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Description
The history of anhanguerids (or at least closely related toothed pterosaurs) can be traced back to the dawn of paleontology, with taxa like Ornithocheirus and Coloborhynchus being described from Britain during the latter half of the 19th century but these animals are only known from highly fragmentary remains and it wasn't until over a hundred years later that we finally had a breakthrough in regards to our understanding of these toothy, keel-crested and generally large-bodied pterodactyloids, thanks numerous and remarkably complete finds from Brazil in the 1970s-1980s, which shed new light on the anatomy these once elusive piscivores.

Dating to the late Aptian-early Albian (115-110 mya), this locality in the Araripe Basin was initially known as the Santana Formation but it has recently been promoted to the Santana Group, with the lower, late Aptian strata becoming the Crato Formation and the upper, early Albian strata the Romualdo Formation, the latter of which has far more fossils belonging to anhanguerid pterosaurs, as well as several different types of tapejarids, including the type species; the modestly sized Tapejara wellnhoferi, and a few theropods such as the spinosaurid Irritator.

Though certain paleontologists (namely Europeans) have tried to synonymize various Santana anhanguerids with old British taxa from the Cambridge Greensand and Wealden Group, such as the aforementioned Ornithocheirus and Coloborhynchus, their arguments hold little water these days because the British genera were erected from such fragmentary remains that they are essentially undiagnostic, on top of having been used as wastebasket taxa. Romualdo has produced well over a dozen distinct anhanguerid genera who vary in size from a modest 4-4.5-meter wingspan to the gigantic Tropeognathus mesembrinus, who stretched an estimated 8.2-8.7 meters from wing tip to wing tip, making it the second largest pterosaur known to us next to the giant azhdarchids of the Late Cretaceous.

Other Romualdo anhanguerids had wingspans between 4-6 meters, including Anhanguera (whose exact number of species remains controversial), Santanadactylus (same), Cearadactylus, and the more recently named Maaradactylus (2014). Maaradactylus kellneri was described from two neck vertebrae and a well-preserved skull stretching around 80 cm long (one of the biggest from Santana) and likely had a wingspan of around 6 meters, possibly making it the second largest anhanguerid at Romualdo. The generic name is based on Maara, the daughter of a chief in Cariri mythology, who was transformed into a river monster with long teeth, devouring fishermen...which honestly would better suit a local spinosaurid like Irritator or Oxalaia.

A second species recently attributed to the genus is M. spielbergi, who was originally described as "Coloborhynchus spielbergi" in 2003 based on a reasonably complete skeleton, its skull around 65 cm long, before later being redescribed as Anhanguera spielbergi by Alexander Kellner in 2006, and finally as Maaradactylus spielbergi in 2019. And if it's not obvious, its specific name is honoring Stephen Spielberg, after missing out on the chance to have Utahraptor named after him back in 1993.

Besides Brazil, well-preserved anhanguerid fossils have been found in Liaoning (such as Liaoningopterus and Guidraco) and various fragmentary fossils are known from North Africa, Queensland, Britain and Iberia, and at least one from Texas (Uktenadactylus), showing that these animals had a global distribution during the Early-Mid Cretaceous and might have been the most populous coastal pterosaurs prior to the Turonian extinction, after which they were replaced by the toothless pteranodontians.

Description
The history of anhanguerids (or at least closely related toothed pterosaurs) can be traced back to the dawn of paleontology, with taxa like Ornithocheirus and Coloborhynchus being described from Britain during the latter half of the 19th century but these animals are only known from highly fragmentary remains and it wasn't until over a hundred years later that we finally had a breakthrough in regards to our understanding of these toothy, keel-crested and generally large-bodied pterodactyloids, thanks numerous and remarkably complete finds from Brazil in the 1970s-1980s, which shed new light on the anatomy these once elusive piscivores.

Dating to the late Aptian-early Albian (115-110 mya), this locality in the Araripe Basin was initially known as the Santana Formation but it has recently been promoted to the Santana Group, with the lower, late Aptian strata becoming the Crato Formation and the upper, early Albian strata the Romualdo Formation, the latter of which has far more fossils belonging to anhanguerid pterosaurs, as well as several different types of tapejarids, including the type species; the modestly sized Tapejara wellnhoferi, and a few theropods such as the spinosaurid Irritator.

Though certain paleontologists (namely Europeans) have tried to synonymize various Santana anhanguerids with old British taxa from the Cambridge Greensand and Wealden Group, such as the aforementioned Ornithocheirus and Coloborhynchus, their arguments hold little water these days because the British genera were erected from such fragmentary remains that they are essentially undiagnostic, on top of having been used as wastebasket taxa. Romualdo has produced well over a dozen distinct anhanguerid genera who vary in size from a modest 4-4.5-meter wingspan to the gigantic Tropeognathus mesembrinus, who stretched an estimated 8.2-8.7 meters from wing tip to wing tip, making it the second largest pterosaur known to us next to the giant azhdarchids of the Late Cretaceous.

Other Romualdo anhanguerids had wingspans between 4-6 meters, including Anhanguera (whose exact number of species remains controversial), Santanadactylus (same), Cearadactylus, and the more recently named Maaradactylus (2014). Maaradactylus kellneri was described from two neck vertebrae and a well-preserved skull stretching around 80 cm long (one of the biggest from Santana) and likely had a wingspan of around 6 meters, possibly making it the second largest anhanguerid at Romualdo. The generic name is based on Maara, the daughter of a chief in Cariri mythology, who was transformed into a river monster with long teeth, devouring fishermen...which honestly would better suit a local spinosaurid like Irritator or Oxalaia.

A second species recently attributed to the genus is M. spielbergi, who was originally described as "Coloborhynchus spielbergi" in 2003 based on a reasonably complete skeleton, its skull around 65 cm long, before later being redescribed as Anhanguera spielbergi by Alexander Kellner in 2006, and finally as Maaradactylus spielbergi in 2019. And if it's not obvious, its specific name is honoring Stephen Spielberg, after missing out on the chance to have Utahraptor named after him back in 1993.

Besides Brazil, well-preserved anhanguerid fossils have been found in Liaoning (such as Liaoningopterus and Guidraco) and various fragmentary fossils are known from North Africa, Queensland, Britain and Iberia, and at least one from Texas (Uktenadactylus), showing that these animals had a global distribution during the Early-Mid Cretaceous and might have been the most populous coastal pterosaurs prior to the Turonian extinction, after which they were replaced by the toothless pteranodontians.