Tapejarids

Tapejaridae (from a Tupi word meaning 'the lord of the ways') is a family of azhdarchoid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.

Description

Tapejarids were small to medium-sized pterosaurs with several unique, shared characteristics, mainly relating to the skull. Most tapejarids possessed a bony crest arising from the snout (formed mostly by the premaxillary bones of the upper jaw tip). In some species, this bony crest is known to have supported an even larger crest of softer, fibrous tissue that extends back along the skull. Tapejarids are also characterized by their large nasoantorbital fenestra, the main opening in the skull in front of the eyes, which spans at least half the length of the entire skull in this family. Their eye sockets were small and pear-shaped. Studies of tapejarid brain cases show that they had extremely good vision, more so than in other pterosaur groups, and probably relied nearly exclusively on vision when hunting or interacting with other members of their species. Tapejarids had unusually reduced shoulder girdles that would have been slung low on the torso, resulting in wings that protruded from near the belly rather than near the back, a "bottom decker" arrangement reminiscent of some planes.

Biology

Tapejarids appear to have been arboreal, having more curved claws than other azhdarchoid pterosaurs and occurring more commonly in fossil sites with other arboreal flying vertebrates such as early birds. Tapejarids have long been speculated as having been frugivores or omnivores, based on their parrot-like beaks. Direct evidence for plant-eating is known in a specimen of '' Sinopterus'' that preserves seeds in the abdominal cavity. The Barremian-
Aptian distribution of some tapejarids may even be partially associated with the first radiation phase of the angiosperms, especially of the genus Klitzschophyllites which represents a more basal angiosperm.

Classification

Tapejaridae was named and defined by Brazilian paleontologist Alexander Kellner in 1989 as the clade containing both '' Tapejara'' and ''Tupuxuara'', plus all descendants of their most recent common ancestor. In 2007, Kellner divided the family: Tapejarinae, consisting of ''Tapejara'' and its close relatives, and Thalassodrominae, consisting of ''Thalassodromeus'' and ''Tupuxuara''. A 2011 study subsumed the family Chaoyangopterinae in as the subfamily Chaoyangopterinae, something not followed by future authors. Kellner's concept of a Tapejaridae consisting of Tapejarinae and Thalassodrominae would be the basis for numerous subsequent phylogenetic analyses.Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' 9: 4940

Various opposing studies have arose challenging Kellner's concept of Tapejaridae. The 2003 model of paleontologist David Unwin found ''Tupuxara'' and ''Thalassodromeus'' to be more distantly related to ''Tapejara'' and therefore outside of Tapejaridae, instead being related to Azhdarchidae. Later, in 2006, British paleontologists David Martill