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Limnonectes kuhlii (Tschudi, 1838)

Class: Amphibia > Order: Anura > Family: Dicroglossidae > Subfamily: Dicroglossinae > Genus: Limnonectes > Species: Limnonectes kuhlii

Rana kuhlii Tschudi, 1838, Classif. Batr.: 40. Syntypes: "Mus. Lugd." (= RMNH) as originally designated although the number of specimens was not; surviving syntypes are RMNH 4297 (2 specimens) according to Gassó Miracle, van den Hoek Ostende, and Arntzen, 2007, Zootaxa, 1482: 54. In addition, Guibé, 1950 "1948", Cat. Types Amph. Mus. Natl. Hist. Nat.: 37, regarded MNHNP 4469 (1 specimen on exchange from RMNH) as the holotype (lectotype by implication) but he regarded the author of the name to be Schlegel in Duméril and Bibron, 1841, Erp. Gen., 8: 384, when this is apparently a subsequent usage as noted by M. S. Hoomoed in Frost, 1985, Amph. Species World: 498. Type locality: "Java", Indonesia.

Rana palmata Tschudi, 1838, Classif. Batr.: 83. Name attributed to Kuhl and provided as a synonym of Rana kuhlii Tschudi.

Rana conspicillata Günther, 1872, Proc. Zool. Soc. London, 1872: 597. Syntypes: BMNH ("many spec." according to Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 20); a total of 8 in the BMNH according Matsui, Dubois, and Ohler, 2013, Asian Herpetol. Res., Ser. 2, 4: 187, who designated BMNH 1947.2.29.20 as lectotype and noted that the remaining paralectotypes are of various other species, including Limnonectes khasianus and Limnonectes. Type locality: "Matang", Sarawak (Borneo), Malaysia. Synonymy by Günther, 1874, Zool. Rec., 9: 79; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 20. See comment. 

Rana paradoxa Mocquard, 1890, Nouv. Arch. Mus. Natl. Hist. Nat. Paris, Ser. 3, 2: 148. Syntypes: MNHNP (6 specimens); including MNHNP 1889.223-225, 1889.245-246 according to Guibé, 1950 "1948", Cat. Types Amph. Mus. Natl. Hist. Nat.: 38, of which MNHNP 1889.222 was designated lectotype by Matsui, Dubois, and Ohler, 2013, Asian Herpetol. Res., Ser. 2, 4: 187. Type locality: "Kina Balu", Sabah (Borneo), Malaysia. Synonymy by Boulenger, 1891, Ann. Mag. Nat. Hist., Ser. 6, 7: 342, 344; Van Kampen, 1923, Amph. Indo-Austral. Arch.: 178. Primary homonym of Rana paradoxa Linnaeus, 1758 (= Pseudis paradoxa). See comment. 

Rana (Rana) kuhliBoulenger, 1920, Rec. Indian Mus., 20: 7; Van Kampen, 1923, Amph. Indo-Austral. Arch.: 178.

Dicroglossus kuhliiDeckert, 1938, Sitzungsber. Ges. Naturforsch. Freunde Berlin, 1938: 138.

Rana (Limnonectes) kuhliiDubois, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 239, by implication.

Limnonectes (Limnonectes) kuhliiDubois, 1987 "1986", Alytes, 5: 63.

Limnonectes kuhliiFei, Ye, and Huang, 1990, Key to Chinese Amph.: 153; Fei, 1999, Atlas Amph. China: 204.

Limnonectes mocquardi Matsui, Dubois, and Ohler, 2013, Asian Herpetol. Res., Ser. 2, 4: 187. Replacement name for Rana paradoxa Mocquard, 1890, preoccupied by Rana paradoxa Linnaeus, 1759. See comment. 

English Names

Kuhl's Wart Frog (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 100).

Kuhl's Creek Frog (Iskandar, 1998, Amph. Java Bali: 74; Das and Dutta, 1998, Hamadryad, 23: 65; Lue, Tu, and Hsiang, 1999, Atlas Taiwan Amph. Rept.: 70; Das, 2007, Amph. Rept. Brunei: 49).

Kuhl's Fanged Frog (Lue, Tu, and Hsiang, 1999, Atlas Taiwan Amph. Rept.: 70).

Large-headed Frog (Fei, 1999, Atlas Amph. China: 204).

Big-headed Mountain Frog (Nutphund, 2001, Amph. Thailand: 132).

Big-headed Frog (Nguyen, Ho, and Nguyen, 2005, Checklist Amph. Rept. Vietnam: 22).

Kuhl's Frog (Nutphund, 2001, Amph. Thailand: 132; Chan-ard, 2003, Photograph. Guide Amph. Thailand: 116; Dinesh, Radhakrishnan, Gururaja, and Bhatta, 2009, Rec. Zool. Surv. India, Occas. Pap., 302: 27).

Distribution

Mountains of Java; populations of closely related, but apparently unnamed species, from Assam (India) to Indochina to the Greater Sundas as far as Sulawesi, Indonesia (see comment).

