View source for Biology:Coloborhynchus

{{short description|Genus of anhanguerid pterosaur from the Cretaceous period}}
{{Automatic taxobox
| name=''Coloborhynchus''
| taxon = Coloborhynchus
| fossil_range = [[Earth:Early Cretaceous|Early Cretaceous]], {{Fossilrange|140|136|latest=93.9}}
| image = Coloborhynchus.png
| image_caption = [[Biology:Holotype|Holotype]] jaw fragment of ''C. clavirostris'' in multiple views
| authority = Owen, 1874
| type_species = {{extinct}}'''''Coloborhynchus clavirostris'''''
| type_species_authority = Owen, 1874
| subdivision_ranks = Other species
| subdivision =
*{{extinct}}''C. capito''? <br><small>(Seeley, 1870)</small>
*{{extinct}}''C. fluviferox''? <br><small>Jacobs ''et al.'', 2019</small>
*{{extinct}}''C. piscator''? <br><small>(Kellner & Tomida, 2000)</small>
*{{extinct}}''C. robustus''? <br><small>(Wellnhofer, 1987)</small>
*{{extinct}}''C. spielbergi''? <br/><small>Veldmeijer, 2003</small>
| synonyms =
{{collapsible list|title=List of synonyms|
{{collapsible list|title=Genus synonymy|
*''"Ptenodactylus"'' <br/><small>Seeley, 1869 (''[[Biology:Ptenodactylus|nomen nudum]]'')</small>
}}
{{collapsible list|title=Synonyms of ''C. capito''|
*''[[Biology:Ornithocheirus|Ornithocheirus]] capito'' <br/><small>Seeley, 1870</small>
*''[[Biology:Nicorhynchus|Nicorhynchus]] capito'' <br/><small>(Seeley, 1870)</small>
}}
{{collapsible list|title=Synonyms of ''C. fluviferox''|
*''Nicorhynchus fluviferox'' <br/><small>(Jacobs ''et al.'', 2019)</small>
}}
{{collapsible list|title=Synonyms of ''C. piscator''|
*''[[Biology:Anhanguera (pterosaur)|Anhanguera]] piscator'' <br/><small>(Kellner & Tomida, 2000)</small>
}}
{{collapsible list|title=Synonyms of ''C. robustus''|
* ''Anhanguera robustus'' <br/><small>(Wellnhofer, 1987)</small>
* ''[[Biology:Tropeognathus|Tropeognathus]] robustus'' <br/><small>Wellnhofer, 1987</small>
* ''Anhanguera piscator''? <br/><small>Kellner & Tomida, 2000</small>
}}
{{collapsible list|title=Synonyms of ''C. spielbergi''|
* ''Anhanguera spielbergi'' <br/><small>(Veldmeijer, 2003)</small>
* ''[[Biology:Maaradactylus|Maaradactylus]] spielbergi'' <br/><small>(Veldmeijer, 2003)</small>
}}
}}
}}

'''''Coloborhynchus''''' is a [[Biology:Genus|genus]] of pterodactyloid [[Biology:Pterosaur|pterosaur]] belonging to the family [[Biology:Anhangueridae|Anhangueridae]], though it has also been recovered as a member of the [[Biology:Ornithocheiridae|Ornithocheiridae]] in some studies. ''Coloborhynchus'' is known from the [[Earth:Lower Cretaceous|Lower Cretaceous]] of {{wipe|England}} ([[Earth:Valanginian|Valanginian]] age, 140 to 136 million years ago),<ref name="Rodrigues & Kellner 2013">{{Cite journal | last1 = Rodrigues | first1 = T. | last2 = Kellner | first2 = A. | doi = 10.3897/zookeys.308.5559 | title = Taxonomic review of the ''Ornithocheirus'' complex (Pterosauria) from the Cretaceous of England | journal = ZooKeys | pages = 1–112 | year = 2013 | issue = 308 | pmid =  23794925| pmc = 3689139| doi-access = free }}</ref> and depending on which species are included, possibly the [[Earth:Albian|Albian]] and [[Earth:Cenomanian|Cenomanian]] ages (113 to 93.9 million years ago) as well. ''Coloborhynchus'' was once thought to be the largest known toothed pterosaur,<ref>{{cite news |url=https://www.bbc.co.uk/nature/15150591 |work=BBC Nature |title='Ugly' fossil is largest toothed pterosaur |archive-url=https://web.archive.org/web/20111006001201/https://www.bbc.co.uk/nature/15150591 |archive-date=6 October 2011 |date=4 October 2011 |first1=Ella |last1=Davies }}</ref> however, a specimen of the closely related ''[[Biology:Tropeognathus|Tropeognathus]]'' is now thought to have had a larger [[Wingspan|wingspan]].<ref name=kellneretal2013/>

