Kaloula

Kaloula is a genus of microhylid frogs found in southern and eastern Asia. They are sometimes known as the Asian narrowmouth toads.[2]

Kaloula
Temporal range: Miocene - Present,[1] 22.3–0 Ma
Banded bullfrog, Kaloula pulchra
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Anura
Family: Microhylidae
Subfamily: Microhylinae
Genus: Kaloula
Gray, 1831
Type species
Kaloula pulchra
Gray, 1831
Diversity
About 19 species (see text)

Species

The genus currently has at least 19 species.[2] Some sources also recognize Kaloula macrocephala Bourret, 1942 as a valid species,[3][4] whereas the Amphibian Species of the World treats it as a synonym of Kaloula pulchra.[2] 5 new species have been described since 2000, and there are unnamed species yet to be described.[2]

Binomial name and authorCommon nameRange
Kaloula assamensis Das, Sengupta, Ahmed, and Dutta, 2005Assam, India
Kaloula aureata Nutphand, 1989Golden bullfrogsouthern Thailand
Kaloula baleata (Müller in Oort & Müller, 1836)Flower pot toadSouth-eastern Asia
Kaloula borealis (Barbour, 1908)North-eastern Asia
Kaloula conjuncta (Peters, 1863)Truncate-toed chorus frogPhilippines
Kaloula ghoshi Cherchi, 1954Brown bullfrogAndaman Islands
Kaloula indochinensis Chan, Blackburn, Murphy, Stuart, Emmett, Ho, and Brown, 2013Indochina
Kaloula kalingensis Taylor, 1922Kalinga narrowmouth toadPhilippines
Kaloula kokacii Ross & Gonzales, 1992Catanduanes narrow-mouthed frognorthern Philippines
Kaloula latidisca Chan, Grismer & Brown, 2014Wide-disked narrow-mouthed frogMalaysia
Kaloula mediolineata Smith, 1917Indochina
Kaloula meridionalis Inger, 1954Philippines
Kaloula nonggangensis Mo, Zhang, Zhou, Chen, Tang, Meng, and Chen, 2013Guangxi, south-western China
Kaloula picta (Duméril & Bibron, 1841)Slender-digit chorus frogPhilippines
Kaloula pulchra Gray, 1831Banded bullfrog, Chubby frogSouthern and south-eastern Asia
Kaloula rigida Taylor, 1922Luzon narrow-mouthed frognorthern Philippines
Kaloula rugifera Stejneger, 1924South-western China
Kaloula verrucosa Boulenger, 1904South-western China
Kaloula walteri Diesmos, Brown & Alcala, 2002northern Philippines

Kaloula pulchra, Kaloula picta, and Kaloula borealis are the most widespread and commonly found varieties, often living near human settlements.

Blackburn, et al. (2013)[5] lists several Kaloula varieties that are likely to be new, previously undescribed species.

Phylogeny

A molecular phylogenetic study by Mo et al. (2013)[6] suggests that the only four Kaloula species found in China, namely Kaloula borealis, Kaloula nonggangensis, Kaloula rugifera, and Kaloula verrucosa, belong to a monophyletic group, termed the K. verrucosa group.

Blackburn, et al. (2013)[5] consider Kaloula species endemic to the Philippines to form a monophyletic group, containing the species Kaloula walteri, Kaloula rigida, Kaloula conjuncta, Kaloula picta, Kaloula kalingensis, and Kaloula kokacii. Kaloula likely colonized the Philippines from Southeast Asia during the Late Miocene, and then radiated into different ecotypes.[5] K. kalingensis and K. kokacii occupy niches as arboreal tree-hole frogs in northern and southern Luzon respectively, while K. rigida and K. walteri are ground frogs in northern and southern Luzon respectively. K. picta, which is a terrestrial ground frog, and K. conjuncta, which is scansorial (climbing) and prefers shrubs, have established themselves throughout the Philippine archipelago. K. picta as well as K. pulchra display the lowest internal genetic diversity, but also have widespread geographical distributions that may have likely occurred be due to human activity.

