Beelzebufo (Fly Lord Toad, named after the Semitic deity that was eventually considered a demon, "Beelzebul", whose name translates to "Lord of the Flies") is a genus of giant frog that lived in the Maevarano Formation of the Mahajanga Province (known as "Majunga" in French) in northwest Madagascar, 70 - 66 million years ago, during the Maastrichtian stage of the Late Cretaceous Period.
Beelzebufo is known from over a hundred fossils collected ever since the first bones were discovered in 2008, enough for paleontologists to reconstruct part of its skeleton, including almost its entire skull. Previously thought to have been 42.5 centimeters (16.7 inches) long,[3] more recent estimates have toned its size down to 23.2 centimeters (9.1 inches), just as big as a modern-day African bullfrog,[1] though a length of 30 centimeters (12 inches, or one foot) still seems possible.[DN 1] Even so, Beelzebufo is still one of the largest frogs ever known. Furthermore, the largest specimens of Beelzebufo have open sutures (fibrous skull joints), indicating that these individuals are not yet mature and thus could have grown even bigger.[1]
A male Beelzebufo about to look for a new pool
“ It had armored skull bones and armored backbone, heavily reinforced, and big, strong jaws. ”
― Marc Jones, Curator, Fossil Reptiles
The huge skull of Beelzebufo has rough surfaces, indicating that its head was partially covered with scutes (bony scales), hence its species name, "ampinga" (which means "shield" in the Malagasy language). This made it look similar to modern-day South American horned frogs, and this lead to studies regarding possible Late Cretaceous biogeographical links between South America and Madagascar.[3][1] However, when further analyzed, most research studies agree that Beelzebufo was not a member of the Ceratophryidae family,[4][5] and, currently, it is, at best, considered a close relative of the horned frogs, with both gaining similar features due to convergent evolution.[6] Overall, while recent studies do support the idea that Beelzebufo was closely related to ceratophryids, Beelzebufo still did not resemble these horned frogs too much, thus, Prehistoric Planet avoided giving Beelzebufo a similar form as the South American horned frogs.[DN 2]
“ Beelzebufo had an especially powerful bite. It would have been able to grab and subdue and kill even relatively large prey up to the size of baby dinosaurs. ”
― Dr. Darren Naish, Lead Scientific Consultant
Bite force calculations based on a Cranwell's horned frog suggest that a Beelzebufo with a skull width of 15.4 centimeters (6.1 inches) could have exerted a bite force of 500 - 2,200 newtons, or 51 - 225 kilograms (112 - 500 pounds), comparable to the bite forces of medium and large mammalian carnivores.[2]
Beelzebufo was featured in the fifth segment of Freshwater, ambushing and devouring a juvenile Masiakasaurus as it ran past his pool. The devil toad reappeared in the third segment of Swamps, where he scares off an adult Masiakasaurus, and spends the rest of the segment trying to call out for mates despite the disturbances caused by a visiting herd of Rapetosaurus. Due to being one of the biggest frogs ever known, capable of being a threat to small dinosaurs, Beelzebufo served as the first focus of the Uncovered segment "What Else Lived Alongside The Dinosaurs?"
“ In Northern Madagascar, conditions are already changing. The first rain in months is reviving the parched land. As pools and channels refill, the animals return. ”
Beelzebufo lived 70 - 66 million years ago in the Maevarano Formation in the Mahajanga Province of Madagascar. The Maevarano Formation is recognized as an alluvial plain where the discharge of rivers fluctuate. The area is also believed to have had reddish soil, with floodplains that supported vegetation able to grow in the relatively dry climate. The region, at times semi-arid (like it is today), at times undergoing strong dry and wet seasons, was eventually claimed by rising sea levels that pushed the shoreline back. This may or may not have been the result of the Cretaceous-Paleogene Extinction Event, a catastrophe that wiped out three-quarters of life on Earth.
Thermoregulation, Hunting Tactics, and Feeding Preferences[]
As an amphibian, Beelzebufo is almost certainly a poikilotherm (equivalent to the outdated concept of "cold-bloodedness"), reliant on its surroundings to regulate its body temperature since it cannot generate its own heat, with slow metabolism to match. This, coupled with its large size (compared to other anurans) as well as its likely poor speed and agility, meant that Beelzebufo is better off relying on ambush, patiently waiting for prey to approach its location, to conserve effort and energy, with its size and its relatively-strong bite force allowing it to attack and subdue even prey as large as juvenile dinosaurs at the crucial moment. Its slow metabolism, however, does allow Beelzebufo to go for long periods without food, the amount of time between meals depending on the size of the prey it consumes.
Mating Habits[]
While sitting in a pool, calling for a mate, a male Beelzebufo is disturbed by a curious Rapetosaurus
“ Female devil toads are fussy, so he needs to find a good spot. Only then will he he able to entice them in. His deep croaks travel far and wide. ”
A male devil toad's chances of attracting a mate are short on time, for it is stated that the females will only mate at the start of the rainy season. The male in the third segment of Swamps is depicted securing a mud pool for himself, a pond from where he could call out to females in the surrounding area with a song of low-pitched, resonating croaks, strong enough to make quite significant vibrations in the water.[DN 3] This is in-line with the behavior of modern-day anurans, some of which are even known to plan further ahead by building special breeding pools for the eggs, constructing channels for their tadpoles to move from one area to another, and even visiting and providing food for their young every day.[DN 4]
Beelzebufo can also prove to be territorial. Territoriality is connected with mating, for, with a place they can call their own, animals can impress attracted females. Furthermore, the larger their personal space, the more likely for them to mate undisturbed, hence, the male Beelzebufo was willing to deter even larger animals that approach him and his chosen turf too closely with hisses and growls (similar to how frogs and toads today ward off threats and foes with a variety of vocalizations ranging from hisses to high-pitched screams), and if the nuisance is small enough (e.g. an adult Masiakasaurus), he is ready to chase them away.
Reproductive Habits[]
Nothing is known regarding the reproductive habits of Beelzebufo. They could have produced small clutches of juveniles and provided active parental care by guarding their tadpoles. Beelzebufo may have also given birth to live young or laid eggs that hatched into fully-developed forms already similar to the adults. Some frogs and toads today even evolved to skip the tadpole phase entirely due to factors like a lack of suitable wet areas for tadpoles to live in, so it is possible Beelzebufo may have had the same strategies to circumvent the semi-arid conditions of the Maevarano Formation.[DN 5]
