ZooKeys 308: 1–112, doi: 10.3897/zookeys.308.5559
Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England
Taissa Rodrigues 1,†, Alexander Wilhelm Armin Kellner 2,‡
1 Department of Biology, Agrarian Sciences Center, Universidade Federal do Espírito Santo. Alto Universitário s/n, Caixa Postal 16, Guararema, CEP 29500–000, Alegre, ES, Brazil
2 Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and Paleontology, Museu Nacional / Universidade Federal do Rio de Janeiro. Quinta da Boa Vista s/n, São Cristóvão, CEP 20940–040, Rio de Janeiro, RJ, Brazil

Corresponding author: Taissa Rodrigues (taissa.rodrigues@gmail.com)

Academic editor: Hans–Dieter Sues

received 20 May 2013 | accepted 21 May 2013 | Published 12 June 2013


(C) 2013 Taissa Rodrigues. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


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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, ‘Ornithocheiruscapito, Camposipterus nasutus comb. n., Camposipterus(?) sedgwickii comb. n., Camposipterus(?) colorhinus comb. n., Cimoliopterus cuvieri comb. n., ‘Ornithocheiruspolyodon, ‘Ornithocheirusplatystomus, ‘Pterodactylusdaviesii, and ‘Ornithocheirusdenticulatus. 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 ‘Cearadactylusligabuei belongs to a different genus than Cearadactylus atrox. Lonchodraconidae fam. n. (more or less equivalent to Lonchodectidae sensu Unwin 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

Pterodactyloidea, Ornithocheiridae, Anhangueridae, Lonchodraconidae, Anhangueria, Cretaceous, Cambridge Greensand

Introduction

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) (Rawson et al. 1978; Unwin et al. 2000; Unwin 2001; Barrett et al. 2008; Witton et al. 2009; Naish et al. 2013). The Cambridge Greensand in itself is one of the richest pterosaur deposits in the world (Wellnhofer 1991a; Kellner 1994), with over 2000 known specimens. Unlike that from most other pterosaur–bearing deposits, the Cambridge Greensand material is not flattened; it is, however, quite fragmentary and found isolated (Hooley 1914; Wellnhofer 1978; Kellner 1990; Wellnhofer 1991a; Unwin 2001). The nature of these fossils, along with decades of competing taxonomic proposals, synonymisations, and misunderstandings, as detailed below, made their taxonomy controversial (see Unwin 2001 and Table 1).

Table 1.

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 Ornithocheiruscapito
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 Pterodactylusdaviesii
Ornithocheirus dentatus Seeley, 1870 Ornithocheirus dentatus Ornithocheirus dentatus Ornithocheirus dentatus Anhanguera cuvieri nomen dubium
Ornithocheirus denticulatus Seeley, 1870 Ornithocheirus denticulatus Ornithocheirus denticulatus Anhanguera cuvieri Ornithocheirusdenticulatus
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 Ornithocheirusplatystomus
Ornithocheirus polyodon Seeley, 1870 Ornithocheirus polyodon Ornithocheirus polyodon Ornithocheirus polyodon Anhanguera fittoni Ornithocheiruspolyodon
Ornithocheirus reedi Seeley, 1870 Ornithocheirus reedi Criorhynchus reedi Criorhynchus reedi Coloborhynchus capito Ornithocheiruscapito
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
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 Ornithocheiruscapito
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 Pterodactylusdaviesii
Ornithocheirus dentatus Seeley, 1870 Ornithocheirus dentatus Ornithocheirus dentatus Ornithocheirus dentatus Anhanguera cuvieri nomen dubium
Ornithocheirus denticulatus Seeley, 1870 Ornithocheirus denticulatus Ornithocheirus denticulatus Anhanguera cuvieri Ornithocheirusdenticulatus
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 Ornithocheirusplatystomus
Ornithocheirus polyodon Seeley, 1870 Ornithocheirus polyodon Ornithocheirus polyodon Ornithocheirus polyodon Anhanguera fittoni Ornithocheiruspolyodon
Ornithocheirus reedi Seeley, 1870 Ornithocheirus reedi Criorhynchus reedi Criorhynchus reedi Coloborhynchus capito Ornithocheiruscapito
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) (International Commission On Zoological Nomenclature 1999). What apparently has gone unnoticed is that the nomenclatural acts concerning the naming of new species in Seeley’s 1869 work were disclaimed and therefore intentionally not available. In the first paragraph of page xv, it can be read in the definition of the work: “An approximate list of the species included in the following Catalogue, with provisional names for new species and reference to the specimens on which they are founded, and to the pages of the Index in which they are described. ” [emphasis added] and also a footnote also on page xv explaining the term “provisional names”: “These names are only intended for the convenience of students using the Museum, and not necessarily to take rank as names of described species” [emphasis added]. Disclaimed acts are recognized in ICZN’s article 8.3, with the result that the names then given for new species were not available.

It is also possible to interpret these sentences (Seeley 1869: xv) as stating that the new names for genera were not disclaimed. Concerning Ornithocheirus, it had a description that, albeit inadequate (“This genus has no teeth anterior to the palate.”), can be regarded as an action that makes the name available. Three species were originally referred to the genus: Ornithocheirus simus, “Ornithocheirus carteri” and “Ornithocheirus platyrhinus”. Since the last two were not available (as explained above), the only available species left on the original description, Ornithocheirus simus, is thus the type species of the genus Ornithocheirus by monotypy (ICZN articles 67.2, 67.2.1 and 68.3) (Unwin and Bakhurina 2000; Unwin 2001). “Ptenodactylus” Seeley, 1869, with 21 referred ‘species’, was not only preoccupied by Ptenodactylus Gray, 1845 (see Unwin 2001), but it was not associated with a description and thus can be considered a nomen nudum.

Seeley (1870) placed 27 species from the Cretaceous of England in the genus Ornithocheirus (Table 2). Although recognizing that Ornithocheirus simus, Ornithocheirus carteri and Ornithocheirus platyrhinus are known by much more massive jaws than the lanceolate tips of the other species, Seeley (1870) mistakenly reinterpreted the holotype of Ornithocheirus simus as a mandible, and thus his 1869 diagnosis of Ornithocheirus, based on the absence of anteriorly directed teeth in the premaxilla, would be invalidated. So he referred all these species to the same genus and re–diagnosed it as “in which teeth are prolonged anterior to the muzzle, and the palate has a longitudinal ridge”. The lumping of all these species into one single genus was the beginning of what can be referred as the Ornithocheirus complex: a wastebasket genus for species of uncertain relationships and represented by fragmentary type material (see Kellner 1994).

Table 2.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Seeley (1870).