Excluding Kaloula taprobanica, the most recent common ancestor of Kaloula likely existed about 22.3 million years ago, around the Oligocene–Miocene boundary. The most recent common ancestor of the terrestrial species K. rigida and K. walteri likely lived 4.8 million years ago during the Early Pliocene, while that of the arboreal species K. kalingensis and K. kokacii likely lived 9.1 million years ago. The most recent common ancestor of K. conjuncta, which currently has four recognized subspecies, likely lived 4.4 million years ago.[5]

Blackburn, et al. (2013)[5] suggests that Kaloula taprobanica might not form a clade with the other Kaloula species, although it does clearly form a clade with Metaphrynella, Ramanella, and the rest of Kaloula.

Other studies relating to Kaloula phylogeny include van Bocxlaer et al. (2007),[1] Matsui et al. (2011),[7] and Trueb et al. (2011).[8]

The following phylogeny has been adapted from Blackburn, et al. (2013).[5] Kaloula assamensis, Kaloula taprobanica, and Kaloula aureata have not been included.

Kaloula 
 

Kaloula verrucosa, Kaloula borealis, Kaloula rugifera, Kaloula nonggangensis[6]

 
 
 

Kaloula pulchra

 
 
 

Kaloula mediolineata

 
 
 

Kaloula baleata, Kaloula indochinensis, Kaloula latidisca[9]

 
(Philippine radiation)[5]
 
 

Kaloula kalingensis

 
 

Kaloula kokacii

 
 
 
 
 

Kaloula conjuncta

 
 

Kaloula picta

 
 
 
 

Kaloula rigida

 
 

Kaloula walteri

 
 
 
 
 
 
 
 
gollark: Really.
gollark: Suuuuuuuuuuure.
gollark: We may need to export all tags from R. Danny prior to its apification.
gollark: Do people in the UK actually *know* English?
gollark: Beware apioforms.

References
  1. van Bocxlaer, I., K. Roelants, S. D. Biju, J. Nagaraju, and F. Bossuyt. 2006. Late Cretaceous vicariance in Gondwanan amphibians. PLoS ONE 1:e74.
  2. Frost, Darrel R. (2014). "Kaloula Gray, 1831". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 21 September 2014.
  3. "Kaloula macrocephala Bourret, 1942". Integrated Taxonomic Information System.
  4. "Microhylidae". AmphibiaWeb: Information on amphibian biology and conservation. [web application]. Berkeley, California: AmphibiaWeb. 2014. Retrieved 21 September 2014.
  5. Blackburn, D. C., Siler, C. D., Diesmos, A. C., McGuire, J. A., Cannatella, D. C. and Brown, R. M. (2013), An adaptive radiation of frogs in a southeast Asian island archipelago. Evolution, 67: 2631–2646.
  6. Mo Y, Zhang W, Zhou S, Chen T, Tang H, Meng Y, Chen W. 2013. A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China. Zootaxa 3710: 165-178.
  7. Matsui, M., A. Hamidy, D. M. Belabut, N. Ahmad, S. Panha, A. Sudin, W. Khonsue, H.-S. Oh, H.-S. Yong, J.-P. Jiang, and K. Nishikawa. 2011. Systematic relationships of Oriental tiny frogs of the family Microhylidae (Amphibia, Anura) as revealed by mtDNA genealogy. Mol. Phylogenet. Evol. 61:167–176.
  8. Trueb, L., R. Diaz, and D. C. Blackburn. 2011. Osteology and chondrocranial morphology of Gastrophryne carolinensis (Anura: Microhylidae), with a review of the osteological diversity of New World microhylids. Phyllomedusa 10:99–135.
  9. Chan KO, Grismer LL, Brown RM. 2014. Reappraisal of the Javanese Bullfrog complex, Kaloula baleata (Mueller, 1836) (Amphibia: Anura: Microhylidae), reveals a new species from Peninsular Malaysia. Zootaxa 3900: 569-580.
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