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Over a decade after the last major review of the Cambridge Greensand pterosaurs, their systematics remains one of the most disputed points in pterosaur taxonomy. Ornithocheiridae is still a wastebasket for fragmentary taxa, and some nomenclatural issues are still a problem. Here, the species from the Cretaceous of England that, at some point, were referred in Ornithocheirus, are reviewed. Investigation of the primary literature confirmed that Criorhynchus should be considered an objective junior synonym of Ornithocheirus. Taxonomic review of more than 30 species known from fragmentary remains showed that 16 of them are undiagnosable (nomina dubia): Palaeornis cliftii, Cimoliornis diomedeus, Pterodactylus compressirostris, Pterodactylus fittoni, Pterodactylus woodwardi, Ornithocheirus brachyrhinus, Ornithocheirus carteri, Ornithocheirus crassidens, Ornithocheirus dentatus, Ornithocheirus enchorhynchus, Ornithocheirus eurygnathus, Ornithocheirus oxyrhinus, Ornithocheirus scaphorhynchus, Ornithocheirus tenuirostris, Ornithocheirus xyphorhynchus, and Pterodactylus sagittirostris. Fourteen species are considered valid, and diagnoses are provided to all of them: Ornithocheirus simus, Lonchodraco giganteus comb. n., Lonchodraco machaerorhynchus comb. n., Lonchodraco(?) microdon comb. n., Coloborhynchus clavirostris, ‘Ornithocheirus’ capito, Camposipterus nasutus comb. n., Camposipterus(?) sedgwickii comb. n., Camposipterus(?) colorhinus comb. n., Cimoliopterus cuvieri comb. n., ‘Ornithocheirus’ polyodon, ‘Ornithocheirus’ platystomus, ‘Pterodactylus’ daviesii, and ‘Ornithocheirus’ denticulatus. These species are referred in the genera Ornithocheirus, Lonchodraco gen. n., Coloborhynchus, Cimoliopterus gen. n., and Camposipterus gen. n., but additional genera are probably present, as indicated by the use of single quotation marks throughout the text. A cladistic analysis demonstrates that Anhangueridae lies within a newly recognized clade, here named Anhangueria, which also includes the genera Cearadactylus, Brasileodactylus, Ludodactylus, and Camposipterus. The anhanguerian ‘Cearadactylus’ ligabuei belongs to a different genus than Cearadactylus atrox. Lonchodraconidae fam. n. (more or less equivalent to Lonchodectidae sensu
Pterodactyloidea, Ornithocheiridae, Anhangueridae, Lonchodraconidae, Anhangueria, Cretaceous, Cambridge Greensand
The Cretaceous of England is exceptionally rich in pterosaur fossils, which are of historical, morphological, and taxonomic importance. Several deposits contain pterosaur remains, among them the Hastings Group (late Berriasian / Valanginian), Wessex Formation (Barremian), Vectis Formation (Barremian / early Aptian), Gault Clay Formation (Albian), Cambridge Greensand (Cenomanian deposit with reworked fossils thought to be Albian in age) and Chalk Formation (Cenomanian / Turonian) (
Abbreviated taxonomic history of the Ornithocheirus complex from the Cretaceous of England. Single quotation marks indicate provisional genera.
Original description | Seeley, 1870 | Hooley, 1914 | Wellnhofer, 1978 | Unwin, 2001 | This work |
---|---|---|---|---|---|
Ornithocheirus brachyrhinus Seeley, 1870 | Ornithocheirus brachyrhinus | Ornithocheirus brachyrhinus | Ornithocheirus brachyrhinus | Anhanguera cuvieri | nomen dubium |
Ornithocheirus capito Seeley, 1870 | Ornithocheirus capito | Criorhynchus capito | Criorhynchus capito | Coloborhynchus capito | ‘Ornithocheirus’ capito |
Ornithocheirus carteri Seeley, 1870 | Ornithocheirus carteri | – | Criorhynchus simus | Ornithocheirus simus | nomen dubium |
Coloborhynchus clavirostris Owen, 1874 | – | Criorhynchus simus | Criorhynchus simus | Coloborhynchus clavirostris | Coloborhynchus clavirostris |
Ornithocheirus colorhinus Seeley, 1870 | Ornithocheirus colorhinus | Ornithocheirus colorhinus | Ornithocheirus colorhinus | Anhanguera cuvieri | Camposipterus(?) colorhinus |
Pterodactylus compressirostris Owen, 1852 | – | Lonchodectes compressirostris | Ornithocheirus compressirostris | Lonchodectes compressirostris | nomen dubium |
Pterodactylus cuvieri Bowerbank, 1852 | Ornithocheirus cuvieri | Ornithocheirus cuvieri | Ornithocheirus cuvieri | Anhanguera cuvieri | Cimoliopterus cuvieri |
Palaeornis cliftii Mantell, 1844 | – | Ornithocheirus clifti [sic] | Ornithocheirus clifti [sic] | – | nomen dubium |
Ornithocheirus crassidens Seeley, 1870 | Ornithocheirus crassidens | Amblydectes crassidens | Criorhynchus crassidens | Coloborhynchus sedgwickii | nomen dubium |
Coloborhynchus clavirostris Owen, 1874 | – | Criorhynchus simus | Criorhynchus simus | Coloborhynchus clavirostris | Coloborhynchus clavirostris |
Pterodactylus curtus Owen, 1874 | – | Ornithocheirus curtus | Ornithocheirus curtus | – | nomen nudum |
Pterodactylus daviesii Owen, 1874 | – | Lonchodectes daviesii | Ornithocheirus daviesi [sic] | Lonchodectes platystomus | ‘Pterodactylus’ daviesii |
Ornithocheirus dentatus Seeley, 1870 | Ornithocheirus dentatus | Ornithocheirus dentatus | Ornithocheirus dentatus | Anhanguera cuvieri | nomen dubium |
Ornithocheirus denticulatus Seeley, 1870 | Ornithocheirus denticulatus | Ornithocheirus denticulatus | – | Anhanguera cuvieri | ‘Ornithocheirus’ denticulatus |
Cimoliornis diomedeus Owen, 1846 | – | Ornithocheirus diomedius [sic] | Ornithocheirus diomedius [sic] | – | nomen dubium |
Ornithocheirus enchorhynchus Seeley, 1870 | Ornithocheirus enchorhynchus | Ornithocheirus enchorhynchus | Ornithocheirus enchorhynchus | Anhanguera cuvieri | nomen dubium |
Ornithocheirus eurygnathus Seeley, 1870 | Ornithocheirus eurygnathus | Amblydectes eurygnathus | Criorhynchus eurygnathus | Coloborhynchus capito | nomen dubium |
Pterodactylus fittoni Owen, 1859 | Ornithocheirus fittoni | Ornithocheirus fittoni | Ornithocheirus fittoni | Anhanguera fittoni | nomen dubium |
Pterodactylus giganteus Bowerbank, 1846 | – | Lonchodectes giganteus | Ornithocheirus giganteus | Lonchodectes giganteus | Lonchodraco giganteus |
Ornithocheirus huxleyi Seeley, 1870 | Ornithocheirus huxleyi | – | Ornithocheirus huxleyi | Lonchodectes microdon | – |
Ornithocheirus machaerorhynchus Seeley, 1870 | Ornithocheirus machaerorhynchus | Lonchodectes machaeorhynchus [sic] | Ornithocheirus machaeorhynchus [sic] | Lonchodectes machaerorhynchus | Lonchodraco machaerorhynchus |
Ornithocheirus microdon Seeley, 1870 | Ornithocheirus microdon | Lonchodectes microdon | Ornithocheirus microdon | Lonchodectes microdon | Lonchodraco(?) microdon |
Ornithocheirus nasutus Seeley, 1870 | Ornithocheirus nasutus | Ornithocheirus nasutus | Ornithocheirus nasutus | Anhanguera fittoni | Camposipterus nasutus |
Pterodactylus nobilis | – | Ornithocheirus nobilis | – | – | nomen nudum |
Ornithocheirus oweni Seeley, 1870 | Ornithocheirus oweni | Lonchodectes oweni | Ornithocheirus oweni | Lonchodectes microdon | Lonchodraco(?) microdon |
Ornithocheirus oxyrhinus Seeley, 1870 | Ornithocheirus oxyrhinus | Ornithocheirus oxyrhinus | Ornithocheirus oxyrhinus | nomen nudum | nomen dubium |
Ornithocheirus platyrhinus Seeley, 1870 | Ornithocheirus platyrhinus | Criorhynchus platyrhinus | Criorhynchus simus | Ornithocheirus simus | Ornithocheirus simus |
Ornithocheirus platystomus Seeley, 1870 | Ornithocheirus platystomus | Amblydectes platystomus | Criorhynchus platystomus | Lonchodectes platystomus | ‘Ornithocheirus’ platystomus |
Ornithocheirus polyodon Seeley, 1870 | Ornithocheirus polyodon | Ornithocheirus polyodon | Ornithocheirus polyodon | Anhanguera fittoni | ‘Ornithocheirus’ polyodon |
Ornithocheirus reedi Seeley, 1870 | Ornithocheirus reedi | Criorhynchus reedi | Criorhynchus reedi | Coloborhynchus capito | ‘Ornithocheirus’ capito |
Pterodactylus sagittirostris Owen, 1874 | – | Lonchodectes sagittirostris | Ornithocheirus sagittirostris | Lonchodectes sagittirostris | nomen dubium |
Ornithocheirus scaphorhynchus Seeley, 1870 | Ornithocheirus scaphorhynchus | Lonchodectes scaphorhynchus | Ornithocheirus scaphorhynchus | Anhanguera cuvieri | nomen dubium |
Pterodactylus sedgwickii Owen, 1859 | Ornithocheirus sedgwicki [sic] | Ornithocheirus sedgwicki [sic] | Ornithocheirus sedgwicki [sic] | Coloborhynchus sedgwickii | Camposipterus(?) sedgwickii |
Pterodactylus simus Owen, 1861 | Ornithocheirus simus | Criorhynchus simus | Criorhynchus simus | Ornithocheirus simus | Ornithocheirus simus |
Ornithocheirus tenuirostris Seeley, 1870 | Ornithocheirus tenuirostris | Lonchodectes tenuirostris | Ornithocheirus tenuirostris | Lonchodectes compressirostris | nomen dubium |
Pterodactylus woodwardi Owen, 1861 | Ornithocheirus woodwardi | Criorhynchus woodwardi | Criorhynchus simus | Coloborhynchus sedgwicki | nomen dubium |
Ornithocheirus xyphorhynchus Seeley, 1870 | Ornithocheirus xyphorhynchus | Ornithocheirus xyphorhynchus | Ornithocheirus xyphorhynchus | Anhanguera cuvieri | nomen dubium |
Pioneer works on this fauna, such as by James Scott Bowerbank (1797–1877) and Richard Owen (1804–1892), initially attributed the pterosaurs from the Cretaceous of England in the genus Pterodactylus, nowadays considered restricted to the Jurassic Solnhofen Limestone of Germany. Harry Govier Seeley (1839–1909) was the first researcher to separate the British forms in new genera.
