# Parapsicephalus

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Genus of rhamphorhynchid pterosaur from the Early Jurassic

Parapsicephalus Temporal range: Toarcian, 182 Ma PreꞒ Ꞓ O S D C P T J K Pg N Fossil skull showing brain endocast (AMNH 1694) Scientific classification Kingdom: Animalia Phylum: Chordata Class: Reptilia Order: †Pterosauria Family: †Rhamphorhynchidae Genus: †Parapsicephalus Arthaber, 1919 Type species †Scaphognathus purdoni Newton, 1888 Species †P. purdoni (Newton, 1888) Synonyms Scaphognathus purdoni Newton, 1888 Dorygnathus purdoni (Newton, 1888)

***Parapsicephalus*** (meaning "beside arch head") is a [genus](/source/Genus) of long-tailed [rhamphorhynchid](/source/Rhamphorhynchid) [pterosaurs](/source/Pterosaur) from the [Lower Jurassic](/source/Lower_Jurassic) [Whitby](/source/Whitby), [Yorkshire](/source/Yorkshire), [England](/source/England). It contains a single species, ***P. purdoni***, named initially as a species of the related rhamphorhynchid *[Scaphognathus](/source/Scaphognathus)* in 1888 but moved to its own genus in 1919 on account of a unique combination of characteristics. In particular, the top surface of the skull of *Parapsicephalus* is convex, which is otherwise only seen in [dimorphodontians](/source/Dimorphodontia). This has been the basis of its referral to the Dimorphodontia by some researchers, but it is generally agreed upon that *Parapsicephalus* probably represents a rhamphorhynchid. Within the Rhamphorhynchidae, *Parapsicephalus* has been synonymized with the roughly contemporary *[Dorygnathus](/source/Dorygnathus)*; this, however, is not likely given the many differences between the two taxa, including the aforementioned convex top surface of the skull. *Parapsicephalus* has been tentatively referred to the [Rhamphorhynchinae](/source/Rhamphorhynchidae) subgrouping of rhamphorhynchids, but it may represent a [basal](/source/Basal_(phylogenetics)) member of the group instead.

## Description

The type skull of *Parapsicephalus*, which is 14 centimetres (5.5 in) long as preserved, suggests that it was of medium size.[1] Comparisons with the related pterosaurs *[Scaphognathus](/source/Scaphognathus)*, *[Dorygnathus](/source/Dorygnathus)*, and *[Jianchangnathus](/source/Jianchangnathus)* indicates that the full skull would have been 18–19.6 centimetres (7.1–7.7 in) long.[1][2][3] Wellnhofer estimated its wingspan at 1 metre (3 ft 3 in);[4] more recently, a referred [humerus](/source/Humerus), which is 10 centimetres (3.9 in) long, has produced wingspan estimates of 1.68–3.26 metres (5 ft 6 in – 10 ft 8 in).[5]

### Skull

Diagram of the skull of *Parapsicephalus purdoni*, viewed from the left

When viewed from the side, the convex top of the skull formed a gentle slope. The elongate frontal process of the [premaxilla](/source/Premaxilla), which extends backwards towards the eyes, may have supported a low crest along its midline.[6] Below the nostril, the premaxilla meets with the [maxilla](/source/Maxilla); their junction is marked by a discontinuity in the surface texture of the bone. Overall, the oval-shaped [antorbital fenestra](/source/Antorbital_fenestra), situated behind and separate from the nostril, measures 45 millimetres (1.8 in) long and 24 millimetres (0.94 in) tall. The maxilla extends backwards in a half-moon shape to encircle the front end of the fenestra, with the top prong of the maxilla forming a 45° angle with the horizontal. The top end of the fenestra is enclosed by the thin, rectangular, and slightly concave [nasal](/source/Nasal_bone), and the [lacrimal](/source/Lacrimal_bone), which is not well-preserved but may have been long, slender, and triangular. On the underside of the skull, the maxilla forms the majority of the palate (not the premaxilla, as previously assumed[7]), extending back from below the external nostrils. Along the midline of the maxilla is situated a thin strip of bone, the [vomer](/source/Vomer), which connects back to join the [pterygoid](/source/Pterygoid_bone).[1]

