Leptobrachium

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: ParaHoxozoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Olfactores
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Batrachomorpha
Classis: Amphibia
Subclassis: Lissamphibia
Superordo: Batrachia
Cladus: Salientia
Ordo: Anura

Familia: Megophryidae
Subfamilia: Leptobrachiinae
Genus: Leptobrachium
Species (38): L. abbotti – L. ailaonicum – L. banae – L. bompu – L. boringii – L. buchardi – L. chapaense – L. guangxiense – L. gunungense – L. hainanense – L. hasseltii – L. hendricksoni – L. huashen – L. ingeri – L. kanowitense – L. kantonishikawai – L. leishanense – L. leucops – L. liui – L. lumadorum – L. lunatum – L. mangyanorum – L. masatakasatoi – L. montanum – L. mouhoti – L. ngoclinhense – L. nigrops – L. promustache – L. pullum – L. rakhinense – L. smithi – L. sylheticum – L. tagbanorum – L. tenasserimense – L. tengchongense – L. waysepuntiense – L. xanthops – L. xanthospilum
Name

Leptobrachium Tschudi, 1838: 81

Type species: Leptobrachium hasseltii Tschudi, 1838, by monotypy.

Synonyms

Septobrachium Tschudi, 1838: 45 [typographical error for Leptobrachium]
Nireus Theobald, 1880: 638
Type species: Nireus pulcherrimus Theobald, 1880, by monotypy.
Vibrissaphora Liu, 1945: 28
Type species: Vibrissaphora boringii Liu, 1945, by monotypy.

References
Primary references

Tschudi, J.J. 1838. Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Petitpierre: Neuchâtel.

Additional references

Hamidy, A. & Matsui, M. 2010. A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia. Zootaxa 2395(1): 34–44. DOI: 10.11646/zootaxa.2395.1.3 Paywall Reference page.
Pawangkhanant, P., Poyarkov, N.A., Duong, T.V., Naiduangchan, M. & Suwannapoom, C. 2018. A new species of Leptobrachium (Anura, Megophryidae) from western Thailand. PeerJ 6: e3584. DOI: 10.7717/peerj.5584 Open access Reference page.
Sondhi, S. & Ohler, A. 2011. A blue-eyed Leptobrachium (Anura: Megophryidae) from Arunachal Pradesh, India. Zootaxa 2912(1): 28–36. DOI: 10.11646/zootaxa.2912.1.2 Paywall Reference page.
Stuart, B.L., Rowley, J.J.L., Tran, D.T.A., Le, D.T.T. & Hoang, H.D. 2011. The Leptobrachium (Anura: Megophryidae) of the Langbian Plateau, southern Vietnam, with description of a new species. Zootaxa 2804(1): 25–40. DOI: 10.11646/zootaxa.2804.1.3 Open access Reference page.
Wogan, G.O.U. 2012. A new species of Leptobrachium from Myanmar (Anura: Megophryidae). Zootaxa 3415(1): 23–36. DOI: 10.11646/zootaxa.3415.1.2 Paywall Reference page.
Hamidy, A., Matsui, M., Nishikawa, K. & Belabut, D.M. 2012. Detection of cryptic taxa in Leptobrachium nigrops (Amphibia, Anura, Megophryidae), with description of two new species. Zootaxa 3398(1): 22–39. DOI: 10.11646/zootaxa.3398.1.2 Paywall. ResearchGate Open access. Reference page.
Stuart, B.L., Phimmachak, S., Seateun, S. & Sivongxay, N. 2012. A new Leptobrachium (Anura: Megophryidae) from the highlands of southeastern Laos. Zootaxa 3155(1): 29–37. DOI: 10.11646/zootaxa.3155.1.3 Open access Reference page.
Yang, J-H., Wang, Y-Y. & Chan, B-P. 2016. A new species of the genus Leptobrachium (Anura: Megophryidae) from the Gaoligongshan Mountain Range, China. Zootaxa 4150(2): 133–148. DOI: 10.11646/zootaxa.4150.2.3 Paywall Reference page.