In 1869, Seeley published an index of specimens from the collection of the Woodwardian Museum (now Sedgwick Museum of Earth Sciences) of the University of Cambridge. This index presented 24 named pterosaur species from the Cretaceous of England, divided in two genera, “Ptenodactylus” and Ornithocheirus. At first glance, this work presents nomenclatural problems as all new species lacked descriptions and would, in principle, be considered nomina nuda based on article 12.1 of the International Code of Zoological Nomenclature (ICZN) (
It is also possible to interpret these sentences (
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Ornithocheiridae |
Ornithocheirus brachyrhinus |
Ornithocheirus capito |
Ornithocheirus carteri |
Ornithocheirus colorhinus |
Ornithocheirus crassidens |
Ornithocheirus cuvieri |
Ornithocheirus dentatus |
Ornithocheirus denticulatus |
Ornithocheirus enchorhynchus |
Ornithocheirus eurygnathus |
Ornithocheirus fittoni |
Ornithocheirus huxleyi |
Ornithocheirus machaerorhynchus |
Ornithocheirus microdon |
Ornithocheirus nasutus |
Ornithocheirus oweni |
Ornithocheirus oxyrhinus |
Ornithocheirus platystomus |
Ornithocheirus platyrhinus |
Ornithocheirus polyodon |
Ornithocheirus reedi |
Ornithocheirus scaphorhynchus |
Ornithocheirus sedgwicki |
Ornithocheirus simus |
Ornithocheirus tenuirostris |
Ornithocheirus woodwardi |
Ornithocheirus xyphorhynchus |
In 1874,
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Family uncertain |
Ornithochirus [sic] compressirostris |
Ornithochirus [sic](?) clifti [sic] |
Ornithochirus [sic](?) curtus |
Ornithochirus [sic] cuvieri |
Ornithochirus [sic] daviesi [sic] |
Ornithochirus [sic] diomedius [sic] |
Ornithochirus [sic] fittoni |
Ornithochirus [sic](?) giganteus |
Ornithochirus [sic] hlavatschi |
Ornithochirus [sic] nobilis |
Ornithochirus [sic] sedgwicki [sic] |
Ornithochirus [sic](?) simus |
=(?) Pterodactylus woodwardi |
Ornithochirus [sic](?) sp. |
Ornithochirus [sic] validus |
= ? Pterodactylus macrurus |
= Doratorhynchus validus |
The first major review of the Ornithocheirus complex was provided by
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Doratohynchus validum [sic] |
=Pterodactylus macrurus |
Ornithocheirus brachyrhinus |
Ornithocheirus capito |
Ornithocheirus carteri |
Ornithocheirus colorhinus |
Ornithocheirus compressirostris |
Ornithocheirus crassidens |
Ornithocheirus cuvieri |
Ornithocheirus dentatus |
Ornithocheirus denticulatus |
Ornithocheirus enchorhynchus |
Ornithocheirus eurygnathus |
Ornithocheirus fittoni |
Ornithocheirus machaerorhynchus |
Ornithocheirus microdon |
Ornithocheirus nasutus |
Ornithocheirus oweni |
Ornithocheirus oxyrhinus |
Ornithocheirus platyrhinus |
Ornithocheirus platysomus [sic] |
Ornithocheirus polyodon |
Ornithocheirus scaphorhynchus |
Ornithocheirus sedgwicki [sic] |
Ornithocheirus simus |
=Pterodactylus woodwardi |
Ornithocheirus tenuirostris |
Pterodactylus hopkinsi |
Pterodactylus oweni |
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Ornithocheiridae |
Ornithocheirinae |
Ornithocheirus brachyrhinus |
Ornithocheirus clifti [sic] |
Ornithocheirus colorhinus |
Ornithocheirus curtus |
Ornithocheirus cuvieri |
Ornithocheirus dentatus |
Ornithocheirus denticulatus |
Ornithocheirus diomedius [sic] |
Ornithocheirus enchorhynchus |
Ornithocheirus fittoni |
Ornithocheirus nasutus |
Ornithocheirus nobilis |
Ornithocheirus oxyrhinus |
Ornithocheirus polyodon |
Ornithocheirus sedgwicki [sic] |
Ornithocheirus xyphorhynchus |
Lonchodectes compressirostris |
Lonchodectes daviesii |
Lonchodectes giganteus |
Lonchodectes machaeorhynchus [sic] |
Lonchodectes microdon |
Lonchodectes oweni |
Lonchodectes sagittirostris |
Lonchodectes scaphorhynchus |
Lonchodectes tenuirostris |
Criorhynchinae |
Amblydectes crassidens |
Amblydectes eurygnathus |
Amblydectes platysomus [sic] |
Criorhynchus capito |
Criorhynchus carteri |
Criorhynchus platyrhinus |
Criorhynchus reedi |
Criorhynchus simus |
=Coloborhynchus clavirostris |
Criorhynchus woodwardi |
Criorhynchus is a taxonomic problem by itself.
Subsequent authors tended to divide the species of the Ornithocheirus complex in only two genera, Ornithocheirus and Criorhynchus (e.g.,
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Ornithocheiridae |
Ornithocheirus compressirostris |
Ornithocheirus cuvieri |
Ornithocheirus daviesi [sic] |
Ornithocheirus fittoni |
Ornithocheirus giganteus |
Ornithocheirus microdon |
Ornithocheirus sagittirostris |
Ornithocheirus sedgwicki [sic] |
Ornithocheiridae incertae sedis |
Ornithocheirus clifti [sic] |
Ornithocheirus curtus |
Ornithocheirus diomedius [sic] |
Ornithocheirus validus |
Uncertain systematic position |
Ornithocheirus brachyrhinus |
Ornithocheirus colorhinus |
Ornithocheirus dentatus |
Ornithocheirus enchorhynchus |
Ornithocheirus huxleyi |
Ornithocheirus nasutus |
Ornithocheirus oxyrhinus |
Ornithocheirus polyodon |
Ornithocheirus machaeorhynchus [sic] |
Ornithocheirus oweni |
Ornithocheirus scaphorhynchus |
Ornithocheirus tenuirostris |
Ornithocheirus xyphorhynchus |
“Ornithocheirus” bunzeli |
“Ornithocheirus” hilsensis |
“Ornithocheirus” hlavatschi |
cf. Ornithocheirus |
Criorhynchidae |
Criorhynchus simus |
=Coloborhynchus clavirostris |
= ? Criorhynchus woodwardi |
= ? Criorhynchus carteri |
= ? Criorhynchus platyrhinus |
Criorhynchidae incertae sedis |
Criorhynchus eurygnathus |
Criorhynchus capito |
Criorhynchus crassidens |
Criorhynchus platystomus |
Criorhynchus reedi |
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after
Ornithocheiridae |
Ornithocheirus simus |
=Ornithocheirus carteri |
=Ornithocheirus platyrhinus |
Ornithocheirus sp. |
Coloborhynchus clavirostris |
Coloborhynchus capito |
=Ornithocheirus eurygnathus |
=Ornithocheirus reedi |
Coloborhynchus sedgwickii |
=Ornithocheirus crassidens |
=Ornithocheirus woodwardi |
Anhanguera cuvieri |
=Ornithocheirus brachyrhinus |
=Ornithocheirus colorhinus |
=Ornithocheirus dentatus |
=Ornithocheirus denticulatus |
=Ornithocheirus enchorhynchus |
=Ornithocheirus scaphorhynchus |
=Ornithocheirus xyphorhynchus |
Anhanguera fittoni |
=Ornithocheirus nasutus |
=Ornithocheirus polyodon |
Lonchodectidae |
Lonchodectes giganteus |
Lonchodectes compressirostris |
=Ornithocheirus tenuirostris |
Lonchodectes machaerorhynchus |
Lonchodectes microdon |
=Ornithocheirus huxleyi |
=Ornithocheirus oweni |
Lonchodectes platystomus |
=Pterodactylus daviesii |
Lonchodectes sagittirostris |
As part of the unpublished PhD thesis of the first author, a careful study and revision of the species referred to the Anhangueridae, Ornithocheiridae and Lonchodectidae was performed. Based on the results from this work, a review of the species from the so–called Ornithocheirus complex is presented here (Tables 1, 8, 9 and 10). Among these species, Ornithocheirus huxleyi Seeley, 1870 (misspelled huxleyii by
List of taxa of the Ornithocheirus complex from the Cretaceous of England, after the present work. Single quotation marks indicate provisional genera. = indicate synonymies.
Ornithocheiridae Seeley, 1870 |
Ornithocheirus simus (Owen, 1861) |
=Ornithocheirus platyrhinus Seeley, 1870 |
Lonchodraconidae fam. n. |
Lonchodraco giganteus (Bowerbank, 1846) comb. n. |
Lonchodraco machaerorhynchus (Seeley, 1870) comb. n. |
Lonchodraco(?) microdon (Seeley, 1870) comb. n. |
=Ornithocheirus oweni Seeley, 1870 |
Anhangueridae Campos and Kellner, 1985 |
Coloborhynchus clavirostris Owen, 1874 |
‘Ornithocheirus’ capito Seeley, 1870 |
=Ornithocheirus reedi Seeley, 1870 |
Anhangueria incertae sedis |
Camposipterus nasutus (Seeley, 1870) comb. n. |
Camposipterus(?) sedgwickii (Owen, 1859) comb. n. |
Camposipterus(?) colorhinus (Seeley, 1870) comb. n. |
Pteranodontoidea incertae sedis |
Cimoliopterus cuvieri (Bowerbank, 1851) comb. n. |
‘Ornithocheirus’ polyodon Seeley, 1870 |
Pterodactyloidea incertae sedis |
‘Ornithocheirus’ platystomus Seeley, 1870 |
‘Pterodactylus’ daviesii Owen, 1874 |
‘Ornithocheirus’ denticulatus Seeley, 1870 |
List of taxa of the Ornithocheirus complex from the Cretaceous of England here considered nomina dubia.
Palaeornis cliftii Mantell, 1844 |
Cimoliornis diomedeus Owen, 1846 |
Pterodactylus compressirostris Owen, 1851 |
Pterodactylus fittoni Owen, 1859 |
Pterodactylus woodwardi Owen, 1861 |
Ornithocheirus brachyrhinus Seeley, 1870 |
Ornithocheirus carteri Seeley, 1870 |
Ornithocheirus crassidens Seeley, 1870 |
Ornithocheirus dentatus Seeley, 1870 |
Ornithocheirus enchorhynchus Seeley, 1870 |
Ornithocheirus eurygnathus Seeley, 1870 |
Ornithocheirus oxyrhinus Seeley, 1870 |
Ornithocheirus scaphorhynchus Seeley, 1870 |
Ornithocheirus tenuirostris Seeley, 1870 |
Ornithocheirus xyphorhynchus Seeley, 1870 |
Pterodactylus sagittirostris Owen, 1874 |
List of taxa of the Ornithocheirus complex from the Cretaceous of England that are nomina nuda.
Ptenodactylus oweni Seeley, 1869 |
Ptenodactylus polyodon Seeley, 1869 |
Ptenodactylus microdon Seeley, 1869 |
Ptenodactylus scaphorhynchus Seeley, 1869 |
Ptenodactylus macrorhinus Seeley, 1869 |
Ptenodactylus brachyrhinus Seeley, 1869 |
Ptenodactylus crassidens Seeley, 1869 |
Ptenodactylus dentatus Seeley, 1869 |
Ptenodactylus nasutus Seeley, 1869 |
Ptenodactylus tenuirostris Seeley, 1869 |
Ptenodactylus capito Seeley, 1869 |
Ptenodactylus eurygnathus Seeley, 1869 |
Ptenodactylus machaerorhynchus Seeley, 1869 |
Ptenodactylus platystomus Seeley, 1869 |
Ptenodactylus enchorhynchus Seeley, 1869 |
Ptenodactylus colorhinus Seeley, 1869 |
Ptenodactylus oxyrhinus Seeley, 1869 |
Ornithocheirus carteri Seeley, 1869 |
Ornithocheirus platyrhinus Seeley, 1869 |
Pterodactylus curtus Owen, 1874 |
Pterodactylus nobilis Owen, 1874 |
Pterodactylus validus Owen, 1874 |
In this paper, diagnoses are provided for all species and genera considered valid, in addition to photographs and illustrations, which, we hope, will facilitate future discussions about the diversity of pterosaurs in England during the Cretaceous and their relationships with species elsewhere. Nomina nuda are marked with double quotation marks, and single quotation marks around genera names indicate that the species is cited as in its original description, but may belong to a different genus.
Institutional abbreviations: BSP – Bayerische Staatssammlung für Paläontologie und historische Geologie, Munich, Germany; CAMSM – Sedgwick Museum of Earth Sciences, Cambridge, England; IVPP – Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China; MANCH – Manchester Museum, Manchester, England; MN, Museu Nacional / Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; NHMUK – Natural History Museum, London, England; QM – Queensland Museum, Brisbane, Australia; SMNS – Staatliches Museum für Naturkunde, Stuttgart, Germany; SMU, Shuler Museum of Paleontology, Southern Methodist University, Dallas, USA; UERJ – Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil; ZIN – Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.
http://species-id.net/wiki/Ornithocheiridae
Ornithocheirus Seeley, 1869.
Ornithocheirus.
Albian.
Cambridge Greensand, England.
the same as for the type genus.