At the back of the antorbital fenestra is the [jugal](/source/Jugal_bone), which has often been illustrated as a small V-shaped, two-pronged structure,[7] but it is actually large and has four prongs. The lacrimal process extends 20 millimetres (0.79 in) forward and upward from the main body of the jugal, while the more robust postorbital process extends 18 millimetres (0.71 in) backward and upward. Collectively, they enclose the bottom of the eye socket, with angles of 45° on each side. The pear-shaped eye socket measures 34 millimetres (1.3 in) tall and 32 millimetres (1.3 in) wide at the widest point. Behind the jugal is the [quadratojugal](/source/Quadratojugal_bone), which has traditionally been depicted as hypertrophied and occupying the location of the back portion of the jugal;[7] it is actually a small, half moon-shaped bone wedged between the jugal and the [quadrate](/source/Quadrate_bone) and situated below the elongate [infratemporal fenestra](/source/Infratemporal_fenestra). Overall, the infratemporal fenestra is shaped similarly to the eye socket. The quadrates are strap-like, and wrap around from the back to the bottom of the skull. Although mostly obscured, the removal of the [parietal](/source/Parietal_bone) during preparation has exposed part of the [endocast](/source/Endocast) of the brain, which has a large [flocculus](/source/Flocculus_(cerebellar)) and [cerebrum](/source/Cerebrum).[1]

The [frontal](/source/Frontal_bone), which is located on the top of the skull between the premaxilla and the eye socket, takes the shape of a sub-rectangle with a large, [rhombic](/source/Rhombus) process extending forward to meet the nasal. The frontal process of the premaxilla cuts into the frontal along the midline of this rhombus. Contacting the rear ends of both the frontal and jugal is the thin and triangular [postorbital](/source/Postorbital_bone), about 15 millimetres (0.59 in) long along the bottom, extends backwards to separate the infratemporal and [supratemporal fenestrae](/source/Supratemporal_fenestra). The supratemporal fenestra itself is a somewhat four-sided oval. Behind the postorbital and closing off the supratemporal fenestra is the three-pronged [squamosal](/source/Squamosal_bone), which is partially overlapped by the robust, spatula-like paroccipital processes of the [occipital](/source/Occipital_bone). Between the processes is the [foramen magnum](/source/Foramen_magnum), which is a 7 millimetres (0.28 in) oval. Further below is the [basoccipital](/source/Occipital_bone), which forms a rounded plate that encloses the back of the skull.[1]

### Referred pectoral girdle

The [humerus](/source/Humerus) referred to *Parapsicephalus* probably had a slightly deflected deltopectoral crest. It bears a sub-triangular medial process, 7 millimetres (0.28 in) wide and 25 millimetres (0.98 in) long, that originates close to the [humeral head](/source/Upper_extremity_of_humerus). In cross-section, the shaft of the humerus is sub-rectangular, and about 7 millimetres (0.28 in) wide at the midpoint; the bottom third of the shaft is bowed forward. The [scapula](/source/Scapula) and [coracoid](/source/Coracoid) appear to be completely fused; they are respectively 67 millimetres (2.6 in) and 59 millimetres (2.3 in) long, and form an angle of 70° to each other, creating overall a V-shaped structure as in *[Sericipterus](/source/Sericipterus)*.[8] The outer end of the scapula is noticeably wider than the inner end, and the [glenoid](/source/Glenoid_cavity) is positioned entirely on the scapula, with the shaft curving about 15° towards the glenoid. Meanwhile, the portion of the coracoid closest to the glenoid is very expanded.[5]

## Discovery and naming

*Parapsicephalus* is only definitely known from a single partial skull lacking the snout, but including a detailed [endocast](/source/Endocast) of the [brain](/source/Brain). It is catalogued under the specimen number GSM 3166, and is stored at the [British Geological Survey](/source/British_Geological_Survey) in [Keyworth](/source/Keyworth), [Nottinghamshire](/source/Nottinghamshire). It was collected in the 1880s by Reverend D.W. Purdon from the Loftus Alum Shale Quarry, in [Loftus, North Yorkshire](/source/Loftus%2C_North_Yorkshire),[9][10] from which fossils had been discovered as early as the early nineteenth century.[11] The quarry has since become disused since operations ceased in 1860. The exposed rocks, which consist of [pyrite](/source/Pyrite)-rich shales with [calcareous](/source/Calcareous) [concretions](/source/Concretion),[10] are part of the upper Alum Shale Member of the [Whitby Mudstone Formation](/source/Whitby_Mudstone_Formation), which has been dated to about 182 million years ago, or the [Toarcian](/source/Toarcian) stage of the [Jurassic](/source/Jurassic) period.[1]

