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 ningchengensis Anurognathus ammoni Batrachognathus volans Archaeopterodactyloidea Germanodactylidae 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 Ctenochasmatoidea Euctenochasmatia Haopterus gracilis Ornithocheiroidea Pteranodontia  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 Pteranodontoidea 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 Neoazhdarchia Azhdarchoidea Pterodactyloidea Caelidracones Monofenestrata Breviquartossa Novialoidea Macronychoptera

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. 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. Bibcode:2012EESTR.103..383A. 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