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, Owen proposed two new genera for the Cretaceous British pterosaurs, Criorhynchus and Coloborhynchus. Criorhynchus was created as a monotypic genus including only Criorhynchus simus because Owen (1874) considered the name Ornithocheirus inappropriate (ornitho, bird, and cheirus, hand). At the time, the British Association Code allowed such emendations on ‘inappropriate’ names (Dayrat 2010), but the present ICZN has modified that. As both Ornithocheirus Seeley, 1869 and Criorhynchus Owen, 1874 have the same type species, they are objective synonyms, and the former has priority over the later (Principle of Priority, ICZN articles 23.1 and 61.3.3). Coloborhynchus had three species referred to it, and no type species was designated. Many years later, Coloborhynchus clavirostris was subsequently designated as type species by Kuhn (1967).

Lydekker (1888), following the suggestions by Seeley (1870), provided a catalogue in which then other named genera from the Cretaceous of England were synonymized with Ornithocheirus (with the incorrect spelling Ornithochirus) (Table 3). In addition, he included 14 species in Ornithocheirus, as “family uncertain”. He considered that “all the species are known by such fragmentary remains that no accurate diagnosis can be given” (Lydekker 1888: 10) and also that “many of the species are probably invalid” (Lydekker 1888: 11).

Table 3.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Lydekker (1888). = indicate synonymies.

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

Newton (1888) agreed with Lydekker (1888) that these species, based on fragmentary remains, should be included in a single genus, Ornithocheirus. However, he claimed the existence of 40 species, a number much higher than proposed by both Seeley (1870) and Lydekker (1888) but, unfortunately, did not list these species or provide information on how he achieved this number. Newton (1888) pointed out that the discovery of new, more complete specimens would probably at the same time reduce the number of species but increase the number of genera.

Woods (1891) provided a catalogue of the type fossils in the Woodwardian Museum in Cambridge and listed 25 species in the genus Ornithocheirus, similar to Seeley (1870), Lydekker (1888), and Newton (1888) (Table 4).

The first major review of the Ornithocheirus complex was provided by Hooley (1914) (Table 5). He provided an appraisal of previous reviews of the genus, reviewed the species present in the collection of the Sedgwick Museum and in other museums, and divided them in five groups or genera based on morphological characters. His group n. 1 had 16 species and was named Ornithocheirus; group n. 2, with nine species, Lonchodectes; group n. 3 held three species and was named Amblydectes; and group n. 4, with six species, was termed Criorhynchus (including Coloborhynchus clavirostris as a synonym of Criorhynchus simus). He added a fifth group, restricted to the edentulous form Ornithostoma, which was not part of the Ornithocheirus complex used by him.

Table 4.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Woods (1891). = indicate synonymies.

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
Table 5.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Hooley (1914). = indicate synonymies.

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. Seeley (1869) was the first to recognize that Ornithocheirus simus was quite distinct from most other pterosaurs from this complex and introduced the genus Ornithocheirus for its reception, separating this species from the ones known by lanceolate jaws. As noted above, Owen (1874) accepted the distinction but regarded the name inappropriate, and thus assigned Ornithocheirus simus to a different genus (Criorhynchus). Lydekker (1888: 16, footnote) acknowledged that Ornithocheirus simus was the type species of the genus Ornithocheirus and both Lydekker (1888) and Hooley (1914) agreed that Ornithocheirus simus was distinguished by its tall rostrum, whereas most other species then referred in Ornithocheirus had lanceolate jaw tips. In order to avoid confusion, Lydekker (1888) preferred to use Criorhynchus for Ornithocheirus simus and Ornithocheirus for the species with lanceolate tips. Hooley (1914) was of similar opinion and favored the name Criorhynchus for Ornithocheirus simus and other species with tall rostra, and Ornithocheirus for the other taxa.

Subsequent authors tended to divide the species of the Ornithocheirus complex in only two genera, Ornithocheirus and Criorhynchus (e.g., Kuhn 1967; Wellnhofer 1978; Kellner and Tomida 2000), excluding Pterodactylus simus from Ornithocheirus and referring it to Criorhynchus, following Lydekker (1888). One major consequence was the uncertainty regarding a type species for Ornithocheirus. Khozatskii and Yur’ev (1964) referred Pterodactylus compressirostris as type species of Ornithocheirus and Pterodactylus simus, of Criorhynchus, both in the family Ornithocheiridae. They also considered Amblydectes and Lonchodectes as Pterosauria incertae sedis.

Kuhn (1967) recognized the genus Criorhynchus, with Criorhynchus simus as type species, in the family Criorhynchidae, with seven species which, according to him, were almost all indefinable. He also recognized the genus Ornthocheirus [sic], with approximately 25 described speciesand Pterodactylus compressirostris as its type species, in the family Ornithocheiridae and subfamily Ornithocheirinae. He was the first author to define type species for the genera Amblydectes and Lonchodectes, respectively Amblydectes crassidens and Lonchodectes compressirostris (Kuhn 1967: 46, using the term “Genotypus”), but considered both genera indeterminate and deemed as synonyms of Ornithocheirus.

Wellnhofer (1978) published a major reference work on pterosaurs. He discussed the species included in Ornithocheirus based on the diagnoses provided by Seeley (1869, 1870), and drew attention to which was the first species cited in the lists provided by this author, probably trying to elucidate Seeley’s original intentions. He did not recognize Pterodactylus simus as the type species of Ornithocheirus and referred it as the type and only valid species of Criorhynchus, in the family Criorhynchidae. Four species were considered synonymous with Criorhynchus simus, and others were referred as Criorhynchidae incertae sedis. All other species of the Ornithocheirus complex were referred to the family Ornithocheiridae. Following Kuhn (1967), he incorrectly referred Pterodactylus compressirostris as the type species of Ornithocheirus and, from the Ornithocheirus complex, attributed eight species to the genus Ornithocheirus, four to Ornithocheiridae incertae sedis, and considered 17 of uncertain systematic position, among them four non–British species (Table 6).

Table 6.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Wellnhofer (1978). = indicate synonymies.

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
Ornithocheirusbunzeli
Ornithocheirushilsensis
Ornithocheirushlavatschi
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

Unwin (2001) undertook the most recent review. He revised the taxonomic history of the pterosaurs from the Cambridge Greensand and, although more complete specimens of the Ornithocheirus complex from the Cretaceous of England had not been discovered, he compared them with the more complete and more recently described pterosaurs from the Santana Group of Brazil. Unwin (2001) designated Pterodactylus simus as the type species of Ornithocheirus and divided the Cambridge Greensand species in two families and four genera: Ornithocheiridae with a monospecific Ornithocheirus, Coloborhynchus with two species, Anhanguera with two species, and Lonchodectidae comprising six species of the genus Lonchodectes (Table 7). Several named species were synonymized with others but diagnoses for the species considered valid were not presented and thus it is not clear which characters were used for these referrals.

