, the V-shaped stripe absent, tibiotarsal articulation of adpressed hindlimb reaching the level to eyes and white to whitish or cream ventral coloration on throat and belly). The new species differs from M. pulchra by a smaller body size (SVL 19.0– 22.7 mm in males and 22.5–23.0 mm in females), toes with rudimentary webbing at bases and lacking disks and longitudinal grooves on fingers (vs. 23.0–32.0 mm in males, 28.0– 36.5 mm in female, webbing reaching to disks at most toes and disks and longitudinal grooves on fingers present).

Comparison with earlier available names. The new species differs from synonyms of the Microhyla achatina as follows. From Diplopelma disciferum Peters 1867 , the new species differs by having smaller body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. 25 mm in D. disciferum ; sex of type unknown) ( Peters 1867). From Hylaplesia achatina Schlegel 1827 , the new species differs by grayish brown dorsum (vs. light pinkish dorsum) ( Schlegel 1827). The new species differs from earlier synonyms of the Microhyla berdmorei as follows. From Microhyla malcolmi Cochran 1927 , the new species differs by toes with rudimentary webbing at bases (vs. webbing between toes reaching up to the base of disc) ( Cochran 1927). The new species differs from Microhyla fowleri Taylor 1934 , by having smaller body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. 37mm in M. fowleri ; sex of the type unknown), toes with rudimentary webbing at bases (vs. webbing between the toes extensive, reaching almost the tip of toes), two palmar tubercles (vs. three) ( Taylor 1934). The new species differs from Engystoma berdmorei Mason 1882 , by toes with rudimentary webbing at bases (vs. toes fully webbed), throat dark brown (vs. dusky) ( Mason 1882). From Callula natarix Cope 1867 , by having smaller body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. (1.5 inches) 38 mm in C. natarix ), toes with rudimentary webbing at bases (vs. toes fully webbed) ( Cope 1867). The new species differs from Microhyla nepenthicola Das & Haas 2010 ; a synonym of Microhyla borneensis , by having larger body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. 10.6–12.8 mm in M. nepenthicola ) ( Das & Haas 2010). The new species differs from the synonyms of Microhyla butleri in following ways. From Microhyla latastii Boulenger 1920 , by toes with rudimentary webbing at bases (vs. one-third of toes webbed) ( Boulenger 1920). From Microhyla grahami Stejneger 1924 , the new species differs by lacking lateral dermal fold (vs. lateral dermal fold present) ( Stejneger 1924). From Microhyla sowerbyi Stejneger 1924 , by toes with rudimentary webbing at bases (vs. webbing extends to one-third of toes) ( Stejneger 1924). From Microhyla boulengeri Vogt 1913 and Microhyla cantonensis Chen 1929 , the new species differs by locality (both synonyms are recorded only from Hainan & Guangdong, China respectively) (Vogt 1913 & Chen 1929). The new species differs from the synonym of M. fissipes i.e. Microhyla eremita Barbour 1920 , by having toe disc (vs. finger and toe disc absent in M. eremita ) ( Barbour 1920). The new species differs from synonyms of M. okinavensis as follows. It differs from M. undulata Brown 1902 , by toes with rudimentary webbing at bases (vs. webbing almost extending half-length of each toe) ( Brown 1902). The new species differs from the synonyms of Microhyla ornata as follows. From Engystoma malabaricum Jerdon 1853 , by having smaller body size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. 27mm / 1.1 inches in E. malabaricum ; sex of type unknown), having dorsal median longitudinal grooves on the toe disks except for the toe I (vs. absence dorsal median longitudinal grooves on toe disks) ( Jerdon 1853), differs from Engystoma carnaticum Jerdon 1853 , by having dorsal median longitudinal grooves on the toe disks except for the toe I (vs. dorsal median longitudinal grooves on toe disks absent) ( Jerdon 1853). The new species differs from Microhyla niasensis Van Kampen 1915 , a synonym of Microhyla palmipes by toes with rudimentary webbing at bases (vs. toes half webbed) ( Van Kampen 1915), differs from the synonyms of Microhyla pulchra as follows. From Microhyla hainanensis Barbour 1908 , by toes with rudimentary webbing at bases (vs. onethird of toes webbed), disk with dorsal median longitudinal groove present at toe tips except for the toe I (vs. disk and dorsal median longitudinal groove absent at toe tips) ( Barbour 1908). From Microhyla melli Vogt 1914 the new species differs by being smaller in size (SVL 19.0– 22.7 mm in males, 22.5–23.0 mm in females vs. 24 mm in M. melli ; sex of the type unknown), first finger shorter than the second (vs. first finger same length as the second) and from Microhyla major Ahl 1930 , by having dorsal median longitudinal grooves on the toe disks except for the toe I (vs. disks and dorsal median longitudinal grooves on toes absent) ( Vogt 1914; Ahl 1930). The new species differs from the synonym of Microhyla rubra i.e. Copea fulva Steindachner 1864 and Microhyla fulva Miranda-Ribeiro 1926 , by type locality (these synonyms are reported from Brazil) ( Steindachner 1864; Miranda- Ribeiro1926).