==History and classification==
[[File:Coloborhynchus.jpg|thumb|left|Lithograph of the ''C. clavirostris'' holotype]] 

Like many [[Biology:Ornithocheiroidea|ornithocheiroid]] pterosaurs named during the 19th century, ''Coloborhynchus'' has a highly convoluted history of classification. Over the years numerous species have been assigned to it, and often, species have been shuffled between ''Coloborhynchus'' and related genera by various researchers.

In 1874 Richard Owen, rejecting the creation by Harry Govier Seeley of the genus ''[[Biology:Ornithocheirus|Ornithocheirus]]'', named a species ''Coloborhynchus clavirostris'' based on [[Biology:Holotype|holotype]] BMNH 1822, a partial snout from the Hastings Beds of the [[Earth:Wealden Group|Wealden Group]] of East Sussex, {{wipe|England}}. The genus name means "maimed beak", a reference to the damaged and eroded condition of the fossil; the [[Biology:Specific name (zoology)|specific name]] means "key snout", referring to its form in cross-section. Owen also reclassified ''Ornithocheirus cuvieri'' and ''O. sedgwickii'' as species within the genus ''Coloborhycnhus'', though he did not designate any of these three as the [[Biology:Type species|type species]]. Owen considered the defining trait of the genus to be the location of the front tooth pairs high on the side of the upper jaws. However, in 1913 Reginald Walter Hooley concluded that this location was an artefact of the erosion and that the genus was indistinguishable from ''Criorhynchus simus'', the second genus and species Owen erected in 1874. Hooley also ignored Owen's reassignment of the two former ''Ornithocheirus'' species, leaving them in that genus. In 1967, Kuhn agreed with Hooley that ''Coloborhynchus clavirostris'' was a synonym of ''Criorhynchus simus''. Furthermore, Kuhn was the first to formally designate ''C. clavirostris'' as the type species of the genus, rather than one of the ''Ornithocheirus'' species. Most later researchers followed these opinions, regarding ''Coloborhynchus'' as invalid relative to ''Criorhynchus''.<ref name=rodrigues&kellner2008/><ref name="Hooley">{{cite journal |last1=Hooley |first1=R. W. |title=LXI.— On the Ornithosaurian genus ''Ornithocheirus'', with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge |journal=Annals and Magazine of Natural History |date=1914 |volume=13 |issue=78 |pages=529–557 |doi=10.1080/00222931408693521 |url=https://www.biodiversitylibrary.org/page/18641288#page/541/mode/1up}}</ref>

This changed in 1994 when Yuong-Nam Lee named ''Coloborhynchus wadleighi'' for a snout found in 1992 in the [[Earth:Albian|Albian]] age [[Earth:Paw Paw Formation|Paw Paw Formation]] {{wipe|Texas}}. The revival of the genus meant that of several related species, then assigned to other genera, had to be re-evaluated to determine whether or not they actually belonged to ''Coloborhynchus''. In 2008, Taissa Rodrigues and Alexander Kellner re-formulated the key features of ''Coloborhynchus'', again based mainly on the unique positions of the tooth sockets. Rodrigues and Kellner argued that Lee's ''C. wadleighi'', which possessed some differences in the skull and teeth from ''C. clavirostris'', and from an earlier time period, belonged in its own genus, which they named ''[[Biology:Uktenadactylus|Uktenadactylus]]''.<ref name=rodrigues&kellner2008/>

A partial lower jaw originally named ''Tropeognathus robustus'' from the Romualdo Member of the Santana Formation in {{wipe|Brazil}} was assigned to ''Coloborhynchus'' in 2001 by Fastnacht, as ''Coloborhynchus robustus''. In 2002, David Unwin supported this position, and also synonymized the more well-known species ''[[Biology:Anhanguera (pterosaur)|Anhanguera]] piscator'' with ''C. robustus''. Rodrigues and Kellner disagreed with this classification, however, noting that both did not possess the unique straightened crest beginning at the snout tip, or sideways pointed teeth, of ''C. clavirostris''. Instead, Rodrigues and Kellner regarded both ''Anhanguera robustus'' and ''Anhanguera piscator'' as valid species of ''Anhanguera''.<ref name=rodrigues&kellner2008/>