Vernacular names
English: Eastern Spadefoot Toads

Eastern spadefoot toads (Leptobrachium, also known as large-eyed litter frogs[1]) comprise a genus of the family Megophryidae in the order Anura, and are found in southern China, northeast India, southeast Asia, and islands of the Sunda Shelf as well as the Philippines.[2] They are characterized by a stocky body with slender, short hindlimbs.[3] In identifying species, iris colour is a valuable diagnostic morphological characteristic (see Leptobrachium bompu for an example of a blue-eyed species); the iris has uniform colour in some species, whereas in other species the upper half is coloured and the lower half is dark.[4]
Leptobrachium boringii male at the height of the breeding season. Nuptial spines are clearly visible.[5] This character has been used to define subgenus Vibrissaphora.

The sister taxon of Leptobrachium is a clade that includes Scutiger and Oreolalax.[2][6]
Subgenera

Two subgenera, Leptobrachium and Vibrissaphora, are recognized; the latter was originally described as a genus, with Vibrissaphora boringii as the type species.[2] These subgenera (or genera) were originally separated by presence (in Vibrissaphora, hence the common name moustache or spiny toad) and absence of nuptial spines on the upper labium in males during the breeding season (in Leptobrachium). Later genetic analyses have not supported this original separation, but still indicate the presence of two distinct clades. These clades can be referred to as subgenera Leptobrachium and Vibrissaphora, but their contents differ from the earlier, purely morphological definition (Vibrissaphora contains all spiny species, but also non-spiny ones). Subgenus Vibrissaphora thus defined is distributed in southern China and Indochina, and subgenus Leptobrachium in the Malay Peninsula and the Malay Archipelago northwest of the Wallace Line.[4] No morphological character that could uniquely separate the subgenera has yet been identified.[7]
Species

As of mid-2019, there were 36 recognized species:[2] and one proposed species

Leptobrachium abbotti (Cochran, 1926)
Leptobrachium ailaonicum (Yang, Chen, and Ma, 1983)
Leptobrachium banae Lathrop, Murphy, Orlov & Ho, 1998
Leptobrachium bompu Sondhi & Orlov, 2011[8]
Leptobrachium boringii (Liu, 1945)
Leptobrachium buchardi Ohler, Teynié & David, 2004[9]
Leptobrachium chapaense (Bourret, 1937)
Leptobrachium guangxiense Fei, Mo, Ye & Jiang, 2009
Leptobrachium gunungense Malkmus, 1996
Leptobrachium hainanense Ye & Fei In Ye, Fei & Hu, 1993
Leptobrachium hasseltii Tschudi, 1838
Leptobrachium hendricksoni Taylor, 1962
Leptobrachium huashen Fei and Ye, 2005
Leptobrachium ingeri Hamidy, Matsui, Nishikawa, and Belabut, 2012
Leptobrachium kanowitense Hamidy, Matsui, Nishikawa, and Belabut, 2012
Leptobrachium kantonishikawai Hamidy and Matsui, 2014
Leptobrachium leishanense (Liu and Hu, 1973)
Leptobrachium leucops Stuart, Rowley, Tran, Le & Hoang, 2011[10]
Leptobrachium liui (Pope, 1947)
Leptobrachium lumadorum Brown, Siler, Diesmos & Alcala, 2010
Leptobrachium mangyanorum Brown, Siler, Diesmos & Alcala, 2010
Leptobrachium masatakasatoi Matsui, 2013[7]
Leptobrachium montanum Fischer, 1885
Leptobrachium mouhoti Stuart, Sok & Neang, 2006
Leptobrachium ngoclinhense Orlov, 2005
Leptobrachium nigrops Berry & Hendrickson, 1963
Leptobrachium promustache (Rao, Wilkinson, and Zhang, 2006)
Leptobrachium pullum (Smith, 1921)
Leptobrachium rakhinensis Wogan, 2012
Leptobrachium smithi Matsui, Nabhitabhata & Panha, 1999
Leptobrachium sylheticum Al-Razi, Maria, and Poyarkov, 2021[11]
Leptobrachium tagbanorum Brown, Siler, Diesmos & Alcala, 2010
Leptobrachium tenasserimense Pawangkhanant, Poyarkov, Duong, Naiduangchan, and Suwannapoom, 2018
Leptobrachium tengchongense Yang, Wang, & Chan, 2016
Leptobrachium waysepuntiense Hamidy & Matsui, 2010[12]
Leptobrachium xanthops Stuart, Phimmachak, Seateun & Sivongxay, 2012[13]
Leptobrachium xanthospilum Lathrop, Murphy, Orlov & Ho, 1998