Seeley erected the name Ornithocheirae in 1870, including only the genus Ornithocheirus. It is corrected to Ornithocheiridae Seeley, 1870 following the article 11.7.1.3 of the ICZN.
http://species-id.net/wiki/Ornithocheirus
Pterodactylus simus Owen, 1861, by monotypy.
Albian.
Cambridge Greensand, England.
As for the type species.
http://species-id.net/wiki/Ornithocheirus_simus
Figs 1–3CAMSM B54428, anterior portion of the rostrum (Fig. 1A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B54552 (Fig. 1E–H), CAMSM B54429 (Fig. 2A–D), CAMSM B54677 (Fig. 2E–H), MANCH L.10832 (Fig. 3A–D), and NHMUK PV 35412 (Fig. 3E–H) (all from the Cambridge Greensand).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): tall rostrum*; first pair of premaxillary teeth directed ventrally*; first pair of upper alveoli slightly displaced posteriorly from the anterior margin of the premaxilla*; ventral margin of the palate straight; rostrum not expanded anteriorly.
Ornithocheirus simus was first described on the basis of a fragmentary anterior portion the premaxillae and maxillae (CAMSM B54428), which remains the best preserved specimen undoubtly referable to this species. This fragment has, in lateral view, a rounded profile, and all preserved tooth sockets are oriented ventrally.
Ornithocheirus simus lacks an anterior expansion of the rostrum. As noticed by
In the original description and illustration, CAMSM B54428 had a tooth preserved in the first left alveolus (
Aside from the taxonomic and nomenclatural problems surrounding Ornithocheirus simus, its basic structure is controversial. Several authors considered it a long–snouted animal with a robust premaxillary crest (e.g.,
Ornithocheirus simus. A–D holotype CAMSM B54428 (Albian, Cambridge Greensand), anterior part of the rostrum A left lateral view B respective line drawing C ventral view D respective line drawing E–H referred specimen CAMSM B54552 (Albian, Cambridge Greensand), anterior part of the rostrum E anterior view F respective line drawing G left lateral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
Ornithocheirus simus. A–D referred specimen CAMSM 54429 (Albian, Cambridge Greensand), anterior part of the rostrum A right lateral view B respective line drawing C anterior view D respective line drawing E–H referred specimen CAMSM 54677 (Albian, Cambridge Greensand), anterior part of the rostrum E right lateral view F respective line drawing G anterior view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
Ornithocheirus simus. A–D referred specimen MANCH L10832 (Albian, Cambridge Greensand), anterior part of the rostrum A left lateral view B respective line drawing C anterior view D respective line drawing E–H referred specimen NHMUK PV 35412 (Albian, Cambridge Greensand), anterior part of the rostrum E right lateral view F respective line drawing G anterior view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. Photos E and G courtesy of The Natural History Museum.
As detailed above, the taxonomic history of the genus Ornithocheirus and of the species Ornithocheirus simus is quite complex. To summarize, CAMSM B54428 was first described by
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Lonchodraco gen. n.
Lonchodraco.
Albian to Cenomanian / Turonian.
Cambridge Greensand and Chalk Formation, England.
the same as for the type genus.
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Derived from the Greek lonchos, meaning lance, and Latin draco, meaning dragon.
Pterodactylus giganteus Bowerbank, 1846.
Lonchodraco giganteus, Lonchodraco machaerorhynchus, and Lonchodraco(?) microdon.
Albian to Cenomanian / Turonian.
Cambridge Greensand and Chalk Formation, England.
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): comparatively small alveoli (up to 4 mm in diameter) in the anterior portions of the upper and lower jaws; alveoli of the anterior portions of the upper and lower jaws without significant variation in size; alveoli placed in an elevation in relation to the palate and to the dorsal margin of the mandible*; deep palatal ridge; mandibular crest present; spacing between alveoli roughly equivalent to their diameters (modified from
Latter workers did not accept Lonchodectes as a valid genus.
In the present work, Lonchodectes compressirostris is considered a nomen dubium (see below) and, therefore, a new genus, Lonchodraco, is here erected to include three of the species previously referred to Lonchodectes: Lonchodraco giganteus, Lonchodraco machaerorhynchus, and Lonchodraco(?) microdon.
http://species-id.net/wiki/Lonchodraco_giganteus
Fig. 4NHMUK PV 39412, anterior portions of the rostrum and mandible, incomplete scapulocoracoid, proximal ends of the humerus and ulna, and a partial wing phalanx (Fig. 4A–G).
Near Maidstone, Burham, Kent, England.
Chalk Formation (Cenomanian / Turonian).
Lonchodraconid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): anterior portion of the premaxillae rounded; anterior portion of the dentaries rounded; divergent alveolar margins of the anterior end of the upper and lower jaws; presence of a premaxillary crest; short, low, blade–like dentary crest*; approximately6 alveoli per 3 cm of jaw margin*.
Lonchodraco giganteus was briefly described by
Lonchodraco giganteus has a complex taxonomic history. The species was named Pterodactylus giganteus by
Lonchodraco giganteus comb. n. Lectotype NHMUK PV 39412 (Cenomanian / Turonian, Chalk Formation). A–F articulated anterior parts of the rostrum and mandible A right lateral view B respective line drawing C left lateral view D respective line drawing E anterior view F respective line drawing G associated scapulocoracoid in posterior view. Abbreviations: ch – choanae, cor – coracoid, d – dentary, dcr – dentary crest, m – maxillae, pl – palatine, pm – premaxillae, pmcr – premaxillaery crest, prid – palatal ridge, sca – scapula. Arrows indicate alveoli or teeth. Scale bar = 10 mm. Photos courtesy of The Natural History Museum.
http://species-id.net/wiki/Lonchodraco_machaerorhynchus
Fig. 5CAMSM B54855, partial mandibular symphysis (Fig. 5A–F).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Lonchodraconid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): deep dentary crest*; ventral margin of the mandible posterior to the dentary crest ascending in lateral view*; ventral depression located posteriorly to the dentary crest*; wide mandibular groove*; approximately 4.5 alveoli per 3 cm of jaw margin.
CAMSM B54855 consists of a fragment of the posterior portion of the mandibular symphysis.
In addition to a deep mandibular groove, CAMSM B54855 shares with Lonchodraco giganteus small and well–spaced alveoli, without significant size variation. However, it differs from this species in having straight alveolar margins in dorsal view (
Lonchodraco machaerorhynchus comb. n. Holotype CAMSM B54855 (Albian, Cambridge Greensand), fragment of the mandibular symphysis A right lateral view B respective line drawing C dorsal view D respective line drawing E ventral view F respective line drawing. Abbreviations: d – dentary, dcr – dentary crest, sul – sulcus. Arrows indicate alveoli or teeth. Scale bar = 10 mm.
CAMSM B54486, anterior portion of the rostrum (Fig. 6A–F).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B 54439 (Fig. 6G–L) (also from the Cambridge Greensand).
Lonchodraconid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): premaxillary crest absent; dorsal margin of the premaxillae rounded; deep palatal ridge*; palate between the elevation of the alveolar margins and the palatal ridge concave; spacing between alveoli larger than their diameters*; approximately 4.5 alveoli per 3 cm of jaw margin.
Lonchodraco(?) microdon was described by
Ornithocheirus oweni was described in the same work (
Lonchodraco(?) microdon has small (approximately 3 mm diameter) and evenly spaced alveoli as in Lonchodraco giganteus and Lonchodraco machaerorhynchus. However, it is distinct from Lonchodraco giganteus, lacking a premaxillary crest, having spacing between the alveoli larger than their diameters, and with parallel alveolar margins. These features constrast with those of the type species of the genus. Lonchodraco(?) microdon shares with Lonchodraco machaerorhynchus the parallel alveolar margins and the same alveolar density (4.5 alveoli per 3 cm of jaw margin), but they differ in the spacing between the alveoli, larger in the first. Due to these differences, it is possible that Lonchodraco(?) microdon actually represents a distinct genus. However, in the absence of further evidence, we refrain from naming a new genus and assign the species tentatively to Lonchodraco.
Lonchodraco(?) microdon comb. n. A–F holotype CAMSM B54486 (Albian, Cambridge Greensand), anterior fragment of the rostrum A ventral view B respective line drawing C anterior view D respective line drawing E posterior view F respective line drawing. In B dashed lines indicate the portion lost since the original description G–L referred specimen CAMSM B 54439 (Albian, Cambridge Greensand), anterior fragment of the rostrum G ventral view H respective line drawing I anterior view J respective line drawing K posterior view L respective line drawing. Abbreviation: prid – palatal ridge. Arrows indicate alveoli or teeth. Scale bar = 10 mm.
All pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.
Camposipterus, Cearadactylus, Ludodactylus, and Anhangueridae.
Late Berriasian / Valanginian to Cenomanian.
Hastings Group, England; Wessex Formation, England; Elrhaz Formation, Niger; Jiufotang Formation, China; Khuren–Dukh, Dzun–Bayin Formation, Mongolia; Romualdo Formation, Brazil; Paw Paw Formation, USA; Cambridge Greensand, England; Kem Kem beds, Morocco.
(18.1) Presence of an anterior expansion of the premaxillary tip with the jaw end tall, and (48.1) larger teeth located at the tip of the rostrum (see “Phylogenetic affinities of the species of the Ornithocheirus complex”, below).
http://species-id.net/wiki/Anhangueridae
Anhanguera Campos and Kellner, 1985.
Anhanguera, Caulkicephalus, Coloborhynchus, Liaoningopterus, Siroccopteryx, Tropeognathus, and Uktenadactylus.
Late Berriasian / Valanginian to Cenomanian.
Hastings Group, England; Wessex Formation, England; Elrhaz Formation, Niger; Jiufotang Formation, China; Khuren–Dukh, Dzun–Bayin Formation, Mongolia; Romualdo Formation, Brazil; Paw Paw Formation, USA; Cambridge Greensand, England; Kem Kem beds, Morocco (Table 11).
Deposits where anhanguerid fossils have been found.
Late Berriasian/Valanginian | Hastings Group, England (Coloborhynchus clavirostris; see |
Barremian | Wessex Formation, England (Caulkicephalus trimicrodon; see |
Aptian | Elrhaz Formation, Niger ( Jiufotang Formation, China (Liaoningopterus gui; see |
Aptian/Albian | Khuren–Dukh, Dzun–Bayin Formation, Mongolia ( |
Albian | Romualdo Formation, Brazil (Tropeognathus mesembrinus and several species of Anhanguera; e.g., Paw Paw Formation, USA (Uktenadactylus wadleighi; see Cambridge Greensand |
Cenomanian | Kem Kem beds, Morocco (Siroccopteryx moroccensis; see |
* this deposit is Cenomanian but the fossils are Albian in age.
Coloborhynchus clavirostris Owen, 1874, by monotypy.
Late Berriasian / Valanginian.
Hastings Group, England.
As for the type species.
http://species-id.net/wiki/Coloborhynchus_clavirostris
Fig. 7NHMUK PV R 1822, anterior portion of the rostrum (Fig. 7A–D).
St.–Leonards–on–Sea, East Sussex, England.
Hastings Group (late Berriasian / Valanginian).
Anhanguerid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): oval depression beneath the first pair of alveoli*; second, third and fourth pairs of alveoli located laterally*; fifth and sixth pairs of alveoli located more medially than the preceding alveoli on the base of the palatal ridge*; anterior part of the palatal ridge bordered by two shallow longitudinally elongated depressions* (from
The holotype of Coloborhynchus clavirostris (NHMUK PV R 1822) is a fragment of the premaxillae and maxillae, and has previously been described in detail (
Coloborhynchus clavirostris. Holotype, NHMUK PV R 1822 (late Berriasian / Valanginian, Hastings Group), anterior part of the rostrum A anterior view B respective line drawing C left lateral view D respective line drawing. Abbreviations: dep – depression, m – maxillae, pm – premaxillae, pmcr – premaxillaery crest, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. Photos courtesy of The Natural History Museum.
CAMSM B 54625, anterior portion of the rostrum (Fig. 8A–F).
Chesterton, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Holotype of Ornithocheirus reedi Seeley, 1870 (Fig. 8G–J) (from the Cambridge Greensand; current whereabouts unkown).
