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.

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 (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.[1]

The matrix includes 100 valid pterosaur species plus a single outgroup (Euparkeria capensis). This represents 70.4% of 142 known pterosaur species. 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]).[2] 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)).[1] 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.[2] A paper focusing on the pterosaur phylogeny (Andres, in press) will be published in an upcoming book named "The Pterosauria".[3] An updated and more resolved version of this phylogeny was published formally by Andres and Myers (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).[4]

Pterosauria 



Preondactylus buffarinii



Austriadactylus cristatus





Peteinosaurus zambellii


 Eudimorphodontidae 
 Raeticodactylinae 

Caviramus schesaplanensis



Raeticodactylus filisurensis





Eudimorphodon cromptonellus




Eudimorphodon ranzii



Eudimorphodon rosenfeldi







 Macronychoptera 


Dimorphodon macronyx



Parapsicephalus purdoni



 Novialoidea 


Campylognathoides liasicus



Campylognathoides zitteli



 Breviquartossa 
 Rhamphorhynchidae 


Scaphognathus crassirostris



 Rhamphorhynchinae 

Dorygnathus banthensis





Cacibupteryx caribensis



Nesodactylus hesperius



Rhamphorhynchus muensteri





Harpactognathus gentryii




Angustinaripterus longicephalus



Sericipterus wucaiwanensis









Sordes pilosus


 Monofenestrata 
 Wukongopteridae 

Pterorhynchus wellnhoferi




Darwinopterus modularis



Wukongopterus lii






Changchengopterus pani


 Caelidracones 
 Anurognathidae 

Dendrorhynchoides curvidentatus




Jeholopterus ninchengensis




Anurognathus ammoni



Batrachognathus volans





 Pterodactyloidea 

Archaeopterodactyloidea (see below)




Haopterus gracilis



Ornithocheiroidea (see below)












Archaeopterodactyloidea 
 Germanodactylidae 

Normannognathus wellnhoferi




Germanodactylus cristatus



Germanodactylus rhamphastinus




 Euctenochasmatia 


Pterodactylus antiquus



Pterodactylus kochi





Ardeadactylus longicollum


 Ctenochasmatoidea 
 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











 Ornithocheiroidea 
 Pteranodontia 
 Nyctosauridae 

Muzquizopteryx coahuilensis



"Nyctosaurus" lamegoi



Nyctosaurus gracilis





Alamodactylus byrdi


 Pteranodontoidea 


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















 Azhdarchoidea 


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











 Neoazhdarchia 

 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.









 Thalassodromidae 

Thalassodromeus sethi




Tupuxuara leonardii



Tupuxuara longicristatus




 Dsungaripteridae 


Domeykodactylus ceciliae



Dsungaripterus weii





Noripterus complicidens



Noripterus parvus








See also

References

  1. 1 2 Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  2. 1 2 Richard J. Butler, Stephen L. Brusatte, Brian B. Andres and 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.
  3. Andres, Brian Blake (2014). "A review of pterosaur phylogeny". In Martill, D., Unwin, D., & Loveridge, R. F. (eds). The Pterosauria. Cambridge University Press.
  4. Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh: 1. doi:10.1017/S1755691013000303.
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