''Kaloula'' is a genus of

microhylid The Microhylidae, commonly known as narrow-mouthed frogs, are a geographically widespread family (biology), family of frogs. The 683 species are in 63 genera and 11 subfamilies, which is the largest number of genera of any frog family. Evolution ...

frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...

s found in southern and eastern Asia. They are sometimes known as the Asian narrowmouth toads.

Species

The genus currently has at least 19 species. Some sources also recognize ''Kaloula macrocephala'' Bourret, 1942 as a valid species, whereas the

Amphibian Species of the World ''Amphibian Species of the World 6.1: An Online Reference'' (ASW) is a herpetology database. It lists the names of frogs, salamanders and other amphibians, which scientists first described each species and what year, and the animal's known range. ...

treats it as a synonym of ''

Kaloula pulchra The banded bullfrog (''Kaloula pulchra'') is a species of frog in the narrow-mouthed frog family Microhylidae. Native to Southeast Asia, it is also known as the Asian painted frog, digging frog, Malaysian bullfrog, common Asian frog, and painte ...

''. 5 new species have been described since 2000, and there are unnamed species yet to be described. ''

'', '' Kaloula picta'', and '' Kaloula borealis'' are the most widespread and commonly found varieties, often living near human settlements. Blackburn, et al. (2013) lists several ''Kaloula'' varieties that are likely to be new, previously undescribed species. *''Kaloula

sp. nov. In biological taxonomy, a ' (plural: '; abbreviation: ' plural abbreviation: ') is a new species. The phrase is Latin, and is used after a binomial name that is being published for the first time. An example is the species of miniature frog, ''P ...

Palawan'': Closely related to '' Kaloula baleata''. Recorded in

Palawan Palawan (), officially the Province of Palawan ( cyo, Probinsya i'ang Palawan; tl, Lalawigan ng Palawan), is an archipelagic province of the Philippines that is located in the region of Mimaropa. It is the largest province in the country in ...

. *''Kaloula sp. nov. Sulawesi'': Closely related to '' Kaloula baleata''. Recorded in the Togian Islands (

Batudaka Island Batudaka is an island of the Central Sulawesi province, located in the Gulf of Tomini. The largest island in the Togian archipelago, it has a land area of over 110 square kilometers with smaller islands) and is divided administratively into villa ...

Central Sulawesi Central Sulawesi (Indonesian: ''Sulawesi Tengah'') is a province of Indonesia located at the centre of the island of Sulawesi. The administrative capital and largest city is located in Palu. The 2010 census recorded a population of 2,635,009 for ...

, and

North Sulawesi North Sulawesi ( id, Sulawesi Utara) is a province of Indonesia. It is located on the Minahasa Peninsula of Sulawesi, south of the Philippines and southeast of Sabah, Malaysia. It borders the Philippine province of Davao Occidental and Soccsks ...

(

Bogani Nani Wartabone National Park Bogani Nani Wartabone National Park is a 2,871 km2 (1,108 mi2) national park on Minahassa Peninsula on Sulawesi island, Indonesia. Formerly known as Dumoga Bone National Park, it was established in 1991 and was renamed in honour of Nan ...

). *''Kaloula sp. nov. Sibuyan'': Closely related to '' Kaloula conjuncta''. Recorded on

Sibuyan Island Sibuyan is a crescent-shaped island, the second largest in an archipelago comprising Romblon Province, Philippines. Located in the namesake Sibuyan Sea, it has an area of and has a total population of 62,815 as of 2020 census. The island has t ...

and

Mount Baloy Mount is often used as part of the name of specific mountains, e.g. Mount Everest. Mount or Mounts may also refer to: Places * Mount, Cornwall, a village in Warleggan parish, England * Mount, Perranzabuloe, a hamlet in Perranzabuloe parish, C ...

Panay Panay is the sixth-largest and fourth-most populous island in the Philippines, with a total land area of and has a total population of 4,542,926 as of 2020 census. Panay comprises 4.4 percent of the entire population of the country. The City o ...