Anhanguerid pterosaur with the following combination of characters that distinguishes it from other members of the clade: anteriorly located and tall premaxillary crest; anterior margin of the premaxillary crest concave in lateral view; first pair of upper teeth positioned on the anterior margin of the rostrum.
‘Ornithocheirus’ capito is known from a fragmentary holotype, which has a tall, anteriorly located premaxillary crest with a concave anterior margin. Due to its fragmentary state, the presence of an anterior expansion cannot be confirmed but is suggested by the structure of the preserved right side of the specimen. These features allow its placement in Anhangueridae.
A second specimen referable to this species is the holotype of Ornithocheirus reedi. It has a median groove that extends along the height of the crest; such groove is usually considered a sign of abrasion among Cambridge Greensand pterosaurs, but
‘Ornithocheirus’ capito differs from Coloborhynchus clavirostris (from a distinct, older deposit, the late Berriasian–Valanginian Hastings Group of the Wealden Supergroup) in the absence of a flat anterior margin of the rostrum.
‘Ornithocheirus’ capito can also be distinguished from Ornithocheirus simus, and from Ornithocheirus, by presence of the first pair of teeth in the anterior margin of the premaxillae. In Ornithocheirus simus, the first pair of alveoli is directed ventrally and not located at the tip of the snout. The combination of features seen in ‘Ornithocheirus’ capito is also absent in Lonchodraco, Cimoliopterus, Camposipterus (see below), or any other known pterosaur, and is diagnostic for the present species even though this species has no autapomorphies.
Here, we recognize that ‘Ornithocheirus’ capito possibly represents a new genus, but we refrain from naming a new one until better material comes to light. Therefore, we refer to it by the name given in its original description, as ‘Ornithocheirus’ capito.
‘Ornithocheirus’ capito. A–F holotype CAMSM B 54625 (Albian, Cambridge Greensand), anterior part of the rostrum A right lateral view B respective line drawing C ventral view D respective line drawing E anterior view F respective line drawing. G–J referred specimen, whereabout unknown, holotype of Ornithocheirus reedi (Albian, Cambridge Greensand), anterior part of the rostrum G anterior view H respective line drawing I right lateral view J respective line drawing. Abbreviations: dep – depression, m – maxillae, pm – premaxillae, pmcr – premaxillary crest. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. G and I from
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After the Brazilian paleontologist Diogenes de Almeida Campos, who made valuable contributions to the knowledge of pterosaur diversity in Brazil and was a major influence to us, and pterus, from the Greek pteron, meaning wing.
Ornithocheirus nasutus Seeley, 1870.
Camposipterus nasutus, Camposipterus(?) sedgwickii and Camposipterus(?) colorhinus.
Albian.
Cambridge Greensand, England.
Pterodactyloid pterosaurs with the following combination of characters that distinguishes it from other members of the clade: anterior tip of the premaxillae and maxillae round in lateral view; premaxillary crest absent; anterior expansion of the rostrum present; palate curving dorsally; first pair of alveoli located anteriorly.
http://species-id.net/wiki/Camposipterus_nasutus
Fig. 9CAMSM B 54556, anterior portion of the rostrum (Fig. 9A–D).
Haslingfield, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): dorsal margin of the rostrum straight to gently concave in lateral view; palatal ridge extends anteriorly until just posterior to the second pair of alveoli; spacing between alveoli irregular, with the anterior alveoli closer and the posterior ones more distant from each other; density of almost 3 alveoli each 3 cm anteriorly and 2, 5 alveoli each 3 cm posteriorly*; tip of the rostrum dorsoventrally flattened, wider than high in anterior view*; second and third alveoli face lateroventrally; anterior portion of the premaxillae slightly expanded.
Camposipterus nasutus was originally described by
Camposipterus nasutus comb. n. Holotype CAMSM B 54556 (Albian, Cambridge Greensand), anterior part of the rostrum A left lateral view B respective line drawing C ventral view D respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B54422, anterior part of the rostrum (Fig. 10A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): tall rostrum; anterior expansion of the rostrum ends abruptly behind the third pair of alveoli*; palatal ridge extending posteriorly to the level of the third pair of alveoli; third pair of alveoli much larger than fourth*.
The third alveoli have different sizes on the left and right sides; this could be explained on taphonomic grounds (e.g.,
It is noteworthy that the drawing of CAMSM B54422 was reversed in
A–D Camposipterus(?) sedgwickii comb. n., holotype CAMSM B54422 (Albian, Cambridge Greensand), anterior part of the rostrum. A right lateral B respective line drawing C ventral view D respective line drawing. E–H Camposipterus(?) colorhinus comb. n., syntype CAMSM B54431 (Albian, Cambridge Greensand), anterior part of the rostrum E left lateral view F respective line drawing G ventral view H respective line drawing. Abbreviations: dep – depression, m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B54431 (Fig. 10E–H) and CAMSM B54432, anterior parts of the rostrum.
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): developed anterior expansion, lacking a marked constriction; presence of a depression above the first pair of alveoli; anterior depression faces anteroventrally*; second and third alveoli very large in size; fourth pair of alveoli much smaller than the second and third.
The syntypes, CAMSM B54431 (the more complete one) and CAMSM B54432, both are anterior portions of upper jaws. They are quite incomplete and abraded, but they are identical where comparable and demonstrate that their features are valid morphological characters rather than taphonomic artifacts.
This lunate area is a depression above the first pair of alveoli; a depression in the same location is also present in Uktenadactylus wadleighi but in the latter the anterior margin of the rostrum faces anteriorly (
Camposipterus(?) colorhinus shares with Camposipterus nasutus and Camposipterus(?) sedgwickii an anterior expansion of the rostrum and a round profile, and thus is tentatively referred to Camposipterus. As the syntypes are incomplete, it is uncertain if this taxon had a crest. It further shares with Camposipterus(?) sedgwickii the presence of a tall rostrum and robust anterior alveoli. However, Camposipterus(?) colorhinus represents a much larger and more robust pterosaur in comparison with the latter species.
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From the Greek kimolia, chalk, and pteron, wing.
Pterodactylus cuvieri Bowerbank, 1851.
Cimoliopterus cuvieri.
Cenomanian / Turonian.
Chalk Formation, England.
As for the type–species.
http://species-id.net/wiki/Cimoliopterus_cuvieri
Fig. 11NHMUK PV 39409, anterior portion of the rostrum (Fig. 11A–D).
Burham, Kent, England.
Chalk Formation (Cenomanian / Turonian).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): premaxillary crest present; premaxillary crest begins posteriorly (at the seventh pair of alveoli) but before the nasoantorbital fenestra*; palatal ridge extending anteriorly up to the third pair of alveoli; second and third alveoli similar in size and larger than the fourth; spacing between alveoli irregular, with the anterior alveoli more closely spaced and the posterior ones more widely separated from each other; almost 3 alveoli per 3 cm of jaw margin anteriorly and 2 alveoli each 3 cm posteriorly*; anterior expansion absent; palate dorsally curved.
NHMUK PV 39409 was originally reported as having a tooth preserved in the first right alveolus. During examination of the holotype in 2007 and 2009, the tooth was no longer preserved with the holotype and could not be found.
Cimoliopterus cuvieri differs from Coloborhynchus clavirostris in the lack of an anteriorly flat rostrum, premaxillary crest at the tip of the rostrum, anterior expansion, or the other diagnostic characters of that species (
Cimoliopterus cuvieri. Holotype NHMUK PV 39409 (Cenomanian / Turonian, Chalk Formation), anterior part of the rostrum A right lateral view B respective line drawing C ventral view D respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, pmcr – premaxillary crest, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. Photos courtesy of The Natural History Museum.
CAMSM B54440, anterior portion of the rostrum (Fig. 12A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): premaxillary crest absent; anterior expansion absent; palate dorsally curved; moderately developed palatal ridge; palate between the elevated alveolar rims and the palatal ridge concave; palatal ridge extending anteriorly up to the second pair of alveoli; alveoli ventrolaterally oriented; spacing between alveoli less than their diameters; approximately 5.5 alveoli per 3 cm of jaw margin*.
‘Ornithocheirus’ polyodon can be excluded from Ornithocheirus because it lacks a tall rostrum. It cannot be referred to Cimoliopterus because the alveolar spacing is quite distinct; in Cimoliopterus cuvieri, only the first three pairs of alveoli are more closely positioned. It can also be excluded from Camposipterus because it lacks an anterior expansion of the rostrum.
‘Ornithocheirus’ polyodon shares with Lonchodraconidae the presence of small alveoli at the tip of the rostrum, which do not vary significantly in size. However, it differs from members of this clade in lacking an elevated alveolar margin or a prominent palatal ridge, as in Lonchodraco(?) microdon. However, it is possible that such elevation would get deeper posteriorly, but this cannot be confirmed in the holotype. Furthermore, the spacing between the alveoli being smaller than their diameters is not present in other lonchodraconids.
It is suggested here that ‘Ornithocheirus’ polyodon might represent a new genus. As the known material is quite incomplete, we refrain from naming this taxon at the present time and use its originally proposed binomen.
A–D ‘Ornithocheirus’ polyodon, holotype CAMSM B54440 (Albian, Cambridge Greensand), anterior part of the rostrum. A right lateral view B respective line drawing C ventral view D respective line drawing. E–H ‘Ornithocheirus’ platystomus, holotype CAMSM B54835 (Albian, Cambridge Greensand), anterior part of the rostrum E right lateral view F respective line drawing G ventral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B54835, anterior portion of the rostrum (Fig. 12E–H).
Horningsea, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): alveoli small (about 4 to 5 mm in diameter); dorsal margin of the rostrum forms an angle of 27° with the ventral margin*.
‘Ornithocheirus’ platystomus is known from a partial premaxilla and maxillae. As
NHMUK PV 43074, partial mandibular symphysis (Fig. 13A–D).
Folkestone, Kent, England.
Gault Clay Formation (Albian).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade (autapomorphies are marked with an asterisk): anterior expansion absent; mandibular crest absent; mandibular groove about 2.5 cm wide; mandibular groove with elevated margins; mandibular groove extends until the second pair of alveoli; alveoli of the anterior portion of the mandible without significant variation in size; alveoli equally spaced; first pair of teeth face anterodorsally; transverse section ‘V–shaped’; approximately 4 alveoli per 3 cm of jaw margin*.
‘Pterodactylus’ daviesii is known from a partial mandibular symphysis from the Gault Clay Formation. In his description,
It shares with Lonchodraconidae the presence of alveoli of the anterior portion of the mandible without significant variation in size, but can be confidently excluded from this clade because its alveoli are slightly larger and are not located on elevated alveolar margins. Although the dentary sulcus is relatively wide, it is narrower than in Lonchodraco machaerorhynchus. ‘Pterodactylus’ daviesii is also distinct from Lonchodraco giganteus. In the latter, the mandible is very wide, with divergent margins, and rounded anteriorly. Although not comparable to Ornithocheirus simus, it is unlikely that it represents this much larger pterosaur.
‘Pterodactylus’ daviesii thus possibly represents a distinct taxon from the Gault Clay Formation. However, we refrain from naming it until more complete material becomes available and refer it using its original designation in single quotation marks.
A–D ‘Pterodactylus’ daviesii, holotype NHMUK PV 43074 (Albian, Gault Clay Formation), anterior part of the mandibular symphysis. A right lateral view B respective line drawing C dorsal view D respective line drawing. E–H ‘Ornithocheirus’ denticulatus, holotype ?CAMSM B 54794 (Albian, Cambridge Greensand), anterior part of the rostrum E ventral view F respective line drawing G right lateral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, prid– palatal ridge, sul – sulcus. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. E and G from
Anterior portion of the rostrum (?CAMSM B 54794) (Fig. 13E–H)
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Pterodactyloid pterosaur with the following combination of characters that distinguishes it from other members of the clade: anterior expansion absent; palate dorsally curved; moderate palatal ridge; palatal ridge extending anteriorly up to the third pair of alveoli; spacing between alveoli approximately equal to their diameters; approximately 4.5 alveoli per 3 cm of jaw margin.
