Ornithocheiridae

Members that belong to this group lived from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 90 million years ago.

Ornithocheirids were excellent fish hunters, using various flight techniques to catch their prey, and were also capable of flying great distances without flapping constantly.

Paleontologists suspect that ornithocheirids were the ancestors of the pteranodontians; this is due to many shared aspects, such as unique flying techniques, capability of long-distance flights, and most of their diet, which mainly consisted of fish.

Most of the ornithocheirid fossil record consists of isolated teeth, as well as fragmentary bones, reaching hundreds or even thousands of remains in some localities.

Therefore, to avoid confusion, Lydekker preferred to use the name Criorhynchus for O. simus, and Ornithocheirus for the species with lanceolate jaw tips, a concept later favored by paleontologist Reginald Walter Hooley in 1914.

[6] Later, in 2001, paleontologist David Unwin revised the taxonomic history of the Cambridge Greensand pterosaurs, and divided Ornithocheiridae into three genera: Ornithocheirus, Coloborhynchus and Anhanguera.

[12][13][14] Other studies such as the ones by Kellner and colleagues in 2019 have used a different concept, classifying Coloborhynchus, Tropeognathus, as well as several other close relatives such as Ludodactylus and Caulkicephalus within the Anhangueridae, which, along with the family Hamipteridae, forms the larger group Anhangueria.

The rostrum in S. moroccensis lacks a constriction that is posterior to the anterior rosette, a feature also shared by O. simus and T. mesembrinus, therefore another synapomorphy supported by these three species.

Yet another feature shared by these three species is that the teeth are short, straight, and relatively uniform in size, something that is not present in other ornithocheirans such as Coloborhynchus and Anhanguera.

[13] In Coloborhynchus, the teeth were found to have been heterodont, elongated, recurved and caniniform, which is similar to those seen in another ornithocheirid (though also recovered as an anhanguerid)[16][18] called Caulkicephalus.

The tails of ornithocheirids are poorly known, though they appear to be composed of at least eleven short vertebrae, and become relatively circular in cross section toward the end of the series.

[26] The family Ornithocheiridae has had a controversial and very confusing taxonomic history; paleontologists who have studied this group seem to have had a different opinion on the composition of ornithocheirid taxonomy.

A term called Anhangueridae was coined by Diogenes de Almeida Campos and Kellner in 1985 to refer to pterosaurs that belong in this family.

[27] The original term Ornithocheirae by Seeley had been redefined as the least inclusive clade containing Anhanguera blittersdorffi and Ornithocheirus simus,[10] therefore it is recovered as a larger group comprising the families Anhangueridae and Ornithocheiridae in recent analysis.

[12][27] In 2019, Pêgas and colleagues have found Aetodactylus, along with two other pterosaurs (Camposipterus and Cimoliopterus), within the clade Targaryendraconia, more specifically placed within the family Cimoliopteridae in a polytomy.

[27][28] In 2014, Andres and colleagues created the subfamily Ornithocheirinae to contain Coloborhynchus and Ornithocheirus, as a sister taxon to Tropeognathus, and altogether formed the family Ornithocheiridae.

[10] In 2018, Longrich and colleagues had included the genus Siroccopteryx in their phylogenetic analysis, specifically as a member of the ornithocheirine subfamily, sister taxon to Coloborhynchus.

[12] In 2019, a study performed by Adele Pentland and colleagues had found the Ornithocheiridae to comprise more genera; while the typical Tropeognathus, Coloborhynchus and Ornithocheirus clade was included in their analysis, Pentland and colleagues found the genera Ferrodraco and Mythunga to belong the Ornithocheiridae as well, specifically sister taxa within the Ornithocheirinae, closely related to Ornithocheirus.

The close relationship between Siroccopteryx, Ornithocheirus and Tropeognathus is supported by several synapomorphies, such the teeth being short, straight, and relatively uniform in size.

However, as noted by Jacobs and colleagues, support for some of these arrangements is relatively weak due to the limited number of characters that can be scored, and the levels of homoplasy are very high.

Large, forward-facing eyes and well-developed flocculi are ideal for dip-feeding as well, which permits effective spotting of prey as well as judgement of distances when striking at them; as such, it seems likely that at least several ornithocheirids were efficient dip feeders.

[22] The first true ornithocheirid specimens were uncovered in the Cambridge Greensand of England, belonging to the infamous genus Ornithocheirus, and dated back to the Albian stage of the Early Cretaceous.

The Romualdo Formation is found to contain a variety of ornithocheirids (or anhanguerids), including Tropeognathus,[21] Coloborhynchus[43] and Araripesaurus,[44] the targaryendraconian Barbosania[45] as well as close relatives such as Anhanguera and Maaradactylus.

[59] Ornithocheirids were also partially distributed in North America, and several specimens are thought to belong to the genus Uktenadactylus (originally Coloborhychus wadleighi).

[61][62][63] Within the fossil site, several specimens of ammonoids were thought to belong to the genera Turrilites and Scaphites,[64] and along with these, remains of the shark Leptostyrax were also found.

Holotype specimens of Ornithocheirus simus (A to D) and junior synonym O. platyrhinus (E to H)
Comparison between the holotypes of Ornithocheirus (A and C) and Tropeognathus (B and D)
Skull comparisons between different anhanguerians , notice the ornithocheirids Caulkicephalus (D), Tropeognathus (E and F) and Uktenadactylus (I and J)
Restoration of three Tropeognathus in flight, notice their high aspect ratio
Holotype skull and mandible of the ornithocheirids Ferrodraco (A) and Mythunga (B), and holotype mandible of Aussiedraco (C)
Restoration of Cimoliopterus stealing prey from a Lonchodectes
Geological map of the Araripe Basin of Brazil, with the extent of the Santana Group shown in dark blue