Polypedates leucomystax (Gravenhorst, 1829)

Class: Amphibia > Order: Anura > Family: Rhacophoridae > Subfamily: Rhacophorinae > Genus: Polypedates > Species: Polypedates leucomystax

Hyla leucomystax Gravenhorst, 1829, Delic. Mus. Zool. Vratislav., 1: 26. Type(s): Breslau Mus. (presumably MNHHWU), according to Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157. Type locality: "Java", Indonesia.

Hyla sexvirgata Gravenhorst, 1829, Delic. Mus. Zool. Vratislav., 1: 28. Holotype in Breslau Mus. (MNHHWU?). by original statement and according to Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157. Type locality: "Java", Indonesia. Synonymy by Tschudi, 1838, Classif. Batr.: 75; Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157; Boulenger, 1890, Fauna Brit. India, Rept. Batr.: 474; Boulenger, 1912, in Robinson (ed.), Vert. Fauna of the Malay Peninsula: 248; Van Kampen, 1923, Amph. Indo-Austral. Arch.: 249.

Hyla quadrilineata Wiegmann, 1834, Nova Acta Phys. Med. Acad. Caesar Leopold Carol., Halle, 17: 260. Types: Not stated, although clearly including animal figured in pl. 22, fig. 1 of the original publication. Type locality: "Java"; corrected to Manila, Luzon [Philippines], by Wolf, 1936, Bull. Raffles Mus., 12: 137–217. Synonymy by Tschudi, 1838, Classif. Batr.: 75; Theobald, 1868, J. Asiat. Soc. Bengal, 37: 84; Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157; Boulenger, 1890, Fauna Brit. India, Rept. Batr.: 474; Boulenger, 1912, in Robinson (ed.), Vert. Fauna of the Malay Peninsula: 248; Van Kampen, 1923, Amph. Indo-Austral. Arch.: 249; Wolf, 1936, Bull. Raffles Mus., 12: 178; and Inger, 1954, Fieldiana, Zool., 33: 382.

Polypedates leucomystaxTschudi, 1838, Classif. Batr.: 34, 75; Duméril and Bibron, 1841, Erp. Gen., 8: 520; Siedlecki, 1909, Biol. Zentralbl., 29: 704; Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 184.

Hyla leucopogon Tschudi, 1838, Classif. Batr.: 75. Types: RMNH, by original designation. Type locality: Not stated. RMNH jar label name proposed in synonymy of Hyla leucomystax Gravenhorst, 1829.

Hyla quadrivirgata Tschudi, 1838, Classif. Batr.: 75. Types: RMNH, by original designation. Type locality: Not stated. RMNH jar label name proposed in synonymy of Hyla leucomystax Gravenhorst, 1829.

Polypedates rugosus Duméril and Bibron, 1841, Erp. Gen., 8: 520. Syntypes: MNHNP. Type locality: "Java", Indonesia, and "Manille" (= Manila), Philippines. Synonymy by Boulenger, 1890, Fauna Brit. India, Rept. Batr.: 475; Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157; Ahl, 1931, Das Tierreich, 55: 134; Wolf, 1936, Bull. Raffles Mus., 12: 178.

Polypedates quadrilineatusGünther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 79; Günther, 1864, Rept. Brit. India: 429.

Limnodytes celebensis Fitzinger, 1861 "1860", Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 42: 413. Types: Presumably NHMW, but not mentioned in recent type lists. Type locality: "Celebes", Indonesia. Nomen nudum. Synonymy (with Polypedates quadrilineatus) by Steindachner, 1864, Verh. Zool. Bot. Ges. Wien, 14: 253. Considered a junior synonym of Rhacophorus maculatus var. quadrilineata [sic] by Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 84.

Rhacophorus maculatus var. quadrilineataBoulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 84.

