Parapsicephalus

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Parapsicephalus
Temporal range:
Ma
Fossil skull showing brain endocast (
AMNH
1694)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Family: Rhamphorhynchidae
Genus: Parapsicephalus
Arthaber, 1919
Type species
Scaphognathus purdoni
Newton, 1888
Species
  • P. purdoni
    (Newton, 1888)
Synonyms

Parapsicephalus (meaning "beside arch head") is a

dimorphodontians. 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; 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 subgrouping of rhamphorhynchids, but it may represent a basal
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, Dorygnathus, and 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, 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, 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; their junction is marked by a discontinuity in the surface texture of the bone. Overall, the oval-shaped 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, and the lacrimal, 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, which connects back to join the pterygoid.[1]

At the back of the antorbital fenestra is the

flocculus and cerebrum.[1]

The

supratemporal fenestrae. The supratemporal fenestra itself is a somewhat four-sided oval. Behind the postorbital and closing off the supratemporal fenestra is the three-pronged squamosal, which is partially overlapped by the robust, spatula-like paroccipital processes of the occipital. Between the processes is the foramen magnum, which is a 7 millimetres (0.28 in) oval. Further below is the basoccipital, which forms a rounded plate that encloses the back of the skull.[1]

Referred pectoral girdle

The

glenoid 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

Whitby Mudstone Formation, which has been dated to about 182 million years ago, or the Toarcian stage of the Jurassic period.[1]

GSM 3166 was described by Edwin Tulley Newton, who loaned it from Rev. Purdon, in 1888. He described it as a species of 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, and expressed that it was not as close to Scaphognathus as Newton had presumed.[12] Later, in 1919, Gustav von Arthaber, based on the shape of the top of the skull, the elongated nostrils and prefrontal bones, the large antorbital fenestra and eye socket, the deep jugal, and the presence of seven teeth in the 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,

dimorphodontians). 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, which also belongs to the Alum Shale Member.[1] It consists of a scapula, coracoid, and 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, Bavaria, 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

dimorphodontian, is reproduced below.[22]

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

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 is not as vertical; the 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, 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.[1]

However, there are some factors that complicate a rhamphorhynchine position. In particular, the pear-shaped

infratemporal fenestra and the overall size of the antorbital fenestra are more similar to scaphognathines than rhamphorhynchines. Additionally, like more basal non-rhamphorhynchid pterosaurs, there is a half moon-shaped process of the 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[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

See also

References

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  10. ^ a b c Simms, M.J. (2004). "British Lower Jurassic stratigraphy: an introduction" (PDF). British Lower Jurassic Stratigraphy. Geological Conservation Review. Vol. 30. Peterborough: Joint Nature Conservation Committee. pp. 3–45.
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  12. ^ Plieninger, F. "Campylognathus zittelli, ein neuer Flugsaurier aus dem Oberen Lias Schwabens" [Campylognathus zitteli, a new pterosaur from the Upper Lias of Swabia]. Palaeontographica (in German). 41: 193–222.
  13. ^ a b Arthaber, G. (1919). "Studien über Flugsaurier auf Grund der Bearbeitung des Wiener exemplars von Dorygnathus banthensis Theod sp" [Studies in pterosaurs from the processing of Viennese examples of Dorygnathus banthensis Theod sp.] (PDF). Thoughts from the Mathematical-Natural Science Section of the Imperial Academy of Science, Vienna. 97: 391–464.
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