Phylogeny of pterosaurs

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.

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

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. Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  3. 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.
  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. S2CID 84617119.
  • 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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