Novialoidea | |
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Fossil specimen of Campylognathoides liasicus , a basal novialoid. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Clade: | † Macronychoptera |
Clade: | † Novialoidea Kellner, 2003 |
Subgroups | |
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Synonyms | |
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Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian age [3] ), their fossils having been found on all continents except Antarctica. [4]
Novialoidea was named by paleontologist Alexander Wilhelm Armin Kellner in 2003 as a node-based taxon consisting of the last common ancestor of Campylognathoides , Quetzalcoatlus and all its descendants. This name was derived from Latin novus "new", and ala, "wing", in reference to the wing synapomorphies that the members of the clade possess. [5]
Paleontologist David Unwin in 2003 had named the group Lonchognatha in the same issue of the journal that published Novialoidea (Geological Society of London, Special Publications 217) and defined it as Eudimorphodon ranzii , Rhamphorhynchus muensteri , their most recent common ancestor and all its descendants (as a node-based taxon). [6] Under Unwin's and Kellner's phylogenetic analyses (where Eudimorphodon and Campylognathoides form a family that is basal to both Rhamphorhynchus and Quetzalcoatlus), and because Novialoidea was named first (in pages 105–137, while Lonchognatha was named in pages 139–190), Lonchognatha is an objective junior synonym of the former. However, other analyses find Lonchognatha to be valid (Andres et al., 2010), [7] or synonymous with the Pterosauria (Andres, 2010). [8]
Below is a cladogram showing the phylogenetic analysis conducted by Brian Andres and colleagues in 2014. Based on the analysis, Novialoidea contains the genus Campylognathoides as well as the group Breviquartossa. [9]
Novialoidea | |
This phylogeny of pterosaurs entails the various phylogenetic trees used to classify pterosaurs throughout the years and varying views of these animals. Pterosaur phylogeny is currently highly contested and several hypotheses are presented below.
Dorygnathus was a genus of rhamphorhynchid pterosaur that lived in Europe during the Early Jurassic period, when shallow seas flooded much of the continent. It had a short wingspan around 1.5 meters, and a relatively small triangular sternum, which is where its flight muscles attached. Its skull was long and its eye sockets were the largest opening therein. Large curved fangs that "intermeshed" when the jaws closed featured prominently at the front of the snout while smaller, straighter teeth lined the back.
Germanodactylus is a genus of germanodactylid pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen Limestone. Its specimens were long thought to pertain to Pterodactylus. The head crest of Germanodactylus is a distinctive feature.
Campylognathoides is an extinct genus of pterosaur discovered in the Württemberg Lias deposits of Germany; this first specimen however, consisted only of wing fragments. Further better preserved specimens were found in the Holzmaden shale; based on these specimens, Felix Plieninger erected a new genus.
Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.
Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii. A second species, E. yixianensis, was named in 2023, but its validity has been questioned.
Lonchodectes was a genus of lonchodectid pterosaur from several formations dating to the Turonian of England, mostly in the area around Kent. The species belonging to it had been assigned to Ornithocheirus until David Unwin's work of the 1990s and 2000s. Several potential species are known; most are based on scrappy remains, and have gone through several other generic assignments. The genus is part of the complex taxonomy issues surrounding Early Cretaceous pterosaurs from Brazil and England, such as Amblydectes, Anhanguera, Coloborhynchus, and Ornithocheirus.
Lonchodectidae or Lonchodraconidae is a group of pterosaurs within the clade Pterodactyloidea. It has variously been considered to be within Ctenochasmatoidea, Azhdarchoidea and Pteranodontoidea. They are notable for their high, conical tooth sockets and raised alveolar margins.
Ornithocheiroidea is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods, with fossil remains found all over the world except Antarctica.
Ctenochasmatoidea is a group of early pterosaurs within the suborder Pterodactyloidea. Their remains are usually found in what were once coastal or lake environments. They generally had long wings, long necks, and highly specialized teeth.
Dsungaripteroidea is a group of pterosaurs within the suborder Pterodactyloidea.
Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. Other notable pterosaur families belonging to the Azhdarchoidea are the Tapejaridae, the Thalassodromidae, and the Chaoyangopteridae. This group contains some of the most advanced members of the pterosaur order.
Rhamphorhynchidae is a group of early pterosaurs named after Rhamphorhynchus, that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley. Members of the group possess no more than 11 pairs of teeth in the rostrum, a deltopectoral crest that is constricted at the base but expanded at the distal end, and a bent phalange on the fifth toe.
Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings, and were commonly interpreted as durophagous and possibly piscivorous pterosaurs. Fossils have been discovered from Early Cretaceous deposits in Asia, South America and possibly Europe.
Eupterodactyloidea is an extinct group of pterodactyloid pterosaurs that existed from the latest Late Jurassic to the latest Late Cretaceous periods. Eupterodactyloids have been found on all continents except Antarctica.
Archaeopterodactyloidea is an extinct clade of pterodactyloid pterosaurs that lived from the middle Late Jurassic to the latest Early Cretaceous periods of Africa, Asia, Europe and North America. It was named by Alexander Wilhelm Armin Kellner in 1996 as the group that contains Germanodactylus, Pterodactylus, the Ctenochasmatidae and the Gallodactylidae. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Pterodactylus, Ctenochasma and Gallodactylus and all its descendants. Although phylogenetic analyses that based on David Unwin's 2003 analysis do not recover monophyletic Archaeopterodactyloidea, phylogenetic analyses that based on Kellner's analyses, or the analyses of Brian Andres recover monophyletic Archaeopterodactyloidea at the base of the Pterodactyloidea.
Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family that contains Ornithocheirus and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an -idae suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants. Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner and Andres that these families should not be synonymized based on their original phylogenetical definitions. However, subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria and Targaryendraconia.
Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic of Europe. It was named by Peter Wellnhofer in 1978 to include Eudimorphodon ranzii. Some phylogenetic analyses suggested that Eudimorphodontidae is a junior synonym of Campylognathoididae, however more comprehensive analyses found Eudimorphodontidae to be basal to Macronychoptera that includes Campylognathoididae and more derived pterosaurs (Breviquartossa). Wang et al. (2009) found Eudimorphodontidae to include six species, but they didn't defined the clade. Brian Andres define Eudimorphodontidae and found Peteinosaurus to be most closely related to it. Furthermore, he found monophyletic Eudimorphodon clade, and defined two subfamilies within Eudimorphodontidae. The Eudimorphodontinae includes all taxa more closely related to Eudimorphodon ranzii than to Raeticodactylus filisurensis while the Raeticodactylinae includes all taxa more closely related to Raeticodactylus filisurensis than to Eudimorphodon ranzii. More recently, Raeticodactylus and Caviramus were moved into their own family, Raeticodactylidae. The below cladogram follows that analysis.
Arcticodactylus is a genus of basal pterosaur living during the Late Triassic in the area of present-day Greenland. Its only species was previously attributed to Eudimorphodon, and its closest relatives may have been Eudimorphodon or Austriadraco.
Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.