Novialoidea

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Novialoidea
Temporal range:
Early Jurassic - Late Cretaceous, 182–66  Ma
Campylognathoides-Ghedo.JPG
Fossil specimen of Campylognathoides liasicus , a basal novialoid.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Clade: Macronychoptera
Clade: Novialoidea
Kellner, 2003
Subgroups
Synonyms
  • Lonchognatha? Unwin, 2003

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]

History and classification

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 

Campylognathoides liasicus

Campylognathoides zitteli

Breviquartossa
  Rhamphorhynchidae  

Scaphognathus crassirostris

Rhamphorhynchinae

  Pterodactylomorpha  

Sordes pilosus

Monofenestrata
Darwinoptera

Pterorhynchus wellnhoferi

Wukongopteridae

Pterodactyliformes

Changchengopterus pani

Caelidracones

Related Research Articles

<i>Eudimorphodon</i> Genus of eudimorphodontid pterosaur from the Late Triassic

Eudimorphodon was a pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic, making Eudimorphodon one of the oldest pterosaurs known. It had a wingspan of about 100 centimeters (3.3 ft) and at the end of its long bony tail may have been a diamond-shaped flap like in the later Rhamphorhynchus. If so, the flap may have helped it steer while maneuvering in the air. Eudimorphodon is known from several skeletons, including juvenile specimens.

<span class="mw-page-title-main">Phylogeny of pterosaurs</span>

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.

<i>Dorygnathus</i> Genus of rhamphorhynchid pterosaur from the Early Jurassic

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, 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. Having two or more morphs of teeth, a condition called heterodonty, is rare in modern reptiles but more common in basal ("primitive") pterosaurs. The heterodont dentition in Dorygnathus is consistent with a piscivorous (fish-eating) diet. The fifth digit on the hindlimbs of Dorygnathus was unusually long and oriented to the side. Its function is not certain, but the toe may have supported a membrane like those supported by its wing-fingers and pteroids. Dorygnathus was according to David Unwin related to the Late Jurassic pterosaur Rhamphorhynchus and was a contemporary of Campylognathoides in Holzmaden and Ohmden.

<i>Campylognathoides</i> Genus of campylognathoidid pterosaur from the Early Jurassic

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.

<span class="mw-page-title-main">Rhamphorhynchoidea</span>

The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhynchoidea as opposed to a more distant common ancestor. Because it is not a completely natural grouping, Rhamphorhynchoidea is not used as a formal group in most scientific literature, though some pterosaur scientists continue to use it as an informal grouping in popular works, such as The Pterosaurs: From Deep Time by David Unwin, and in some formal studies. Rhamphorhynchoids were the first pterosaurs to have appeared, in the late Triassic Period. Unlike their descendants, the pterodactyloids, most rhamphorhynchoids had teeth and long tails, and most species lacked a bony crest, though several are known to have crests formed from soft tissue like keratin. They were generally small, with wingspans rarely exceeding 2.5 meters, though one specimen alluded to by Alexander Stoyanow would be among the largest pterosaurs of all time with a wingspan of 10 meters, comparable to the largest azhdarchids. However, this alleged giant Jurassic pterosaur specimen is not recorded anywhere outside the original Time article. Nearly all rhamphorhynchoids had become extinct by the end of the Jurassic Period, though some anurognathids persisted to the early Cretaceous. The family Wukongopteridae, which shows a mix of rhamphorhynchoid and pterodactyloid features, is known from the Daohugou Beds which are most commonly dated to the Jurassic, but a few studies give a Cretaceous date. Furthermore, remains of a non-pterodactyloid from the Candeleros Formation extend the presence of basal pterosaurs into at least the early Late Cretaceous.

<span class="mw-page-title-main">Pterodactyloidea</span> Suborder of monofenestratan pterosaurs

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.

<span class="mw-page-title-main">Lonchodectidae</span> Family of pteranodontoid pterosaurs

Lonchodectidae or Lonchodraconidae is a group of pterosaurs within the clade Pterodactyloidea. It has variously been considered to be within Ctenochasmatoidea, Azhdarchoidea and Pteranodontia. They are notable for their high, conical tooth sockets and raised alveolar margins.

<span class="mw-page-title-main">Ornithocheiroidea</span> Clade of pterodactyloid pterosaurs

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.

<span class="mw-page-title-main">Ctenochasmatoidea</span> Superfamily of archaeopterodactyloid pterosaurs

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.

<span class="mw-page-title-main">Dsungaripteroidea</span> Superfamily of reptiles (fossil)

Dsungaripteroidea is a group of pterosaurs within the suborder Pterodactyloidea.

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of ornithocheiroid pterosaurs

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. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.

<span class="mw-page-title-main">Dimorphodontidae</span> Family of basal macronychopteran pterosaurs

Dimorphodontidae is a group of early "rhamphorhynchoid" pterosaurs named after Dimorphodon, that lived in the Late Triassic to Early Jurassic. While fossils that can be definitively referred to the group are rare, dimorphodontids may have had a broad distribution, with fossils known from the UK, the southwest United States, and possibly Antarctica.

<span class="mw-page-title-main">Dsungaripteridae</span> Family of ornithocheiroid pterosaurs

Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings.

<span class="mw-page-title-main">Archaeopterodactyloidea</span> Infraorder of pterodactyloid pterosaurs

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.

<span class="mw-page-title-main">Ornithocheirae</span> Clade of pteranodontoid pterosaurs

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.

<span class="mw-page-title-main">Eudimorphodontidae</span> Family of eopterosaurs from the Late Triassic

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 Greenland. Its only species was previously attributed to Eudimorphodon, and its closest relatives may have been Eudimorphodon or Austriadraco.

<span class="mw-page-title-main">Tapejaroidea</span> Clade of ornithocheiroid pterosaurs

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.

<i>Seazzadactylus</i> Genus of austriadraconid pterosaur from the Late Triassic

Seazzadactylus is a basal pterosaur genus that during the late Triassic lived in the area of present Italy.

References

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  2. Codorniú, Laura; Paulina Carabajal, Ariana; Pol, Diego; Unwin, David; Rauhut, Oliver W.M. (2016). "A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium". PeerJ. 4: e2311. doi: 10.7717/peerj.2311 . PMC   5012331 . PMID   27635315.
  3. Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. Zitteliana, 61-107.
  4. Richard J. Butler; Stephen L. Brusatte; Brian B. Andres; Roger B. J. Benson (2012). "How do geological sampling biases affect studies of morphological evolution in deep time? A case study of the Pterosauria (Reptilia: Archosauria)". Evolution. 66 (1): 147–162. doi:10.1111/j.1558-5646.2011.01415.x. PMID   22220871. S2CID   205783384.
  5. Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
  6. Unwin, D. M., (2003): On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
  7. Brian Andres, James M. Clark & Xu Xing (2010) A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs, Journal of Vertebrate Paleontology, 30:1, 163-187, DOI: 10.1080/02724630903409220
  8. Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  9. Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology . 24 (9): 1011–6. doi: 10.1016/j.cub.2014.03.030 . PMID   24768054.