Hylorana longipes Fischer, 1885, Arch. Naturgesch., 51: 47. Holotype: BMNH 1947.2.8.37 (formerly 1885.6.15.1), according to Dubois, 1987 "1986", Alytes, 5: 81. Type locality: "Pagat, Distrikt Batang Alai", southeastern Borneo, Indonesia. Synonymy by Boulenger, 1886, Zool. Rec., 22: 23 (with Rhacophorus maculatus quadrilineata); by Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 157; Van Kampen, 1923, Amph. Indo-Austral. Arch.: 246; Wolf, 1936, Bull. Raffles Mus., 12: 178; and Dubois, 1987 "1986", Alytes, 5: 81. Not mentioned by Inger, 1966, Fieldiana, Zool., 52: 307.

Polypedates maculatus quadrilineatusFischer, 1885, Arch. Naturgesch., 51: 43.

Rhacophorus leucomystaxBoulenger, 1889, Proc. Zool. Soc. London, 1889: 29; Taylor and Elbel, 1958, Univ. Kansas Sci. Bull., 38: 1068.

Rhacophorus leucomystax leucomystaxMocquard, 1890, Nouv. Arch. Mus. Natl. Hist. Nat. Paris, Ser. 3, 2: 122, 150.

Rhacophorus leucomystax quadrilineatusMocquard, 1890, Nouv. Arch. Mus. Natl. Hist. Nat. Paris, Ser. 3, 2: 122, 150.

Rhacophorus leucomystax var. sexvirgataBoettger, 1894, in Semon (ed.), Zool. Froschungsreisen Aust. Malay. Archipel., 5: 113.

Rhacophorus leucomystax quadrilineataWerner, 1903, Abh. Math. Physik. Cl. Bayer. Akad. Wiss., 22: 369.

Rhacophorus maculatus leucomystaxAnnandale, 1912, Rec. Indian Mus., 8: 14.

Rhacophorus maculatus himalayensis Annandale, 1912, Rec. Indian Mus., 8: 14, 24. Types: Not stated, although presumably originally including specimens in ZSIC. Type locality: "The Eastern Himalayas, Assam, western China". Synonymy by Ahl, 1931, Das Tierreich, 55: 127. See comment under Polypedates teraiensis.

Hyla wirzi Roux, 1927, Rev. Suisse Zool., 34: 121. Holotype: NHMB 3898 according to Forcart, 1946, Verh. Naturforsch. Ges. Basel, 57: 132. Type locality: "Région du lac Sentani, Nouvelle-Guinee hollandaise septentrionale [= northern Dutch New Guinée, near Djayapura, formerly Hollandia], West Irian, Indonesia. Considered a likely synonym of Polypedates leucomystax by Tyler, 1968, Zool. Verh., Leiden, 96: 7. Menzies, 2006, Frogs New Guinea & Solomon Is.: 271, regarded the synonymy as correct and the type locality possible.

Rhacophorus (Polypedates) leucomystaxBourret, 1927, Fauna Indochine, Vert., 3: 264.

Rhacophorus (Polypedates) quadrilineatusBourret, 1927, Fauna Indochine, Vert., 3: 264.

Rhacophorus kampeni Ahl, 1927, Sitzungsber. Ges. Naturforsch. Freunde Berlin, 1927: 43. Holotype: ZMB. Type locality: "Sumatra", Indonesia. Synonymy by Wolf, 1936, Bull. Raffles Mus., 12: 178.

Rhacophorus (Rhacophorus) kampeniAhl, 1931, Das Tierreich, 55: 116.

Rhacophorus (Rhacophorus) leucomystaxAhl, 1931, Das Tierreich, 55: 134; Dubois, 1987 "1986", Alytes, 5: 77.

Rhacophorus (Rhacophorus) himalayanusAhl, 1931, Das Tierreich, 55: 137.

Rhacophorus (Rhacophorus) leucomystax leucomystax — Wolf, 1936, Bull. Raffles Mus., 12: 178.

Rhacophorus (Rhacophorus) wirziForcart, 1946, Verh. Naturforsch. Ges. Basel, 57: 132.

Rhacophorus leucomystax quadrilineatusInger, 1954, Fieldiana, Zool., 33: 382.

Polypedates leucomystaxDutta, 1997, Amph. India Sri Lanka: 93.

