Nyctosaurids are characterized by their lack of all but the wing finger. In most pterosaurs, the hand has four fingers, with the fourth elongated to support the wing, and the remaining three are usually small, clawed, and used in walking or climbing. The lack of functional fingers in nyctosaurids may suggest that they spent almost all of their time in the air, rarely walking on the ground. Nyctosaurids also possessed a distinctively enlarged crest for muscle attachment on their upper arm bone, or humerus, the deltopectoral crest, hatchet shaped like in the unrelated rhamphorhynchids.[3] Nyctosaurids are generally characterized as specialized, pelagic soarers like frigatebirds; however, the Alcione species appear to have had shorter wings and possibly have been divers like some modern piscivorous birds.[4]
Nyctosaurids have occasionally been included in the similar family Pteranodontidae, though researchers including Christopher Bennett and Alexander Kellner have both concluded that they belonged to a separate lineage.[5] Analyses by David Unwin did indicate a close relationship between Pteranodon and Nyctosaurus, though he used the name Pteranodontia for the clade containing both genera. Both opinions were published before the discovery of the second definitively known nyctosaurid, Muzquizopteryx, in 2006.[6]
Most nyctosaurid fossils have been found in formations dating to the late Cretaceous period of the western United States and Mexico. Nyctosaurus dates from 85-84.5 million years ago, in the Niobrara Formation of Kansas. Muzquizopteryx is the oldest nyctosaurid known from definitive remains, dating to the Turonian-Coniacian boundary in Coahuila.[7] A possible nyctosaurid specimen from Europe has been reclassified as an azhdarchoid.[8]
Before the late 2010s, three forms were known from the Maastrichtian: a single potentially nyctosaurid humerus (upper arm bone) from Mexico, "Nyctosaurus" lamegoi from Brazil,[9][10] and a nyctosaurid complete wing-phalanx1, a claw (digit phalanx manus), and a partial ulna from Jordan. The Jordan specimen is of particular interest as it is the first record of a nyctosaurid from the Old World and represents the latest record of the family (uppermost Maastrichtian).[11] Beginning in 2016, Nicholas Longrich, David Martill, and Brian Andres presented evidence of several nyctosaurid and pteranodontid species from the latest Maastrichtian age of north Africa, suggesting that these lineages went through an evolutionary radiation in the Old World shortly before the K-Pg extinction event. Three of these pterosaurs were named in 2018, and were called Alcione, Barbaridactylus, and Simurghia.[4]
Classification
In 2022, Fernandes et al. described Epapatelo as a new pteranodontian from Angola. Including Epapatelo in the phylogenetic analysis of Longrich et al. (2018), they recovered a new clade, Aponyctosauria, composed of the Nyctosauridae, Alcione, Simurghia, and Epapatelo.[12]
In 2024, Alcione, Simurghia, and Epapatelo were referred to as members of Nyctosauridae based on phylogenetic analysis, and "N." lamegoi was included as a species of Simurghia.[1]
↑ Nicholson, H.A. and Lydekker, R. (1889). A manual of palaeontology for the use of students: with a general introduction on the principles of palæontology, Volume II. Blackwood, 1889.
↑ Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN0691150613,
↑ Bennett, S. C. (1994). "Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea)", Occasional Papers of the Museum of Natural History, University of Kansas, Lawrence, 169: 1-70
↑ Frey, E., Buchy, M.-C., Stinnesbeck, W., González, A. G. & di Stefano, A. (2006). "Muzquizopteryx coahuilensis n.g., n. sp., a nyctosaurid pterosaur with soft tissue preservation from the Coniacian (Late Cretaceous) of northeast Mexico (Coahuila)." Oryctos, 6: 19-39.
↑ Price, L. I. 1953. A presença de Pterosáuria no Cretáceo superior do Estada da Paraiba. Divisão de Geologia e Mineralogia Notas Preliminares e Estudos, 71, 1-10.
↑ Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN0691150613,
↑ Kaddumi H. F. 2009. On the remains of the first pterosaur (Ornithocheiroidea:Nyctosauridae) from the Muwaqqar Chalk Marl Formation of Harrana. In: Fossils of the Harrana Fauna and the Adjacent Areas. Publications of the Eternal River Museum of Natural History, Amman, pp 241-247.
The Maastrichtian is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from 72.1 to 66 million years ago. The Maastrichtian was preceded by the Campanian and succeeded by the Danian.