Comment

In the Limnonectes (Limnonectes) kuhlii group of Dubois, 1987 "1986", Alytes, 5: 63. See Boulenger, 1920, Rec. Indian Mus., 20: 62; Taylor, 1962, Univ. Kansas Sci. Bull., 43: 408-412; Okada, 1931, Tailless Batr. Japan. Empire: 160-162; Bourret, 1942, Batr. Indochine: 278-282; Pope, 1931, Bull. Am. Mus. Nat. Hist., 61: 495-499; and Inger, 1966, Fieldiana, Zool., 52: 196-202, for discussion and review. See also Berry, 1975, Amph. Fauna Peninsular Malaysia: 71-72. Inger and Tan, 1996, Raffles Bull. Zool., 44: 559, cite a personal communication from Iskandar that Bornean populations are not conspecific with Javan populations, and that the Bornean population may be a composite, and Iskandar, 1998, Amph. Java Bali: 74-75, discussed the species and considered it restricted to the mountains of Java. Fei, 1999, Atlas Amph. China: 204-205, provided a brief account for the Chinese population, map, and figure. Emerson, Inger, and Iskandar, 2000, Mol. Phylogenet. Evol., 16: 131-142, suggested that nominal Limnonectes kuhlii is composed of at least four species which likely are not even each others' closest relatives: Taiwan (populations now assigned to Limnonectes fujianensis), Sabah plus Brunei, Endau plus Thailand, and Kuala Lumpur. On the basis of an mtDNA study of phylogeny, Zhang, Nie, Peng, Ge, Wang, Xu, and Tang, 2005, Acta Zool. Sinica, 51: 354-359, placed populations from Borneo most closely related to Limnonectes asperatus, the population from Taiwan most closely related to Limnonectes fujianensis, and another, genetically remote population in Yunnan. Inasmuch as the type locality is Java, it is unclear whether the name applies to any of these entities. Yang, 1991, Amph. Fauna of Yunnan: 129-131, discussed a frog allied to this species from China. Chanda, 2002, Handb. Indian Amph.: 116-119, provided a brief account (as Rana kuhlii). See brief account and photo by Manthey and Grossmann, 1997, Amph. Rept. Südostasiens: 95-96. Huang, 1990, Fauna Zhejiang, Amph. Rept.: 61- 62, provided an account (as Rana kuhlii) for Zhejiang populations. See also brief account (as Rana kuhlii) of a similar species by Zhao and Yang, 1997, Amph. Rept. Hengduan Mountains Region: 27. Malkmus, Manthey, Vogel, Hoffmann, and Kosuch, 2002, Amph. Rept. Mount Kinabalu: 142-144, provided an account for the Bornean population. Stuart, 1999, in Duckworth et al. (eds.), Wildlife in Lao PDR: 45, reported the species in Laos along with a concern for existing taxonomic confusion. Chan-ard, 2003, Photograph. Guide Amph. Thailand: 116-117, provided a very brief account, map for Thailand, and photograph (although the account presumably included Limnonectes megastomias, unnamed at the time). Nguyen, Ho, and Nguyen, 2005, Checklist Amph. Rept. Vietnam: 22, provided specific localities for Vietnam and (p. 148) a photograph. Stuart, Sok, and Neang, 2006, Raffles Bull. Zool., 54: 138, provided a record for Cambodia. Lue, Tu, and Hsiang, 1999, Atlas Taiwan Amph. Rept.: 70-71, provided a brief account for Taiwan (as Rana kuhlii). Sarkar and Ray, 2006, In Alfred (ed.), Fauna of Arunachal Pradesh, Part 1: 296, provided a brief account (as Rana kuhlii) for Arunachal Pradesh, northeastern India. See comment under Limnonectes bannaensis. Das, Jankowski, Makmor, and Haas, 2007, Mitt. Hamburg. Zool. Mus. Inst., 104: 150-151, provided a brief description.  McLeod, 2008, Zootaxa, 1807: 26-26, named Limnonectes megastomias from within nominal Limnonectes kuhlii and provided additional evidence for cryptic species under this name. Yang, 2008, in Yang and Rao (ed.), Amph. Rept. Yunnan: 68-69, provided a brief account (as Rana kuhlii) for Yunnan, China. Das, 2007, Amph. Rept. Brunei: 49, provided a photograph and brief account. Mathew and Sen, 2010, Pict. Guide Amph. NE India: 37-38, provided a brief characterization and photograph of the species that goes under this name in northeastern India. McLeod, 2010, Mol. Phylogenet. Evol., 56: 991-1000, suggested that nominal Limnonectes kuhlii is composed of an aggregation of not-particularly-closely-related species, implying that application of this name to any population outside of Java is likely to be in error. Thai populations were allocated to other species (Limnonectes jarujini, Limnonectes taylori) by Matsui, Panha, Khonsue, and Kuraishi, 2010, Zootaxa, 2615: 1-22, and who discussed the results of McLeod (2010) insofar as they applied to Thai populations. See Limnonectes siskdagu, which is another recent segregate.  Matsui, Dubois, and Ohler, 2013, Asian Herpetol. Res., Ser. 2, 4: 187-189, discussed nomenclatural issues and suggested that Rana paradoxa Mocquard, 1889, and Rana conspicillata Gunther, 1872, were not conspecific with each other or with Rana kuhlii Tschudi, 1838 (sensu stricto), but did not address the systematics in a detailed fashion; clearly Limnonectes kuhlii is a composite, will be dissected in the future, and the nomenclatural care this paper represents will make that considerably easier. 

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