Coloborhynchus
Coloborhynchus | |
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Holotype jaw fragment of C. clavirostris in multiple views | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | †Pterosauria |
Suborder: | †Pterodactyloidea |
Clade: | †Ornithocheirae |
Genus: | †Coloborhynchus Owen, 1874 |
Type species | |
†Coloborhynchus clavirostris Owen, 1874
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Other species | |
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Synonyms | |
List of synonyms
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Coloborhynchus is a
History and classification
Like many 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
This changed in 1994 when
A partial lower jaw originally named
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
In 2001, Unwin also reassigned the two other species from the 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 (=Lonchodectes).[4]
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.
List of species and synonyms
Species which have been assigned to Coloborhynchus by various scientists over the years include:
- C. clavirostris Owen, 1874, the type species
- ?C. ligabuei (Dalla Vecchia, 1993) = Cearadactylus ligabuei Dalla Vecchia, 1993 [also classified as Anhanguera ligabuei]
- ?C. piscator[9] (Kellner & Tomida, 2000) = Anhanguera piscatorKellner & Tomida, 2000 [also classified as Anhanguera or C. robustus]
- C. cuvieri (Bowerbank, 1851) = Ornithocheirus cuvieri (Newton, 1888) = Pterodactylus cuvieri Bowerbank, 1851 [now classified as Cimoliopterus]
- C. sedgwicki (Owen 1859) = Ornithocheirus sedgwicki (Newton, 1888) = Pterodactylus sedgwickii Owen, 1859 [now classified as Aerodraco]
- ?C. robustus(Wellnhofer, 1987) = Tropeognathus robustus Wellnhofer, 1987 [now seen as a nomen dubium]
- C. wadleighi Lee, 1994 [also classified as Uktenadactylus]
- C. moroccensis (Mader & Kellner, 1999) = Siroccopteryx moroccensis Mader & Kellner 1999
- C. piscator[9] (Kellner & Tomida, 2000) = Anhanguera piscatorKellner & Tomida, 2000
- C. spielbergi[9] Veldmeijer, 2003 [now classified as Maaradactylus]
- C. capito (Seeley, 1870) = Ornithocheirus capito Seeley, 1870 = "Ptenodactylus capito" Seeley, 1869 [now classified as Nicorhynchus]
- C. fluviferox Jacobs, Martill, Ibrahim & Longrich, 2018[6] [now classified as Nicorhynchus]
Description
The
Like the related Anhanguera and 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.[6]
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 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.[4]
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.[4]
The possible species Coloborhynchus capito represents the second largest known ornithocheirid (after a Tropeognathus specimen),[3] and indeed the largest toothed pterosaur known. A referred specimen from the Cambridge Greensand of 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 cm tall and 5.6 cm wide, with teeth up to 1.3 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 cm, leading to an estimated wingspan of 7 metres (23 ft).[11] However, this species may belong to a different genus.[1]
A rostrum fragment diagnostic to Coloborhynchus sp. is known from the Wessex Formation of England.[12]
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 Ornithocheirae.[13] 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.[14] 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.[6] However, many subsequent analyses made in 2019 and 2020 have recovered Coloborhynchus within the family Anhangueridae, more specifically within the subfamily Coloborhynchinae.[15][16][17] The cladogram on the left shows a phylogenetic analysis made by Borja Holgado and Rodrigo Pêgas in 2020.[7]
Topology 1: Pentland et al. (2019).[14]
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Topology 2: Holgado & Pêgas (2020).[7]
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Paleobiology
Similar to most anhanguerids, Coloborhynchus is traditionally thought to be an oceanic piscivore.[18] However one study found it among the carnivorous pterosaur taxa, as opposed to several other piscivore and insectivore species.[19]
See also
References
- ^ PMID 23794925.
- ^ Davies, Ella (4 October 2011). "'Ugly' fossil is largest toothed pterosaur". BBC Nature. Archived from the original on 6 October 2011.
- ^ PMID 23538956.
- ^ a b c d e f g h Rodrigues, Taiss; Kellner, Alexander W. A. (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana. 28: 219–228.
- .
- ^ S2CID 134439172.
- ^ .
- ISSN 1867-6812.
- ^ a b c d Veldmeijer, A. J. (2002). "Description of Coloborhynchus spielbergi sp. nov. (Pterodactyloidea) from the Albian (Lower Cretaceous) of Brazil". Scripta Geology. 125 (3003): 35–139.
- PMID 28484676.
- .
- .
- S2CID 84617119.
- ^ PMID 31582757.
- PMID 31784545.
- S2CID 209595986.
- .
- PMID 33116130.