Table 7.

List of taxa of the Ornithocheirus complex from the Cretaceous of England, after Unwin (2001). = indicate synonymies.

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 Seeley [1881]) has never been figured and the holotype could not be located. Wellnhofer (1978) listed this species under the name Ornithocheirus, but among the species that, according to him, had uncertain systematic positions. Unwin (2001) recently synonymized it with Lonchodectes microdon (but see below). Here, we note these referrals but refrain from providing a discussion about it.

Table 8.

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
Ornithocheiruscapito 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.
Ornithocheiruspolyodon Seeley, 1870
Pterodactyloidea incertae sedis
Ornithocheirusplatystomus Seeley, 1870
Pterodactylusdaviesii Owen, 1874
Ornithocheirusdenticulatus Seeley, 1870
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
Ornithocheiruscapito 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.
Ornithocheiruspolyodon Seeley, 1870
Pterodactyloidea incertae sedis
Ornithocheirusplatystomus Seeley, 1870
Pterodactylusdaviesii Owen, 1874
Ornithocheirusdenticulatus Seeley, 1870
Table 9.

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
Table 10.

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. Rodrigues and Kellner (2008) reviewed the genus Coloborhynchus and its presence in the Santana Group, and the species from the Cambridge Greensand that were briefly cited in that work are also examined here.

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.

Systematic palaeontology
Pterosauria Kaup, 1834 Pterodactyloidea Plieninger, 1901
Ornithocheiridae Seeley, 1870

http://species-id.net/wiki/Ornithocheiridae

Ornithocheiridae Seeley: Seeley 1870: p. 110
Criorhynchidae Hooley: Hooley 1914: p. 557
Type genus.

Ornithocheirus Seeley, 1869.

Included genus.

Ornithocheirus.

Recorded temporal range.

Albian.

Recorded stratigraphic range.

Cambridge Greensand, England.

Diagnosis.

the same as for the type genus.

Remarks.

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.

Ornithocheirus Seeley, 1869

http://species-id.net/wiki/Ornithocheirus

Ornithocheirus Seeley: Seeley 1869: p. xvi
Ornithocheirus Seeley: Seeley 1870: p. 112
Criorhynchus Owen: Owen 1874: p. 7
Ornithochirus [sic] Seeley: Lydekker 1888: p. 10
Criorhynchus Owen: Kuhn 1967: 38
Type species.

Pterodactylus simus Owen, 1861, by monotypy.

Recorded temporal range.

Albian.

Recorded stratigraphic range.

Cambridge Greensand, England.

Diagnosis.

As for the type species.

Ornithocheirus simus (Owen, 1861)

http://species-id.net/wiki/Ornithocheirus_simus

Figs 13
Pterodactylus simus Owen: Owen 1861: p. 2, pl. I, fig. 1–5
Ornithocheirus simus (Owen): Seeley 1869: p. xvi
Ornithocheirus simus (Owen): Seeley 1870: p. 127
Criorhynchus simus (Owen): Owen 1874: p. 7
Ornithochirus [sic](?) simus (Owen): Lydekker 1888: p. 16
Criorhynchus simus (Owen): Hooley 1914: p. 536
Criorhynchus simus (Owen): Arthaber 1922: p. 18, fig. 7a, b
Criorhynchus simus (Owen): Kuhn 1967: 38
Criorhynchus simus (Owen): Wellnhofer 1978: p. 60, fig. 8, 29
Ornithocheirus simus (Owen): Unwin 2001: p. 194, table 1
Ornithocheirus platyrhinus Seeley: Seeley 1869: p. xvii [disclaimed]
Ornithocheirus platyrhinus Seeley: Seeley 1870: p. 128
Criorhynchus platyrhinus (Seeley): Hooley 1914: p. 536
Criorhynchus simus (Owen): Wellnhofer 1978: p. 60 [synonymy]
Ornithocheirus simus (Owen): Unwin 2001: fig. 7, table 1 [synonymy]
Holotype.

CAMSM B54428, anterior portion of the rostrum (Fig. 1A–D).

Type locality.

Cambridge, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Referred specimens.

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).

Diagnosis.

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.

Description.

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. Owen (1861) described the specimen in detail and noted its very large size, that its first pair of alveoli were directed downwards, and that the anterior margin of the rostrum is tall above the first pair of alveoli. Owen (1861) thus separated it from Pterodactylus sedgwickii [=Camposipterus(?) sedgwickii, see below], in which the first pair of alveoli opens on the anterior margin of the rostrum, facing somewhat forwards, and the anterior margin of the rostrum was not as tall. Owen (1861) also noted that the anterior depression present in the holotype of Ornithocheirus simus was not as marked in another specimen referable to this species. Personal observations of several rostra referable to Ornithocheirus simus (for example, CAMSM B54429, B54552, and B54677, MANCH L.10832, and NHMUK PV 35412) suggest that this depression could have been produced by postmortem abrasion and should be avoided as a character (contra Fastnacht 2001).

Ornithocheirus simus lacks an anterior expansion of the rostrum. As noticed by Owen (1861), there is matrix adhered on the right side of the specimen, which can give the false impression that the palate is broader at the fourth pair of alveoli. Another interesting feature noted by Owen (1861) is the separation between the alveoli of the first pair, equivalent to the largest diameter of the alveolus, and that the bone between these two alveoli projects below from the level of the palate, but not forming an elongated structure. Owen (1861) observed that no median ridge is preserved.

In the original description and illustration, CAMSM B54428 had a tooth preserved in the first left alveolus (Owen 1861: table I, figures 1 to 5). Unwin (2001) suggested that the tooth was possibly glued in this position. During examination of the holotype in 2007, it was observed that the tooth was not preserved with the holotype anymore and could not be found.

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., Wellnhofer 1987, 1991; Fastnacht 2001; Unwin 2001; Veldmeijer 2006), whereas others have suggested that it was actually a short–snouted pterosaur with a tall and massive rostrum (e.g., Hooley 1914; Arthaber 1922; Kuhn 1967; Kellner 1990). References to a reconstruction as a longirostrine pterosaur with a thick premaxillary crest are based on the alleged similarities between Ornithocheirus simus and the more complete holotype of Tropeognathus mesembrinus Wellnhofer, 1987. As the holotype and the several rostra referable to Ornithocheirus simus are fragmentary, it is very difficult to assess which view is correct. Therefore, we refrained from using the presence or absence of a crest in the diagnosis, but several other features (e.g., tall rostrum, position of the first pair of premaxillary teeth) support the distinctiveness of this species among the Cambridge Greensand pterosaur assemblage and in comparison with Tropeognathus mesembrinus (see Taxa from other deposits, below), leading us to propose here that Ornithocheiridae should be restricted to Ornithocheirus simus.