GSM 3166 was described by [Edwin Tulley Newton](/source/Edwin_Tulley_Newton), who loaned it from Rev. Purdon, in 1888. He described it as a species of *[Scaphognathus](/source/Scaphognathus)*, *S. purdoni*, named after Purdon; he did not include it in the type species *S. crassirostris* due to differences in the curvature of the top of the skull, as well as the midline channel on the top of the skull. In his description of the braincase, he noted its intermediate morphology between that of lizards and birds, which he considered evidence of a close relationship between birds, pterosaurs, and "reptiles".[7] F. Plieninger subsequently compared GSM 3166 to *[Campylognathoides](/source/Campylognathoides)*, and expressed that it was not as close to *Scaphognathus* as Newton had presumed.[12] Later, in 1919, [Gustav von Arthaber](https://en.wikipedia.org/w/index.php?title=Gustav_von_Arthaber&action=edit&redlink=1), based on the shape of the top of the skull, the elongated nostrils and [prefrontal bones](/source/Prefrontal_bone), the large [antorbital fenestra](/source/Antorbital_fenestra) and eye socket, the deep [jugal](/source/Jugal_bone), and the presence of seven teeth in the [maxilla](/source/Maxilla), referred GSM 3166 to the new genus *Parapsicephalus*.[1][13]

### Possible synonymy with *Dorygnathus*

Restoration of *Dorygnathus banthensis*; *Parapsicephalus* has been considered a member of the same genus

In 2003, [David Unwin](https://en.wikipedia.org/w/index.php?title=David_Unwin&action=edit&redlink=1), [Kenneth Carpenter](/source/Kenneth_Carpenter), and others suggested that *Parapsicephalus* was actually closer to the roughly contemporaneous *[Dorygnathus](/source/Dorygnathus)* from German deposits. Unwin formally renamed it to the new combination *Dorygnathus purdoni*.[14][15][16] This renaming was adopted by some researchers[17][18] but not others.[2][19][20][21] However, a 2017 redescription of GSM 3166 noted a number of ways in which *Dorygnathus* differed from *Parapsicephalus*: the greater angle of the lacrimal process of the maxilla, the more reduced maxillary process of the [premaxilla](/source/Premaxilla), the broader angle of the jugal beneath the eye socket, the overall thinner jugal relative to skull height, the more rounded the eye socket, the more oval-shaped [supratemporal fenestra](/source/Supratemporal_fenestra), and the convex top of the skull (which is unique among pterosaurs except for [dimorphodontians](/source/Dimorphodontia)). On the basis of these characteristics, the study recognized *Parapsicephalus* as a distinct genus.[1]

### Possible additional specimens

The specimen NHMUK PV R36634 was found in 2011 within a concretion in [Saltwick Bay](/source/Saltwick_Bay), which also belongs to the Alum Shale Member.[1] It consists of a [scapula](/source/Scapula), [coracoid](/source/Coracoid), and [humerus](/source/Humerus); the head of the humerus was broken off during excavation as a result of the concretion being hammered open (which is the usual method for exposing ammonites preserved in concretions). Although it is impossible to refer this specimen to *Parapsicephalus* with confidence, its provenance and similarity to *Dorygnathus* were the basis of the tentative identification of the specimen as belonging to this genus.[5] An additional possible specimen is a skull collected in 1994 from [Altdorf](/source/Altdorf_bei_N%C3%BCrnberg), [Bavaria](/source/Bavaria), [Germany](/source/Germany), which bears great similarity to GSM 3166 and also preserves some additional elements. It is currently held by a private collector, but will soon be donated to an institution in the UK.[1]

## Classification

Although Newton originally considered *Parapsicephalus* as being a species of *[Scaphognathus](/source/Scaphognathus)*,[7] Arthaber remarked that it was actually more similar to *[Dimorphodon](/source/Dimorphodon)* instead.[13] The prevailing view since Arthaber's renaming of *Parapsicephalus* as a distinct genus has been that *Parapsicephalus* represents some kind of [rhamphorhynchid](/source/Rhamphorhynchidae),[14][19] although several phylogenies by Brian Andres and colleagues supported Arthaber's hypothesis of it being closely affiliated with *Dimorphodon*.[21][22] Characters which support this placement include the convex top of the skull, the pear-shaped eye socket, the angle of the [quadrate](/source/Quadrate_bone), and the thick [jugal](/source/Jugal_bone).[21] The topology recovered by Andres and Myers in 2013, showing *Parapsicephalus* as a [dimorphodontian](/source/Dimorphodontia), is reproduced below.[22]

