Nyctosauridae (meaning "night lizards" or "bat lizards") is a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

of specialized soaring

pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...

s of the late

Cretaceous Period The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 77.1 million years, it is the ninth and longest geologi ...

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

, and possibly other continents including

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

. It was named in 1889 by Henry Alleyne Nicholson and

Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...

.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. 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. Nyctosaurids are generally characterized as specialized,

pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...

soarers like frigatebirds; however, the '' Alcione'' species appear to have had shorter wings and possibly have been divers like some modern piscivorous birds. Nyctosaurids have occasionally been included in the similar family

Pteranodontidae The Pteranodontidae are a Family (biology), family of large pterosaurs from the Late Cretaceous of North America and possibly other continents including Europe and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a ...

, though researchers including Christopher Bennett and Alexander Kellner have both concluded that they belonged to a separate lineage. Analyses by

David Unwin David (; , "beloved one") was a king of ancient Israel and Judah and the third king of the United Monarchy, according to the Hebrew Bible and Old Testament. The Tel Dan stele, an Aramaic-inscribed stone erected by a king of Aram-Damas ...

did indicate a close relationship between ''

Pteranodon ''Pteranodon'' (; from and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...

'' and ''

Nyctosaurus ''Nyctosaurus'' (meaning "night lizard") 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 ...

'', though he used the name

Pteranodontia Pteranodontia is an extinct group of ornithocheiroid pterodactyloid pterosaurs. It lived during the Late Cretaceous (Turonian to Maastrichtian stages) of North America, South America, Europe and Africa. They were some of the most advanced pterosa ...

for the clade containing both genera. Both opinions were published before the discovery of the second definitively known nyctosaurid, ''

Muzquizopteryx ''Muzquizopteryx'' is a genus of nyctosaurid pterodactyloid pterosaur from the Late Cretaceous period (early Coniacian stage) of what is now Coahuila, Mexico. Discovery and naming In the 1990s, José Martínez Vásquez, a worker at the chalk q ...

'', in 2006.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. Most nyctosaurid fossils have been found in formations dating to the late Cretaceous period of the western

United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...

and

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

. ''Nyctosaurus'' dates from 85-84.5 million years ago, in the

Niobrara Formation The Niobrara Formation , also called the Niobrara Chalk, is a geologic formation in North America that was deposited between 87 and 82 million years ago during the Coniacian, Santonian, and Campanian stages of the Late Cretaceous. It is com ...

Kansas Kansas ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...

. ''Muzquizopteryx'' is the oldest nyctosaurid known from definitive remains, dating to the

Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...

Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by ...

boundary in

Coahuila Coahuila, formally Coahuila de Zaragoza, officially the Free and Sovereign State of Coahuila de Zaragoza, is one of the 31 states of Mexico. The largest city and State Capital is the city of Saltillo; the second largest is Torreón and the thi ...

.Schmidt, H., Buchy, M.-C., Vega, F.J., Smith, K.T., Ifrim, C., Frey, E., Keller, G., Rindfleisch, A., González, A.H.G., Lionel Cavin, L. and Stinnesbeck, W. (2006).
A new lithographic limestone deposit in the Upper Cretaceous Austin Group at El Rosario, county of Múzquiz, Coahuila, northeastern Mexico
." ''

Revista Mexicana de Ciencias Geológicas The ''Revista Mexicana de Ciencias Geológicas'' is a triannual peer-reviewed open-access scientific journal published by the National Autonomous University of Mexico, Sociedad Geológica Mexicana, Instituto Nacional de Geoquímica, and Socieda ...

'', 22(3): 401-418. A possible nyctosaurid specimen from Europe has been reclassified as an azhdarchoid. Before the late 2010s, three forms were known from the

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

: a single potentially nyctosaurid humerus (upper arm bone) from Mexico, ''"Nyctosaurus" lamegoi'' from Brazil, 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).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. 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 ''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 '' Simurghia''.

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''. 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''.

Notes

{{Taxonbar, from=Q2440698 Pteranodontia Coniacian first appearances Maastrichtian extinctions Pterosaur families