In the original description,
‘Ornithocheirus’ denticulatus does not share the combination of characters present in Ornithocheirus, Lonchodraco, Cimoliopterus and Camposipterus and thus cannot be referred to any of these genera. In particular, ‘Ornithocheirus’ denticulatus lacks the alveolar pattern present in Cimoliopterus cuvieri. ‘Ornithocheirus’ denticulatus somewhat resembles ‘Pterodactylus’ daviesii in that both lack an anterior expansion of the jaw, sagittal crests, and have alveoli without significant variation in size and equally spaced. They differ in the alveolar density, which is slightly higher in ‘Ornithocheirus’ denticulatus. Unfortunately, the known material of both species is not directly comparable and their taxonomic identity cannot be confirmed.
Collection data provided by the curators at the Sedgwick Museum of Earth Sciences in Cambridge list the specimen CAMSM B 54794 as the holotype of ‘Ornithocheirus’ denticulatus. However, the specimen could not be found during review of the collection in October 2009. The specimen was, however, one of the few figured by
In the lack of more complete material, we here refrain from naming a new genus based on the present specimen, and refer to it using the binomen in which it was originally proposed.
NHMUK PV 2353 and 2353a, partial left humerus.
Cuckfield, Sussex, England.
Upper Tunbridge Wells Sand Formation (
This species was first described as a bird (
NHMUK PV 39418, distal end of a wing metacarpal.
Chesterton, Cambridgeshire, England.
Chalk Formation (Cenomanian / Turonian).
The two known pterosaur clades from the Chalk Formation are the Lonchodraconidae and Cimoliopterus cuvieri, whose metacarpals are unknown. This material is quite fragmentary, and its structure does not allow species or genus–level identification. Therefore, Cimoliornis diomedeus is considered a nomen dubium.
NHMUK PV 39410, partial rostrum and mandible (Fig. 14A–H).
Burham, Kent, England.
Chalk Formation (Cenomanian / Turonian).
The holotype of Pterodactylus compressirostris consists of the middle portion of the rostrum (Fig. 14E–H), without the anterior end of the rostrum, and by a mandibular fragment that is strongly compressed mediolaterally (Fig. 14A–D). The rostrum has been subject to some distortion. The symphyseal fragment has been considered part of the upper jaw since its original description (
The lectotype of Pterodactylus compressirostris does not have a mandibular crest or raised alveoli, excluding it from Lonchodraconidae. It is distinctly compressed, incomplete, and non–diagnostic. Regarding the the cranial portion, few comparisons to Lonchodraco(?) microdon are possible as the specimens are not directly comparable, but they differ in the depth of the palatal ridge, which is lower in Pterodactylus compressirostris. The cranial fragment has small and widely spaced alveoli, reminiscent of Lonchodraconidae, but it is not possible to evaluate whether this feature extended to the tips of the jaws, as in Lonchodraco giganteus, Lonchodraco machaerorhynchus, and Lonchodraco(?) microdon, or if the anteriormost alveoli showed size variation, as in Anhangueridae. The referred specimen shares with Anhangueridae small alveoli on raised alveolar margins in the posterior portion of the maxillae, with the spacing between them roughly equivalent to their diameters. It differs from Anhangueridae in the lack of a premaxillary crest. However, crucial information is lacking due to the absence of the anterior portion of the rostrum in this specimen, a portion very diagnostic for toothed pteranodontoids, and upon which the taxonomy of the group is largely based. Both specimens upon which the species is based are uninformative. Therefore, we here consider Pterodactylus compressirostris a nomen dubium.
Pterodactylus compressirostris was until recently involved in a taxonomic problem.
To further complicate the taxonomy of this species,
Pterodactylus compressirostris, holotype NHMUK PV 39410 (Cenomanian / Turonian, Chalk Formation). A–D proposed lectotype, fragment of the mandibular symphysis A left lateral view B respective line drawing C dorsal view D respective line drawing. E–H referred specimen, portion of the rostrum E left lateral view F respective line drawing G ventral view H respective line drawing. Abbreviations: ch – choanae, d – dentary, m – maxillae, naof – nasoantorbital fenestra, pl – palatine, pm – premaxillae, prid – palatal ridge, sul– sulcus. Arrows indicate alveoli or teeth. Scale bar = 10 mm. Photos courtesy of The Natural History Museum.
CAMSM B54423, anterior portion of the rostrum (Fig. 15A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B54423 is a fragment of the anterior portion of the premaxillae and maxillae, lacking the anteriormost end. It is likely that the first pair of alveoli is not preserved (
A–D Pterodactylus fittoni, holotype CAMSM B54423 (Albian, Cambridge Greensand), anterior part of the rostrum. A right lateral view B respective line drawing C ventral view D respective line drawing. E–H Ornithocheirus woodwardi, holotype CAMSM B 54433a (Albian, Cambridge Greensand), anterior part of the rostrum E anterior view F respective line drawing G right lateral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B 54433a, anterior portion of the rostrum (Fig. 15E–H).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
The holotype of Pterodactylus woodwardi comprises a fragment of the anterior portion of the tip of the rostrum. It is quite incomplete, consisting mostly of a transverse section. The first pair of teeth is located anteriorly, and the second pair faces anteroventrally. There is no premaxillary crest at the anteriormost tip of the rostrum, but the presence of a more posteriorly located crest cannot be ruled out. The specimen is very fragmentary and several important characters cannot be observed on it. Therefore, it is considered a nomen dubium.
Pterodactylus woodwardi was listed as Ornithocheirus woodwardi by
CAMSM B54443, anterior portion of the rostrum (Fig. 16A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Ornithocheirus brachyrhinus is known from the tip of a snout, with a dorsally curved palate and lacking an anteriorly located crest. It shares with Cimoliopterus cuvieri features such as the curved palate, the anterior end being higher than wide, lack of an anterior expansion, and absence of an anterior crest. The structure of Ornithocheirus brachyrhinus corresponds perfectly to the tip of the snout of Cimoliopterus cuvieri and it is possibly referable to that species, as
A–D Ornithocheirus brachyrhinus, holotype CAMSM B54443 (Albian, Cambridge Greensand), anterior part of the rostrum. A right lateral view B respective line drawing C ventral view D respective line drawing. E–H Ornithocheirus carteri, holotype CAMSM B 54437 (Albian, Cambridge Greensand), anterior part of the rostrum E anterior view F respective line drawing G left lateral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B 54437, anterior portion of the rostrum (Fig. 16E–H).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B 54499, anterior portion of a jaw (Fig. 17A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Description: CAMSM B 54499 is a very fragmentary specimen, in which much of the left side, especially the oral surface, was not preserved.
The taxonomy of this species is controversial. It was placed in the genus Ornithocheirus by
A–D Ornithocheirus crassidens, holotype CAMSM B 54499 (Albian, Cambridge Greensand), anterior fragment of the rostrum. A right lateral view B respective line drawing C ventral view D respective line drawing. E–H Ornithocheirus dentatus, holotype CAMSM B 54544 (Albian, Cambridge Greensand), anterior part of the rostrum E right lateral view F respective line drawing G ventral view H respective line drawing. Abbreviations:: m – maxillae, pm – premaxillae, prid – palatal ridge. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B 54544, anterior portion of the rostrum (Fig. 17E–H).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B 54444, anterior portion of the rostrum (Fig. 18A–D).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B 54444 is a fragment of the anterior portion of the premaxillae and maxillae, including three pairs of alveoli. The first pair of alveoli is located anteriorly, separated by a thin wall of bone from the second pair. The spacing between the second and third pairs is larger but still smaller than the diameter of the alveoli. Such spacing is common within species of the Ornithocheirus complex which share dorsal curvature of the palate, as is in the present specimen.
A–D Ornithocheirus enchorhynchus, holotype CAMSM B 54444 (Albian, Cambridge Greensand), anterior part of the rostrum. A right lateral view B respective line drawing C ventral view D respective line drawing. E–H Ornithocheirus eurygnathus, holotype CAMSM B54644 (Albian, Cambridge Greensand), anterior part of the rostrum E ?right lateral view F respective line drawing G ?ventral view H respective line drawing. Abbreviations: m – maxillae, pm – premaxillae. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
CAMSM B54644, anterior fragment of an ?upper jaw (Fig. 18E–H).
Ditton, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B54644 was tentatively identified by
CAMSM B 54612, anterior fragment of an upper jaw (Fig. 19A–B).
Smithswashing, Coton, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Ornithocheirus oxyrhinus was described by
A–B Ornithocheirus oxyrhinus, holotype CAMSM B 54612 (Albian, Cambridge Greensand), anterior part of the rostrum. A ventral view B respective line drawing. C–E Ornithocheirus scaphorhynchus, holotype CAMSM B 54441 (Albian, Cambridge Greensand), anterior part of the rostrum C ventral view D respective line drawing E line drawing in posterior view. F–I Ornithocheirus tenuirostris, holotype CAMSM B 54584 (Albian, Cambridge Greensand), anterior part of the rostrum F right lateral view G respective line drawing H ventral view I respective line drawing. J–K Ornithocheirus xyphorhynchus, holotype (Albian, Cambridge Greensand), anterior part of the rostrum J lateral view K dorsal view. Abbreviations: m – maxillae, pm – premaxillae, prid – palatal ridge, sul – sulcus. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm. J and K from
CAMSM B 54441, anterior portion of the rostrum (Fig. 19C–E).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
CAMSM B 54584, anterior portion of the rostrum (Fig. 19F–I).
Coton, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
Ornithocheirus tenuirostris was described by
anterior portion of the rostrum (collection data could not be recovered) (Fig. 19J–K).
Cambridge, Cambridgeshire, England.
Cambridge Greensand (Cenomanian; fossils Albian in age).
The holotype of Ornithocheirus xyphorhynchus was illustrated by
The species was based on a fragmentary mandible, lacking the tip. It did not have a dentary crest (
NHMUK PV R 1823, partial mandibular rami (Fig. 20A–E).
St.–Leonards–on–Sea, Sussex. England.
Hastings Group, Wealden (late Berriasian / Valanginian).
The holotype of Pterodactylus sagittirostris consists of partial associated mandibular rami. The specimen is not comparable to any of the species referred in Lonchodectes by
Pterodactylus sagittirostris, holotype NHMUK PV R 1823 (upper Berriasian / Valangianian, Hastings Group), part of the mandibular rami. A right lateral view B respective line drawing C left lateral view D respective line drawing E ventral view. Abbreviation: d – dentary. Arrows indicate alveoli or teeth. Scale bar = 10 mm. Photos courtesy of The Natural History Museum.
As explained above, in 1869 Seeley created several names for pterosaur taxa from the Cretaceous of England, but these nomenclatural acts were disclaimed and, therefore, intentionally unavailable.
With the genus Ornithocheirus used as a wastebasket for the Cambridge Greensand species with unknown relationships, fragmentary specimens from other regions of the world have ended up being referred to Ornithocheirus as well. Among them is ‘Ornithocheirus’ wiedenrothi Wild, 1990 (holotype SMNS 56628; Fig. 21A–D), known from two pieces of a mandibular symphysis, a right articular, and fragments of wing bones from the Hauterivian of Germany (
‘Ornithocheirus’ wiedenrothi, holotype SMNS 56628 (Hauterivian, Engelbostel clay pit, Hannover), anterior part of the mandibular symphysis. A dorsal view B respective line drawing C left lateral view D respective line drawing. Abbreviations: d – dentary, sul – sulcus. Arrows and numbers indicate alveoli or teeth and their respective position. Scale bar = 10 mm.
