Phylogeny of pterosaurs

Highly simplified pterosaur phylogeny – topology based on that recovered by Unwin[1]

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.

Unwin (2003)

The matrix includes 19 pterosaur groups (most of which are supra-specific) plus a single outgroup (Euparkeria capensis). The taxa were coded for 60 characters.

Pterosauria 

Preondactylus

 Macronychoptera 

Dimorphodontidae

 Caelidracones 

Anurognathidae

 Lonchognatha 

Campylognathoididae

 Breviquartossa 
 Rhamphorhynchidae 

Rhamphorhynchinae

Scaphognathinae

 Pterodactyloidea 
 Ornithocheiroidea 

Istiodactylus

 Euornithocheira 

Ornithocheiridae

 Pteranodontidae 

Nyctosaurus

Pteranodontinae

 Lophocratia 
 Ctenochasmatoidea 

Gallodactylidae

 Euctenochasmatia 

Pterodactylus

Lonchodectes

Ctenochasmatidae

 Dsungaripteroidea 

Germanodactylus

Dsungaripteridae

 Azhdarchoidea 

Tapejara

Neoazhdarchia

Tupuxuara

Azhdarchidae

Kellner (2003)

The matrix includes 39 valid pterosaur species, although Rhamphorhynchus longicaudus and Nyctosaurus bonneri are usually considered to be synonymous with R. muensteri and N. gracilis respectively, plus a three outgroup species (Ornithosuchus longidens, Herrerasaurus ischigualastensis and Scleromochlus taylori). The taxa were coded for 74 characters.

Pterosauria 
 Anurognathidae 

Anurognathus ammoni

 Asiaticognathidae 

Batrachognathus volans

Dendrorhynchoides curvidentatus

Sordes pilosus

Scaphognathus crassirostris

Preondactylus buffarinii

Dorygnathus banthensis

Dimorphodon macronyx

Peteinosaurus zambellii

"Eudimorphodon" rosenfeldi

 Novialoidea 
 Campylognathoididae 

Campylognathoides liasicus

Eudimorphodon ranzii

 Rhamphorhynchidae 

Rhamphorhynchus longicaudus

Rhamphorhynchus muensteri

 Pterodactyloidea 
 Archaeopterodactyloidea 

Pterodactylus antiquus

Pterodactylus kochi

 Germanodactylidae 

Germanodactylus cristatus

Germanodactylus rhamphastinus

 Ctenochasmatidae 

Ctenochasma

Pterodaustro guinazui

 Gallodactylidae 

Cycnorhamphus suevicus

Gallodactylus canjuersensis

 Dsungaripteroidea 

Nyctosaurus bonneri

Nyctosaurus gracilis

 Ornithocheiroidea 
 Pteranodontoidea 

Pteranodon longiceps

Istiodactylus latidens

Coloborhynchus clavirostris

 Anhangueridae 

Tropeognathus mesembrinus

Anhanguera blittersdorffi

Anhanguera piscator

Anhanguera santanae

 Tapejaroidea 
 Dsungaripteridae 

Dsungaripterus weii

Noripterus complicidens

"Phobetor" parvus

 Azhdarchoidea 
 Tapejaridae 

Tupuxuara leonardii

Tapejara wellnhoferi

Tapejara imperator

 Azhdarchidae 

Azhdarcho lancicollis

Quetzalcoatlus sp

Andres and Myers (2013)

In 2010, Brian Blake Andres wrote a review of pterosaur phylogeny in his dissertation. His phylogenetic analysis combined data mainly from three different matrixes: Kellner's original analysis (2003) and its updates (Kellner (2004), Wang et al. (2005) and Wang et al. (2009)), Unwin's original analysis (2003) and its updates (Unwin (2002), Unwin (2004), Lu et al. (2008) and Lu et al. (2009)) and previous analyses by Andres et al. (2005), Andres and Ji (2008) and Andres et al. (2010). Additional characters are taken from DallaVecchia (2009), Bennett' analyses (1993–1994) and various older, non-phylogenetic, papers.[2]

The matrix includes 100 valid pterosaur species plus a single outgroup (Euparkeria capensis). This represents 70.4% of 142 known pterosaur species back then. These were scored for 183 morphological characters (compared to 3 outgroups plus 57 ingroups which were scored for 89 characters of Wang et al. 2009 [the latest version of Kellner's analysis] and to 1 outgroups plus 59 ingroups which were scored for 117 characters of Lu et al. 2012 [the latest version of Unwin's analysis]).[3] The resultant topology is well supported and more resolved than previous analyses. Furthermore, it codes only species as terminal taxa, (unlike some analyses, e.g., Unwin (2003) who used mainly families) and uses the holotype specimens for the codings (unlike some analyses, e.g., Kellner (2003)).[2] This phylogenetic analysis was used by Richard J. Butler, Stephen L. Brusatte, Brian B. Andres and Roger B. J. Benson (2012) to assess the morphological diversity and fossil sampling biases of the Pterosauria.[3] A paper focusing on the pterosaur phylogeny (Andres) was published in a book named "The Pterosauria".[4] An updated and more resolved version of this phylogeny was published formally by Andres and Timothy Myers in 2013, containing 185 characters and 109 ingroup taxa. Below is a cladogram showing these results after the exclusion of three taxa that can be coded only for one character (clade names follow Andres & Myers, 2013).[5]

