''Ornithocheirus'' (from

Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...

"ὄρνις", meaning bird, and "χεῖρ", meaning hand) is a

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 ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

known from fragmentary fossil remains uncovered from sediments in the

United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...

and possibly

Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...

. Several species have been referred to the genus, most of which are now considered as dubious species, or members of different genera, and the genus is now often considered to include only the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

, ''Ornithocheirus simus''. Species have been referred to ''Ornithocheirus'' from the mid-

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

period of

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 ...

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

and

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 ...

, but ''O. simus'' is known mostly from the

, though a specimen referred to ''O.'' cf. ''simus'' is also known from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

Kem Kem Group The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group (stratigraphy), group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two ...

. Because ''O. simus'' was originally named based on poorly preserved fossil material, the genus ''Ornithocheirus'' has suffered enduring problems of

zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...

. Fossil remains of ''Ornithocheirus'' have been recovered mainly from the

Cambridge Greensand The Cambridge Greensand is a geological unit in England whose strata are earliest Cenomanian in age. It lies above the erosive contact between the Gault Formation and the Chalk Group in the vicinity of Cambridgeshire, and technically forms the l ...

England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...

, dating to the beginning of the

Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...

stage of the early Cretaceous period, about 110 million years ago. Additional fossils from the

Santana Formation The Santana Group is a geologic group, formerly included as the middle part of the Araripe Group, in the Araripe Basin of northeastern Brazil. The group comprises the Crato, Ipubi and Romualdo Formations and is dated to the Aptian to Albian s ...

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

are sometimes classified as species of ''Ornithocheirus'', but have also been placed in their own genera, most notably ''

Tropeognathus ''Tropeognathus'' (meaning "keel jaw") is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within a ...

''.

Discovery and naming

During the 19th century, in England many fragmentary pterosaur fossils were found in the

, a layer from the early Cretaceous, that had originated as a sandy seabed. Decomposing pterosaur cadavers, floating on the sea surface, had gradually lost individual bones that sank to the bottom of the sea. Water currents then moved the bones around, eroding and polishing them, until they were at last covered by more sand and fossilised. Even the largest of these remains were damaged and difficult to interpret. They had been assigned to the genus ''

Pterodactylus ''Pterodactylus'' (from ) is a genus of extinct pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying reptile and one of the first prehis ...

'', as was common for any pterosaur species described in the early and middle 19th century.Unwin D.M., 2001, "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England", ''Mitteilungen aus dem Museum für Naturkunde in Berlin'', Geowissenschaftliche Reihe 4: 189–221 Young researcher

Harry Govier Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fa ...

was commissioned to bring order to the pterosaur collection of the

Sedgwick Museum The Sedgwick Museum of Earth Sciences, is the geology museum of the University of Cambridge. It is part of the Department of Earth Sciences and is located on the university's Downing Site in Downing Street, central Cambridge, England. The Sedg ...

Cambridge Cambridge ( ) is a List of cities in the United Kingdom, city and non-metropolitan district in the county of Cambridgeshire, England. It is the county town of Cambridgeshire and is located on the River Cam, north of London. As of the 2021 Unit ...

. He soon concluded that it was best to create a new genus for the Cambridge Greensand material that he named ''Ornithocheirus'' (meaning "bird hand"), as he in this period still considered pterosaurs to be the direct ancestors of birds, and assumed the hand of the genus to represent a transitional stage in the evolution towards the bird hand. To distinguish the best pieces in the collection, and partly because they had already been described as species by other scientists. Between the years 1869 and 1870, Seeley each gave them a separate species name: ''O. simus'', ''O. woodwardi'', ''O. oxyrhinus'', ''O. carteri'', ''O. platyrhinus'', ''O. sedgwickii'', ''O. crassidens'', ''O. capito'', ''O. eurygnathus'', ''O. reedi'', ''O. cuvieri'', ''O. scaphorhynchus'', ''O. brachyrhinus'', ''O. colorhinus'', ''O. dentatus'', ''O. denticulatus'', ''O. enchorhynchus'', ''O. xyphorhynchus'', ''O. fittoni'', ''O. nasutus'', ''O. polyodon'', ''O. tenuirostris'', ''O. machaerorhynchus'', ''O. platystomus'', ''O. microdon'', ''O. oweni'' and ''O. huxleyi'', thus 27 in total. As yet Seeley did not designate a

. When Seeley published his conclusions in his 1870 book ''The Ornithosauria'', this provoked a reaction by the leading British paleontologist of his day, Sir

Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...