Another Brazilian species from the Romualdo Member was named ''Coloborhynchus spielbergi'' by Veldmeijer in 2003. It shares one or two characters in common with ''C. clavirostris'' (such as a flattened upper surface of the snout), though Rodrigues and Kellner regarded them as dubious, and noted that they are also present in other genera. Kellner recombined it as ''Anhanguera spielbergi'' in 2006. Similarly, Kellner excluded ''C. araripensis'' (formerly assigned to the genus ''[[Biology:Santanadactylus|Santanadactylus]]'' from the genus, based on lack of comparable diagnostic features. Unwin, in 2001, assigned the species ''[[Biology:Siroccopteryx|Siroccopteryx moroccensis]]'' to ''Coloborhynchus'', based on its similarity to ''C. wadleighi'' (type species of ''Uktenadactylus''). Kellner, who regarded ''Uktenadactylus'' as a distinct genus in 2008, also regarded ''Siroccopteryx'' as distinct, and noted that like the other species assigned to ''Coloborhynchus'', lacked its unique characteristics of the tooth row, a position also supported by Fastnacht in 2001.<ref name=rodrigues&kellner2008/>

In 2001, Unwin also reassigned the two other species from the [[Earth:Cambridge Greensand|Cambridge Greensand]] to ''Coloborhynchus'': ''C. capito'' and ''C. sedgwickii'', the second of which being one of the original members of the genus according to Richard Owen in 1874. According to Kellner, ''C. capito'' is too incomplete to fully compare to ''C. clavirostris'', and its precise classification is open to debate. He noted that ''C. sedgwicki'' does not possess the unique features of ''C. clavirostris'' (in fact it lacks a crest altogether), and may instead belong to the same genus as "''Ornithocheirus''" ''compressirostris'' (=''[[Biology:Lonchodectes|Lonchodectes]]'').<ref name=rodrigues&kellner2008/>

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''.<ref name="Rodrigues & Kellner 2013"/> In 2018, Jacobs ''et al.'' named a new species of ''Coloborhynchus'', ''C. fluviferox'' from the Ifezouane Formation of the [[Earth:Kem Kem Group|Kem Kem Group]] of Morocco based on a partial snout, and also tentatively referred another specimen from the same locality to a different, unnamed species.<ref name=J2019/> A 2020 review of Coloborhynchinae by Holgado and Pêgas moved both ''C. capito'' and ''C. fluviferox'' to a new genus, ''[[Biology:Nicorhynchus|Nicorhynchus]]'', and also referred the unnamed Ifezouane Formation coloborhynchine to ''N. fluviferox''.<ref name="holgado2020">{{cite journal |last1=Holgado |first1=B. |last2=Pêgas |first2=R.V. |year=2020 |title=A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae |journal=Acta Palaeontologica Polonica |volume=65 |doi=10.4202/app.00751.2020 |doi-access=free }}</ref> However, a 2023 review of Kem Kem pterosaurs found the traits that distinguish ''Nicorhynchus'' from ''Coloborhynchus'' to be subtle enough to justify their synonymy, stating that the material was damaged and fragmentary enough to support this.<ref>{{Cite journal |last1=Smith |first1=Roy E. |last2=Ibrahim |first2=Nizar |last3=Longrich |first3=Nicholas |last4=Unwin |first4=David M. |last5=Jacobs |first5=Megan L. |last6=Williams |first6=Cariad J. |last7=Zouhri |first7=Samir |last8=Martill |first8=David M. |date=2023-02-04 |title=The pterosaurs of the Cretaceous Kem Kem Group of Morocco |journal=PalZ |volume=97 |issue=3 |pages=519–568 |language=en |doi=10.1007/s12542-022-00642-6 |issn=1867-6812|doi-access=free }}</ref>

