''Nicorhynchus'' (meaning "

knucker Knucker is a dialect word for a sort of water dragon, living in ''knuckerholes'' in Sussex, England. "The word comes from the Old English '' nicor'' which means "water monster" and is used in the poem ''Beowulf''. It may also be related to the w ...

snout", in reference to its likely ecology) is a genus of

anhanguerid Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is 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 ...

from the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...

period. It contains two species, the

type species In 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 species that contains the biological type specime ...

, ''N. capito'', 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 lo ...

of England, and ''N. fluviferox'' from the

Kem Kem Group The Kem Kem Group (commonly known as the Kem Kem beds) is a geological 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 geological formations ...

of Morocco. These species were previously assigned to ''

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 Engl ...

''.

History

The genus ''

'' has had a convoluted history, with many species having been assigned to the genus. In 2013, Rodrigues and Kellner considered ''Coloborhynchus'' to be monotypic, containing only ''C. clavirostris'', and placed most other species in other genera, or declared them '' nomina dubia''. One of these species was ''Coloborhynchus capito'', which was originally named by

Harry 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 fat ...

in 1870 as a species of ''

Ornithocheirus ''Ornithocheirus'' (from Ancient Greek "ὄρνις", meaning ''bird'', and "χεῖρ", meaning ''hand'') is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the UK and possibly Morocco. Several species have ...

''. Its

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

is CAMSM B 54625, a snout. In 2001, this species was moved to ''Coloborhynchus'' by

David Unwin David (; , "beloved one") (traditional spelling), , ''Dāwūd''; grc-koi, Δαυΐδ, Dauíd; la, Davidus, David; gez , ዳዊት, ''Dawit''; xcl, Դաւիթ, ''Dawitʿ''; cu, Давíдъ, ''Davidŭ''; possibly meaning "beloved one". w ...

, who also synonymized ''Ornithocheirus reedi'' (known from a lost specimen) with it. Rodrigues and Kellner recognized that the species was distinct from ''Coloborhynchus'', but did not give it a new name pending the discovery of more complete material. Nicorhynchus fluviferox

In 2018, Jacobs ''et al.'' named a new species of ''Coloborhynchus'', ''C. fluviferox'' from the Ifezouane Formation of the

Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria t ...

based on a partial snout, and also tentatively referred another specimen from the same locality to a different, unnamed species. A 2020 review of a subfamily called

Coloborhynchinae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...

by Borja Holgado and Rodrigo Pêgas moved both ''C. capito'' and ''C. fluviferox'' to a new genus, ''Nicorhynchus'', and also associated the unnamed Ifezouane Formation coloborhynchine to ''N. fluviferox''. The name ''Nicorhynchus'' is derived from the Old English ''nicor'' (

, a kind of water dragon) and the Ancient Greek ''rhynchos'' ("snout"), in reference to its likely ecology as a fish-eating, flying reptile found in river and marine deposits.

Description

Coloborhynchus fluviferox by Nick Longrich

The species ''N. capito'' represents the second largest known

(after a ''

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 ...

'' specimen), and indeed the second largest toothed pterosaur known after ''

''. A referred specimen from the

England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...

described in 2011 consists of a very large upper jaw tip which displays the tooth characteristics that distinguish ''N. capito'' from other species. The jaw tip is nearly tall and wide, with teeth up to in base diameter. If the proportions of this specimen were consistent with species of ''Coloborhynchus'', the total skull length could have been up to , leading to an estimated wingspan of .

Classification

The describers of the genus, Holgado and Pêgas, had recovered ''Nicorhynchus'' within the subfamily Coloborhynchinae, which in turn was within the family Anhangueridae, sister taxon to ''

Uktenadactylus ''Uktenadactylus'' is a genus of anhanguerid pterodactyloid pterosaurs from the Lower Cretaceous Paw Paw Formation of Texas, United States and the Wessex Formation on the Isle of Wight, England. Fossil remains of ''Uktenadactylus'' dated back to ...

''. Their cladogram is shown below.

References

{{Taxonbar, from=Q99753010 Pteranodontoids Albian life Cenomanian life Cretaceous pterosaurs of Africa Cretaceous Morocco Fossils of Morocco Fossil taxa described in 2020