Restoration of *Dimorphodon macronyx*, considered a close relative of *Parapsicephalus* by some authors

Pterosauria Preondactylus buffarinii Austriadactylus cristatus Peteinosaurus zambellii Eudimorphodontidae Macronychoptera Dimorphodon macronyx Parapsicephalus purdoni Novialoidea

A 2017 analysis of *Parapsicephalus* found little support for it being placed in the Dimorphodontia: its skull was comparatively longer; the snout is slightly upturned with outward-projecting teeth, as in rhamphorhynchids; the [quadrate](/source/Quadrate_bone) is not as vertical; the [antorbital fenestra](/source/Antorbital_fenestra) is offset below the nostril instead of being at the same level; and, although the top of the skull is convex in both, the condition in *Parapsicephalus* is not quite as extreme. Thus, affinities with the Rhamphorhynchidae were considered more probable. Within the Rhamphorhynchidae, unlike the [scaphognathines](/source/Rhamphorhynchidae), the antorbital fenestra is more than twice as long as it is tall,[8][14] and has a concave back margin; the angle of the quadrate is also more than 120°.[23] This implies that *Parapsicephalus* is a member of the [Rhamphorhynchinae](/source/Rhamphorhynchidae).[1]

However, there are some factors that complicate a rhamphorhynchine position. In particular, the pear-shaped [infratemporal fenestra](/source/Infratemporal_fenestra) and the overall size of the antorbital fenestra are more similar to scaphognathines than rhamphorhynchines. Additionally, like more [basal](/source/Basal_(phylogenetics)) non-rhamphorhynchid pterosaurs, there is a half moon-shaped process of the [premaxilla](/source/Premaxilla) extending beneath the nostril. It is thus possible that *Parapsicephalus* represents a basal rhamphorhynchid that is not in either group, which is not unexpected given its temporal context. The contemporary *[Allkaruen](/source/Allkaruen)*[24] is also a potentially viable subject of comparison, although its material and that of *Parapsicephalus* do not readily overlap.[1]

## Paleoecology

The Alum Shale Member of the [Whitby Mudstone Formation](/source/Whitby_Mudstone_Formation) was probably deposited in an oxygen-poor, shallow-water environment.[10] A number of marine reptiles are known from this locality: the [ichthyosaurs](/source/Ichthyosauria) *[Stenopterygius](/source/Stenopterygius)*, *[Temnodontosaurus](/source/Temnodontosaurus)*, and *[Eurhinosaurus](/source/Eurhinosaurus)*; the [plesiosaurs](/source/Plesiosaur) *[Eretmosaurus](/source/Eretmosaurus)*, *[Sthenarosaurus](/source/Sthenarosaurus)*, and *[Microcleidus](/source/Microcleidus)*; and the [thalattosuchians](/source/Thalattosuchia) *[Steneosaurus](/source/Steneosaurus)* and *[Pelagosaurus](/source/Pelagosaurus)*. Indeterminate theropod remains have also been found.[1][9]

## See also

- [List of pterosaur genera](/source/List_of_pterosaur_genera)

- [Timeline of pterosaur research](/source/Timeline_of_pterosaur_research)

## References

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1. **[^](#cite_ref-carpenter_16-0)** Carpenter, K.; Unwin, D.; Cloward, K.; Miles, C.; Miles, C. (January 2003). ["A new scaphognathine pterosaur from the Upper Jurassic Morrison Formation of Wyoming, USA"](http://sp.lyellcollection.org/content/217/1/45.abstract). *Geological Society, London, Special Publications*. **217** (1): 45–54. [Bibcode](/source/Bibcode_(identifier)):[2003GSLSP.217...45C](https://ui.adsabs.harvard.edu/abs/2003GSLSP.217...45C). [doi](/source/Doi_(identifier)):[10.1144/GSL.SP.2003.217.01.04](https://doi.org/10.1144%2FGSL.SP.2003.217.01.04). [S2CID](/source/S2CID_(identifier)) [129087640](https://api.semanticscholar.org/CorpusID:129087640).