===List of species and synonyms===
[[File:Ornithocheirus capito.jpg|upright|thumb|Holotype and referred specimen of the possible species ''C. capito'']]
Species which have been assigned to ''Coloborhynchus'' by various scientists over the years include:
*''C. clavirostris'' Owen, 1874, the [[Biology:Type species|type species]]
*?''C. ligabuei'' (Dalla Vecchia, 1993) = ''[[Biology:Cearadactylus|Cearadactylus]] ligabuei'' Dalla Vecchia, 1993 <small>[also classified as ''Anhanguera ligabuei'']</small>
*?''C. piscator''<ref name=spielbergi/> (Kellner & Tomida, 2000) = ''[[Biology:Anhanguera (pterosaur)|Anhanguera piscator]]'' Kellner & Tomida, 2000 <small>[also classified as ''Anhanguera'' or ''C. robustus'']</small>
Species assigned to ''Coloborhynchus'' in the past include:
*''C. cuvieri'' (Bowerbank, 1851) = ''[[Biology:Ornithocheirus|Ornithocheirus]] sedgwicki'' (Newton, 1888) = ''Pterodactylus cuvieri'' Bowerbank, 1851 <small>[now classified as ''[[Biology:Cimoliopterus|Cimoliopterus]]'']</small>
*''C. sedgwicki'' (Owen 1859) = ''[[Biology:Ornithocheirus|Ornithocheirus]] sedgwicki'' (Newton, 1888) = ''Pterodactylus sedgwickii'' Owen, 1859 <small>[now classified as ''[[Biology:Camposipterus|Camposipterus]]'']</small>
*''C. araripensis''<ref name=spielbergi/> (Wellnhofer, 1985) = ''Santanadactylus araripensis'' Wellnhofer, 1985 <small>now seen as a ''nomen dubium''</small><ref>{{cite journal|last1=Pinheiro|first1=Felipe L.|last2=Rodrigues|first2=Taissa|date=4 May 2017|title=''Anhanguera'' taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?|journal=PeerJ|volume=5|pages=e3285|doi=10.7717/peerj.3285|pmc=5420195|pmid=28484676 |doi-access=free }}</ref>
*?''C. robustus'' (Wellnhofer, 1987) = ''Tropeognathus robustus'' Wellnhofer, 1987 <small>[now seen as a ''nomen dubium'']</small>
*''[[Biology:Uktenadactylus|C. wadleighi]]'' Lee, 1994 <small>[also classified as ''[[Biology:Uktenadactylus|Uktenadactylus]]'']</small>
*''C. moroccensis'' (Mader & Kellner, 1999) = ''[[Biology:Siroccopteryx|Siroccopteryx moroccensis]]'' Mader & Kellner 1999
*''C. piscator''<ref name=spielbergi/> (Kellner & Tomida, 2000) = ''[[Biology:Anhanguera (pterosaur)|Anhanguera piscator]]'' Kellner & Tomida, 2000
*''C. spielbergi''<ref name="spielbergi">{{cite journal|last1=Veldmeijer|first1=A. J.|date=2002|title=Description of ''Coloborhynchus spielbergi'' sp. nov. (Pterodactyloidea) from the Albian (Lower Cretaceous) of Brazil|url=http://resolver.tudelft.nl/uuid:4d3f32fa-d5ec-4ca2-9f57-42e487cfdb3a|journal=Scripta Geology|volume=125|issue=3003|pages=35–139}}</ref> Veldmeijer, 2003 <small>[now classified as ''[[Biology:Maaradactylus|Maaradactylus]]'']</small>
*''C. capito'' (Seeley, 1870) = ''[[Biology:Ornithocheirus|Ornithocheirus]] capito'' Seeley, 1870 = "Ptenodactylus capito" Seeley, 1869 <small>[now classified as ''[[Biology:Nicorhynchus|Nicorhynchus]]'']</small>
*''C. fluviferox'' Jacobs, Martill, Ibrahim & Longrich, 2018<ref name=J2019>{{cite journal|last1=Jacobs |first1=M.L. |last2=Martill |first2=D.M. |last3=Ibrahim |first3=N. |last4=Longrich |first4=N. |year=2019 |title=A new species of ''Coloborhynchus'' (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa |journal=Cretaceous Research |volume=95 |pages=77–88 |doi=10.1016/j.cretres.2018.10.018 |s2cid=134439172 |url=https://researchportal.port.ac.uk/portal/files/12269480/A_new_species_of_Coloborhynchus.pdf }}</ref> <small>[now classified as ''[[Biology:Nicorhynchus|Nicorhynchus]]'']</small>

