Ornithocheiromorpha

Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ago.

[3] When ornithocheiromorphs first appeared, they were initially scavengers, consisting in a more terrestrial setting, but their success had made them the top predators of the skies, as well as the most common type of fish-eating pterosaur throughout the early Late Cretaceous.

[6] P. compressirostris later became the type species of a newly created genus called Lonchodectes (meaning "lance biter") in a review by English paleontologist Reginald Walter Hooley in 1914.

[11] In 2013, Brazilian paleontologists Taissa Rodrigues & Alexander Kellner made a deeper analysis on the species Pterodactylus cuvieri.

[14] Later, Reginald Hooley discussed O. latidens in detail, based on specimens he had found, which led Ornithodesmus to be placed within a new family called Ornithodesmidae.

[16] In 1993, the British paleontologists Stafford C. Howse and Andrew C. Milner concluded that the holotype sacrum and only specimen of O. cluniculus didn't belong to a pterosaur, but instead to a maniraptoran theropod dinosaur.

They also pointed out that no detailed attempts had been made to compare the sacrum of O. cluniculus with those of pterosaurs, and that O. latidens had in effect been treated as the type species of the genus Ornithodesmus.

The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations.

[33] Other anhanguerids like Cearadactylus had its first preparations with many serious mistakes: the front of the snout and the lower jaws were confused leading to a reconstruction in which the anterior part of the head was upside down.

With this arrangement, the maxilla was kinked, and its interlocking teeth suggested that Cearadactylus had a piscivourous diet, allowing the animal to keep hold of slippery fish.

The skull is very elongated however, and a hollow profile is seen, but not very pointed, as the upper edge and the line of the jaw run nearly parallel over most of their length.

[43] Anhanguera, another well-known genus, had a shorter tail, with broad caudal vertebrae that bore a "duplex" cross-section similar to Pteranodon.

The hip joint of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity.

[48] Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae, and Anhangueridae to form a group in 2014, which he called Lanceodontia.

[2] In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha.

Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants.

Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of Ornithocheirus species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.

[38] The description of Iberodactylus in Spain also made some paleontologists reclassify the genus Hamipterus in a newly named family called Hamipteridae.

[58] The previously recovered basal eupterodactyloid Haopterus was reclassified due to the description of Mimodactylus, and is placed in a new family called Mimodactylidae.

[59] Many recent analyses have also recovered several ornithocheirids, including Tropeognathus, Coloborhynchus, and Caulkicephalus within the family Anhangueridae, meaning that they were more closely related to Anhanguera than to Ornithocheirus.

[52][38][59][53][31] Later, in 2019, a different phylogenetic analysis, this time conducted by Borja Holgado and colleagues, focused on derived pterodactyloids with particular emphasis on anhanguerians.

[56] Serradraco Aussiedraco Hongshanopterus Targaryendraco Lonchodectes Lonchodraconidae Istiodactylidae Boreopteridae Hamipterus Brasileodactylus Barbosania Cearadactylus Cimoliopteridae Ornithocheiridae Anhangueridae In their description of Lingyuanopterus in 2022, Xu Yizhi and colleagues recovered the following phylogeny based on the earlier work of Borja Holgado and Rodrigo Pêgas.

Lonchodectidae Mimodactylidae Hongshanopterus Istiodactylidae Boreopteridae Ornithocheirus Targaryendraconia Anhangueridae Hamipteridae Ornithocheiromorphs were originally regarded as piscivorous creatures, feeding mainly on small and mid-sized fish.

[15] In 1991, Peter Wellnhofer compared the jaw endings of Istiodactylus with those of a duck, but he then noticed that it wasn't a "duck-billed pterosaur" or anything similar, even though it was popularly called that way.

[28] Another ornithocheiromorph that possessed similar features to Istiodactylus is Liaoxipterus, which is known from a skull with several unique traits, including numerous peg-like teeth.

[66] This would likely have required them to use unique modes of locomotion compared to other pterosaurs, this can already be seen in earlier evolutions such as Istiodactylus and Nurhachius, with powerful musculature attachments and well-developed pectoral and upper arm bones.

Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontians Ludodactylus[37] and Brasileodactylus, as well as the ornithocheirid Arthurdactylus.

The formation also contains other creatures such as the enantiornithine Cratoavis, the neosuchian Susisuchus, and several species of fish, including Belonostomus, Calamopleurus,[73] Cladocyclus,[74] Dastilbe[75] and Lepidotes.

[35] Another important fossil site is the Wessex Formation in the Isle of Wight, near the coast of England, which dates back around 140 and 125 million years ago (Berriasian to Barremian stages).

[77] 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.

Associated and referred bones of Pterodactylus giganteus , now considered as Lonchodraco giganteus
Holotype of Pterodactylus cuvieri , now known as Cimoliopterus
Skull fragments of Ornithodesmus latidens , which is currently known as Istiodactylus latidens
Comparison between the holotypes of Tropeognathus mesembrinus and Ornithocheirus simus
Size of Istiodactylus latidens compared to a human
Skull comparison between different species of ornithocheiromorphs; notice their different structures
Skeleton of Anhanguera showing neck and torso vertebrae
A rotated pelvis of Anhanguera santanae , showing its right side
Skeletal reconstruction of Mimodactylus , a mimodactylid istiodactyliform
Life reconstruction of Haliskia , an anhanguerid
Life reconstruction of Mimodactylus , one of many new ornithocheiromorphs recognized in 2019
Life reconstruction of Lingyuanopterus , an istiodactylid
A hypothetical recreation of a group of Istiodactylus feeding on a carcass of a stegosaur
Cimoliopterus (right) stealing prey from a Lonchodectes (left), both were derived members of this group and possible fish hunters
Restoration of a flying Istiodactylus ; notice its high aspect ratio
Geological map of the Araripe Basin , with the extent of the Santana Group shown in dark blue
Mounted skeleton of Irritator with a possible anhanguerid in its jaws
Reconstruction of the skeleton of Ferrodraco on a diagram showing the known material, and based on the related Tropeognathus mesembrinus