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 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.

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 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.

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 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.

Remarks.

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 Owen (1861) as Pterodactylus simus. Upon recognition that Ornithocheirus simus differed from the species of Pterodactylus, Seeley (1869, 1870) referred it to Ornithocheirus, whereas Owen (1874) subsequently transferred it to Criorhynchus. Both Ornithocheirus and Criorhynchus are based on the same type species (Ornithocheirus simus), and are therefore objective synonyms.

Seeley (1870: 128) named the species Ornithocheirus platyrhinus based on CAMSM B54552, an anterior portion of the rostrum (Fig. 1E–H), with the description: “another fragment, with the area very long, is marked Ornithocheirus platyrhinus”. The area to which Seeley (1870) referred is the tall rostrum. Even this a short characterization makes the name available. CAMSM B54552 is quite incomplete but shows features diagnostic of Ornithocheirus simus: tall rostrum; first pair of premaxillary teeth ventral; first pair of upper dental alveoli slightly placed back from the anterior margin of the premaxilla and ventral profile of the palate straight. Due to the fragmentary nature of this material, it cannot be determined if the anterior expansion of the rostrum was also absent or if the other alveoli have the same sizes and spacing as in CAMSM B54428. Both specimens differ slightly in size, CAMSM B54552 being approximately 7.5 cm high anteriorly and CAMSM B54428 is approximately 6.5 cm high. This difference may be due to ontogenetic or individual variation. Hence, we follow Unwin (2001) in considering Ornithocheirus platyrhinus a subjective junior synonym of Ornithocheirus simus.

Type genus.

Lonchodraco gen. n.

Included genus.

Lonchodraco.

Recorded temporal range.

Albian to Cenomanian / Turonian.

Recorded stratigraphic range.

Cambridge Greensand and Chalk Formation, England.

Diagnosis.

the same as for the type genus.

Etymology.

Derived from the Greek lonchos, meaning lance, and Latin draco, meaning dragon.

Type species.

Pterodactylus giganteus Bowerbank, 1846.

Recorded temporal range.

Albian to Cenomanian / Turonian.

Recorded stratigraphic range.

Cambridge Greensand and Chalk Formation, England.

Diagnosis.

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 Unwin 2001).

Remarks.

Hooley (1914) created the genus Lonchodectes, to which he assigned nine species. Six of them were listed in alphabetical order: Lonchodectes compressirostris, Lonchodectes machaeorhynchus [sic], Lonchodectes microdon, Lonchodectes oweni, Lonchodectes scaphorhynchus and Lonchodectes tenuirostris. Other three were cited latter and referred to the genus, still in the same publication: Lonchodectes daviesii, Lonchodectes giganteus and Lonchodectes sagittirostris (Hooley 1914). There was no designation of a type species.

Latter workers did not accept Lonchodectes as a valid genus. Kuhn (1967) and Wellnhofer (1978) considered it synonymous with Ornithocheirus. Kuhn (1967: 46) designated Lonchodectes compressirostris as the type species of the genus, using the term “Genotypus” (see above). This is a valid subsequent designation under Article 69 of ICZN. Unwin (2001), while reviewing the Cambridge Greensand pterosaurs, re–instated the genus and concluded that, of the nine species referred by Hooley (1914), only five were valid: Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon, Lonchodectes giganteus and Lonchodectes sagittirostris. He also added another species to the genus, Lonchodectes platystomus, which Hooley (1914) had placed in the genus Amblydectes.

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.

Lonchodraco giganteus (Bowerbank, 1846), comb. n.

http://species-id.net/wiki/Lonchodraco_giganteus

Fig. 4
Pterodactylus giganteus Bowerbank: Bowerbank 1846: p. 8, fig. 1, 2, 5.
Pterodactylus giganteus Bowerbank: Bowerbank 1848: pl. I, fig. 1.
Pterodactylus conirostris Owen: Owen in Dixon 1850: p. 401, pl. XXXVIII, fig. 4–7
Pterodactylus giganteus Bowerbank: Bowerbank 1851: p. 19
Cimoliornis diomedaeus [sic] (Gervais): Owen 1851b: p. 21
Pterodactylus giganteus Bowerbank: Owen 1851a: p. 91, pl. XXXI, fig. 1–9, 12–13
Ornithochirus [sic](?) giganteus (Bowerbank): Lydekker 1888: p. 12
Lonchodectes giganteus (Bowerbank): Hooley 1914: p. 538
Ornithodesmus(?) giganteus (Bowerbank): Arthaber 1922: p. 20, fig. 10
Ornithocheirus giganteus (Bowerbank): Wellnhofer 1978: p. 57, fig. 28
Lonchodectes giganteus (Bowerbank): Unwin 2001: p. 210
Lectotype.

NHMUK 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).

Type locality.

Near Maidstone, Burham, Kent, England.

Type horizon.

Chalk Formation (Cenomanian / Turonian).

Diagnosis.

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*.

Description.

Lonchodraco giganteus was briefly described by Bowerbank (1846), and then in more detail by Owen (1851a). The lectotype, NHMUK PV 39412, includes the anterior parts of the rostrum and mandible preserved, and, contra Bowerbank (1846) and subsequent authors (Wellnhofer 1978; Martill 2011), does not include the anterior portion of the nasoantorbital fenestra because what appears to be the anterior margin of the fenestra is not present on both sides of the specimen and most likely represents breakage. The lectotype of Lonchodraco giganteus is readily distinguishable from pterosaurs from other Cretaceous deposits in Britain. Owen (in Dixon 1850) described it as deep–jawed and cone–beaked. The tips of the jaws are dorsoventrally flattened, and there is no upward curvature of the palate. The alveolar margins of the upper and lower jaw are divergent even in their anterior portions. The premaxilla is tall and triangular in lateral view, indicating the presence of a crest. The crest is not thin as seen in Anhanguera or thick as in Coloborhynchus (Fastnacht 2001; Rodrigues and Kellner 2008). The mandibular symphysis also has a distinctive crest from that in anhanguerids because it does not start at the tip of the mandible. The crest is blade–like, short, and located medially in the relatively wide symphysis. Unfortunately, incomplete preparation of the specimen precludes more detailed observations of its oral region, including the palatal ridge. The mandibular groove appears to be deep but cannot be accurately measured. The teeth are conical and elongated, smaller than the ones in anhanguerids; similarly, the alveoli are small and oval to round. Lonchodraco giganteus has a shorter mandibular crest and a larger tooth density than Lonchodraco machaerorhynchus and a tall rostrum as opposed to the elongated premaxillae and maxillae in Lonchodraco(?) microdon and ‘Ornithocheiruspolyodon.