Pterosauria 

Preondactylus buffarinii

Austriadactylus cristatus

Peteinosaurus zambellii

 Eudimorphodontidae 
 Raeticodactylinae 

Caviramus schesaplanensis

Raeticodactylus filisurensis

Eudimorphodon cromptonellus

Eudimorphodon ranzii

Eudimorphodon rosenfeldi

Dimorphodon macronyx

Parapsicephalus purdoni

Campylognathoides liasicus

Campylognathoides zitteli

 Rhamphorhynchidae 

Scaphognathus crassirostris

 Rhamphorhynchinae 

Dorygnathus banthensis

Cacibupteryx caribensis

Nesodactylus hesperius

Rhamphorhynchus muensteri

Harpactognathus gentryii

Angustinaripterus longicephalus

Sericipterus wucaiwanensis

Sordes pilosus

 Wukongopteridae 

Pterorhynchus wellnhoferi

Darwinopterus modularis

Wukongopterus lii

Changchengopterus pani

 Anurognathidae 

Dendrorhynchoides curvidentatus

Jeholopterus ninchengensis

Anurognathus ammoni

Batrachognathus volans

Archaeopterodactyloidea

Normannognathus wellnhoferi

Germanodactylus cristatus

Germanodactylus rhamphastinus

Pterodactylus antiquus

Pterodactylus kochi

Ardeadactylus longicollum

 Gallodactylidae 

Boreopterus cuiae

Feilongus youngi

Cycnorhamphus suevicus

Gallodactylus canjuersensis

 Ctenochasmatidae 
 Gnathosaurinae 

Kepodactylus insperatus

Elanodactylus prolatus

Huanhepterus quingyangensis

Plataleorhynchus streptophorodon

Gnathosaurus subulatus

Gnathosaurus macrurus

 Ctenochasmatinae 

Ctenochasma elegans

Ctenochasma roemeri

Pterodaustro guinazui

Eosipterus yangi

Beipiaopterus chenianus

Gegepterus changi

Haopterus gracilis

Ornithocheiroidea
 Nyctosauridae 

Muzquizopteryx coahuilensis

"Nyctosaurus" lamegoi

Nyctosaurus gracilis

Alamodactylus byrdi

Pteranodon longiceps

Pteranodon sternbergi

 Istiodactylidae 

Longchengpterus zhaoi

Nurhachius ignaciobritoi

Liaoxipterus brachyognathus

Istiodactylus latidens

Istiodactylus sinensis

Lonchodectes compressirostris

Aetodactylus halli

Cearadactylus atrox

Brasileodactylus araripensis

Ludodactylus sibbicki

 Anhangueridae 

Liaoningopterus gui

Anhanguera araripensis

Anhanguera blittersdorffi

Anhanguera piscator

Anhanguera santanae

 Ornithocheiridae 

Tropeognathus mesembrinus

Ornithocheirus simus

Coloborhynchus clavirostris

Coloborhynchus wadleighi

Bennettazhia oregonensis

Nemicolopterus crypticus

"Sinopterus" gui

 Tapejaridae 

Huaxiapterus jii

Eopteranodon lii

Sinopterus dongi

"Huaxiapterus" corollatus

"Huaxiapterus" benxiensis

Bakonydraco galaczi

Tapejara wellnhoferi

Tupandactylus navigans

Tupandactylus imperator

 Chaoyangopteridae 

Eoazhdarcho liaoxiensis

Shenzhoupterus chaoyangensis

Chaoyangopterus zhangi

Jidapterus edentus

 Azhdarchidae 

Radiodactylus langstoni

 Azhdarchinae 

Azhdarcho lancicollis

TMM 42489

Zhejiangopterus linhaiensis

Arambourgiania philadelphiae

Quetzalcoatlus northropi

Quetzalcoatlus sp.

 Thalassodrominae 

Thalassodromeus sethi

Tupuxuara leonardii

Tupuxuara longicristatus

 Dsungaripteridae 

Domeykodactylus ceciliae

Dsungaripterus weii

Noripterus complicidens

Noripterus parvus

See also

References

  1. ^ Naish D, Simpson M, Dyke G (2013) A New Small-Bodied Azhdarchoid Pterosaur from the Lower Cretaceous of England and Its Implications for Pterosaur Anatomy, Diversity and Phylogeny. PLoS ONE 8(3): e58451. https://doi.org/10.1371/journal.pone.0058451
  2. ^ a b Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  3. ^ a b 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.
  4. ^ Andres, Brian Blake (2014). "A review of pterosaur phylogeny". In Martill, D.; Unwin, D.; Loveridge, R. F. (eds.). The Pterosauria. Cambridge University Press.
  5. ^ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 383–398. doi:10.1017/S1755691013000303.
  • Pterosauria @ Mikko's Phylogeny Archive
  • 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.) (2003): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1–347
  • Peters, D., (2007): The origin and radiation of the Pterosauria — in Flugsaurier: The Wellnhofer pterosaur meeting, Munich, 2007, 27–28
  • Unwin, D. M., 2003: On the phylogeny and evolutionary history of pterosaurs. pp. 139–190. — in Buffetaut, E. & Mazin, J.-M., (eds.) (2003): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1–347
  • Wellnhofer, P., (1991): The Illustrated Encyclopedia of Pterosaurs. Salamander Books Ltd., London, pp. 192

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