This family includes only two genera, Barbourula and Bombina. Barbourula occurs in the Philippine Islands (Barbourula busuangensis) and Borneo (Barbourula kalimantanensis). The approximately eight species of Bombina are found in Europe, western Asia, China, Vietnam, and Korea.
Bombina are warty, aquatic toads, and tend to be gregarious. The vocal behavior of some Bombina is unusual in that the call is produced during inhalation rather than exhalation as in other frogs. They lay pigmented eggs in ponds. The tadpoles have beaks, two upper and three lower rows of denticles, and a median spiracle (typical Orton Type 3).
Species of Bombina have bright red or yellow mottling on the venter. The German word Unke means toad. Bombina exhibit an "unken reflex" when bothered. The animal will arch its back and limbs to expose the bright belly, and may turn over on its back. This acts as a warning to predators. Not surprisingly, the skin toxins of Bombina are distasteful.
Barbourula are not as brightly colored but have webbed fingers in addition to webbed toes. Tadpoles of Barbourula are unknown. Both genera have flattened bodies.
Fossil Bombina are known from the Pliocene to the Pleistocene; there are no fossils of Barbourula.
Relationships to Outgroups
Prior to 1985, Bombina and Barbourula were generally placed in the Discoglossidae, with Alytes and Discoglossus, although the dissimilarity of Alytes and Discoglossus, on one hand, and Bombina on the other has often been noted (e.g., Lanza et al., 1976).
Cannatella (1985) presented a phylogenetic analysis indicating Discoglossidae was paraphyletic. Ford and Cannatella (1993) formalized this by defining Bombinatoridae to be the node-based name for the most recent common ancestor of Bombina and Barbourula, and all its descendants.
In contrast to Cannatella (1985), Hay et al. (1995) found Bombina orientalis and Discoglossus pictus to be closest relatives; this supports the monophyly of Discoglossidae in the traditional sense. Roelants and Bossuyt (2005) found a similar result, and continued recognition of Bombinatoridae and Discoglossidae.
Griffiths (1963) stated that the diagnostic morphological feature of Discoglossidae sensu lato (including Bombinatoridae and Discoglossidae) is a triradiate sternum. This type of sternum is also present in Leiopelma.
Evolution and Systematics
Discussion of Phylogenetic Relationships
Morphological synapomorphies of Bombinatoridae include an expanded flange of the quadratojugal and the presence of endochondral ossifications in the hyoid plate; these are the same as parahyoid bones, which are also present (Cannatella, 1985; Clarke, 1987).
Yu et al. (2007) estimated the phylogeny of most of the species of Bombina using several mitochondial genes; they recovered a largely European clade (subgenus Bombina) and a largely Asian clade (subgenus Grobina).
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:160
Specimens with Barcodes:139
Species With Barcodes:16
The Bombinatoridae are often referred to as fire-bellied toads because of their brightly colored ventral sides, which show they are highly toxic. This family includes two genera, Barbourula and Bombina, both of which have flattened bodies.
Bombina species are warty, aquatic toads about 7 cm (2.8 in) in length, and most noted for their bright bellies. They often display the unken reflex when disturbed; the animal will arch its back and limbs to expose the bright belly, and may turn over on its back. This acts as a warning to predators. The vocal behavior of some Bombina sp. is unusual in that the call is produced during inhalation rather than exhalation as in other frogs. They lay pigmented eggs in ponds.
Barbourula species occur in the Philippine Islands and Borneo, while Bombina species are found throughout Eurasia. They are slightly less colored than Bombina, and possess webbed fingers in addition to webbed toes. Characteristics of tadpoles of Barbourula are unknown.
Barbourula was considered to be situated intermediate between Discoglossus and Bombina, but closer to the latter, and was therefore also added to the Bombinatoridae when that family was split from the Discoglossidae.
- Genus Barbourula
- Genus Bombina
- B. bombina (Linnaeus, 1761) – European fire-bellied toad
- B. lichuanensis Ye and Fei, 1993 – Lichuan bell toad
- B. fortinuptialis Tian & Wu, 1978 – large-spined bell toad or Guangxi firebelly toad
- B. microdeladigitora Liu, Hu & Yang, 1960 – Hubei firebelly toad, considered a synonym of B. maxima
- B. maxima (Boulenger, 1905) – Yunnan firebelly toad
- B. orientalis (Boulenger, 1890) – Oriental fire-bellied toad
- B. pachypus (Bonaparte, 1838) – Apennine yellow-bellied toad
- B. variegata (Linnaeus, 1758) – yellow-bellied toad
- San Mauro, Diego; Mario Garcia-Paris and Rafael Zardoya (December 2004). "Phylogenetic relationships of discoglossid frogs (Amphibia:Anura:Discoglossidae) based on complete mitochondrial genomes and nuclear genes". Gene 343 (2): 357–66. doi:10.1016/j.gene.2004.10.001. PMID 15588590.
- San Mauro, Diego; Miguel Vences, Marina Alcobendas, Rafael Zardoya and Axel Meyer (May 2005). "Initial diversification of living amphibians predated the breakup of Pangaea" ([dead link] – Scholar search). American Naturalist 165 (5): 590–9. doi:10.1086/429523. PMID 15795855.
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