Remarks.

Lonchodraco giganteus has a complex taxonomic history. The species was named Pterodactylus giganteus by Bowerbank (1846). He referred several specimens to the species, including both cranial and postcranial material. Some of these specimens were found associated (NHMUK PV 39412), whereas others were not found associated but came from the same locality as the associated material; additional material was collected at different localities. It is unclear which specimen was considered the holotype. Bowerbank (1848) described the paleohistology of some bones that he referred to Pterodactylus giganteus, including the cranial material (NHMUK PV 39412; Bowerbank 1848: fig. (1). Owen (in Dixon 1850) proposed a new name, Pterodactylus conirostris, for NHMUK PV 39412, because he argued that the specimen was not gigantic in size and thus deemed the specific epithet giganteus inappropriate. Bowerbank (1851) responded that at the time of the description larger pterosaurs were unknown, that modifications of the names of species based on them being inappropriate would cause much instability, and refused to adopt Pterodactylus conirostris. Bowerbank (1851) cited the Law of Priority of the British Association Code (also known as the Strickland Code, published in 1843), which was approved by a committee that included Owen (International Commission on Zoological Nomenclature 1999; Dayrat 2010; see also Martill 2010). The Law of Priority stated that the first name of a species should be the one considered valid. Owen (1851a, b) answered Bowerbank that he had understood that the name Pterodactylus giganteus was proposed for a bone from the Chalk Formation that he (Owen 1842) had previously described as avian. He also pointed out that, among the material described and referred as Pterodactylus giganteus by Bowerbank (1846), there were at least two individuals, of very different size, the smaller one (NHMUK PV 39412) being osteologically mature (based on the fusion between scapula and coracoid), and the other one much larger. Owen (1851a, b) assumed the larger individual to be the one referred as Pterodactylus giganteus and thus designated Pterodactylus conirostris for the cranial material and the bones associated with it. He also brought up several rules of the British Association Code on which he based his designations, including exceptions to the Law of Priority in relation to inappropriate names (Owen 1851a, b; Dayrat 2010), but finally accepted the name Pterodactylus giganteus for the material (Owen 1851a). The name Pterodactylus conirostris has never been used since, but the question as to which material was the holotype of Pterodactylus giganteus remained overlooked for several years. Hooley (1914: 538) reviewed the species based only on the cranial material (NHMUK PV 39412). Finally, Wellnhofer (1978: 57), in his review, designated NHMUK PV 39412 as the lectotype of Pterodactylus giganteus, citing only the skull material and not the associated bones. Pterodactylus giganteus Bowerbank, 1846 and Pterodactylus conirostris Owen, 1850 clearly are objective synonyms because they are founded on the same type specimen, and the former binomen has priority over the latter.

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 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.

Lonchodraco machaerorhynchus (Seeley, 1870), comb. n.

http://species-id.net/wiki/Lonchodraco_machaerorhynchus

Fig. 5
Ptenodactylusmachaerorhynchus Seeley: Seeley 1869: p. xvi [disclaimed]
Ornithocheirus machaerorhynvhus Seeley: Seeley 1870: p. 113, pl. XII, fig. 1–2
Lonchodectes machaeorhynchus [sic] (Seeley): Hooley 1914: p. 535
Lonchodectes machaerorhynchus (Seeley): Unwin 2001: p. 195, fig. 12D–E, table 1
Holotype:

CAMSM B54855, partial mandibular symphysis (Fig. 5A–F).

Type locality.

Cambridge, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Diagnosis.

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.

Description.

CAMSM B54855 consists of a fragment of the posterior portion of the mandibular symphysis. Seeley (1870) described the mandible as narrow, with parallel alveolar margins, deep mandibular sulcus, spacing between alveoli equal to the size of their diameters, and with the lateral margins forming a sharp keel, which Unwin (2001) interpreted as a dentary crest. Seeley (1870) mentioned the presence of a suture with the angular bone, which would almost reach the tip of the mandible, as a unique character for this species in comparison to the pterosaurs from Germany. We here interpret this structure as the posterior margin of a dentary crest rather than a suture.

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 (Seeley 1870), whereas the margins diverge in Lonchodraco giganteus. The mandibular crest of Lonchodraco machaerorhynchus is deep and its apex coincident with its terminus, with a ventral depression posterior to it. In Lonchodraco giganteus, the mandibular crest is short and blade–like, being more restricted to the length of the symphysis; the presence of a depression cannot be determined.

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 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.

Lonchodraco(?) microdon (Seeley, 1870), comb. n. Fig. 6
Ptenodactylusmicrodon Seeley: Seeley 1869: p. xvi [disclaimed]
Ornithocheirus microdon Seeley: Seeley 1870: p. 116, pl. XII, fig. 6–7
Lonchodectes microdon (Seeley): Hooley 1914: p. 535
Ornithocheirus microdon Seeley: Wellnhofer 1978: p. 58
Lonchodectes microdon (Seeley): Unwin 2001: p. 211, table 1
Ornithocheirus oweni Seeley: Seeley 1870: p. 115
Lonchodectes oweni (Seeley): Hooley 1914: p. 535
Lonchodectes microdon (Seeley): Unwin 2001: p. 195, fig. 11C–D, table 1 [synonymy]
Holotype.

CAMSM B54486, anterior portion of the rostrum (Fig. 6A–F).

Type locality.

Cambridge, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Referred specimen.

CAMSM B 54439 (Fig. 6G–L) (also from the Cambridge Greensand).

Diagnosis.

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.

Description.

Lonchodraco(?) microdon was described by Seeley (1870) on the basis of CAMSM B54486, a fragmentary upper jaw with a prominent palatal ridge. Seeley (1870) noted the presence of a concavity on the palate between the elevation of the alveolar margins and the palatal ridge, that the palatal ridge becomes more prominent than the alveolar borders posteriorly, and that the alveolar margins are compressed and rounded. Seeley (1870) also noted a small tip of jaw “associated” with the specimen, but this possibly does not represent the same individual. CAMSM B54486 was recently sampled for histological analysis (M. Riley, pers. comm.). The transverse section of the area that was cut is wider than high, whereas the opposite end is higher than wide, suggesting that the sampled area was located near the anterior end of the snout.

