Santanadactylus
Santanadactylus is a potentially dubious genus of pterodactyloid pterosaur from the Albian-age Romualdo Member of the Upper Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará State, Brazil. Four species have been named, but today are not considered congeneric with each other. It was a rather large pterosaur.
History and species
The genus was named in 1980 by the Dutch paleontologist Paul de Buisonjé. The type species is S. brasilensis, the specific name referring to Brazil. It is based on holotype UvA M 4894, an upper part of the right humerus and a right scapulacoracoid. RGM.631745, consisting of two cervical vertebrae from a different individual, was assigned as a paratype, referred because it was found in the same lot of 25 chalk nodules bought from collectors. However, Thomas and McDavid recovered the paratype RGM.631745 within Azhdarchomorpha in 2025, suggesting that specimen cannot be confidently referred to Santanadactylus. Additional remains, including a notarium were subsequently referred to the taxon by Wellnhofer et al. and Wellnhofer.In 1985, Peter Wellnhofer, a German paleontologist who has written numerous scientific publications on pterosaurs, named three additional species: S. araripensis, S. pricei, and S. spixi. S. araripensis, named after the Araripe Plateau, was a large species based on BSP 1982 I 89, remains including a partial skull and arms; the preserved skull section had no crest. S. pricei, named after Llewellyn Ivor Price, was the smallest of the three species; it was based on BSP 1980 I 122, a left wing from the elbow down, and additional arm material has been referred to it over the years. S. spixi, intermediate in size, was based on BSP 1980 I 121, another left wing, the name honoring Johann Baptist von Spix.
Over the years, the species of this taxon have been reassessed. Chris Bennett suggested that the hypodigm of S. brasilensis was a chimera of a pteranodontid and something else, S. araripensis and S. pricei were pteranodontids, and S. spixi was a dsungaripterid. Wellnhofer removed S. spixi from the genus as well. Kellner and Campos agreed that S. spixi was not congeneric with S. brasiliensis, but suggested that it was a tapejarid. Unwin proposed that "Santanadactylus" spixi was a species of Tupuxuara based on comparison with specimens of this genus. Averianov considered Santanadactylus spixi a nomen dubium probably synonymous with Tupuxuara longicristatus, and he also indicated that the paratype of S. brasiliensis was likely referable to T. longicristatus. Kellner renamed S. araripensis to Anhanguera araripensis, followed by Wang et al., though Veldmeijer included it in Coloborhynchus. Recent study, however, considers S. araripensis dubious.