page_title: Timeline of pterosaur research
section_title: 1990s
hierarchical_section_title: Timeline of pterosaur research / 20th century / 1990s
caption_attribution_description: Tupandactylus imperator (formerly classified as Tapejara imperator).
context_page_description: This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs went extinct millions of years before humans evolved, humans have coexisted with pterosaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about thunderbirds told by the Native Americans of the modern western United States may have been influenced by observations of Pteranodon fossils. These thunderbirds were said to have warred with water monsters, which agrees well with the co-occurrence of Pteranodon and the ancient marine reptiles of the seaway over which it flew.
  The formal study of pterosaurs began in the late 18th century when naturalist Cosimo Alessandro Collini of Mannheim, Germany published a description of an unusual animal with long arms, each bearing an elongated finger.
context_section_description: 1990
  Michael Everhart discovered his first Pteranodon specimen in Kansas. In life it would have had a roughly 14 foot wingspan.
  Stewart reported that Nyctosaurus and Pteranodon made their first appearances in the stratigraphic column in the middle of the Smoky Hill Chalk, which dates back to the Santonian.
  Bennett published a study of the biostratigraphy of Pteranodon. He found that the type specimen of P. sternbergi was discovered in the lower chalk, while the type specimen of P. longiceps was discovered in the upper portion of the chalk.
  1991
  Peter Wellnhofer published The Illustrated Encyclopedia of Pterosaurs. Wellnhofer's book was only the second serious book about pterosaurs ever published. In it, he argued that Othniel Charles Marsh was correct to reconstruct the tail vane of Rhamphorhynchus with a vertical orientation based on its asymmetry and also provided additional evidence for this orientation based on the shape of its tail vertebrae. Wellnhofer also observed that pterosaurs had large, birdlike brains. Wellnhofer also argued in favor of quadrupedal pterosaurs.
  Chatterjee described the new genus and species Protoavis texensis.
  1992
  Bennett published a study of sexual dimorphism in Pteranodon using 400 specimens. He found there to be two size based morphs, a larger form with a larger crest and narrow pelvis and a small form with a small crest and wide pelvis. He concluded that the larger form was male and the smaller form was female. He found that female Pteranodon outnumbered male Pteranodon by 2:1. Bennett argued that since large crests were only associated with one size morph, that it functioned purely as display. He also hypothesized that Pteranodon sternbergi was the direct ancestor of Pteranodon longiceps.
  Bennett argued that the purported pterosaur footprints reported by Gillette and Thomas were actually produced by crocodylians.
  Lockley and others reported the presence of Early Cretaceous pterosaur tracks in Spain.
  1993
  Moratalla reported the presence of Early Cretaceous pterosaur tracks in Spain.
  1994
  Bennett studied the Pteranodon remains curated by the Yale Peabody Museum. His research debunked several claims originally made by O. C. Marsh. The teeth Marsh originally referred to Pteranodon Bennett attributed to the fish genus Xiphactinus. Bennet also argued that none of the Pteranodon species Marsh named after P. longiceps were actually distinct from it. Bennett criticized conclusions drawn by Miller and Harksen as well. Bennett criticized the former's 1971 classification scheme for Pteranodon species and the latter's 1966 reconstruction of Pteranodon sternbergi as having excessively long jaws. Nyctosaurus also received some attention in this paper. Bennett concluded that N. bonneri was a junior synonym of N. gracilis.
  Unwin and Bakhurina  published a paper arguing that much of the supposed soft tissue impressions of the Sordes type specimen were not the remains of a furry body covering. Instead, they seemed to be the remains of the fibrous tissue that reinforced the wing membrane.
  Frey and Martill described the new genus and species Arthurdactylus conandoylei.
  Cai and Wei described the new genus and species Zhejiangopterus linhaiensis.
  Kellner and Campos described the new species Tupuxuara leonardii.
  Lee described the new species Coloborhynchus wadleighi.
  Logue reported the discovery of a new fossil track site featuring Pteraichnus-like traces.
  Moratalla and others reported fossil pterosaur footprints in Europe.
  1995
  Jean-Michel Mazin and others reported the discovery of footprints left behind by during the Late Jurassic in what is now Crayssac, France which they attributed to pterosaurs. This paper has been regarded as the conclusion of the controversy regarding the type of gait pterosaurs utilized on the ground.
  Lockley and others described the new ichnospecies Pteraichnus stokesi.
  Howse and A. R. Milner described the new genus and species Plataleorhynchus streptophorodon.
  Padian, de Ricqlés and Horner described the new genus and species Montanazhdarcho minor.
  The putative pterosaurian origins of
caption: Life restoration of Tapejara imperator
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