Ornithocheirus oweni was described in the same work (Seeley 1870) on the basis of CAMSM B 54439 (Fig. 6G–L), also an upper jaw. This species is quite similar to Lonchodraco(?) microdon. Both holotypes share a rostrum with parallel alveolar margins, absence of a premaxillary crest, rounded dorsal margin of the premaxillae and maxillae, deep palatal ridge and the concave palate between the alveolar margins and the palatal ridge. Seeley (1870) pointed out that Ornithocheirus oweni differs from Lonchodraco(?) microdon in that its jaw margins are not round but flattened, by the presence of rough interspaces between the alveoli, teeth (alveoli?) circular instead of oval, and the dorsal margin of the premaxillae and maxillae rounded instead of having a sharp keel. However, examination of CAMSM B 54439 established that the alveolar margins are poorly preserved so that the number, shape, and spacing of the alveoli cannot be seen. Furthermore, Lonchodraco(?) microdon has a rounded dorsal margin of the rostrum, as can be observed in anterior and posterior transverse sections of the holotype. As both species share the same morphological features and come from the same deposit they are here considered conspecific.

Remarks.

Unwin (2001) synonymized Ornithocheirus oweni with Lonchodraco(?) microdon and, acting as the First Reviser (ICZN’s article 24.2.2), gave Lonchodraco(?) microdon priority.

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.

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 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.

Anhangueria new clade
Branch–based definition.

All pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.

Recorded temporal range.

Late Berriasian / Valanginian to Cenomanian.

Recorded stratigraphic range.

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.

Synapomorphies.

(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).

Anhangueridae Campos & Kellner, 1985

http://species-id.net/wiki/Anhangueridae

Anhangueridae Campos & Kellner: Campos and Kellner 1985: p. 459
Ornithocheiridae Seeley: Unwin 2001: p. 205
Type genus.

Anhanguera Campos and Kellner, 1985.

Recorded temporal range.

Late Berriasian / Valanginian to Cenomanian.

Recorded stratigraphic range.

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).

Table 11.

Deposits where anhanguerid fossils have been found.

Late Berriasian/Valanginian Hastings Group, England (Coloborhynchus clavirostris; see Owen 1874)
Barremian Wessex Formation, England (Caulkicephalus trimicrodon; see Steel et al. 2005)
Aptian Elrhaz Formation, Niger (Blackburn 2002)
Jiufotang Formation, China (Liaoningopterus gui; see Wang and Zhou 2003)
Aptian/Albian Khuren–Dukh, Dzun–Bayin Formation, Mongolia (Bakhurina and Unwin 1995) (spelled Hüren–Dukh, Züünbayan Formation by Unwin and Bakhurina 2000)
Albian Romualdo Formation, Brazil (Tropeognathus mesembrinus and several species of Anhanguera; e.g., Campos and Kellner 1985)
Paw Paw Formation, USA (Uktenadactylus wadleighi; see Lee 1994)
Cambridge Greensand, England (see Unwin 2001; the present work)
Cenomanian Kem Kem beds, Morocco (Siroccopteryx moroccensis; see Mader and Kellner 1999)
Late Berriasian/Valanginian Hastings Group, England (Coloborhynchus clavirostris; see Owen 1874)
Barremian Wessex Formation, England (Caulkicephalus trimicrodon; see Steel et al. 2005)
Aptian Elrhaz Formation, Niger (Blackburn 2002)
Jiufotang Formation, China (Liaoningopterus gui; see Wang and Zhou 2003)
Aptian/Albian Khuren–Dukh, Dzun–Bayin Formation, Mongolia (Bakhurina and Unwin 1995) (spelled Hüren–Dukh, Züünbayan Formation by Unwin and Bakhurina 2000)
Albian Romualdo Formation, Brazil (Tropeognathus mesembrinus and several species of Anhanguera; e.g., Campos and Kellner 1985)
Paw Paw Formation, USA (Uktenadactylus wadleighi; see Lee 1994)
Cambridge Greensand*, England (see Unwin 2001; the present work)
Cenomanian Kem Kem beds, Morocco (Siroccopteryx moroccensis; see Mader and Kellner 1999)

* this deposit is Cenomanian but the fossils are Albian in age.

* this deposit is Cenomanian but the fossils are Albian in age.

Type species:

Coloborhynchus clavirostris Owen, 1874, by monotypy.

Recorded temporal range.

Late Berriasian / Valanginian.

Recorded stratigraphic range.

Hastings Group, England.

Diagnosis.

As for the type species.

Coloborhynchus clavirostris Owen, 1874

http://species-id.net/wiki/Coloborhynchus_clavirostris

Fig. 7
Coloborhynchus clavirostris Owen: Owen 1874: p. 6, pl. I, fig. 1–4
Criorhynchus simus (Owen): Hooley 1914: p. 537
Criorhynchus clavirostris (Owen): Arthaber 1922: p. 18, fig. 7c
Criorhynchus simus (Owen): Kuhn 1967: 38
Criorhynchus simus (Owen): Wellnhofer 1978: p. 60
Coloborhynchus clavirostris Owen: Lee 1994: p. 756, fig. 4
Coloborhynchus clavirostris Owen: Unwin 2001: p. 206
Coloborhynchus clavirostris Owen: Veldmeijer 2003: 42
Coloborhynchus clavirostris Owen: Rodrigues and Kellner 2008: p. 220, fig. 1.1, 2.1, 3.1
Coloborhynchus clavirostris Owen: Martill, Sweetman and Witton 2011: p. 380, fig. 25.8
Holotype.

NHMUK PV R 1822, anterior portion of the rostrum (Fig. 7A–D).

Type locality.

St.–Leonards–on–Sea, East Sussex, England.

Type horizon.

Hastings Group (late Berriasian / Valanginian).

Diagnosis.

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 Rodrigues and Kellner 2008).

Description.

The holotype of Coloborhynchus clavirostris (NHMUK PV R 1822) is a fragment of the premaxillae and maxillae, and has previously been described in detail (Owen 1874; Veldmeijer 2003; Rodrigues and Kellner 2008). It has a flattened anterior margin, where the first pair of tooth sockets is located. The second, third and fourth pairs of alveoli face laterally, and the fifth and sixth pairs are situated closer to the midline. Coloborhynchus clavirostris also has an anterior expansion and a strong palatal ridge in addition a robust premaxillary crest, which begins at the tip of the rostrum.

Remarks.

Rodrigues and Kellner (2008) reviewed the taxonomic history of the genus Coloborhynchus and the species Coloborhynchus clavirostris, and thus only the main points are repeated here. The genus and species were erected by Owen (1874). Later authors such as Hooley (1914), Kuhn (1967) and Wellnhofer (1978) regarded it synonymous with Ornithocheirus simus [=Criorhynchus simus]. Lee (1994) revalidated Coloborhynchus, and Unwin (2001) not only considered it a valid genus, but also referred additional species to the genus. Rodrigues and Kellner (2008), noting several unique characters of Coloborhynchus clavirostris, restricted the genus to the type species. This view is followed here.

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 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.