Advertisement calls. Five advertisement calls of Microhyla fanjingshanensis sp. nov. were recorded in the Fanjing Mountain, Yinjiang County, Guizhou Province, China on 25 April 2018 between 22:00 23:00. The call description is based on recordings of the holotype CIBFJS20180425013 ( Fig. 7 ) from the ridge of a paddy field, and the ambient air temperature was 17.9 °C. The call sounds like a quacking duck and each call consists of three notes. Each note had 10–12 pulses (mean 10.73 ± 0.884, n = 15). Call duration was 1.238 –2.157 second (mean 1.881 ± 0.456, n = 5). Call interval was 0.167 –0.614 second (mean 0.342 ± 0.209, n = 4). Each note had a duration of 0.307 – 0.660 seconds (mean 0.493 ± 0.100, n = 15) and the intervals between notes had a duration of 0.172 –0.407 seconds (mean 0.256 ± 0.070, n = 10). Notes were repeated in series at a rate of 1.52–3.26 (mean 2.116 ± 0.467, n = 14) per second. Amplitude modulation within note was apparent, beginning with moderately high energy pulses then decreasing towards the end of each note. The average dominant frequency was 7680.83 ± 33.43 (7720 7782 Hz, n = 15).

Ecology. Microhyla fanjingshanensis sp. nov. is only known from the Fanjing Mountain, Yinjiang County, Guizhou Province, China. The species inhabits paddy fields near evergreen broad-leaf forest ( Fig. 8A ) at an elevation about 1300 m a. s. l.. Amplexed individuals could be found on the ridge of paddy field or in the paddy field ( Fig. 8B ). Two sympatric amphibian species, Hyla gongshanensis wulingensis and Polypedates megacephalus were also found in the type locality of the new species.

Etymology. The specific epithet “ fanjingshanensis ” refers to the distribution of this species, Fanjing Mountain in Yinjiang County, Guizhou Province, China. For the common name, we suggest Fanjing Mountain Pygmy frog (English) and Fan jing shan ji wa (Chinese).

Figures

FIGURE 4. The holotype specimen CIBFJS20180425013 of Microhyla fanjingshanensis sp. nov.. A, dorsal view. B, ventral view. C, ventral view of hand. D, ventral view of foot.

FIGURE 5. Photos of the holotype CIBFJS20180425013 of Microhyla fanjingshanensis sp. nov. in life.A, dorsal view. B, ven- tral view. C, dorsolateral view showing the single external subgular vocal sac. D, ventral view of hand. E, ventral view of foot.

FIGURE 6. Color variation in Microhyla fanjingshanensis sp. nov.. A, dorsal view of the male specimen CIBFJS20180425002. B, ventral view of the male specimen CIBFJS20180425002. C, dorsolateral view of the male specimen CIBFJS20180425006. D, ventral view of the male specimen CIBFJS20180425006. E, dorsal view of the female specimen CIBFJS20180425011..

FIGURE 7. Advertisement call of the holotype CIBFJS20180425013 of Microhyla fanjingshanensis sp. nov.. A, waveform showing one note. B, sonogram showing one note. C, waveform showing three notes of one advertisement call. D, sonogram showing three notes of one call.

FIGURE 8. Habitats of Microhyla fanjingshanensis sp. nov. in the type locality, Fanjing Mountain, Yinjiang County, Guizhou Province, China. A, landscape of montane forests in the type locality. B, a mountain stream occupied by the species in its type locality; insert: mating couples of the species.

Tables

TABLE 1. Morphological comparison of Microhyla fanjingshanensis sp. nov. (MF) with M. mixtura (MM) and M. beilunensis (MB). All units in mm. P-values are at 95% significance. Morphometric characters are explained in the methods section.

