''Beelzebufo'' ( or ) (meaning "devil toad") is an extinct genus of hyloid

frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...

from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

Berivotra Berivotra is a rural municipality in Madagascar. It belongs to the district of Maevatanana, which is a part of Betsiboka Region. The population of the municipality was 14,675 in 2018. Infrastructure It is situated on the Route Nationale 4 fro ...

and Maevarano Formations, approximately 70 million years ago in what is now

Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...

. The type species is ''B. ampinga'', and common names assigned by the popular media to it include devil frog, devil toad, and the frog from hell.

Discovery and naming

The first

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

bones were found in 1993 by David W. Krause of New York's

Stony Brook University Stony Brook University (SBU), officially the State University of New York at Stony Brook, is a public university, public research university in Stony Brook, New York, United States, on Long Island. Along with the University at Buffalo, it is on ...

, and the species ''Beelzebufo ampinga'' was named and described by Evans, Jones & Krause (2008). The holotype is specimen UA 9600, consisting of a fused cervical and second presacral centra. These specimens were then described in more detail by Evans, Jones, and Krause (2014). Some 100 fossil isolated partial bones have been found between 1993 and 2011. Some portions of articulated skull are also known: specimen FMNH PR 2512 (a specimen discovered in 2010 which preserves most of the braincase, part of the palate, part of the skull roof and several pre-sacral vertebrae) and specimen FMNH PR 2512 (which preserves one of the posterior flanges). Researchers have been able to reconstruct parts of the frog's

skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...

, including nearly the entire skull.

Description

In early studies, it is suggested that ''Beelzebufo'' had snout-vent lengths of up to 42.5 cm (16.7 in). However in later studies, animals of this species are estimated to have grown to at least in snout-vent length, which is around the size a modern

African bullfrog The African bullfrog (''Pyxicephalus adspersus''), also known as the giant bullfrog or the South African burrowing frog, is a species of frog in the family Pyxicephalidae. It is also known as the pixie frog due to its scientific name. It is foun ...

can reach. The head of ''Beelzebufo'' was very big, and bones of the

skull roof The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In com ...

show a rugous external surface, indicating at least parts of the head may have borne bony scales, called

scute A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "Scutum (shield), shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of Bird anatomy#Scales, birds. The ter ...

s. The skull sutures are open in even the largest specimens of ''Beelzebufo'', showing that it might have grown even bigger.

Classification

Within the superfamily

Hyloidea Hyloidea is a superfamily of frogs.R.Alexander Pyron, John J.Wiens, 2011, ''A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians'' Hyloidea accounts for 54% ...

, the definitive phylogenetic position of ''Beelzebufo'' is uncertain. While initially placed as a member of the family

Ceratophryidae The Ceratophryidae, also known as common horned frogs, are a family of frogs found in South America. It is a relatively small family with three extant genera and 12 species. Despite the common name, not all species in the family have the horn-like ...

, a 2018 study suggested that ''Beelzebufo'', and other extinct frogs with ceratophryid-like traits such as '' Baurubatrachus'', were instead part of a more ancient group of

Neobatrachia The Neobatrachia (Neo-Latin ''neo-'' ("new") + ''batrachia'' ("frogs")) are a suborder of the Frog, Anura, the order (biology), order of frogs and toads. This suborder is the most advanced and apomorphic of the three anuran suborders alive toda ...

, distantly related to horned frogs. A 2022 study recovered '' Baurubatrachus'' and ''Beelzebufo'' as sister genera, with the clade formed by the two genera in turn being the sister clade to extant Ceratophyridae. A 2023 phylogenetic study instead recovered ''Beelzebufo'' as a sister taxon of the hylid ''

Ranoidea australis ''Ranoidea australis'', commonly known as the giant frog, northern snapping frog, or round frog, is a burrowing species of frog native to Australia. It occurs from western Queensland through to northern Western Australia. Description These frogs ...

''.

Paleobiology

''Beelzebufo'' most likely was a predator whose expansive mouth allowed it to eat relatively large prey, perhaps even juvenile

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

s. Bite force measurements from a growth series of Cranwell's horned frog (''Ceratophrys cranwelli''), suggest that the bite force of a large ''Beelzebufo''—skull width —may have been between .

Paleobiogeography

The fossils of ''Beelzebufo'' are from Madagascar, which, while still attached to India, separated from the coast of Somalia in the earliest stage of the Late

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

. While its definitive taxonomic placement is uncertain, ''Beelzebufo'' has been suggested to resemble horned frogs (Ceratophryidae) of

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

, which raised the possibility of a close biogeographic link between Madagascar and South America during the Cretaceous. The initial description of ''Beelzebufo'' hypothesis reignited interest and research into skeletal variation among living members of the Ceratophyridae. These investigations suggest several of the similarities between ''Beelzebufo'' and horned frogs may have evolved by convergence.

References

External links

{{Taxonbar, from=Q814066 Cretaceous frogs Late Cretaceous amphibians Maevarano fauna Fossil taxa described in 2008 Maastrichtian life Beelzebub Neobatrachia Prehistoric animals of Madagascar Fossils of Madagascar