1. **[^](#cite_ref-hone_17-0)** Hone, D.W.E.; Benton, M.J. (2007). ["An evaluation of the phylogenetic relationships of the pterosaurs among archosauromorph reptiles"](https://web.archive.org/web/20170222053210/http://www.davehone.co.uk/wp-content/uploads/2014/01/Hone-Benton-2007-Archosauromorphs.pdf) (PDF). *Journal of Systematic Palaeontology*. **5** (4): 465–469. [doi](/source/Doi_(identifier)):[10.1017/S1477201907002064](https://doi.org/10.1017%2FS1477201907002064). [S2CID](/source/S2CID_(identifier)) [86145645](https://api.semanticscholar.org/CorpusID:86145645). Archived from [the original](http://www.davehone.co.uk/wp-content/uploads/2014/01/Hone-Benton-2007-Archosauromorphs.pdf) (PDF) on 2017-02-22. Retrieved 2017-02-21.

1. **[^](#cite_ref-bar_18-0)** Barrett, P.M.; Butler, R.J.; Edwards, N.P.; Milner, A.R. (2008). ["Pterosaur distribution in time and space: an atlas"](https://epub.ub.uni-muenchen.de/12007/1/zitteliana_2008_b28_05.pdf) (PDF). *Zitteliana*. **B28**: 61–107. [ISSN](/source/ISSN_(identifier)) [1612-4138](https://search.worldcat.org/issn/1612-4138).

1. ^ [***a***](#cite_ref-cacibu_19-0) [***b***](#cite_ref-cacibu_19-1) Gasparini, Z.; Fernandez, M.; de la Fuente, M. (2004). ["A New Pterosaur from the Jurassic of Cuba"](https://doi.org/10.1111%2Fj.0031-0239.2004.00399.x). *Palaeontology*. **47** (4): 919–927. [doi](/source/Doi_(identifier)):[10.1111/j.0031-0239.2004.00399.x](https://doi.org/10.1111%2Fj.0031-0239.2004.00399.x).

1. **[^](#cite_ref-palate_20-0)** Osi, A.; Prondvai, E.; Frey, E.; Pohl, B. (2010). ["New Interpretation of the Palate of Pterosaurs"](https://doi.org/10.1002%2Far.21053). *Anatomical Record*. **293** (2): 243–258. [doi](/source/Doi_(identifier)):[10.1002/ar.21053](https://doi.org/10.1002%2Far.21053). [PMID](/source/PMID_(identifier)) [19957339](https://pubmed.ncbi.nlm.nih.gov/19957339). [S2CID](/source/S2CID_(identifier)) [31314805](https://api.semanticscholar.org/CorpusID:31314805).

1. ^ [***a***](#cite_ref-andres2014_21-0) [***b***](#cite_ref-andres2014_21-1) [***c***](#cite_ref-andres2014_21-2) Andres, B.; Clark, J.; Xu, X. (2014). ["The Earliest Pterodactyloid and the Origin of the Group"](https://doi.org/10.1016%2Fj.cub.2014.03.030). *Current Biology*. **24** (9): 1011–1016. [doi](/source/Doi_(identifier)):[10.1016/j.cub.2014.03.030](https://doi.org/10.1016%2Fj.cub.2014.03.030). [PMID](/source/PMID_(identifier)) [24768054](https://pubmed.ncbi.nlm.nih.gov/24768054).

1. ^ [***a***](#cite_ref-LoneStarPterosaurs_22-0) [***b***](#cite_ref-LoneStarPterosaurs_22-1) Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". *Earth and Environmental Science Transactions of the Royal Society of Edinburgh*. **103** (3): 383–398. [doi](/source/Doi_(identifier)):[10.1017/S1755691013000303](https://doi.org/10.1017%2FS1755691013000303). [S2CID](/source/S2CID_(identifier)) [84617119](https://api.semanticscholar.org/CorpusID:84617119).

1. **[^](#cite_ref-qinglong_23-0)** Lu, J.; Unwin, D.M.; Zhao, B.; Gao, C.; Shen, C. (2012). ["A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae) from the Middle/Upper Jurassic of Qinglong, Hebei Province, China"](http://www.mapress.com/zootaxa/2012/f/z03158p019f.pdf) (PDF). *Zootaxa*. **3158**: 1–19. [doi](/source/Doi_(identifier)):[10.11646/zootaxa.3158.1.1](https://doi.org/10.11646%2Fzootaxa.3158.1.1).

