Editing Pterosaur (section)

==Classification and phylogeny==
{{further|Phylogeny of pterosaurs}}
[[File:Eudimorphodon_ranzii.jpg|thumb|Fossil of ''[[Eudimorphodon]]'', one of the most primitive pterosaurs]]
In [[phylogenetic taxonomy]], the [[clade]] Pterosauria has usually been defined as node-based and anchored to several extensively studied taxa as well as those thought to be primitive. One 2003 study defined Pterosauria as "The most recent common ancestor of the Anurognathidae, ''[[Preondactylus]]'' and ''Quetzalcoatlus'' and all their descendants."<ref name="kellner2003">{{cite journal | last1 = Kellner | first1 = A. W. | year = 2003 | title = Pterosaur phylogeny and comments on the evolutionary history of the group | journal = Geological Society, London, Special Publications | volume = 217 | issue = 1| pages = 105–37 | doi=10.1144/gsl.sp.2003.217.01.10 | bibcode = 2003GSLSP.217..105K| s2cid = 128892642 }}</ref> However, these types of definition would inevitably leave any related species that are slightly more primitive out of the Pterosauria. To remedy this, a new definition was proposed that would anchor the name not to any particular species but to an anatomical feature, the presence of an enlarged fourth finger that supports a wing membrane.<ref name=earlyarchosaurs>Nesbitt, S.J., Desojo, J.B., & Irmis, R.B. (2013). ''Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and Their Kin''. Geological Society of London. {{ISBN|1862393613}}</ref> This apomorphy-based definition was adopted by the [[PhyloCode]] in 2020 as "[T]he clade characterized by the apomorphy fourth manual digit hypertrophied to support a wing membrane, as inherited by ''Pterodactylus'' (originally ''Ornithocephalus'') ''antiquus'' (Sömmerring 1812)".<ref>{{cite book |editor1-last=de Queiroz |editor1-first=K. |editor2-last=Cantino |editor2-first=P. D. |editor3-last=Gauthier |editor3-first=J. A. |year=2020 |title=Phylonyms: A Companion to the PhyloCode |publisher=CRC Press Boca Raton, FL |page=2072 |isbn=9780429821202 |url=https://books.google.com/books?id=McHgDwAAQBAJ&pg=PA2072 }}</ref> A broader clade, '''Pterosauromorpha''', has been defined as all [[ornithodira]]ns more closely related to pterosaurs than to [[dinosaur]]s.<ref name=padian1997>Padian, K. (1997). "Pterosauromorpha", pp. 617–18 in Currie, P.J. and Padian, K. ''The Encyclopedia of Dinosaurs''. Academic Press. {{ISBN|0122268105}}.</ref>

The internal [[biological classification|classification]] of pterosaurs has historically been difficult, because there were many gaps in the [[fossil record]]. Starting from the 21st century, new discoveries are now filling in these gaps and giving a better picture of the evolution of pterosaurs. Traditionally, they were organized into two [[suborders]]: the [[Rhamphorhynchoidea]], a "primitive" group of long-tailed pterosaurs, and the [[Pterodactyloidea]], "advanced" pterosaurs with short tails.<ref name=DU06b>{{cite book |last=Unwin |first=David M. |title=The Pterosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=978-0-13-146308-0 |page=246}}</ref> However, this traditional division has been largely abandoned. Rhamphorhynchoidea is a [[paraphyletic]] (unnatural) group, since the pterodactyloids evolved directly from them and not from a common ancestor, so, with the increasing use of [[cladistics]], it has fallen out of favor among most scientists.<ref name=WittonPycnofibres/><ref name=luetal2008>{{cite journal |author1=Lü J. |author2=Unwin D.M. |author3=Xu L. |author4=Zhang X. | year = 2008 | title = A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution | journal = Naturwissenschaften | volume = 95 | issue = 9| pages = 891–97 | doi = 10.1007/s00114-008-0397-5 | pmid = 18509616 | bibcode = 2008NW.....95..891L|s2cid=13458087 }}</ref>

Within pterosaurs, several smaller clades have been named.  The clade '''Novialoidea''' was named by paleontologist [[Alexander Kellner|Alexander Wilhelm Armin Kellner]] in [[2003 in paleontology|2003]] as a [[node-based taxon]] consisting of the last common ancestor of ''[[Campylognathoides]]'', ''[[Quetzalcoatlus]]'' and all its descendants. This name was derived from [[Latin]] ''novus'' "new", and ''ala'', "wing", in reference to the wing [[synapomorphy|synapomorphies]] that the members of the clade possess.<ref name=Kellner03>Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. &mdash; ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347</ref>