English Names

Java Whipping Frog (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 113; Schleich, Anders, and Kästle, 2002, in Schleich and Kästle (eds.), Amph. Rept. Nepal: 80).

Common tree Frog (Kelaart, 1853, Prodr. Faunae Zeylan., 1: 193; Lim and Lim, 1992, Guide Amph. Rept. Singapore: 38).

Brown Tree Frog (Karsen, Lau, and Bogadek, 1986, Hong Kong Amph. Rept.: 31).

Malayan House Frog (Kiew, 1987, Malayan Nat. J., 41: 421).

Four-lined Tree Frog (Lim and Lim, 1992, Guide Amph. Rept. Singapore: 38).

Four-lined Treefrog (Grismer, 2012, Field Guide Amph. Rept. Seribuat Arch.: 62).

White-lipped Tree Frog (Polypedates leucomystax leucomystax: Maeda and Matsui, 1990, Frogs Toads Japan, Ed. 2: 161; Goris and Maeda, 2004, Guide Amph. Rept. Japan: 115).

Malayan Tree Frog (Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 133).

Brown Tree Frog (Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 133).

Bamboo Tree Frog (Shrestha, 2001, Herpetol. Nepal: 91).

House Tree Frog (Nutphund, 2001, Amph. Thailand: 138).

Jar Tree Frog (Nutphund, 2001, Amph. Thailand: 138).

Stripe Tree Frog (Nutphund, 2001, Amph. Thailand: 140).

Common Treefrog (Chan-ard, 2003, Photograph. Guide Amph. Thailand: 152).

Four-lined Tree Frog (Das, Jankowski, Makmor, and Haas, 2007, Mitt. Hamburg. Zool. Mus. Inst., 104: 162).

Distribution

Nominally from eastern India and eastern Nepal (although these are unlikely to be canonical Polypedates leucomystax; see comments under Polypedates teraiensis and Polypedates megacephalus) to Myanmar and to southern China (see comment under Polypedates megacephalus) to Philippines, and throughout Thailand to Cambodia, peninsular Malaysia, Sumatra, Sulawesi, and Borneo (Sarawak, Malaysia, East Timor, and Brunei) (distribution highly provisional); the southern Ryukyus (Okinawajima and Miyakojima), Japan. On the basis of genetically confirmed specimens and localities within that set reasonably assumed to be Polypedates leucomystax, the restricted range would be peninsular Myanmar, peninsular Thailand, Malaya and Singapore and Sundaland (Indonesia, Brunei, East Timor, Malaysia) to the Philippines and the southern Ryukyu Is. (Japan); possibly to Cambodia. 

