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 (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. 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, in press) will be published in an upcoming book named "The Pterosauria".[4] 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).[5]

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     Eupterodactyloidea (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

Eupterodactyloidea

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

• List of pterosaurs
• Pterosaur
• Graphical timeline of pterosaurs

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.
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