Nyctosaurus is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time Nyctosaurus was alive, was covered in an extensive shallow sea. Some remains belonging to a possible Nyctosaurus species called "N." lamegoi have been found in Brazil, though it likely belongs to a different genus Simurghia. The genus Nyctosaurus has had numerous species referred to it, though how many of these may actually be valid requires further study. At least one species possessed an extraordinarily large antler-like cranial crest.
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head. The spectacularly-crested Nyctosaurus is sometimes included in this family, though usually placed in its own family, the Nyctosauridae.
Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.
Beipiaopterus is a genus of ctenochasmatid pterosaur from the Early Cretaceous period of what is now the Yixian Formation of the China.
Muzquizopteryx is a genus of nyctosaurid pterodactyloid pterosaur from the Late Cretaceous period of what is now Coahuila, Mexico.
Kepodactylus is an extinct genus of ctenochasmatid pterodactyloid pterosaur from the Kimmeridgian-Tithonian-age Upper Jurassic Morrison Formation of Colorado, United States.
Ornithocheiroidea is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods, with fossil remains found all over the world except Antarctica.
Cretornis is a pterosaur genus from the late Cretaceous period of what is now the Jizera Formation in the Czech Republic, dating to about 92 million years ago. It only contains a single species, Cretornis hlavaci.
Elanodactylus is a genus of ctenochasmatid pterodactyloid pterosaur from the Early Cretaceous period of what is now the Yixian Formation of Liaoning, China.
Pteranodontia is an extinct group of ornithocheiroid pterodactyloid pterosaurs that lived during the Late Cretaceous period of North America and Africa. They were some of the most advanced pterosaurs, and possessed highly specialized cranial crests that may have served as sexual attraction, with males having a much larger crest.
Eupterodactyloidea is an extinct group of pterodactyloid pterosaurs that existed from the latest Late Jurassic to the latest Late Cretaceous periods. Eupterodactyloids have been found on all continents except Antarctica.
Pteranodontoidea is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous of Asia, Africa, Europe, North America and South America. It was named by Alexander Wilhelm Armin Kellner in 1996. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Anhanguera, Pteranodon and all its descendants. The clade Ornithocheiroidea is sometimes considered to be the senior synonym of Pteranodontoidea, however it depends on its definition. Brian Andres in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.
Alamodactylus is an extinct genus of pteranodontian pterosaur known from the Late Cretaceous of Texas, southern United States. It contains a single species, Alamodactylus byrdi.
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.
Tethydraco is a genus of pterodactyloid pterosaur that lived during the Late Cretaceous period of what is now the area of present Morocco, about 66 million years ago. Tethydraco was originally assigned to the family Pteranodontidae. Some researchers argued that subsequently described material suggests that it may have been an azhdarchid, and possibly synonymous with Phosphatodraco, though this has been disputed. The type and only species is T. regalis.
Alcione is a genus of pterosaur from the Ouled Abdoun Basin of Morocco, dating back to the Maastrichtian stage of the Late Cretaceous period. Only one species, A. elainus, is known. This pterosaur lived in a marine environment alongside several other pterosaurs, including Simurghia and Barbaridactylus.
Barbaridactylus is a nyctosaurid pterosaur from the Ouled Abdoun Basin of Morocco, a basin that dates back to the Maastrichtian stage of the Late Cretaceous. It was published in 2018 by paleontologists Nicholas R. Longrich, David M. Martill, and Brian Andres. In the same publication, two other pterosaurs from the same basin were described: Alcione and Simurghia. The type and only species is B. grandis.
Simurghia is a genus of pterosaur from the Ouled Abdoun Basin of Morocco, a basin that dates to the Maastrichtian stage of the Late Cretaceous period, about 66 million years ago. It was published in 2018 by paleontologists Nicholas R. Longrich, David M. Martill, and Brian Andres, along with two other pterosaurs from the same basin: Alcione and Barbaridactylus. The type species is S. robusta. A possible second species, S. ("Nyctosaurus") lamegoi, is known from the Campanian-Maastrichtian age strata of Brazil.
Epapatelo is a genus of pteranodontian pterosaur from the late Cretaceous Mocuio Formation of Namibe Province, Angola. The genus contains a single species, E. otyikokolo, known from partial left limb bones.
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