Comment

The names Rhacophorus leucomystax or Polypedates leucomystax have been applied at various times to populations of frogs from Nepal and India to Japan, southern China and Taiwan, the Philippines, Thailand, Myanmar, and Borneo. But, with the taxonomic exclusion of Polypedates mutus, Polypedates impresus, Polypedates braueri, and Polypedates megacephalus, the distribution of Polypedates leucomystax (sensu stricto) started to come into view. Hence, literature before the appearance of Pan, Dang, Wang, Zheng, Rao, and Li, 2013, Asian Herpetol. Res., 4: 124−133, Rujirawan, Stuart, and Aowphol, 2013, Zootaxa, 3702: 545–565, Yuan, Deng, Jiang, Klaus, Orlov, Yang, and Li, 2021, Zool. Res., Kunming, 42: 116–123,  Kuraishi, Matsui, Hamidy, Belabut, Ahmad, Panha, Sudin, Yong, Jiang, Ota, Thong, and Nishikawa, 2013, Zool. Scripta, 42: 317−325, and Buddhachat and Suwannapoom, 2018, PeerJ, 6 (e4263): 1–13, should be employed with caution. In the Rhacophorus (Rhacophorus) leucomystax group of Dubois, 1987 "1986", Alytes, 5: 77; see comment under Polypedates. See Bourret, 1942, Batr. Indochine: 425–430, for review (also as Rhacophorus leucomystax) of southeastern Asia populations (now partitioned into several species of the Polypedates leucomystax). Liu, 1950, Fieldiana, Zool. Mem., 2: 370, provided an account for Chinese population (as Rhacophorus leucomystax; now Polypedates braueri, Polypedates impresus, Polypedates megacephalus, and Polypedates mutus). See also Inger, 1954, Fieldiana, Zool., 33: 376–384, for Philippine population. See Taylor, 1962, Univ. Kansas Sci. Bull., 43: 496–505, for an account of the aggregate Thai populations (as Rhacophorus leucomystax, although now partitioned into several species). Heyer, 1971, Fieldiana, Zool., 58: 61–82, reported on the call from Thailand, although the identification is suspect based on geography and likely applies to Polypedates megacephalus. See also Berry, 1975, Amph. Fauna Peninsular Malaysia: 101–102, for account. The attribution of the name Hyla leucomystax to either Boie or Kuhl in Gravenhorst is in error. Gravenhorst used the label (= manuscript) name of Boie as provided by the Curator De Haan. As Gravenhorst is responsible for the description, he is the author, under the International Code of Zoological Nomenclature (1984) according to M.S. Hoogmoed In Inger, 1985, in Frost (ed.), Amph. Species World: 541. Discussed by Dubois, 1987 "1986", Alytes, 5: 79–81. See account for Philippine population by Brown and Alcala, 1994, Proc. California Acad. Sci., Ser. 4, 48: 203. See account by Ye, Fei, and Hu, 1993, Rare and Economic Amph. China: 329 (now Polypedates braueri, Polypedates impresus, and Polypedates megacephalus). See account by Inger, Orlov, and Darevsky, 1999, Fieldiana, Zool., N.S., 92: 25–26, regarding Vietnam population (now Polypedates braueri, Polypedates mutus, Polypedates impresus, and Polypedates megacephalus). Two call types have been detected in Sabah, Malaysia, suggesting that this species is a composite according to Stuebing in Inger and Tan, 1996, Raffles Bull. Zool., 44: 566. Orlov, Lathrop, Murphy, and Ho, 2001, Russ. J. Herpetol., 8: 35–36, provided an account for the Vietnam population and noted the likely occurrence of five species masquerading under this name in that country alone (none of which is canonical Polypedates leucomystax). Iskandar, 1998, Amph. Java Bali: 89–90, provided a brief account for the Javan population. Lim and Lim, 1992, Guide Amph. Rept. Singapore: 38, provided a brief account. Maeda and Matsui, 1990, Frogs Toads Japan, Ed. 2: 158–161, provided an account for the Ryukyu, Japan, population, and suggested that the population in Okinawa is introduced from Indochina. Kuraishi, Matsui, and Ota, 2009, Pacific Sci., 63: 317–325, discussed the origin of the Ryukyu population and on the basis of molecular evidence suggested that the source population is in the Philippines. See brief account and photo by Manthey and Grossmann, 1997, Amph. Rept. Südostasiens: 128–129, and (p. 138; without discussion regarded Rhacophorus kampeni as distinct). Huang, 1990, Fauna Zhejiang, Amph. Rept.: 86–88, provided