==Description==
[[File:Coloborhynchus clavirostris holotype with reconstructed margin.jpg|thumb|left|1914 illustration of the ''C. clavirostris'' holotype with dotted lines reconstructing worn away jaw margin]]
The type specimen of ''Coloborhynchus'' is known only from a partial upper jaw. Therefore, according to Rodrigues and Kellner's 2008 re-evaluation on ''Coloborhynchus clavirostris'', it can only be differentiated from its relatives based on its unique combination of tooth socket positions. In ''Coloborhynchus'', the two front teeth pointed forward and were higher on the jaw than the other teeth, while the next three pairs of teeth pointed to the sides. The final two (preserved) pairs of teeth pointed downward. Finally, a unique oval depression was located below the first pair of teeth.<ref name="rodrigues&kellner2008">{{cite journal|last1=Rodrigues|first1=Taiss|last2=Kellner|first2=Alexander W. A.|year=2008|title=Review of the pterodactyloid pterosaur ''Coloborhynchus''|url=https://epub.ub.uni-muenchen.de/12017/1/zitteliana_2008_b28_15.pdf|journal=Zitteliana|volume=28|pages=219–228}}</ref>

Like the related ''[[Biology:Anhanguera (pterosaur)|Anhanguera]]'' and ''[[Biology:Uktenadactylus|Uktenadactylus]]'', the tip of the snout flared out into a wider rosette, in contrast to the narrow posterior jaws. However, whereas the rosettes of species typically assigned to ''Anhanguera'' were rounded and spoon-shaped, those of ''Coloborhynchus'' were robust and box-shaped.<ref name=J2019/>

Also like its close relatives, ''Coloborhynchus'' had a keel-shaped crest on the front of its jaws, though it was broad and thinned from base to top, rather than the uniformly thin crests of its relatives. This kind of thickened crest is also seen in ''[[Biology:Siroccopteryx|Siroccopteryx moroccensis]]'', which may be its closest relative or a member of the same genus. It also had a straight, rather than curved, front margin, unlike its relatives, and begins at the tip of the snout, rather than further back as in other species.<ref name=rodrigues&kellner2008/>

A second specimen showing all of these same unique features was reported to Brazilian paleontologist Alexander Kellner by Darren Naish in 2007, and likely represents a second specimen of ''C. clavirostris'', though it has not yet been described.<ref name=rodrigues&kellner2008/>
[[File:Coloborhynchus fluviferox by Nick Longrich.jpg|thumb|upright|Life restoration of the possible species ''C. fluviferox'']]
The possible species ''Coloborhynchus capito'' represents the second largest known ornithocheirid (after a ''[[Biology:Tropeognathus|Tropeognathus]]'' specimen),<ref name="kellneretal2013">{{cite journal|last1=Kellner|first1=Alexander W. A.|last2=Campos|first2=Diogenes A.|last3=Sayão|first3=Juliana M.|last4=Saraiva|first4=Antônio A.F.|last5=Rodrigues|first5=Taissa|last6=Oliveira|first6=Gustavo|last7=Cruz|first7=Lilian A.|last8=Costa|first8=Fabiana R.|last9=Silva|first9=Helder P.|last10=Ferreira|first10=Jennyfer S.|date=March 2013|title=The largest flying reptile from Gondwana: a new specimen of ''Tropeognathus'' cf. ''T. mesembrinus'' Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil|journal=Anais da Academia Brasileira de Ciências|volume=85|issue=1|pages=113–135|doi=10.1590/S0001-37652013000100009|pmid=23538956|doi-access=free}}</ref> and indeed the largest toothed pterosaur known. A referred specimen from the [[Earth:Cambridge Greensand|Cambridge Greensand]] of {{wipe|England}} described in 2011 consists of a very large upper jaw tip which displays the tooth characteristics that distinguish ''C. capito'' from other species. The jaw tip is nearly 10&nbsp;cm tall and 5.6&nbsp;cm wide, with teeth up to 1.3&nbsp;cm in base diameter. If the proportions of this specimen were consistent with other known species of ''Coloborhynchus'', the total skull length could have been up to 75&nbsp;cm, leading to an estimated wingspan of {{convert|7|m|ft}}.<ref name=martill&unwin2011>{{cite journal |last1=Martill |first1=David M. |last2=Unwin |first2=David M. |title=The world's largest toothed pterosaur, NHMUK R481, an incomplete rostrum of ''Coloborhynchus capito'' (Seeley, 1870) from the Cambridge Greensand of England |journal=Cretaceous Research |date=April 2012 |volume=34 |pages=1–9 |doi=10.1016/j.cretres.2011.09.003 }}</ref> However, this species may belong to a different genus.<ref name="Rodrigues & Kellner 2013"/>