Another occurrence of the Ornithocheirus complex is ‘Ornithocheirus’ sp. A, based on NHMUK PV R 11958, a mandible from the Purbeck Limestone Formation of England (Berriasian) (
ZIN PNT–S50–1, a partial mandibular symphysis from the Cenomanian of Lysaya Gora Hill, Saratov district, in the southern European part of Russia, has been referred either as Ornithocheirus(?) sp. (
Additional remains from the territories of the former Soviet Union referred as ornithocheirids come from different localities in Russia and in Uzbekistan and include cranial and postcranial elements and isolated teeth. These records include a partial tip of the rostrum (specimen ZIN PH no. 50/44), referred as Ornithocheirus sp., from the Khodzhakul locality in Uzbekistan (upper Albian or lower Cenomanian;
QM F10613, a mandibular symphysis from Albian Toolebuc Formation of Australia, has been referred to aff. Ornithocheirus sp. (
Aetodactylus halli, from the Tarrant Formation (middle Cenomanian) of Texas, is known only from its holotype, SMU 76383. This specimen consists of a complete mandible, whose tip bears resemblances to those from the Ornithocheirus complex (
Lastly, the species of the Ornithocheirus complex from England have been compared with the anhanguerids and related taxa from the Romualdo and Crato formations of Brazil. As shown above, no species from the Cambridge Greensand or the Chalk Formation shows a combination of characters only found in the genus Anhanguera: the presence of an anterior expansion of the rostrum, a premaxillary crest that begins near the tip of the rostrum, and the fifth and sixth upper alveoli smaller than the fourth and seventh (
Comparison between Ornithocheirus simus and Tropeognathus mesembrinus. A and C Ornithocheirus simus, holotype CAMSM B54428 (Albian, Cambridge Greensand), anterior part of the rostrum A anterior view C ventral view B and D Tropeognathus mesembrinus, holotype BSP 1987 I 46 (Aptian / Albian, Romualdo Formation), anterior part of the rostrum B anterior view D ventral view. Arrows and numbers mark the position of the first pair of alveoli. Scale bar = 10 mm.
The phylogenetic position of the species of the Ornithocheirus complexwithin Pterodactyloidea has been poorly studied.
The phylogenetic studies of
The cladistic matrix of
In addition, the strict consensus tree from this reanalysis shows some differences in relation to the one published by
Strict consensus tree of the reanalysis of the matrix by
Identical topology to the one found in the strict consensus tree of the reanalysis using PAUP* was recovered analyzing the same matrix in TNT (
It is interesting to note that the analyses of
The analyses of
A second dispute involves the use of postcranial material in the analyses of
The cervical vertebrae that
Therefore, reanalyses using the humeri and cervical vertebrae as separate OTUs were undertaken, using the data matrices of
Strict consensus tree of the reanalysis of the matrix by
Strict consensus tree of the reanalysis of the matrix by
In the case of the matrix by
Strict consensus tree of the reanalysis of the matrix by
In order to access the phylogenetic relationships of the species of the Ornithocheirus complex, we used a slightly modified version of the character matrix of
The run with all OTUs (with a total of 81 taxa, three of them outgroups, all characters treated as unordered) resulted in 30 most parsimonious trees with a length of 246 steps each. Several nodes were collapsed in the strict consensus tree, including some non–pterodactyloid taxa (as noted by
Strict consensus tree of the analysis including the species of the Ornithocheirus complex (see text for details).
The low resolution of the consensus tree is likely due to only a few taxa whose position changes greatly between different input trees (e.g.,
The agreement subtree of our analysis had only 46 taxa of the original 81 and presented a monophyletic Anhanguera, with Tropeognathus mesembrinus, Cearadactylus atrox, the clade comprising Ludodactylus sibbicki and ‘Cearadactylus’ ligabuei, and ‘Ornithocheirus’ polyodon as successive sister groups (Fig. 28). The recovery of Anhanguera piscator and Anhanguera santanae as sister groups, and Anhanguera spielbergi as sister group to the both, is an artifact: the synapomorphy of both species is the loss of the notarium, but this structure is present only in adult specimens, while Anhanguera piscator and Anhanguera santanae (AMNH 22555) are known by only immature individuals. The recovery of Cearadatylus atrox as the proximate sister group of Anhangueridae was first shown by
Agreement subtree of the analysis including the species of the Ornithocheirus complex.
A new analysis was undertaken and, in comparison to the agreement subtree, five additional species were not pruned from the analysis: Anhanguera araripensis, Anhanguera blittersdorffi, Caulkicephalus trimicrodon, Camposipterus nasutus, and Cimoliopterus cuvieri. The resulting strict consensus tree (Fig. 29) recovered Pteranodon longiceps as the sister group to all other pteranodontoids, and Istiodactylidae as sister group to the remaining ones, which formed a monophyletic group, supported by the presence of a ridge on the palate and a sulcus on the mandible (ch. 35.1). This clade is composed of Cimoliopterus cuvieri, ‘Ornithocheirus’ polyodon, and a clade with the remaining pteranodontoids, in a trichotomy. The latter, more restricted clade, is supported by the presence of an anterior expansion of the premaxillary tip with the jaw end high (ch. 18.1) and larger teeth located at the tip of the rostrum (ch. 48.1), and includes all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri. This clade was also recovered when no taxa were pruned, and is here named Anhangueria (see above). Among anhanguerians, Cearadactylus was recovered as polyphyletic, but it is worthy of notice that this information could never have been retrieved if not for the complete preparation and subsequent redescription of the holotype of Cearadactylus atrox by
Agreement subtree of the analysis including the species of the Ornithocheirus complex with additional five taxa not pruned.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with additional 11 taxa not pruned.
The agreement subtree plus Ornithocheirus simus recovers this taxon in a polytomy at the base of Pterodactyloidea + Rhamphorhynchidae (Fig. 31). The same happens when Lonchodraconidae is not pruned in the agreement subtree (Fig. 32). This is possibly an artifact caused by the incompleteness of the known specimens, and they most likely nest closer to anhanguerians. New characters and more complete specimens, besides perhaps preparation (either mechanical or virtual) of the holotype and only known specimen of Lonchodraco giganteus, are needed to better evaluate this question.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with Ornithocheirus simus not pruned.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with the Lonchodraconidae (more or less equivalent to the Lonchodectidae sensu
The species of the Ornithocheirus complex have been reviewed by several authors (e.g.,
One would expect that, having so many species based on fragmentary material, several would prove to be non–diagnosable. However,
Some species regarded by
Another major difference between the present work and the review done by
The pterosaur assemblage from the Cretaceous of England possesses a high diversity. One reviewer pointed out that this may be an artifact caused by lack of knowledge of intraspecific variation. This may well be true, but, as previously noted (e.g.,
The Cambridge Greensand is a remanié deposit (
Analysis of the phylogenetic relationships of these species, especially with the ones from the more or less coeval Romualdo and Crato formations of the Santana Group in Brazil, proved to be challenging, as expected. Using a modified version of the matrix by
We would like to acknowledge the editor, Hans–Dieter Sues, and Darren Naish and Alexander Averianov for their thorough review and constructive criticism. Thanks also to Juliana Sayão, Leonardo Avilla, Marcelo Britto and Ulisses Caramaschi for their valuable comments on the original version, Brian Andres for sharing his pterosaur bibliography, Richard Butler for discussion on pruning, Lorna Steel for photographs, information, and permission to photograph the specimens at the NHMUK, André Veldmeijer for photographs of Anhanguera spielbergi and discussion on taxonomy, Miguel Monné for help with synonymy lists, and the Willi Hennig Society for making TNT freely available. TR wishes to thank Oliver Rauhut for the supervision during her studies in Munich, Germany. We also thank the following curators for access to specimens: Oliver Rauhut (Bayerische Staatssammlung für Paläontologie und historische Geologie), Angela Milner, Sandra Chapman and Lorna Steel (Natural History Museum), Dan Pemberton and Matt Riley (Sedgwick Museum of Earth Sciences), Eberhard ‘Dino’ Frey (Staatliches Museum für Naturkunde Karlsruhe), Martina Kölbl–Ebert (Jura Museum Eichstätt), Rainer Schoch (Staatliches Museum für Naturkunde Stuttgart), Daniela Schwarz–Wings (Museum für Naturkunde Berlin), Gilles Cuny, Bent Lindow and Niels Bonde (Geological Museum, Natural History Museum of Denmark), Stephen Hutt (Dinosaur Isle Museum), David Gelsthorpe (Manchester Museum), Urs Oberli (Sankt Gallen), Jon de Vos (Nationaal Natuurhistorisch Museum, Naturalis), Mark Norell (American Museum of Natural History), Fabio Dalla Vecchia (Institut Català de Paleontologia), Mauro Bon (Museo di Storia Naturale di Venezia), Wang Xiaolin (Institute of Vertebrate Paleontology and Paleoanthropology), and Lü Junchang (Institute of Geology, Chinese Academy of Geological Sciences). Funding was provided by the Conselho Nacional de Desenvolvimento Tecnológico e Científico (CNPq – grants 130556/2005–0, 140407/2007–3 and 290019/2008–7 to TR and 304965/2006–5 to AWAK), Fundação Carlos Chagas Filho de Amparo à Pesquisa do Rio de Janeiro (FAPERJ – grant 26/102.779/2008 to AWAK), Deutscher Akademischer Austauschdienst (DAAD – grant A / 08 71633 to TR), Jackson School of Geosciences Student Member Travel Grant (grant to TR), Richard Gilder Graduate School Collection Study Grant (grant to TR). This research also received support from the SYNTHESYS Project (http://www.synthesys.info), which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program (grant NL–TAF–91 to TR).
Character list (modified from Wang et al. 2009).
1. Dorsal margin of the skull:
0 – straight or curved downward; 1 – concave; 2 – only rostrum curved upward
2. Upper and lower jaw:
0 – laterally compressed; 1 – comparatively broad
3. Rostral part of the skull anterior to the external nares (modified):
0 – reduced; 1 – elongated
4. Rostral end of premaxillae/maxillae downturned:
0 – absent; 1 – present
5. Anterior tip of the rostrum with a dorsally reflected palatal surface (added from Andres and Ji 2008):
0 – absent; 1 – present
6. Position of the external naris:
0 – above the premaxillary tooth row; 1 – displaced posterior to the premaxillary tooth row
7. Process separating the external nares:
0 – broad; 1 – narrow
8. Dorsoventrally compressed and elongated naris:
0 – absent; 1 – present
9. Naris size relative to the antorbital fenestra:
0 – naris smaller than antorbital fenestra; 1 – naris larger than antorbital fenestra; 2 – both very reduced (slit–like)
10. Naris and antorbital fenestra:
0 – separated; 1 – confluent, shorter than 45% of the skull length; 2 – confluent, longer than 45% of the skull length
11. Antorbital fenestra, shape:
0 – eliptical or ovoid; 1 – triangular, with base and height subequal; 2 – triangular with height larger than base; 3 – very elongated anteroposteriorly
12. Orbit comparatively small and positioned very high in the skull:
0 – absent; 1 – present
13. Orbit pear–shaped:
0 – absent; 1 – present
14. Position of the orbit relative to the nasoantorbital fenestra (naris + antorbital fenestra):
0 – same level or higher; 1 – orbit lower than the dorsal rim of the nasoantorbital fenestra
15. Suborbital opening:
0 – absent; 1 – present
16. Premaxillary sagittal crest, position (modified):
0 – absent; 1 – confined to the anterior portion of the skull, beginning near or at the tip of the rostrum; 2 – starting anterior to the anterior margin of the nasoantorbital fenestra, not reaching the skull roof above the orbit; 3 – starting anterior to the anterior margin of the nasoantorbital fenestra, extending beyond occipital region; 4 – starting at about the anterior margin of the nasoantorbital fenestra, reaching the skull roof above the orbit but not extending over the occipital region; 5 – starting close or at the anterior portion of the skull and extended over the occipital region; 6 – starting close or at the anterior portion of the skull, reaching orbit but not extended over the occipital region; 7 – starting at the posterior half of the nasoantorbital fenestra; 8 – located at the anterior portion of the skull, beginning more posteriorly but before the anterior margin of the nasoantorbital fenestra.