Paleontologist [[David Unwin]] in 2003 had named the group '''Lonchognatha''' in the same issue of the journal that published Novialoidea ([[Geological Society of London]], Special Publications 217) and defined it as ''[[Eudimorphodon ranzii]]'', ''[[Rhamphorhynchus muensteri]]'', their most recent common ancestor and all its descendants (as a node-based taxon).<ref name=Unwin03>Unwin, D. M., (2003): On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. &mdash; ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347</ref> Under Unwin's and Kellner's [[phylogenetic]] analyses (where ''[[Eudimorphodon]]'' and ''Campylognathoides'' form a family that is basal to both ''[[Rhamphorhynchus]]'' and ''Quetzalcoatlus''), and because Novialoidea was named first (in pages 105–137, while Lonchognatha was named in pages 139–190), Lonchognatha is an objective [[junior synonym]] of the former. However, other analyses find Lonchognatha to be valid (Andres ''et al.'', 2010),<ref>Brian Andres, James M. Clark & Xu Xing (2010) A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs, Journal of Vertebrate Paleontology, 30:1, 163-187, DOI: 10.1080/02724630903409220</ref> or synonymous with the [[Pterosauria]] (Andres, 2010).<ref name=BBA10>{{cite book |last=Andres |first=Brian Blake |year=2010 |title=Systematics of the Pterosauria |url=http://gradworks.umi.com/34/40/3440534.html |publisher=Yale University |pages=366}} [https://www.proquest.com/docview/846734901 A preview that shows the cladogram without clade names]</ref>

The precise relationships between pterosaurs is still unsettled. Many studies of pterosaur relationships in the past have included limited data and were highly contradictory. However, newer studies using larger data sets are beginning to make things clearer. The [[cladogram]] (family tree) below follows a [[phylogenetic]] analysis presented by Longrich, Martill and Andres in 2018, with clade names after Andres ''et al.'' (2014).<ref name=kryptodrakon/><ref name=longrichetal2018>{{cite journal | last1 = Longrich | first1 = N.R. | last2 = Martill | first2 = D.M. | last3 = Andres | first3 = B. | year = 2018 | title = Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary | journal = PLOS Biology | volume = 16 | issue = 3| page = e2001663 | doi = 10.1371/journal.pbio.2001663 | pmid = 29534059 | pmc = 5849296 | doi-access = free }}</ref>
<div style="overflow:auto">
{{clade| style=font-size:90%; line-height:100%;
|label1='''Pterosauria'''
|1={{clade
   |1=[[Eopterosauria]][[File:Eudimorphodon BW.jpg|30px]]
   |label2=[[Macronychoptera]]
   |2={{clade
      |1=[[Dimorphodontia]][[File:Dimorphodon2DB white background.jpg|30px]]
      |sublabel2={{clade label|[[Novialoidea]] |right=0.7em |top=-1.8em}}
      |2={{clade
         |1=[[Campylognathoididae]][[File:Campylogn DB.jpg|30px]]
         |sublabel2={{clade label|[[Breviquartossa]] |right=0.7em |top=-1.8em}}
         |2={{clade
            |1=[[Rhamphorhynchidae]][[File:Rhamphorhynchus DB.jpg|30px]]
            |sublabel2={{clade label|[[Pterodactylomorpha]] |right=0.7em |top=-1.8em}}
            |2={{clade
               |1=''[[Sordes]]''[[File:SordesDB flipped.jpg|30px]]
               |sublabel2={{clade label|[[Monofenestrata]] |right=0.7em |top=-1.8em}}
               |2={{clade
                  |1=[[Darwinoptera]]
                  |sublabel2={{clade label|[[Pterodactyliformes]] |right=0.7em |top=-1.8em}}
                  |2={{clade
                     |1=''[[Changchengopterus]]''
                     |sublabel2={{clade label|[[Caelidracones]] |right=0.7em |top=-1.8em}}
                     |2={{clade
                        |1=[[Anurognathidae]][[File:AnurognathusDB white background.jpg|25px]]
                        |label2=[[Pterodactyloidea]]
                        |2={{clade
                           |1=''[[Kryptodrakon]]''[[File:Kryptodrakon.jpeg|30px]]
                           |label2=[[Lophocratia]]
                           |2={{clade
                              |label1=[[Archaeopterodactyloidea]]
                              |1={{clade
                                 |1=[[Germanodactylidae]][[File:Altmuehlopterus DB.jpg|30px]]
                                 |label2=[[Euctenochasmatia]]
                                 |2={{clade
                                    |1=''[[Pterodactylus]]''[[File:Pterodactylus BMMS7 life.png|30px]]
                                    |2=[[Ctenochasmatoidea]][[File:Plataleorhynchus.jpg|35px]] 
                                    }} 
                                 }}
                              |label2=[[Eupterodactyloidea]]
                              |2={{clade
                                 |1=''[[Haopterus]]''
                                 |label2=[[Ornithocheiroidea]]
                                 |2={{clade
                                    |label1=[[Azhdarchoidea]]
                                    |1={{clade
                                       |1=[[Tapejaromorpha]][[File:Bakonydraco as tapejarid DB.jpg|30px]]
                                       |2=[[Neoazhdarchia]][[File:Quetzalcoatlus07.jpg|30px]]
                                       }}
                                    |2={{clade
                                       |1=''[[Piksi]]''
                                       |label2=[[Pteranodontoidea]]
                                       |2={{clade
                                          |1=[[Pteranodontia]][[File:Pteranodon longiceps mmartyniuk wiki.png|30px]]
                                          |2=[[Ornithocheiromorpha]][[File:LiaoningopterusDB flipped.jpg|25px]] 
                                          }}
                                       }}
                                    }}
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }} 
}}
</div>