. *''Kaloula sp. nov. Samar & Leyte'': Related to '' Kaloula conjuncta'' and '' Kaloula picta''. Recorded in

Taft, Eastern Samar Taft, officially the Municipality of Taft ( war, Bungto han Taft; tl, Bayan ng Taft), is a 4th class municipality in the province of Eastern Samar, Philippines. According to the 2020 census, it has a population of 18,786 people. It is bounded o ...

, and Danao and Baybay in

Leyte Leyte ( ) is an island in the Visayas group of islands in the Philippines. It is eighth-largest and sixth-most populous island in the Philippines, with a total population of 2,626,970 as of 2020 census. Since the accessibility of land has be ...

. *''Kaloula sp. nov. Panay'': Related to '' Kaloula kalingensis'' and ''

Kaloula kokacii The Bicol narrowmouth toad or Catanduanes narrow-mouthed frog (''Kaloula kokacii'') is a species of frog in the family Microhylidae. It is endemic to the Philippines, where it is found in the Bicol Peninsula and Catanduanes Island. Its natural h ...

''. Recorded in

Sibalom Sibalom, officially the Municipality of Sibalom, ( krj, Banwa kang Sibalom; hil, Banwa sang Sibalom; tl, Bayan ng Sibalom), is a 2nd class municipality in the province of Antique, Philippines. According to the 2020 census, it has a population ...

Antique Province Antique, officially the Province of Antique (; krj, Probinsya kang Antique; hil, Probinsya sang Antique; tl, Probinsya ng Antique), is a province in the Philippines located in the Western Visayas region. Its capital is San Jose de Buenavista, t ...

, Panay. *''Kaloula sp. nov. East Luzon'': Related to '' Kaloula kalingensis'' and ''

''. Recorded on

Mount Makiling Mount Makiling (also known as Mount Maquiling), is an inactive stratovolcano located in the provinces of Laguna and Batangas on the island of Luzon, Philippines. The mountain rises to an elevation of above mean sea level and is the highest fe ...

, Sierra Madres, and

Aurora Memorial National Park Aurora Memorial National Park is a protected area of the Philippines located within the Sierra Madre mountain range between the provinces of Nueva Ecija and Aurora in Central Luzon. The Aurora Memorial National park covers an area of 5,676 hectar ...

Phylogeny

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

study by Mo et al. (2013) suggests that the only four ''Kaloula'' species found in China, namely '' Kaloula borealis'', '' Kaloula nonggangensis'', '' Kaloula rugifera'', and '' Kaloula verrucosa'', belong to a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

group, termed the ''K. verrucosa'' group. Blackburn, et al. (2013) consider ''Kaloula'' species

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

to the

Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...

to form a

group, containing the species '' Kaloula walteri'', '' Kaloula rigida'', '' Kaloula conjuncta'', '' Kaloula picta'', '' Kaloula kalingensis'', and ''

''. ''Kaloula'' likely colonized the Philippines from Southeast Asia during the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

, and then radiated into different

ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species, ...

s. ''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 Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose num ...

(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 The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...

–

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

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 Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...

, 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. Blackburn, et al. (2013) suggests that '' Kaloula taprobanica'' might not form a clade with the other ''Kaloula'' species, although it does clearly form a clade with '' Metaphrynella'', ''

Ramanella ''Uperodon'' is a genus of microhylid frogs. They occur in South Asia (Pakistan, India, Sri Lanka, Bhutan, Nepal, and Bangladesh) and Myanmar. ''Uperodon'' reached its current composition in 2016 when the genus Ramanella was brought into its syn ...

'', and the rest of ''Kaloula''. Other studies relating to ''Kaloula'' phylogeny include van Bocxlaer et al. (2007), Matsui et al. (2011), and Trueb et al. (2011).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. The following phylogeny has been adapted from Blackburn, et al. (2013). '' Kaloula assamensis'', '' Kaloula taprobanica'', and '' Kaloula aureata'' have not been included.

References

External links

AmphibiaWeb
{{Taxonbar, from=Q2456224 Microhylidae Amphibians of Asia Amphibian genera Taxa named by John Edward Gray