. Owen was not an evolutionist and he therefore considered the name ''Ornithocheirus'' to be inappropriate; he also thought it was possible to distinguish two main types within the material, based on differences in snout form and tooth position — the best fossils consisted of jaw fragments. In 1874, he created two new genera: ''

Coloborhynchus ''Coloborhynchus'' is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. ''Coloborhynchus'' is known from the Lower Cretaceous of Engla ...

'' and ''Criorhynchus''. ''Coloborhynchus'' (meaning "maimed beak") which comprised a new type species called ''Coloborhynchus clavirostris'', as well as two other species reassigned from ''Ornithocheirus'': ''C. sedgwickii'' and ''C. cuvieri''. ''Criorhynchus'' (meaning "ram beak") consisted entirely of former ''Ornithocheirus'' species: the type species, ''Criorhynchus simus'', and furthermore such as ''C. eurygnathus'', ''C. capito'', ''C. platystomus'', ''C. crassidens'' and ''C. reedi''. Ornithocheirus MANCH L10832

Seeley did not accept Owen's position. In 1881 he designated ''O. simus'' the type species of ''Ornithocheirus'' and named a new separate species called '' O. bunzeli''. In 1888,

Edwin Tulley Newton Edwin Tulley Newton (4 May 1840 – 28 January 1930) was a British paleontologist. Newton originally worked at handicrafts, but was able to attend Thomas Henry Huxley's lectures and by 1865, was appointed as his assistant. In 1882, he beca ...

reassigned several existing species names into ''Ornithocheirus'', which created new combinations: ''O. clavirostris'', ''O. daviesii'', ''O. sagittirostris'', ''O. validus'', ''O. giganteus'', '' O. clifti'', ''O. diomedeus'', ''O. nobilis'', ''O. curtus'', ''O. macrorhinus'' and ''O. hlavaci''. He also reassigned the species ''O. umbrosus'' and ''O. harpyia'' into ''Ornithocheirus'', which were formerly species given to the genus ''

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 ...

'' by

Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...

back in 1872. In 1914

Reginald Walter Hooley Reginald Walter Hooley (5 September 1865 – 5 May 1923) was a businessman and amateur paleontologist, collecting on the Isle of Wight. He is probably best remembered for describing the dinosaur ''Iguanodon atherfieldensis'', now ''Mantellisaurus ...

made a new attempt to structure the large number of species. Hooley synonymized Owen's ''Criorhynchus'' to ''Ornithocheirus'', in which he also sunk ''Coloborhynchus'' into that genus, meaning that the only generic name he kept was ''Ornithocheirus''. To allow for a greater differentiation, Hooley created two new genera, again based on jaw form: ''

Lonchodectes ''Lonchodectes'' (meaning "lance biter") was a genus of lonchodectid pterosaur from several formations dating to the Turonian (Late Cretaceous) of England, mostly in the area around Kent. The species belonging to it had been assigned to ''Orni ...

'' and ''

Amblydectes ''Amblydectes'' is a genus of pterosaur known from jaw fragments. It apparently had a jaw flattened towards the tip and triangular in cross-section. It has at times been synonymized with ''Coloborhynchus'', ''Criorhynchus'', ''Lonchodectes'', ...

''. The genus ''Lonchodectes'' (meaning "lance biter") consisted of the former species ''Pterodactylus compressirostris'', and ''Pterodactylus giganteus'', which were reassigned as ''Lonchodectes compressirostris'', the type species, and ''Lonchodectes giganteus'', in addition, Hooley also named a new separate species called ''L. daviesii''. The genus ''Amblydectes'' (meaning "blunt biter") also consisted of three species: ''A. platystomus'', ''A. crassidens'' and ''A. eurygnathus''. Hooley's classification however, was rarely applied later in the century, and therefore paleontologists weren't aware of it, and kept subsuming all the poorly preserved and confusing material under the name ''Ornithocheirus''. In 1964, a Russian-language overview of Pterosauria designated the species ''Lonchodectes compressirostris'', which was identified as ''Pterodactylus compressirostris'' in the overview, as the type species of ''Ornithocheirus'', which was then followed by Kuhn in 1967, and Wellnhofer in 1978, yet those authors weren't aware that back in 1881, Seeley made already made the species ''P. simus'' as the type species of ''Ornithocheirus'', in which defined the new combination of ''O. simus''. Ornithocheirus CAMSM 54429