17. Premaxillary sagittal crest shape (modified):
0 – striated, low with a nearly straight dorsal margin; 1 – striated, high with a nearly straight dorsal margin; 2 – striated, high, spike–like; 3 – round dorsal margin, bladeshaped; 4 – smooth, moderately expanded anteriorly and forming a low rod–like extension posteriorly; 5 – smooth, very expanded anteriorly and forming a low rod–like extension posteriorly; 6 – smooth, starting low anteriorly and very expanded posteriorly; 7 – smooth, with a straight anterior margin, forming a triangular lateral outline
18. Expansion of the premaxillary tip:
0 – absent; 1 – present, with jaw end high; 2 – present, with the jaw end dorsoventrally flattened
19. Posterior ventral expansion of the maxilla:
0 – absent; 1 – present
20. Maxilla–nasal contact:
0 – broad; 1 – narrow; 2 – absent
21. Nasal process:
0 – absent; 1 – placed laterally, long, straight, and directed ventrally (not fused with maxillae); 2 – placed laterally, reduced; 3 – placed medially, long; 4 – placed medially, reduced; 5 – placed laterally, short and directed anteriorly
22. Foramen on nasal process:
0 – absent; 1 – present
23. Anterior process of jugal rodlike and deflected dorsally
0 – absent; 1 – present
24. Lacrimal process of the jugal:
0 – broad; 1 – thin, subvertical; 2 – thin, strongly inclined posteriorly
25. Bony frontal crest:
0 – absent; 1 – low and blunt; 2 – low and elongated; 3 – high and expanded posteriorly
26. Bony parietal crest:
0 – absent; 1 – present, blunt; 2 – present, laterally compressed and posteriorly expanded, with a rounded posterior margin; 3 – present, constituting the base of the posterior portion of the cranial crest
27. Posterior region of the skull rounded with the squamosal displaced ventrally:
0 – absent; 1 – present
28. Position of the quadrate relative to the ventral margin of the skull:
0 – vertical or subvertical; 1 – inclined about 120° backwards; 2 – inclined about 150° backwards
29. Position of the articulation between skull and mandible:
0 – under the posterior half of the orbit or further backwards; 1 – under the middle part of the orbit; 2 – under the anterior half of the orbit
30. Helical jaw joint:
0 – absent; 1 – present
31. Supraoccipital:
0 – does not extend backwards; 1 – extends backwards
32. Foramen pneumaticum piercing the supraoccipital:
0 – absent; 1 – present
33. Expanded distal ends of the paroccipital processes:
0 – absent; 1 – present
34. Basisphenoid:
0 – short; 1 – elongated
35. Palate with ridge and mandible with sulcus (added from Bennett 1994; Andres and Ji 2008):
0 – absent; 1 – present
36. Palatal ridge (modified):
0 – discrete, tapering anteriorly; 1 – strong, tapering anteriorly; 2 – strong, confined to the posterior portion of the palate
37. Maxilla excluded from the internal naris:
0 – absent; 1 – present
38. Opening between pterygoids and basisphenoid (interpterygoid opening):
0 – absent or very reduced; 1 – present and larger than subtemporal fenestra; 2 – present but smaller than subtemporal fenestra
39. Large distinct foramina (cup–shaped structures) on the lateral side anterior portion of the dentary:
0 – absent; 1 – present
40. Mandibular symphysis:
0 – absent or very short; 1 – present, at least 30% of mandible length
41. Expansion of the dentary tip (new):
0 – absent; 1 – present, with jaw end high; 2 – present, with the jaw end dorsoventrally flattened
42. Anterior tip of the dentary downturned:
0 – absent; 1 – present
43. Tip of the dentary projected anteriorly:
0 – absent; 1 – present
44. Dentary bony sagittal crest, presence:
0 – absent; 1 – present
45. Dentary bony sagittal crest, shape:
0 – blade–like and short, placed anteriorly; 1 – massive and deep
46. Distinctively elongated and posteriorly oriented articular and retroarticular process:
0 – absent; 1 – present
47. Position and presence of teeth:
0 – teeth present, evenly distributed along the jaws; 1 – teeth absent from the anterior portion of the jaws; 2 – teeth confined to the anterior part of the jaws; 3 – jaws toothless
48. Teeth size distribution (modified):
0 – all teeth roughly the same size; 1 – larger teeth located at the tip of the rostrum; 2 – larger teeth located posteriorly
49. Variation in the size of the anterior teeth with the 5th and 6th smaller than the 4th and 7th:
0 – absent; 1 – present
50. Teeth with a broad and oval base:
0 – absent; 1 – present
51. Multicusped teeth:
0 – absent; 1 – present
52. Teeth finely serrated (modified):
0 – present; 1 – absent
53. Peg–like teeth:
0 – absent; 1 – present, 15 or less on each side of the upper jaws; 2 – present, more than 15 on each side of the upper jaws
54. Long slender teeth:
0 – absent or less than 150; 1 – present, more than 150
55. Laterally compressed and triangular teeth:
0 – absent; 1 – present
56. Alveoli located on an elevation at the anterior end of the upper and lower jaws (“parapet–like” alveoli) (new, based on Unwin 2001):
0 – absent; 1 – present
57. Notarium:
0 – absent; 1 – present
58. Atlas and axis:
0 – unfused; 1 – fused
59. Postexapophyses on cervical vertebrae:
0 – absent; 1 – present
60. Lateral pneumatic foramen on the centrum of the cervical vertebrae:
0 – absent; 1 – present
61. Midcervical vertebrae:
0 – short, sub–equal in length; 1 – elongated; 2 – extremely elongated
62. Cervical ribs on midcervical vertebrae:
0 – present; 1 – absent
63. Neural spines of the midcervical vertebrae:
0 – tall, blade–like; 1 – tall, spike–like; 2 – low, blade–like; 3 – extremely reduced or absent
64. Number of caudal vertebrae:
0 – more than 15; 1 – 15 or less
65. Caudal vertebrae with elongated zygapophyses forming rod–like bony processes:
0 – absent; 1 – present
66. Length of the scapula:
0 – subequal or longer than coracoid; 1 – scapula shorter than coracoid (1 > sca/cor > 0.80); 2 – substantially shorter than coracoid (sca/cor ≤ 0.80)
67. Proximal surface of scapula:
0 – elongated; 1 – sub–oval
68. Shape of scapula:
0 – elongated; 1 – stout, with constructed shaft
69. Coracoidal contact surface with sternum:
0 – no developed articulation surface; 1 – articulation surface flattened, lacking posterior expansion; 2 – articulation surface oval, with posterior expansion
70. Deep coracoidal flange:
0 – absent; 1 – present
71. Broad tubercle on ventroposterior margin of coracoid:
0 – absent; 1 – present
72. Cristospine:
0 – absent; 1 – shallow and elongated; 2 – deep and short
73. Proportional length of the humerus relative to the metacarpal IV (hu/mcIV):
0 – hu/mcIV > 2.50; 1 – 1.50 < hu/mcIV < 2.50; 2 – 0.40 < hu/mcIV < 1.50; 3 – hu/mcIV < 0.40
74. Proportional length of the humerus relative to the femur (hu/fe):
0 – hu/fe ≤ 0.80; 1 – 1.4 > hu/fe > 0.80; 2 – hu/fe > 1.40
75. Proportional length of the humerus plus ulna relative to the femur plus tibia (hu+ul/fe+ti):
0 – humerus plus ulna about 0.80% or less of femur plus tibia length (hu+ul/fe+ti ≤ 0.80); 1 – humerus plus ulna larger than 0.80% of femur plus tibia length (hu+ul/fe+ti > 0.80)
76. Pneumatic foramen on the ventral side of the proximal part of the humerus:
0 – absent; 1 – present
77. Pneumatic foramen present on dorsal side of the proximal part of the humerus:
0 – absent; 1 – present
78. Deltopectoral crest of the humerus:
0 – reduced, positioned close to the humerus shaft; 1 – enlarged, proximally placed, with almost straight proximal margin; 2 – subrectangular, extending down the humerus shaft for at least 30% of humerus length; 3 – distally expanded; 4 – enlarged, hatchet shaped, proximally placed; 5 – enlarged, hatched shaped, positioned further down the humerus shaft; 6 – enlarged, warped; 7 – long, proximally placed, curving ventrally
79. Medial (= ulnar) crest of the humerus:
0 – absent or reduced; 1 – present, directed posteriorly; 2 – present, massive, with a developed proximal ridge
80. Distal end of the humerus:
0 – oval or D–shaped; 1 – subtriangular
81. Proportional length of the ulna relative to the metacarpal IV (ul/mcIV):
0 – ulna 3.6 times longer than metacarpal IV (ul/mcIV > 3.6); 1 – length of ulna between 3.6 and two times the length of metacarpal IV (3.6 > ul/mcIV > 2); 2 – ulna less than two times the length of metacarpal IV (ul/mcIV < 2)
82. Diameter of radius and ulna:
0 – subequal; 1 – diameter of the radius about half that of the ulna; 2 – diameter of the radius less than half that of the ulna
83. Distal syncarpals:
0 – unfused; 1 – fused in a rectangular unit; 2 – fused in a triangular unit
84. Pteroid:
0 – absent; 1 – shorter than half the length of the ulna; 2 – longer that half the length of the ulna
85. Metacarpals I–III:
0 – articulating with carpus; 1 – metacarpal III articulates with carpus, metacarpals I and II reduced; 2 – not articulating with carpus
86. Proportional length of the first phalanx of manual digit IV relative to the metacarpal IV (ph1d4/mcIV):
0 – both small and reduced; 1 – both enlarged with ph1d4 over four times the length of mcIV (ph1d4/mcIV> 4.0); 2 – both enlarged with ph1d4 between two and four times the length of mcIV (4.0 > ph1d4/mcIV≥ 2.0); 3 – both enlarged with ph1d4 less than two times the length of mcIV (ph1d4/mcIV< 2.0)
87. Proportional length of the first phalanx of manual digit IV relative to the tibiotarsus (ph1d4/ti):
0 – ph1d4 reduced; 1 – ph1d4 elongated and less than twice the length of ti (ph1d4/ti smaller than 2.00); 2 – ph1d4 elongated about or longer than twice the length of ti (ph1d4/ti subequal/larger than 2.00)
88. Proportional length of the second phalanx of manual digit IV relative to the first phalanx of manual digit IV (ph2d4/ph1d4):
0 – both short or absent; 1 – elongated with second phalanx about the same size or longer than first (ph2d4/ph1d4 larger than 1.00); 2 – elongated with second phalanx up to 30% shorter than first (ph2d4/ph1d4 between 0.70 – 1.00); 3 – elongated with second phalanx more than 30% shorter than first (ph2d4/ph1d4 smaller than 0.70)
89. Proportional length of the third phalanx of manual digit IV relative to the first phalanx of manual digit IV (ph3d4/ph1d4):
0 – both short or absent; 1 – ph3d4 about the same length or larger than ph1d4; 2 – ph3d4 shorter than ph1d4
90. Proportional length of the third phalanx of manual digit IV relative to the second phalanx of manual digit IV (ph3d4/ph2d4):
0 – both short or absent; 1 – ph3d4 about the same size or longer than ph2d4; 2 – ph3d4 shorter than ph2d4
91. Proportional length of the femur relative to the metacarpal IV (fe/mcIV):
0 – femur about twice or longer than metacarpal IV (fe/ mcIV ≥ 2.00); 1 – femur longer but less than twice the length of metacarpal IV (1.00 < fe/mcIV < 2.00); 2 – femur about the same length or shorter than metacarpal IV (fe/mcIV ≤ 1.00)
92. Length of metatarsal III:
0 – more than 30% of tibia length; 1 – less than 30% of tibia length
93. Fifth pedal digit:
0 – with four phalanges; 1 – with 2 phalanges; 2 – with 1 or no phalanx (extremely reduced)
94. Last phalanx of pedal digit V:
0 – reduced or absent; 1 – elongated, straight; 2 – elongated, curved; 3 – elongated, very curved (boomerang shape)
Data matrix (modified from
Ornithosuchus longidens
0000000000 000000–000 0–00000000 00000–0000 0000–00000 0000000000 0000000000 0000000000 0000000000 0000
Scleromochlus taylori
000?00?000 00?000–000 0–0?00000? ????????0? 00?0–000?0 0?00?00??? 00?00????? ???00??0?? ?0?0?????? ????
Herrerasaurus ischigualastensis
0000000000 000000–000 0–00000000 00000–0000 0000–00000 0000000000 0000000000 0000000000 0000000000 0020
Anurognathus ammoni
010000100? ??0??0–00? 0–?00????? ????????00 0000–00000 0110000??? 0??10????? ??011??100 00?1011??? 001?
Jeholopterus ningchengensis
01000010?? ?????0–00? ????00??0? ????????0? 0000–00000 0110000??? 01?1000010 0?021??1?? 00?1011222 1011
Dendrorhynchoides curvidentatus
010000100? ?????0–00? 0–??0????? ????????0? 0000–00000 0110000??? 00?100?010 0?021??1?? 00?1011222 00??
Batrachognathus volans
?10000100? ?????0–00? 0–??0????? ??????0?00 0000–00000 011000??0? 0?0???001? ???21??10? ?????????? ????
