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Taxonomic Review of the Ornithocheirus Complex (Pterosauria) From the Cretaceous of England

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Taxonomic Review of the Ornithocheirus Complex (Pterosauria) From the Cretaceous of England

Taissa Rodrigues et al. Zookeys.

Abstract

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. Lonchodraconidaefam. n. (more or less equivalent to LonchodectidaesensuUnwin 2001) is a monophyletic entity, but its exact phylogenetic position remains uncertain, as is the case of Ornithocheirus simus. Therefore, it is proposed that Ornithocheiridae should be constricted to its type species and thus is redundant. Other taxa previously referred as "ornithocheirids" are discussed in light of the revised taxonomy.

Keywords: Anhangueria; Anhangueridae; Cambridge Greensand; Cretaceous; Lonchodraconidae; Ornithocheiridae; Pterodactyloidea.

Figures

Figure 1.
Figure 1.
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.
Figure 2.
Figure 2.
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.
Figure 3.
Figure 3.
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.
Figure 4.
Figure 4.
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.
Figure 5.
Figure 5.
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.
Figure 6.
Figure 6.
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.
Figure 7.
Figure 7.
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.
Figure 8.
Figure 8.
‘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 Seeley 1881.
Figure 9.
Figure 9.
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.
Figure 10.
Figure 10.
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.
Figure 11.
Figure 11.
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.
Figure 12.
Figure 12.
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.
Figure 13.
Figure 13.
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 Seeley 1870. Photos A and C courtesy of The Natural History Museum.
Figure 14.
Figure 14.
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.
Figure 15.
Figure 15.
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.
Figure 16.
Figure 16.
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.
Figure 17.
Figure 17.
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.
Figure 18.
Figure 18.
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.
Figure 19.
Figure 19.
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 Seeley 1881.
Figure 20.
Figure 20.
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.
Figure 21.
Figure 21.
‘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.
Figure 22.
Figure 22.
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.
Figure 23.
Figure 23.
Strict consensus tree of the reanalysis of the matrix by Lü et al. (2010). Lonchodectes is used sensu Unwin (2001).
Figure 24.
Figure 24.
Strict consensus tree of the reanalysis of the matrix by Lü et al. (2010), with the codings of the humeri and cervical vertebrae in Lonchodectes (sensu Unwin 2001) as separate OTUs.
Figure 25.
Figure 25.
Strict consensus tree of the reanalysis of the matrix by Lü et al. (2010), with the OTU Lonchodectes (sensu Unwin 2001) restricted to codings of skull and scapulocoracoid characters.
Figure 26.
Figure 26.
Strict consensus tree of the reanalysis of the matrix by Wang et al. (2008), with the humeri and cervical vertebrae referred to Lonchodectes by Lü et al. (2010) included as separate OTUs.
Figure 27.
Figure 27.
Strict consensus tree of the analysis including the species of the Ornithocheirus complex (see text for details).
Figure 28.
Figure 28.
Agreement subtree of the analysis including the species of the Ornithocheirus complex.
Figure 29.
Figure 29.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with additional five taxa not pruned.
Figure 30.
Figure 30.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with additional 11 taxa not pruned.
Figure 31.
Figure 31.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with Ornithocheirus simus not pruned.
Figure 32.
Figure 32.
Agreement subtree of the analysis including the species of the Ornithocheirus complex with the Lonchodraconidae (more or less equivalent to the Lonchodectidae sensu Unwin 2001) not pruned.

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