1. **[^](#cite_ref-codorniu2016_24-0)** Codorniú, L.; Carabajal, A.P.; Pol, D.; Unwin, D.; Rauhut, O.W.M (2016). ["A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5012331). *PeerJ*. **4** e2311. [doi](/source/Doi_(identifier)):[10.7717/peerj.2311](https://doi.org/10.7717%2Fpeerj.2311). [PMC](/source/PMC_(identifier)) [5012331](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5012331). [PMID](/source/PMID_(identifier)) [27635315](https://pubmed.ncbi.nlm.nih.gov/27635315).

v t e Pterosauria Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Archosauria Clade: Avemetatarsalia Clade: Pterosauromorpha Avemetatarsalia see Avemetatarsalia Pterosauria see below↓ Pterosauria Pterosauria Daohugoupterus Eotephradactylus Preondactylia Austriadactylus Preondactylus Caviramidae? Carniadactylus? Caviramus Raeticodactylus? Austriadraconidae Arcticodactylus? Austriadraco Seazzadactylus Eopterosauria Austriadraco? Peteinosaurus? Preondactylia? Raeticodactylidae Caviramus? Pachagnathus Raeticodactylus Yelaphomte Eudimorphodontidae Arcticodactylus? Carniadactylus? Eudimorphodon Zambellisauria? Peteinosaurus? Macronychoptera Herbstosaurus Dimorphodontidae Allkaruen? Caelestiventus Dimorphodon Parapsicephalus? Peteinosaurus? Rhamphinion? Lonchognatha? Eudimorphodon? Novialoidea Campylognathoididae Bergamodactylus Campylognathoides Rhamphorhynchidae Angustinaripterus Bellubrunnus Cacibupteryx Dearc Dolicorhamphus? Dorygnathus Harpactognathus Klobiodon Nesodactylus Orientognathus Parapsicephalus? Qinglongopterus Rhamphorhynchus Sericipterus Scaphognathidae? Scaphognathidae? Fenghuangopterus Jianchangnathus Jianchangopterus Scaphognathus Sordes? Pterodactylomorpha see below↓ Pterodactylomorpha Pterodactylomorpha Allkaruen? Sordes? Monofenestrata Archaeoistiodactylus Makrodactylus Melkamter Normannognathus Anurognathidae? Darwinoptera Ceoptera Pterorhynchus Wukongopteridae Archaeoistiodactylus? Cuspicephalus Darwinopterus Douzhanopterus? Kunpengopterus Wukongopterus Pterodactyliformes Changchengopterus? Douzhanopterus Laueropterus Skiphosoura Caelidracones Anurognathidae Anurognathus Batrachognathus Cascocauda Dendrorhynchoides? Jeholopterus? Luopterus Mesadactylus Sinomacrops Vesperopterylus Pterodactyloidea Dermodactylus Eurolimnornis Herbstosaurus? Kryptodrakon Ningchengopterus Pangupterus Propterodactylus Samrukia? Wenupteryx Lophocratia see below↓ Lophocratia Archaeopterodactyloidea Prejanopterus Germanodactylidae Altmuehlopterus? Germanodactylus Normannognathus? Tendaguripterus? Euctenochasmatia Diopecephalus Pterodactylus Ctenochasmatoidea Gallodactylidae Aurorazhdarcho? Cycnorhamphus Normannognathus? Petrodactyle? Moganopterinae? Aurorazhdarchia Aerodactylus Gallodactylidae? Aurorazhdarchidae Ardeadactylus? Aurorazhdarcho Huanhepterus? Ctenochasmatidae Ardeadactylus? Balaenognathus Beipiaopterus Cathayopterus Cratonopterus Ctenochasma Elanodactylus Eosipterus Feilongus Forfexopterus Garudapterus Gegepterus Gladocephaloideus Gnathosaurus Huanhepterus? Kepodactylus Liaodactylus Lusognathus Moganopterus Otogopterus Petrodactyle? Plataleorhynchus Pterodaustro Pterofiltrus Spathagnathus Tacuadactylus Eupterodactyloidea Altmuehlopterus? Ornithocheiroidea see below↓ Ornithocheiroidea