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.

## Contents

* 1 Unwin (2003)
* 2 Kellner (2003)
* 3 Andres and Myers (2013)
* 4 See also
* 5 References

## 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  |  | Template:Clade label |  |  |

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

Template:Clade label
Template:Clade label
|  |  |

Haopterus gracilis

|

Ornithocheiroidea |  | Template:Clade label |  |  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

Template:Clade label
|  |  |  |  |

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

Template:Clade label
Template:Clade label
Template:Clade label
Template:Clade label
Template:Clade label
Template:Clade label
Template:Clade label
Template:Clade label

## 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. ↑ 2.0 2.1 Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. pp. 366. http://gradworks.umi.com/34/40/3440534.html. A preview that shows the cladogram without clade names
3. ↑ 3.0 3.1 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.. The Pterosauria. Cambridge University Press. http://www.easons.com/display.asp?K=9780521518956&sf1=contributor&st1='David Unwin'&m=4&dc=19.
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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