an account  as Polypedates leucomystax (now likely Polypedates braueri as Polypedates leucomystax). Zhang and Wen, 2000, Amph. Guangxi: 127–128, provided an account for Guangxi (now likely Polypedates braueri, Polypedates impresus, and/or Polypedates megacephalus). Orlov, Murphy, Ananjeva, Ryabov, and Ho, 2002, Russ. J. Herpetol., 9: 96, commented on the Vietnamese distribution (of what is now the Polypedates leucomystax complex). See brief account (as Polypedates leucomystax) by Shrestha, 2001, Herpetol. Nepal: 92–93, although the specimens are unlikely to be canonical Polypedates leucomystax. Reported for southwestern Cambodia by Ohler, Swan, and Daltry, 2002, Raffles Bull. Zool., 50: 465–481, although this needs to be confirmed by genetic analysis. Nutphund, 2001, Amph. Thailand: 138–140, provided a brief characterization and photographs for Thailand. Malkmus, Manthey, Vogel, Hoffmann, and Kosuch, 2002, Amph. Rept. Mount Kinabalu: 194–196, provided an account. Lever, 2003, Naturalized Rept. Amph. World: 226, regarded the populations in the Philippines and Ryukyus as introduced. Chan-ard, 2003, Photograph. Guide Amph. Thailand: 152–153, provided a very brief account (as Polypedates leucomystax), map for Thailand, and photograph, which likely covers several species as now understood. Goris and Maeda, 2004, Guide Amph. Rept. Japan: 115–117, provided an account for Japan, map, and photograph. Teynié, David, Ohler, and Luanglath, 2004, Hamadryad, 29: 36, provided a record and comments for a member of this species group for Laos (likely not Polypedates leucomystax). Nguyen, Ho, and Nguyen, 2005, Checklist Amph. Rept. Vietnam: 38, provided specific localities for Vietnam (as Polypedates leucomystax, but likely Polypedates megacephalus as currently understood). Reported for Pulau Langkawi, Kadeh, northwestern West Malaysia, by Grismer, Youmans, Wood, Ponce, Wright, Jones, Johnson, Sanders, Gower, Yaakob, and Lim, 2006, Hamadryad, 30: 61–74. See comment under Polypedates insularisMcKay, 2006, Field Guide Amph. Rept. Bali: 40–41, provided a brief account and photograph. Menzies, 2006, Frogs New Guinea & Solomon Is.: 271–272, regarded an old record for Papua New Guinea (as Hyla wirzi) as possibly correct, although this would need to be confirmed. Das, Jankowski, Makmor, and Haas, 2007, Mitt. Hamburg. Zool. Mus. Inst., 104: 162, provided a brief description for Sarawak, Malaysia (Borneo). Das and Dutta, 2007, Hamadryad, 31: 154–181, noted a number of larval descriptions in the literature for nominal Polypedates leucomystax, which, as currently understood apply to a number of species in the Polypedates leucomystax complex. Das, 2007, Amph. Rept. Brunei: 72, provided a photograph and brief account. Neang and Holden, 2008, Field Guide Amph. Cambodia: 108, provided a photograph, brief account of identification, ecology, and range in Cambodia. Yang, 2008, in Yang and Rao (ed.), Amph. Rept. Yunnan: 110–111, provided a brief account for a species in this species complex (as Polypedates leucomystax) for Yunnan, China. Mathew and Sen, 2010, Pict. Guide Amph. NE India: 103–104, provided a brief characterization and photographs of animals from northeastern India assigned by them to this species although these identifications require confirmation; see comments under Polypedates teraiensisSheridan, Bickford, and Su, 2010, Raffles Bull. Zool., 58: 369–379, reported on significant call differences between populations in Singapore and Thailand showing only small genetic distances. Mahony, Hasan, Kabir, Ahmed, and Hossain, 2009, Hamadryad, 34: 80–94, and Sarker and Howlader, 2012, Herpetol. Rev., 43: 301, provided records assigned to this species from Bangladesh (but these require genetic confirmation; see comments under Polypedates teraiensis). Brown, Linkem, Siler, Sukumaran, Esselstyn, Diesmos, Iskandar, Bickford, Evans, McGuire, Grismer, Supriatna, and Andayani, 2010, Mol. Phylogenet. Evol., 57: 598–619, reported on mtDNA phylogeography, which provisionally suggests that a number of subtaxa may be recognizable within nominal Polypedates leucomystaxLü, Pang, Yang, and Zhang, 2010, Asian