From the seventies onwards many new pterosaur fossils were found in

in deposits slightly older than the Cambridge Greensand, 110 million years old. Unlike the English material, these new finds included some of the best preserved large pterosaur skeletons and several new genera names were given to them, such as ''

Anhanguera Anhanguera may refer to: People * Bartolomeu Bueno da Silva (1672–1740), a bandeirante also known as Anhanguera * Anhangüera I, father of Bartolomeu Bueno da Silva Places in Brazil * Anhanguera, Goiás, a municipality in the state of Goiás ...

''. This situation caused a renewed interest in the ''Ornithocheirus'' material and the validity of the several names based on it, for it might be possible that it could by more detailed studies be established that the Brazilian pterosaurs were actually

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

s of the European types. Several European researchers concluded that this was indeed the case. Unwin revived ''Coloborhynchus'' and Michael Fastnacht ''Criorhynchus'', each author ascribing Brazilian species to these genera. However, in 2000 Unwin stated that ''Criorhynchus'' could not be valid. Referring to Seeley's designation of 1881 he considered ''Ornithocheirus simus'',

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

CAMSM B.54428, to be the type species. This also made it possible to revive ''Lonchodectes'', using as type the former ''O. compressirostris'', which then became ''L. compressirostris''. As a result, though over forty species have been named in the genus ''Ornithocheirus'' over the years, only ''O. simus'' is currently considered valid by all pterosaur researchers. The species ''Tropeognathus mesembrinus'', which was named by Peter Wellnhofer in 1987, was assigned to ''Ornithocheirus'' 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 ...

in 2003, making ''Tropeognathus'' a junior synonym. In 1989 however,

Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...

considered it as an ''Anhanguera mesembrinus'', then as a ''Coloborhynchus mesembrinus'' by

André Veldmeijer André — sometimes transliterated as Andre — is the French and Portuguese form of the name Andrew and is now also used in the English-speaking world. It used in France, Quebec, Canada and other French-speaking countries, as well in Portugal, ...

in 1998 and as a ''Criorhynchus mesembrinus'' by Michael Fastnacht in 2001. Even earlier, in 2001, Unwin had referred the "Tropeognathus" material to ''O. simus'' in which was followed by Veldmeijer; however, Veldmeijer rejected ''O. simus'' as the type species in favor of ''O. compressirostris'' (alternately ''Lonchodectes''), and he used the names ''Criorhynchus simus'' and ''Criorhynchus mesembrinus'' instead.Veldmeijer, A.J. (2006).
Toothed pterosaurs from the Santana Formation (Cretaceous; Aptian-Albian) of northeastern Brazil. A reappraisal on the basis of newly described material
." Tekst. – Proefschrift Universiteit Utrecht.

Formerly assigned species

In 2013, Rodrigues and Kellner found ''Ornithocheirus'' to be monotypic, containing only ''O. simus'', and placed most other species in other genera, or declared them ''

nomina dubia In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...

''. They also considered ''O. platyrhinus'' a junior synonym of ''O. simus''. Misassigned species: *''O. compressirostris'' (Hooley, 1914) = ''Pterodactylus compressirostris'', Owen, 1851 [now classified as ''

''] *''O. crassidens'' (Seeley, 1870) = [now classified as ''

''] *''O. cuvieri'' (Seeley, 1870) = ''Pterodactylus cuvieri'', Bowerbank, 1851 ow classified as ''Cimoliopterus''">Cimoliopterus.html" ;"title="ow classified as ''Cimoliopterus">ow classified as ''Cimoliopterus''/small> *''O. curtus'' (Hooley, 1914) = ''

curtus'', Owen, 1874 *''O. giganteus'' (Richard Owen">Owen Owen may refer to: People and fictional characters * Owen (name), including a list of people and fictional characters with the given name or surname Places United States * Owen, Missouri, a ghost town * Owen, Wisconsin * Owen County, Indiana ...

Discovery and naming

Formerly assigned species

Description

Classification

See also

Notes

References