Ornithocheiroidea Piksi? Tapejaroidea Dsungaripteridae Banguela? Domeykodactylus Dsungaripterus Lonchognathosaurus? Noripterus Ordosipterus Puntanipterus? Tendaguripterus? Azhdarchoidea Inabtanin Ornithostoma Tapejaromorpha Bennettazhia Keresdrakon? Thalassodromidae? Alanqa? Argentinadraco? Banguela? Kariridraco Leptostomia? Thalassodromeus Tupuxuara Xericeps? Tapejaridae Afrotapejara Aymberedactylus Bakonydraco? Caiuajara Caupedactylus Eopteranodon? Europejara Huaxiadraco Keresdrakon? Lacusovagus? Nemicolopterus Sinopterus Tapejara Torukjara Tupandactylus Vectidraco Wightia Neoazhdarchia Dsungaripteromorpha? Microtuban Dsungaripteridae Thalassodromidae? Azhdarchomorpha Chaoyangopteridae Apatorhamphus? Chaoyangopterus Eoazhdarcho Jidapterus Lacusovagus? Meilifeilong Microtuban? Shenzhoupterus Azhdarchiformes Montanazhdarcho Radiodactylus Alanqidae? Alanqa Argentinadraco? Keresdrakon? Leptostomia? Montanazhdarcho? Xericeps? Azhdarchidae Aerotitan? Alanqa? Albadraco? Aralazhdarcho Arambourgiania Azhdarcho Cryodrakon Eurazhdarcho? Galgadraco Gobiazhdarcho Hatzegopteryx Infernodrakon Mistralazhdarcho? Navajodactylus? Nipponopterus Palaeocursornis Phosphatodraco? Quetzalcoatlus Thanatosdrakon Tsogtopteryx Wellnhopterus? Zhejiangopterus? Pteranodontoidea see below↓ Pteranodontoidea Pteranodontoidea Santanadactylus Pteranodontia Pteranodontidae Bogolubovia Dawndraco Pteranodon Tethydraco Volgadraco? Nyctosauromorpha Alamodactylus Cretornis Volgadraco? Aponyctosauria Alcione Epapatelo Simurghia Nyctosauridae Barbaridactylus Muzquizopteryx Nyctosaurus Ornithocheiromorpha Aussiedraco Serradraco Unwindia Lonchodectidae Ikrandraco Lonchodectes Lonchodraco? Lanceodontia Draigwenia Lonchodectidae? Istiodactyliformes Hongshanopterus? Linlongopterus Yixianopterus Lonchodectidae? Mimodactylidae Haopterus Linlongopterus Mimodactylus Istiodactylidae Istiodactylus Liaoxipterus Lingyuanopterus Longchengpterus Luchibang Nurhachius Ornithocheiriformes Barbosania Brasileodactylus Cearadactylus Hamipterus Boreopteridae Boreopterus Zhenyuanopterus Ornithocheirae Ornithocheiridae Aetodactylus? Araripesaurus Arthurdactylus Camposipterus? Caulkicephalus? Cimoliopterus? Coloborhynchus? Draigwenia? Ferrodraco? Guidraco? Haliskia? Ludodactylus? Mythunga? Ornithocheirus Siroccopteryx? Tropeognathus? Uktenadactylus? Targaryendraconia? Saratovia Cimoliopteridae Aetodactylus Camposipterus? Cimoliopterus Targaryendraconidae Aussiedraco? Barbosania? Targaryendraco Anhangueria Brasileodactylus? Ornithocheiridae? Hamipteridae? Hamipterus Iberodactylus Anhangueridae Aerodraco Akharhynchus Anhanguera Amblydectes? Caulkicephalus? Cearadactylus? Coloborhynchus Ferrodraco? Guidraco Haliskia? Liaoningopterus Ludodactylus Maaradactylus Mythunga Nicorhynchus Siroccopteryx? Thapunngaka Tropeognathus Uktenadactylus

Taxon identifiers Parapsicephalus purdoni Wikidata: Q2351702 GBIF: 4818510 IRMNG: 11348283 Open Tree of Life: 4947325 Paleobiology Database: 143942

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Adapted from the Wikipedia article [Parapsicephalus](https://en.wikipedia.org/wiki/Parapsicephalus) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Parapsicephalus?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.