page_title: Ornithocheiromorpha
section_title: Discoveries outside Europe
hierarchical_section_title: Ornithocheiromorpha / History of research / Discoveries outside Europe
caption_attribution_description: Comparison between Ornithocheirus simus and Tropeognathus mesembrinus. A and C Ornithocheirus simus, holotype CAMSM B54428 (Albian, Cambridge Greensand), anterior part of the rostrum A anterior view C ventral view B and D Tropeognathus mesembrinus, holotype BSP 1987 I 46 (Aptian / Albian, Romualdo Formation), anterior part of the rostrum B anterior view D ventral view. Arrows and numbers mark the position of the first pair of alveoli. Scale bar = 10 mm.
context_page_description: Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 94 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by better adapted and more advanced pterosaur species such the pteranodontids and azhdarchoids. The most diverse families of the group, the Ornithocheiridae and Anhangueridae, survived until the early Late Cretaceous with Ferrodraco, Ornithocheirus, Cimoliopterus, Lonchodectes and Lonchodraco being some of the last remaining. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.
  Ornithocheiromorphs are considered to be some of the largest animals to have ever flown.
context_section_description: Other important ornithocheiromorph discoveries include the ornithocheirids Tropeognathus and Anhanguera from the Romualdo Formation in Brazil. Tropeognathus was described with its type species, T. mesembrinus in 1987 by German paleontologist Peter Wellnhofer. The generic name is derived from Greek τρόπις, tropis, meaning "keel", and γνάθος, gnathos, meaning "jaw". The specific name is derived from Koine mesembrinos, "of the noontide", simplified as "southern", in reference to the provenance from the Southern hemisphere. The description then led to an enormous taxonomic confusion. In 1989, Brazilian paleontologist Alexander Kellner considered it an Anhanguera mesembrinus, then a Coloborhynchus mesembrinus by Veldmeijer in 1998, and then a Criorhynchus mesembrinus in 2001 by German paleontologist Michael Fastnacht. T. mesembrinus was then considered a junior synonym of Ornithocheirus simus by British paleontologist David Unwin in 2001, but he then proposed an Ornithocheirus mesembrinus in 2003. In 2013 however, Taissa Rodrigues and Alexander Kellner concluded that Tropeognathus would be valid again, and containing only T. mesembrinus, the type species. 
  The discovery of Anhanguera in 1985 made some authors assign several species of Ornithocheirus to the new genus, this is due to the similarities of Anhanguera and Ornithocheirus. One of which was Ornithocheirus cuvieri, formerly known as Pterodactylus cuvieri, was renamed by David Unwin in 2001 as Anhanguera cuvieri, though in 2013 it has been considered a synonym of Cimoliopterus. Other species of Anhanguera include A. piscator, which is known from a nearly-complete skeleton, and was once proposed to belong to the genus Coloborhynchus, but it has recently been placed back into Anhanguera by Andres and Myers (2013). Another species which is also known from a complete skeleton is A. spielbergi, which was also originally considered as a species of Coloborhynchus in 2003, though in 2018, it has been reassigned to the genus Maaradactylus, specifically to its own species, M. spielbergi.
  A discovery in Asia, specifically northwestern China, was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. The site represented a nesting colony that storm floods had covered with mud. Dozens of individuals could be secured to run into the many hundreds. In 2014, a new species was named and described: Hamipterus tianshanensis. It was named by Wang Xiaolin, Alexander Kellner, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic name Hamipterus combines that of the Hami region, with the word pteron, meaning "wing", and the specific name refers to the provenance from the Tian Shan, a mountain range. These findings represented an impressively large concentration of pterosaur fossils, and together with the Pterodaustro nesting colonies in Argentina, both are regarded as the most diverse fossil concentrations of pterosaurs ever known.
caption: Comparison between the holotypes of Tropeognathus mesembrinus and Ornithocheirus simus
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