Ornithocheirus capito Seeley, 1870 Fig. 8
Ptenodactyluscapito Seeley: Seeley 1869: p. xvi [disclaimed]
Ornithocheirus capito Seeley: Seeley 1870: p. 126
Criorhynchus capito (Seeley): Hooley 1914: p. 536
Criorhynchidae incertae sedis: Wellnhofer 1978: 60
Coloborhynchus capito (Seeley): Unwin 2001: p. 206, table 1
Ornithocheiruscapito Seeley: Rodrigues and Kellner 2008: p. 226, fig. 4.1, 4.3
Ornithocheirus reedi Seeley: Seeley 1870: p. 126
Ornithocheirus reedii [sic] Seeley: Seeley 1881: p. 13, pl. I, fig. 3
Criorhynchus reedi (Seeley): Hooley 1914: p. 538
Criorhynchus reedii [sic] (Seeley): Arthaber 1922: p. 18
Criorhynchidae incertae sedis Wellnhofer 1978: 60
Coloborhynchus capito (Seeley): Unwin 2001: fig. 6C, 8, table 1 [synonymy]
Holotype.

CAMSM B 54625, anterior portion of the rostrum (Fig. 8A–F).

Type locality.

Chesterton, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Referred specimen.

Holotype of Ornithocheirus reedi Seeley, 1870 (Fig. 8G–J) (from the Cambridge Greensand; current whereabouts unkown).

Diagnosis.

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.

Description.

Ornithocheiruscapito 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 Seeley (1881) affirmed that the bone surface is rather smooth, and abrasion is unlikely. Furthermore, a groove in the midline of the crest is present in the holotype of Uktenadactylus wadleighi (SMU 73058) from the Albian Paw Paw Formation of the USA (Lee 1994; Rodrigues and Kellner 2008), which also has a concave anterior margin of the crest. However, ‘Ornithocheiruscapito differs from SMU 73058 in the height of the crest, which is much higher just behind the second pair of alveoli.

Ornithocheiruscapito 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. Rodrigues and Kellner (2008) listed four autapomorphies for Coloborhynchus clavirostris, none of which is present in ‘Ornithocheiruscapito. It also can be distinguished from Uktenadactylus wadleighi in the absence of an oval depression above the first pair of teeth (Lee 1994; Rodrigues and Kellner 2008).

Ornithocheiruscapito 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 ‘Ornithocheiruscapito 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.

Remarks.

Unwin (2001) synonymized ‘Ornithocheirusreedi with ‘Ornithocheiruscapito (as Coloborhynchus capito). ‘Ornithocheirusreedi, described by Seeley (1870), was referred by Hooley (1914) and provisionally by Wellnhofer (1978) to Criorhynchus reedi, and its name was misspelled reedii by Seeley (1881) and readi by Barrett et al. (2008). Its holotype, illustrated by Seeley (1881: pl. I, figs 3a–3b; here as Fig. 8E–H), belonged to the collection of W. Reed of York by the time it was described and could not be found in the collections of the Natural History Museum, the Sedgwick Museum of Earth Sciences or the Manchester Museum during visits in October 2009. Therefore, we base our remarks on the descriptions and illustrations provided by Seeley (1870, 1881).

Seeley (1870) first described Ornithocheirus reedi and interpreted the holotype as an upper jaw. Later Seeley (1881) redescribed the specimen as a lower jaw, based on the presence of a median groove. Curiously, in the same work, it was illustrated oriented as being a part of a snout. Unfortunately, the specimen was not figured in palatal view, but it is very tall and we here interpret it as the tip of the premaxillae. It bears a median crest with a markedly concave anterior margin. As ‘Ornithocheiruscapito and Ornithocheirus reedi share this feature (unknown in other Cambridge Greensand species) and come from the same deposit, we agree with Unwin (2001) that they represent the same taxon. Both were described in the same work (Seeley 1870). Acting as First Reviewer, Unwin (2001) gave priority to the former binomen.

Here, we recognize that ‘Ornithocheiruscapito 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 ‘Ornithocheiruscapito.

Figure 8.

Ornithocheiruscapito. 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 8.

Ornithocheiruscapito. 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.

Anhangueria incertae sedis
Etymology.

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.

Type species.

Ornithocheirus nasutus Seeley, 1870.

Recorded temporal range.

Albian.

Recorded stratigraphic range.

Cambridge Greensand, England.

Diagnosis.

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.

Camposipterus nasutus (Seeley, 1870), comb. n.

http://species-id.net/wiki/Camposipterus_nasutus

Fig. 9
Ptenodactylusnasutus Seeley: Seeley 1869: p. xvi [disclaimed]
Ornithocheirus nasutus Seeley: Seeley 1870: p. 120
Ornithocheirus nasutus Seeley: Hooley 1914: p. 535
Anhanguera fittoni (Owen): Unwin 2001: fig. 10C–E, table 1 [synonymy]
Holotype.

CAMSM B 54556, anterior portion of the rostrum (Fig. 9A–D).

Type locality.

Haslingfield, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Diagnosis.

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.

Description.

Camposipterus nasutus was originally described by Seeley (1870) as Ornithocheirus nasutus. Seeley noted that it has an expansion at the tip of the rostrum, a palatal ridge extending posteriorly to the level of the second pair of alveoli, the first pair of alveoli facing forward, and a dorsoventrally compression of the tip of the rostrum. It differs from Cimoliopterus cuvieri, which possesses a premaxillary crest but no anterior expansion of the rostrum, and which is higher than wide in anterior view, in contrast with the wider than high tip of the rostrum of Camposipterus nasutus.

Remarks.

Unwin (2001) synonymized the species with Anhanguera fittoni [=Pterodactylus fittoni, here considered a nomen dubium, see below]. We do not agree with this view because the holotype of Camposipterus nasutus is dorsoventrally flattened and has an anterior expansion. By contrast, Pterodactylus fittoni is known from a fragmentary rostrum that, although incomplete anteriorly, does not share these features. It can also be excluded from Anhanguera because it does not have a premaxillary crest; furthermore, no species definitely referable to Anhanguera has a dorsoventrally flattened rostrum. It can be expected that the description of new, more complete specimens from the Romualdo Formation of the Santana Group, currently under work by several researchers, will help shed light in its relationships with taxa such as Brasileodactylus araripensis Kellner, 1984 (see Kellner 1984), but so far the dorsoventrally flattened anterior end of the rostrum seems to be diagnostic for Camposipterus nasutus.

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 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.

