page_title: Ornithocheiromorpha
section_title: Paleoecology
hierarchical_section_title: Ornithocheiromorpha / Paleoecology
caption_attribution_description: English: Geologic map of the Araripe Basin in NE Brazil
context_page_description: Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 94 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by better adapted and more advanced pterosaur species such the pteranodontids and azhdarchoids. The most diverse families of the group, the Ornithocheiridae and Anhangueridae, survived until the early Late Cretaceous with Ferrodraco, Ornithocheirus, Cimoliopterus, Lonchodectes and Lonchodraco being some of the last remaining. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.
  Ornithocheiromorphs are considered to be some of the largest animals to have ever flown.
context_section_description: Ornithocheiromorphs were present since the Early Cretaceous (Valanginian stage), and evolved throughout the rest of the period. Studies on the ornithocheirids Tropeognathus and Anhanguera, suggested that ornithocheiromorphs also expanded their distribution, while also evolving specialized features. The significant adaptions of the ornithocheiromorphs had also made them the top pterosaurs in almost every formation by the early Late Cretaceous, including the ones they were rare in, such as the Blesa Formation in Spain, where only Iberodactylus has been found.
  Even though the ornithocheiromorphs were discovered worldwide, most of them were concentrated in specific places. One specific fossil site containing an impressive number of pterosaurs is the Romualdo Formation, a diverse Lagerstätte in the larger geologic group called the Santana Group (sometimes called the Santana Formation), which is located in the Araripe Basin of northeastern Brazil, and dates back 111-108 million years ago during the Albian stage of the Early Cretaceous. The formation includes many species of Anhanguera, several fossil remains of basal ornithocheiromorphs, including Brasileodactylus, Cearadactylus and Unwindia, as well as the ornithocheirids Tropeognathus, Coloborhynchus, Maaradactylus, Araripesaurus and Barbosania. These pterosaur genera were just some of the many recovered from the site, which also include the thalassodromids Tupuxuara and Thalassodromeus, as well as the tapejarid Tapejara, the basal pteranodontoid Santanadactylus and the dubious Araripedactylus. Some other creatures from the formation include the theropods Irritator, Mirischia and Santanaraptor, and the crocodylomorph Araripesuchus. The formation also includes several turtle remains, with some specimens referring to Santanachelys, Cearachelys and Araripemys. A few fish remains were also found within the Romualdo Formation, some of which were referred to Brannerion, Rhinobatos, Rhacolepis, Tharrhias and Tribodus. The Santana Group also consists of another Lagerstätte called the Crato Formation, which is not as diverse as the Romualdo Formation, but its fossil remains are still considered important. This fossil site underlies the Romualdo Formation, and dates back around 115-113 million years ago during the Aptian stage, meaning that its fossil content is of older age. Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontians Ludodactylus and Brasileodactylus, as well as the ornithocheirid Arthurdactylus. Other pterosaur genera include the tapejarids Tupandactylus and Aymberedactylus, as well as the chaoyangopterid Lacusovagus. The formation also contains other creatures such as the enantiornithine Cratoavis, the neosuchian Susisuchus, and several species of fish, including Belonostomus, Calamopleurus, Cladocyclus, Dastilbe and Lepidotes. These fish genera were suggested to be prey for the pterosaurs that lived in the formation, but fossil remains are limited, so the subject is still controversial.
  Another important fossil site is the Wessex Formation in the Isle of Wight, near the coast of England, which dates back 140-125 million years ago (Berriasian to Barremian stages). The formation doesn't contain many fossil remains of pterosaurs compared to the Romualdo Formation, but it is still a very important site. Fossil remains of the istiodactylid Istiodactylus, the ornithocheirid Caulkicephalus, and the tapejarid Wightia were found. The formation is also known for several theropods, including the spinosaurid Baryonyx, the tyrannosauroid Eotyrannus, the dromaeosaurid Ornithodesmus, the compsognathid Aristosuchus as well as the allosauroid Neovenator. Differents types of herbivorous dinosaurs like Iguanodon, Polacanthus, Ornithopsis, Mantellisaurus and Hypsilophodon were also found within the fossil site. Some other animals from the formation include the neosuchian Bernissartia, sea turtles such as Helochelydra and Brodiechelys, the cartilaginous fish Hybodus, ray-finned fishes such as Belonostomus, Caturus, Lepidotes and Sche
caption: Geological map of the Araripe Basin, with the extent of the Santana Group shown in dark blue
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