					Measurements						P -value from Mann-Whitney U test
			Male					Female			Male		Female
	MF (n = 10) Range Mean ± SD		MM (n = 8) Range Mean ± SD		MB (n = 37) Range Mean ± SD		MF (n = 2) Range Mean ± SD		MB (n = 4) Range Mean ± SD		MF vs. MM	MF vs. MB	MF vs. MB
SVL	19.0–22.7	21.4 ± 1.0	18.8–25.2	22.2 ± 2.0	19.1–23.7	21.9 ± 1.2	22.5–23.0	22.7 ± 0.4	26.4–28.3	27.6 ± 0.8	0.248	0.232	0.064
HDL	7.3–8.87	8.1 ± 0.6	5.8–6.6	6.2 ± 0.3	5.7–7.2	6.4 ± 0.4	8.6–8.7	8.7 ± 0.1	6.8–8.0	7.4 ± 0.5	0.000	0.000	0.064
HDW	8.7–10.3	9.3 ± 0.5	6.3–8.2	7.3 ± 0.6	7.0–9.4	7.7 ± 0.5	10.2–10.4	10.3 ± 0.1	7.5–8.9	8.4 ± 0.6	0.000	0.000	0.064
SL	2.9–3.5	3.1 ± 0.2	2.7–3.1	2.9 ± 0.1	2.3–3.0	2.7 ± 0.2	3.4–3.5	3.4 ± 0.1	2.8–3.4	3.0 ± 0.2	0.004	0.000	0.064
IND	2.1–2.8	2.4 ± 0.2	1.8–2.2	1.9 ± 0.2	1.9–3.0	2.1 ± 0.2	2.4–2.5	2.5 ± 0.1	2.3–2.4	2.3 ± 0.1	0.000	0.000	0.064
IOD	1.8–2.3	2.1 ± 0.1	2.5–3.1	2.8 ± 0.2	2.1–3.0	2.5 ± 0.3	2.3–2.1	2.3 ± 0.04	2.6–3.1	2.8 ± 0.2	0.000	0.000	1.000
UEW	1.4–1.9	1.7 ± 0.6	1.4–1.7	1.5 ± 0.1	1.3–1.9	1.7 ± 0.1	1.8–2.0	1.9 ± 0.1	1.8–2.1	1.9 ± 0.1	0.003	0.979	0.064
ED	1.9–2.7	2.3 ± 0.2	2.5–2.9	2.7 ± 0.1	2.0–3.3	2.8 ± 0.3	2.6–2.6	2.6 ± 0.01	2.9–3.3	3.1 ± 0.2	0.021	0.000	1.000
LAL	8.0–9.9	9.1 ± 0.5	7.9–10.5	9.6 ± 0.9	7.8–9.4	8.7 ± 0.4	9.0–9.5	9.3 ± 0.4	9.5–10.3	10.0 ± 0.4	0.534	0.001	0.064
LW	1.4–2.0	1.7 ± 0.2	1.07–1.4	1.3 ± 0.1	1.1–1.7	1.4 ± 0.1	1.9–2.1	2.0 ± 0.1	1.4–1.5	1.4 ± 0.02	0.000	0.000	0.064
HAL	5.2–6.0	5.6 ± 0.3	4.7–6.5	5.8 ± 0.5	4.8–6.0	5.3 ± 0.3	6.1–6.2	6.2 ± 0.1	5.8–6.7	6.1 ± 0.4	1.000	0.008	0.064
HLL	36.0–39.7	37.4 ± 0.9	33.0–42.7	38.98 ± 3.1	30.2–37.9	35.1 ± 1.9	40.6–41.0	40.8 ± 0.3	39.8–42.6	41.2 ± 1.5	0.477	0.000	0.064
TL	12.4–14.3	13.3 ± 0.7	11.1–14.4	13.8 ± 1.0	9.0–12.7	11.7 ± 0.7	14.0–14.6	14.3 ± 0.4	12.9–14.5	13.8 ± 0.8	0.131	0.000	0.064
TW	3.7–4.3	4.0 ± 0.2	2.6–3.8	3.48 ± 0.4	3.0–4.0	3.4 ± 0.3	4.5–4.7	4.6 ± 0.2	3.7–4.0	3.9 ± 0.1	0.000	0.000	0.064
TFL	17.2–19.2	18.0 ± 0.6	15.4–20.3	18.5 ± 1.5	13.9–17.7	16.3 ± 0.9	17.4–19.5	18.5 ± 1.5	18.5–20.5	19.5 ± 1.0	0.859	0.000	0.064
FL	12.5–14.0	13.2 ± 0.5	10.9–14.2	12.8 ± 0.98	9.9–12.5	11.5 ± 0.6	14.2–14.4	14.3 ± 0.1	12.9–13.4	13.2 ± 0.3	0.010	0.000	0.064

TABLE 3. Ranges of adult snout-vent lengths (SVL) of members of the genus Microhyla Tschudi, 1838 (data from Bain & Nguyen 2004; Poyarkov et al. 2014; Howlader et al. 2015; Sheshadri et al. 2016; Wijayathilaka et al. 2016; Nguyen et al. 2019; our study). Unit in mm. Species with adult SVL ranges that do not overlap with the new species are marked in bold.