Camposipterus(?) sedgwickii (Owen, 1859), comb. n. Fig. 10A–D
Pterodactylus sedgwickii Owen: Owen 1859: p. 2, pl. I, fig. 1
Ptenodactylussedgwicki [sic] (Owen): Seeley 1869: p. xvi [disclaimed]
Ornithocheirus sedgwicki [sic] (Owen): Seeley 1870: p. 112
Coloborhynchus sedgwickii (Owen): Owen 1874: p. 6
Ornithochirus [sic] sedgwicki [sic] (Owen): Lydekker 1888: p. 15
Ornithocheirus sedgwicki [sic] (Owen): Hooley 1914: p. 535
Ornithocheirus sedgwicki [sic] (Owen): Arthaber 1922: p. 17
Ornithocheirus sedgwicki [sic] (Owen): Wellnhofer 1978: p. 58, fig. 28
Coloborhynchus sedgwickii (Owen): Unwin 2001: p. 194, fig. 9, table 1
Ornithocheirussedgwickii (Owen): Rodrigues and Kellner 2008: p. 226, fig. 4.2, 4.4
Holotype.

CAMSM B54422, anterior part of the rostrum (Fig. 10A–D).

Type locality.

Cambridge, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Diagnosis.

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*.

Description.

Owen (1859) described Pterodactylus sedgwickii based on an upper jaw, CAMSM B54422. He pointed out the presence of the first pair of alveoli at the anterior margin of the rostrum, facing forward; an anterior expansion of the rostrum, where the large second and third pairs of alveoli are located; alveoli four to six approximately same size; spacing between alveoli smaller than their diameter; and obtuse tip of the rostrum.

The third alveoli have different sizes on the left and right sides; this could be explained on taphonomic grounds (e.g., Kellner 2010). On the left side, the bone surrounding the alveolus seems to have been forced outwards, perhaps by phosphate deposition (Seeley 1870). On the other hand, the absence of the fifth alveolus on the left side, as noted by Owen (1859), is a possibly pathological feature seen in other pterosaurs, such as in the holotype of Anhanguera robustus (BSP 1987 I 47) and in a specimen referred to Anhanguera blittersdorffi (Pz–DBAV–UERJ 40).

It is noteworthy that the drawing of CAMSM B54422 was reversed in Owen (1859) and in Wellnhofer (1978) and Unwin (2001), which present illustrations based on Owen’s. Also, there is a breakage and a discontinuity between the anterior, round part of the rostrum and the more posteriorly located dorsal margin, which is not figured by the aforementioned authors.

Owen (1859) referred a lower jaw, CAMSM B54421, to the same species, partially based on a similar alveolar density as in the holotype. CAMSM B54421 is very fragmentary and does not have the tip of the mandible preserved. Furthermore, these two specimens do not fit together, with the upper jaw much broader than the lower one. There is no evidence to support placement of CAMSM B54421 in Camposipterus(?) sedgwickii, and we here restrict the species to its type material.

Remarks.

Hooley (1914) and Wellnhofer (1978) placed Camposipterus(?) sedgwickii in Ornithocheirus. It differs from this genus in the absence of a tall rostrum, the first pair of teeth facing ventrally, and other characters (see above). Owen (1874) and Unwin (2001) referred it to Coloborhynchus. Rodrigues and Kellner (2008) excluded Camposipterus(?) sedgwickii from Coloborhynchus, as it differs from Coloborhynchus clavirostris in that the anterior end of the rostrum is round rather than flat and the spacing between the first and second pairs of alveoli is larger in Coloborhynchus clavirostris. Camposipterus(?) sedgwickii shows none of the autapomorphies listed for Coloborhynchus clavirostris. Therefore, we tentatively place this species in the genus Camposipterus. Camposipterus nasutus and Camposipterus(?) sedgwickii share the presence of an expansion of the rostrum, absence of a premaxillary crest, and round rostrum in lateral view. They differ in the different height of the tip of the rostrum, which is taller than wide in Camposipterus(?) sedgwickii, the extension of the palatal ridge, and the size of the expansion, which is larger in Camposipterus(?) sedgwickii than in Camposipterus nasutus.

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 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.

Camposipterus(?) colorhinus (Seeley, 1870), comb. n. Fig. 10E–H
Ptenodactyluscolorhinus Seeley: Seeley 1869: p. xvi [disclaimed]
Ornithocheirus colorhinus Seeley: Seeley 1870: p. 124
Ornithocheirus colorhinus Seeley: Hooley 1914: p. 535
Anhanguera cuvieri (Bowerbank): Unwin 2001: fig. 10A, B, table 1
Syntypes.

CAMSM B54431 (Fig. 10E–H) and CAMSM B54432, anterior parts of the rostrum.

Type locality.

Cambridge, Cambridgeshire, England.

Type horizon.

Cambridge Greensand (Cenomanian; fossils Albian in age).

Diagnosis.

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.

Description.

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. Seeley (1870) pointed out the presence of a slightly convex median part of the palate (seen in CAMSM B54431), an anterior expansion of the rostrum, and the large, round alveoli, the first of which faces anteroventrally. Seeley (1870) interpreted the presence of a lunate area as attachment for a lip. We verified this observation but question his interpretation, as the tips of the jaws of pterosaurs were covered by a horny beak (Seeley 1901: fig. 20; Kellner and Tomida 2000; Frey et al. 2003; Pinheiro et al. 2011).

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 (Lee 1994; Rodrigues and Kellner 2008), whereas in Camposipterus(?) colorhinus this edge faces anteroventrally and, consequently, the orientation of the depression is also different. Furthermore, the first pair of teeth in Uktenadactylus is located higher and the anterior expansion is more squarish and not round as in the species under discussion.

Remarks.

Unwin (2001) referred to CAMSM B54431 as “a large individual of Anhanguera cuvieri” [=Cimoliopterus cuvieri]. Camposipterus(?) colorhinus can be excluded from Cimoliopterus cuvieri, which lacks an anterior expansion of the rostrum or a depression above the first pair of alveoli.

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.

Pteranodontoidea incertae sedis
Etymology.

From the Greek kimolia, chalk, and pteron, wing.

Type species.

Pterodactylus cuvieri Bowerbank, 1851.

Included species.

Cimoliopterus cuvieri.

Recorded temporal range.

Cenomanian / Turonian.

Recorded stratigraphic range.

Chalk Formation, England.

Diagnosis.

As for the type–species.

Cimoliopterus cuvieri (Bowerbank, 1851), comb. n.

http://species-id.net/wiki/Cimoliopterus_cuvieri

Fig. 11
Pterodactylus cuvieri Bowerbank: Bowerbank 1851: p. 15, pl.IV
Pterodactylus cuvieri Bowerbank: Owen 1851b: p. 29
Pterodactylus cuvieri Bowerbank: Owen 1851a: p. 88, pl. XXVIII, fig. 1–7
Ptenodactyluscuvieri (Bowerbank): Seeley 1869: p. xvi [disclaimed]
Ornithocheirus cuvieri (Bowerbank): Seeley 1870: p. 113
Coloborhynchus cuvieri (Bowerbank): Owen 1874: p. 6