References

Wild Borneo: The Wildlife and Scenery of Sabah, Sarawak, Brunei, and Kalimantan. New Holland Publishers. 2006. p. 69. ISBN 1845373782.
Frost, Darrel R. (2017). "Leptobrachium Tschudi, 1838". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 25 June 2017.
Matsui, Masafumi; Jarujin Nabhitabhata; Somsak Panha (1999). "On Leptobrachium from Thailand with a description of a new species (Anura: Pelobatidae)". Japanese Journal of Herpetology. 18 (1): 19–29. doi:10.5358/hsj1972.18.1_19.
Matsui, M.; Hamidy, A.; Murphy, R. W.; Khonsue, W.; Yambun, P.; Shimada, T.; Ahmad, N.; Belabut, D. M.; Jiang, J. P. (2010). "Phylogenetic relationships of megophryid frogs of the genus Leptobrachium (Amphibia, Anura) as revealed by mtDNA gene sequences". Molecular Phylogenetics and Evolution. 56 (1): 259–272. doi:10.1016/j.ympev.2010.03.014. PMID 20302957.
Hudson, C. M.; Fu, J. (2013). "Male-biased sexual size dimorphism, resource defense polygyny, and multiple paternity in the Emei moustache toad (Leptobrachium boringii)". PLOS ONE. 8 (6): e67502. Bibcode:2013PLoSO...867502H. doi:10.1371/journal.pone.0067502. PMC 3696078. PMID 23840725.
Pyron, A. R.; Wiens, J. J. (2011). "A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians". Molecular Phylogenetics and Evolution. 61 (2): 543–583. doi:10.1016/j.ympev.2011.06.012. PMID 21723399.
Matsui, M. (2013). "A new Leptobrachium (Vibrissaphora) from Laos (Anura: Megophryidae)". Current Herpetology. 32 (2): 182–189. doi:10.5358/hsj.32.182. hdl:2433/216851. S2CID 85577993.
Sondhi, S.; Orlov, A. (2011). "A blue-eyed Leptobrachium (Anura: Megophryidae) from Arunachal Pradesh, India" (PDF). Zootaxa. 2912: 28–36. doi:10.11646/zootaxa.2912.1.2.
Ohler, A.; Teynié, A. & David, P. (204). "A green-eyed Leptobrachium (Anura: Megophryidae) from southern Laos" (PDF). Raffles Bulletin of Zoology. 52 (2): 695–700. Archived from the original (PDF) on 2012-03-16.
Stuart, B. L.; Rowley, J. J.; Tran, Dao T. A.; Le, Duong T. T. & Hoang, H. D. (2011). "The Leptobrachium (Anura: Megophryidae) of the Langbian Plateau, southern Vietnam, with description of a new species" (PDF). Zootaxa. 2804: 25–40. doi:10.11646/zootaxa.2804.1.3.
"Leptobrachium sylheticum Al-Razi, Maria, and Poyarkov, 2021 | Amphibian Species of the World".
Hamidy, A.; Matsui, M (2010). "A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia" (PDF). Zootaxa. 2395: 34–44. doi:10.11646/zootaxa.2395.1.3. Archived from the original (PDF) on 2016-03-03. Retrieved 2011-06-09.
Stuart, B. L.; Phimmachak, S.; Seateun, S.; Sivongxay, N. (2012). "A new Leptobrachium (Anura: Megophryidae) from the highlands of southeastern Laos" (PDF). Zootaxa. 2155: 29–37.