Herpetol. Res., Ser. 2, 1: 86–89, discussed confusion in identification of, and distribution of mtDNA haplotypes in Polypedates megacephalus and Polypedates leucomystax in Yunnan, China. Thong-aree, Chan-ard, Cota, and Makchai, 2011, Thailand Nat. Hist. Mus. J., 5: 99–106, reported the species from Bala Forest, Narathiwat, extreme southern Thailand. Population on Sulawesi now assigned to Polypedates iskandariChan-ard, Cota, and Makchai, 2011, Amph. E. Region Thailand: 86–87, provided a photograph and brief account for eastern Thailand. Gaulke, 2011, Herpetofauna Panay Island: 110–114, provided a brief account for Panay Island, Philippines. Aran, Chuaynkern, Duangjai, and Chuaynkern, 2012, J. Wildlife Thailand, 19: 41–73, described larval morphology including the oral apparatus. O'Shea, Sanchez, Heacox, Kathriner, Carvalho, Ribeiro, Soares, Araujo, and Kaiser, 2012, Asian Herpetol. Res., Ser. 2, 3: 114–126, reported a population on East Timor that might be specifically distinct. See brief account by Grismer, 2012, Field Guide Amph. Rept. Seribuat Arch.: 62–64, for the Seribuat Archipelago, West Malaysia. Kuraishi, Matsui, Hamidy, Belabut, Ahmad, Panha, Sudin, Yong, Jiang, Ota, Thong, and Nishikawa, 2013, Zool. Scripta, 42: 54–70: 1–16, applied molecular evidence to the Polypedates leucomystax group and suggested that it is composed of multiple named and unnamed species and they attempted to delimit these species geographically, although ranges remains substantially provisional. Pan, Dang, Wang, Zheng, Rao, and Li, 2013, Asian Herpetol. Res., 4: 124–133, provided a phylogeny reconstruction of the Polypedates leucomystax complex that suggests that this species is restricted to southern Myanmar and Indonesia and Malaysia; but see the recent naming of Polypedates discantus from peninsular Malaysia and adjacent Thailand. Shahriza and Ibrahim, 2014, Check List, 10: 253–259, provided a photograph and brief natural history observations for a population in Kedah, West Malaysia. Sumarli, Grismer, Anuar, Muin, and Quah, 2015, Check List, 11(4, Art. 1679): 14 reported specimens from Mount Tebu, Lata Belatan, Lata Tembaka, and the base of Mount Lawit,Terregganu, West Malaysia, and briefly reported on their morphology and habitat. Haas, Kueh, Joseph, bin Asri, Das, Hagmann, Schwander, and Hertwig, 2018, Evol. Syst., 2: 89–114, provided a brief account of morphology and natural history for the Sabah population. Chettri, 2017, Das (ed.), Diversity Ecol. Amph. India: 97–98, questioned the identifications for specimens reported from Sikkim, India. Diversity of the Family Rhacophoridae (Amphibia: Anura) in Lai Chau Province, Vietnam. Do, Ngo, and Nguyen, 2017, Hue Univ. J. Sci: Nat. Sci., 126: 91, noted that earlier records Vietnam, are referrable to Polypedates mutus or Polypedates megacephalusBuddhachat and Suwannapoom, 2018, PeerJ, 6 (e4263): 1–13, discussed the molecular evidence for at least five lineages/species in Thailand of the Polypedates leucomystax complex, of which only the southern-most population was identified as Polypedates leucomystax, while from the northern part of the peninsula to the Myanmar and Laos border regions were identified as Polypedates megacephalus, Polypedates impresus, and an unnamed lineageHui, Ngadi, Md-Zain, Md-Zairi, and Abdul-Latiff, 2020, Biodiversitas, 21: 2425–2429, reported the species from Pulau Tinggi, off the southeastern coast of peninsular Malaysia. Sreekumar and Dinesh, 2020, Rec. Zool. Surv. India, 120: 33–40, noted that records from Maharashtra, India, are in error. Fatihah-Syafiq, Badli-Sham, Fahmi-Ahmad, Aqmal-Naser, Rizal, Azmi, Grismer, and Ahmad, 2020, ZooKeys, 985: 143–162, reported the species from Bidong Island, Terengganu, Malaysia. See brief account for East Java by Amin, 2020, Frogs of East Java: 55–77. Makchai, Chuaynkern, Safoowong, Chuachat, and Cota, 2020, Amph. N. Thailand: 161, provided photographs, a brief account for Thailand, and a range map. Hong, Anuar, Grismer, and Quah, 2021, Check List, 17: 798, reported the species from Batu Hampar Recreational Forest, Kedah, West Malaysia.   

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