A rostrum fragment diagnostic to ''Coloborhynchus'' sp. is known from the Wessex Formation of England.<ref name="Martill 2015">{{cite journal |last1=Martill |first1=David M. |title=First occurrence of the pterosaur Coloborhynchus (Pterosauria, Ornithocheiridae) from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England |journal=Proceedings of the Geologists' Association |date=June 2015 |volume=126 |issue=3 |pages=377–380 |doi=10.1016/j.pgeola.2015.03.004 |url=https://researchportal.port.ac.uk/portal/files/3567035/First_occurrence_of_the_pterosaur.pdf }}</ref>

==Classification==
A topology made by Andres and Myers in 2013 placed the species ''Coloborhynchus clavirostris'' and ''C. wadleighi'' (now generally considered as ''Uktenadactylus'') within the family Ornithocheiridae as sister taxa to ''Ornithocheirus simus'', and the family itself is placed within the more inclusive clade [[Biology:Ornithocheirae|Ornithocheirae]].<ref name=LoneStarPterosaurs>{{cite journal |last1=Andres |first1=Brian |last2=Myers |first2=Timothy S. |title=Lone Star Pterosaurs |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |date=September 2012 |volume=103 |issue=3–4 |pages=383–398 |doi=10.1017/S1755691013000303 |s2cid=84617119 }}</ref> Later, in 2019, Pentland ''et al.'' recovered a similar result to the one by Andres and Myers in 2013 where ''Coloborhynchus'' is the sister taxon of ''Ornithocheirus''. Their cladogram is shown on the left.<ref name="Pentland2019">{{Cite journal|last1=Pentland|first1=Adele H.|last2=Poropat|first2=Stephen F.|last3=Tischler|first3=Travis R.|last4=Sloan|first4=Trish|last5=Elliott|first5=Robert A.|last6=Elliott|first6=Harry A.|last7=Elliott|first7=Judy A.|last8=Elliott|first8=David A.|date=December 2019|title=''Ferrodraco lentoni'' gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia|journal=Scientific Reports|volume=9|issue=1|pages=13454|doi=10.1038/s41598-019-49789-4|pmid=31582757|pmc=6776501|bibcode=2019NatSR...913454P }}</ref> In 2018, Jacobs ''et al.'' also recovered a similar cladogram compared to Andres and Myers (2013), where both ''Coloborhynchus'' and ''Siroccopteryx'' were assigned to the Ornithocheiridae, however, unlike the study made by Andres and Myers, they used the name ''Uktenadactylus wadleighi'' to refer to ''C. wadleighi'' in their analysis. They published their conclusion in 2019.<ref name=J2019/> However, many subsequent analyses made in 2019 and 2020 have recovered ''Coloborhynchus'' within the family [[Biology:Anhangueridae|Anhangueridae]], more specifically within the subfamily Coloborhynchinae.<ref name="Holgado2019">Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' '''9''': 4940 {{doi|10.1038/s41598-019-41280-4}}</ref><ref name="Mimodactylus">{{cite journal |last1=Kellner |first1=Alexander W. A. |last2=Caldwell |first2=Michael W. |last3=Holgado |first3=Borja |last4=Vecchia |first4=Fabio M. Dalla |last5=Nohra |first5=Roy |last6=Sayão |first6=Juliana M. |last7=Currie |first7=Philip J. |title=First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity |journal=Scientific Reports |date=2019 |volume=9 |issue=1 |page=17875 |doi=10.1038/s41598-019-54042-z|pmid=31784545 |pmc=6884559 |bibcode=2019NatSR...917875K |doi-access=free }}</ref><ref name="Targaryendraco">{{cite journal |last1=Pêgas |first1=Rodrigo V. |last2=Holgado |first2=Borja |last3=Leal |first3=Maria Eduarda C. |title=On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids |journal=Historical Biology |date=2021 |volume=33 |issue=8 |pages=1266–1280 |doi=10.1080/08912963.2019.1690482|s2cid=209595986 }}</ref> The cladogram on the left shows a phylogenetic analysis made by Borja Holgado and Rodrigo Pêgas in 2020.<ref name="holgado2020"/>