Cacibupteryx caribensis
001??10000 000000–?00 0–00000000 00000–10?? ??????00?? ?????????? ?????????? ?????????? ?????????? ????
Eudimorphodon ranzii
0010010000 000000–000 0–0000010? 0?0?????00 0100–00000 1100000?00 00?0?00010 0?11?00200 11?10????? 1???
‘Eudimorphodon’ cromptonellus
?????????? ?????????? ????00???? ?000?????? ?????0???0 11??000??? ?????????? ??111????? 1???021211 101?
Carniadactylus rosenfeldi
?01??1??00 ????????0? ?????????? ?????????0 ??????0000 1100000?00 0????0???0 0?1110???? 1????21111 00??
Peteinosaurus zambellii
?0???????? ?????????? ?????????? ?????????? ?????000?0 1100000??? ???010001? ??111??100 11?1021111 0011
Dimorphodon macronyx
0010010010 200000–001 0–0000000? 0?0?????00 0000–00000 0100000?00 0000100010 ??11100100 1111021111 1011
Wukongopterus lii
0010010??? ???????00? ??0????1?? ????????00 0000–00000 0120000??0 10201000?0 ??111?01?? 10?1021111 1013
Dorygnathus banthensis
0010010010 000000–000 0–00000100 0?0????101 0010–00000 0100000??? 0000100010 ??11100300 10?1021111 1013
Scaphognathus crassirostris
0010010000 000000–000 0–00000100 0?000–010? 0000–00000 0100000?00 0000100010 0111100100 10?1021111 0013
Preondactylus buffarinii
00100100?0 ?0?000–00? 0–??000?0? ?????????0 0000–?0?00 ??00000??? 0??0?????? ??111??1?? 10?1021111 00??
Sordes pilosus
0010010000 ?00000–00? 0–?000010? 0?0?????00 0000–00000 01?0000?0? 0000100010 0?011??100 00?1021111 0013
Austriadactylus cristatus
0010010010 0000061000 0–000?000? ?????????? ??????0000 000000???? 00?00????0 ?????????? ?0???????1 ????
Angustinaripterus longicephalus
0010010100 3000020101 ??00????0? ????????00 ??????0000 01000????? ?????????? ?????????? ?????????? ????
Harpactognathus gentryii
00100101?0 ?????2?0?? ?????????? ????0–1??? ??????0??? ?????????? ?????????? ?????????? ?????????? ????
Campylognathoides zitteli
0010010010 100000–000 0–1000010? 0???????00 0100–00000 0100000000 0000100010 0?1???12?? 11?1012122 0010
Campylognathoides liasicus
0010010010 100000–000 0–10000100 0?000–1100 0100–00000 0100000000 0000100010 0111100200 11?1012122 1010
Caviramus schesaplanensis
?0???????? ?????????? ?????????? ????????10 ?000–100?0 110000???? ?????????? ?????????? ?????????? ????
Raeticodactylus filisurensis
0010010000 0000022000 0–00??010? ????????10 0001110000 1100000??? 00???????? ???21??20? ??????1211 ?0??
Rhamphorhynchus longicaudus
0010010020 000000–000 0–00000110 0?000–??01 0010–00000 010000000? 0000100010 0111100400 11?1022222 1012
Rhamphorhynchus muensteri
0010010020 000000–000 0–00000110 00000–1101 0010–00000 0100000000 0000100010 0111100400 11?1012222 1012
Pterodactylus antiquus
00100100–1 –00000–002 100000121? 0?0?0–??01 0000–00000 0120000000 1121000010 0121100700 21?1031222 2020
Pterodactylus kochi
00100100–1 –00000–002 100000121? 0?0?0–??01 0000–00000 0120000000 1121000010 0121100700 21?2031222 2020
Germanodactylus ramphastinus
00100100–1 –000040002 100000121? 0???????0? 0000–00000 0120000??? 11??0?001? ??211??7?0 21???31??? 2???
Germanodactylus cristatus
00100100–1 –000040002 100000121? 0?0?????01 0000–00000 0120000??? 11???00010 0?211??7?0 21???31222 21??
Gnathosaurus subulatus
00100100–1 –000020202 500000121? 0?0???0201 2000–00100 010000???? ?????????? ?????????? ?????????? ????
Pterodaustro guinazui
10100100–1 –00000–002 100000121? 0?0?????01 0000–00000 010100???0 11210?001? ??211??7?0 2?1??31222 2020
Ctenochasma gracile
10100100–1 –0?000–002 0–0?00121? 0?0?????01 0000–00000 010100??00 112?0?001? ??211??7?? 21???312?? 2020
Feilongus youngi
10100100–1 –000010002 10000212?0 0?0?0–??0? 0000–02000 010000???? ?????????? ?????????? ?????????? ????
Cycnorhamphus suevicus
10100100–1 –00000–002 200002121? ??0?0–??01 0000–02000 010000?00? 112??0001? ??210??700 21??031222 21??
Gallodactylus canjuersensis
10100100–1 –0?000–002 20??02??1? ??0?????0? 0?00–02000 010000???? ?????0001? ?????????? ??????1222 ?1??
Nyctosaurus bonneri
00100100–1 –00000–002 0–00000?1? 0?0?????01 0000–03–00 0100001?1? 010?0????? ??31???50? ????232222 2???
Nyctosaurus gracilis
00100100–1 –00000–002 0–00000?11 0?010–1201 0000–03–00 0100001110 0101000010 013111050? 2122232222 20??
Nemicolopterus crypticus
00100100–1 –00000–002 6001000?2? ????????01 0000–03–00 0100000??0 01?10000?0 ???11?17?? 2????31?22 2120
Aussiedraco molnari
?0???????? ?????????? ?????????? ????1???01 0000–?00?0 ???000???? ?????????? ?????????? ?????????? ????
‘Ornithocheirus’ polyodon
?010110??? ?????0–0?? ?????????? ????10???? ??????00?0 ???000???? ?????????? ?????????? ?????????? ????
‘Ornithocheirus’ denticulatus
?010110??? ?????0–0?? ?????????? ????10???? ??????0000 ???000???? ?????????? ?????????? ?????????? ????
‘Pterodactylus’ daviesii
?0???????? ?????????? ?????????? ????1???01 0000–?0000 ???000???? ?????????? ?????????? ?????????? ????
‘Ornithocheirus’ platystomus
?010?10??? ?????170?? ?????????? ????10???? ??????0??0 ???000???? ?????????? ?????????? ?????????? ????
Ornithocheirus simus
?010010??? ?????0–0?? ?????????? ????10???? ??????0000 ?100?0???? ?????????? ?????????? ?????????? ????
Cimoliopterus cuvieri
?010110??? ?????830?? ?????????? ????10???? ??????0000 01?000???? ?????????? ?????????? ?????????? ????
Pteranodon longiceps
10100100–1 –00000–002 4000330121 01010–1201 0000–03–00 0100001111 0111011010 0121110611 2122232222 2020
Nurhachius ignaciobritoi
00101100–2 –00000–002 ??02???12? ????????0? 0000–02000 01001010?? 011??1?1?0 ?22110?61? 222?1312?? 21??
Istiodactylus latidens
00101100–2 –?0??0–0?2 0–?2?????0 ????0–???1 0000–?2000 0100101?1? 0????11110 02?1???611 ?2?????1?? ????
Noripterus complicidens
?????????? ?????????? ?????????? ?????????? ?????????1 0?00?0??11 010??????? ??20001720 211??312?? 2120
‘Phobetor’ parvus
00100100–1 –100131012 0–0023012? 1?1?????01 0000–01–01 010000???? ?????????? ?????????? ??1??????? ????
Dsungaripterus weii
20100100–1 –100131012 0–00230121 1111101201 0000–01–01 0100001111 0101000010 ?2200??7?0 211?131222 2120
Lonchodraco(?) microdon
001??10??? ?????0–?0? ?????????? ????11???? ??????0000 ???001???? ?????????? ?????????? ?????????? ????
Lonchodraco machaerorhynchus
?0???????? ?????????? ?????????? ????1???01 ???11?00?0 ???001???? ?????????? ?????????? ?????????? ????
Lonchodraco giganteus
?010010??? ?????170?? ?????????? ????11??01 00010?0000 010001???? ?????????? ?????????? ?????????? ????
Zhejiangopterus linhaiensis
00100100–1 –00100–002 0–000?012? 1???????0? 0000–03–00 01000011?? 213?0000?1 0?200??7?? 21?2?313?? 2???
Azhdarcho lancicollis
?????????? ?????????? ?????????? ?????????? ??????3–00 0100001110 213??????? ?????107?0 ?????????? ????
Quetzalcoatlus sp.
00100100–1 –00107?002 0–00???121 ????0–1?01 0000–03–00 0100001110 213?000011 0?20010720 211??31322 2?20
Tupandactylus imperator
00110100–2 –011055002 ??0123012? 1????????1 ?001103–00 010000???? ?????????? ?????????? ?????????? ????
Tapejara wellnhoferi
00110100–2 –011055002 ??01230120 11110–1201 0001103–00 0100000?11 010?000010 1?20011720 2112?31??? 2120
Thalassodromeus sethi
00100100–2 –011056002 4001230121 1111121201 0000–03–00 010000???? ?????????? ?????????? ?????????? ????
Tupuxuara leonardii
00100100–2 –011056002 ??01230121 1111121201 0000–03–00 0100001111 010??00010 1220010720 211??313?? 2???
‘Cearadactylus’ ligabuei
10101100–? ?????0–102 ?????????? ????101??? ??????0100 010000???? ?????????? ?????????? ?????????? ????
Camposipterus(?) colorhinus
?01011???? ?????0–1?? ?????????? ????10???? ??????01?0 ???000???? ?????????? ?????????? ?????????? ????
Camposipterus(?) sedgwickii
?01011???? ?????0–1?? ?????????? ????10???? ??????0100 ???000???? ?????????? ?????????? ?????????? ????
Camposipterus nasutus
0010110??? ?????0–10? ?????????? ????10???? ??????0100 ???000???? ?????????? ?????????? ?????????? ????
Brasileodactylus araripensis
?0???????? ?????????? ?????????? ????1???01 1000–?01?0 0??000???? ?????????? ?????????? ?????????? ????
Ludodactylus sibbicki
10101100–1 –00000–?02 3100330121 ????101?01 ?000–00100 010000???? ?????????? ?????????? ?????????? ????
Cearadactylus atrox
00101100–1 –00001?102 ??00???121 ????101??1 1000–001?0 010000???? ?????????? ?????????? ?????????? ????
Caulkicephalus trimicrodon
?0101100–? –????13102 ????33???? ????101??? ??????0100 010000???? ?????????? ?????????? ?????????? ????
‘Ornithocheirus’ capito
?010110??? ?????1?1?? ?????????? ?????????? ??????01?0 010000???? ?????????? ?????????? ?????????? ????
Coloborhynchus clavirostris
?010110??? ?????1?1?? ?????????? ????10???? ??????01?0 ??00?0???? ?????????? ?????????? ?????????? ????
Liaoningopterus gui
?0101100–? –000013102 ??00??0121 ????????01 ?0010001?0 010000??11 01???????? ?????????? ?????????? ????
Anhanguera araripensis
00101100–1 –000013102 ??00110121 0101101201 ?????00110 0?0000???? ?????????? ?????01611 ?20??????? ????
Anhanguera spielbergi
00101100–1 –000013102 ?–00110121 0101101201 1001000110 0100001?11 ?????21120 02?1101611 ?2???????? ????
Anhanguera robustus
?0???????? ?????????? ?????????1 ????1???01 10010001?0 010000???? ?????????? ?????????? ?????????? ????
Anhanguera piscator
00101100–1 –000013102 3100110121 0101??1201 1001000110 0100000111 0111021120 0221101611 222?1????? 2120
Anhanguera blittersdorffi
00101100–1 –000013102 ??00110121 0101101201 1001000110 010000???? ?????????? ?????????? ?????????? ????
Anhanguera santanae
00101100–1 –00001?102 3100110121 0101101201 ?00??00110 0100000111 0111?21120 0????01611 ?22?1????? ????
Tropeognathus mesembrinus
00101100–1 –000013102 ??00110121 0101111201 1001000100 010000???? ?????????? ?????????? ?????????? ????