Species	SVL Males Females		Species	SVL Males Females
M. fanjingshanensis sp. nov.	19.0–22.7	22.5–23.0	M. mihintalei	21.7–27.3	24.4
M. achatina	16	23	M. minuta	14.7–15.9	15.7–17.2
M. annamensis	15.2–19.8	18.2–22.6	M. mixtura	20.5–23.7	23.8–26.6
M. annectens	14.5–15.6	18.2–18.4	M. mukhlesuri	16.5–21.0	17.3–18.4
M. arboricola	13.2–15.0	15.9–17.0	M. mymensinghensis	14.2–17.6	15.2–21.3
M. aurantiventris	25.2–27.0	30	M. nanapollexa	–	16.6
M. beilunensis	19.1–23.7	26.4–28.3	M. nilphamariensis	17.4	–
M. berdmorei	23.8–28.9	26.2–45.6	M. okinavensis	22.5–28.2	26.5–30.8
M. borneensis	10.6–12.8	17.9–18.8	M. orientalis	15.8–17.4	15.8–17.4
M. butleri	20.0–25.0	21.0–26.0	M. ornata	13.4–24.9	24.9–26.2
M. chakrapanii M. darevskii M. darreli	22 27.0–32.6 15.0–15.7	– – –	M. palmipes M. perparva M. petrigena	16 10.5–11.9 13.9–16.2	21.8 12.4–14.5 15.1–17.8
M. fissipes	18.0–27.5	20.0–28.0	M. picta	25.5–30.1	27.2–33.4
M. fusca	23	–	M. pineticola	17.2–19.5	18.0–23.0
M. gadjahmadai	18.2–21.3	20.4–25.5	M. pulchra	23.0–32.0	28.0–36.5
M. heymonsi	16.5–22.0	18.0–26.5	M. pulchella	14.7–21.6	18.1–25.8
M. karunaratnei	15.8–19.1	19.6–21.0	M. pulverata	17.5–19.5	18.8–20.2
M. kodial	16.9–17.4	18.0–20.4	M. rubra	20.0–27.5	20.5–29.5
M. laterite	15.3–16.3	18.4	M. sholigari	–	11.0–15.0
M. maculifera	12.0–13.3	11.8	M. superciliaris	–	12
M. malang	18.7–22.2	19.0–23.4	M. taraiensis	20.0–20.3	24.1
M. mantheyi	15.0–29.2	14.8–24.1	M. zeylanica	14.4–18.3	15.8–20.0
M. marmorata	18.8–21.5	21.1–23.2

TABLE 2. Measurements of the adult specimens of Microhyla fanjingshanensis sp. nov., M. mixtura and M. beilunensis. All unit in mm. See abbreviations for the morphological characters in Materials and Methods section.

Species	Locality	Voucher	Sex	SVL	HDL	HDW	SL
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425006	Male	22.7	8.8	10.3	3.4
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425007	Male	21.9	8.8	9.9	3.4
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425008	Male	22.0	7.6	9.1	2.9
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425009	Male	20.8	8.8	9.4	3.0
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425010	Male	20.8	8.0	8.8	3.2
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425004	Male	22.2	7.9	9.1	3.2
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425002	Male	19.0	7.4	8.8	3.0
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425003	Male	21.8	8.5	9.8	3.1
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425011	Male	21.3	7.3	8.7	2.9
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425013	Male	21.7	8.0	9.2	3.1
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425005	Female	23.0	8.6	10.4	3.5
Microhyla fanjingshanensis sp. nov.	Fanjing Mountain Nature Reserve, Yinjiang, Guizhou, China	CIBFJS20180425012	Female	22.5	8.7	10.2	3.4
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97001	Male	22.2	6.6	7.6	2.9
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97002	Male	22.5	6.7	8.3	2.9
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97003	Male	23.1	7.1	8.8	3.0
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97004	Male	23.4	6.7	9.4	2.7
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97005	Male	21.9	6.3	7.5	2.6
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97006	Male	22.1	6.6	7.8	2.7
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97007	Male	22.0	6.5	7.4	2.6
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97008	Male	20.8	6.0	7.8	2.5
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97009	Male	21.4	6.2	7.9	2.7
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97010	Male	19.3	5.8	7.1	2.5
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97011	Male	22.9	6.8	7.8	2.8
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97012	Male	21.2	6.3	7.5	2.4
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97013	Male	19.3	5.7	7.0	2.3
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97014	Male	22.3	6.9	8.0	3.0
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97015	Male	20.0	6.1	7.6	2.6
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97016	Male	20.4	6.5	7.6	2.9
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97017	Male	19.1	5.7	7.1	2.6
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97018	Male	21.8	6.7	7.5	2.7
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97019	Male	21.4	6.0	7.7	2.7
Microhyla beilunensis	Chaiqiao, Beilun, Ningbo, Zhejiang, China	A97020	Male	22.4	6.8	8.1	2.6

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