[[File:Coloborhynchus clavirostris.jpg|thumb|right|''C. clavirostris'' holotype with interpretative line drawings]] 
{{col-begin|width=80%}}
{{col-2}}
'''Topology 1:''' Pentland ''et al.'' (2019).<ref name="Pentland2019"/>
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                                    |label1=[[Biology:Anhangueridae|Anhangueridae]]
|1={{clade
|1=''[[Biology:Liaoningopterus|Liaoningopterus]]''
|2=''[[Biology:Anhanguera (pterosaur)|Anhanguera]]''
}}
                                    |label2=[[Biology:Ornithocheiridae|Ornithocheiridae]]
|2={{clade
|1=''[[Biology:Tropeognathus|Tropeognathus]]''
|label2=Ornithocheirinae
|2={{clade
|1='''''Coloborhynchus'''''
|2={{clade
|1=''[[Biology:Ornithocheirus|Ornithocheirus]]''
|2={{clade
|1=''[[Biology:Ferrodraco|Ferrodraco]]''
|2=''[[Biology:Mythunga|Mythunga]]''
}} }} }} }} }} }} }} }} }} }}
{{col-2}}
'''Topology 2:''' Holgado & Pêgas (2020).<ref name="holgado2020"/>
{{clade|style=font-size:90%; line-height:90%
| label1=[[Biology:Anhangueridae|Anhangueridae]]
| 1={{clade
| label1=[[Biology:Tropeognathinae|Tropeognathinae]]
| 1={{clade
| 1={{clade
| 1=''[[Biology:Siroccopteryx|Siroccopteryx]]''
| 2=''[[Biology:Tropeognathus|Tropeognathus]]''
}}
| 2={{clade
| 1=''[[Biology:Mythunga|Mythunga]]''
| 2=''[[Biology:Ferrodraco|Ferrodraco]]''
}}
}}
| label2=Coloborhynchinae
| 2={{clade
| 1=''[[Biology:Aerodraco|Aerodraco]]''
| 2={{clade
| 1='''''Coloborhynchus'''''
| 2={{clade
| 1=''[[Biology:Nicorhynchus|Nicorhynchus]]''
| 2=''[[Biology:Uktenadactylus|Uktenadactylus]]''
}}
}}
}}
| label3=Anhanguerinae
| 3={{clade
| 1={{clade
| 1=''[[Biology:Caulkicephalus|Caulkicephalus]]''
| 2=''[[Biology:Guidraco|Guidraco]]''
| 3=''[[Biology:Ludodactylus|Ludodactylus]]''
}}
| 2={{clade
| 1=''[[Biology:Liaoningopterus|Liaoningopterus]]''
| 2={{clade
| 1=''[[Biology:Cearadactylus|Cearadactylus]]''
| 2=''[[Biology:Maaradactylus|Maaradactylus]]''
}}
| 3=''[[Biology:Anhanguera (pterosaur)|Anhanguera]]''
}}
}}
}}
}}
{{col-end}}

==Paleobiology==
Similar to most anhanguerids, ''Coloborhynchus'' is traditionally thought to be an oceanic piscivore.<ref>{{cite conference |last1=Myers |first1=Timothy S. |title=Diet of ornithocheiroid pterosaurs inferred from stable carbon isotope analysis of tooth enamel |book-title=Abstracts with Programs |volume=49 |publisher=Geological Society of America |conference=GSA Annual Meeting in Seattle, Washington, USA |date=2017 |page=305496 |doi=10.1130/abs/2017AM-305496 }}</ref> However one study found it among the carnivorous pterosaur taxa, as opposed to several other piscivore and insectivore species.<ref>{{cite journal |last1=Bestwick |first1=Jordan |last2=Unwin |first2=David M. |last3=Butler |first3=Richard J. |last4=Purnell |first4=Mark A. |title=Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis |journal=Nature Communications |date=28 October 2020 |volume=11 |issue=1 |page=5293 |doi=10.1038/s41467-020-19022-2 |pmid=33116130 |pmc=7595196 |bibcode=2020NatCo..11.5293B |doi-access=free }}</ref>

==See also==
* [[Biology:List of pterosaur genera|List of pterosaur genera]]
* [[Biology:Timeline of pterosaur research|Timeline of pterosaur research]]

==References==
{{Reflist}}
[https://web.archive.org/web/20090214101614/http://www.pterosaur.co.uk/species/LCP/LCP.htm The Pterosaur Database (multiple species, scroll down)]
{{Taxonbar|from=Q286251}}

[[Category:Pteranodontoids]]

{{Sourceattribution|Coloborhynchus}}