Campylognathoides-Jurassic Kratts

Here's a look at Campylognathoides, a pterosaur from Jurassic Kratts-Dimorphodon Rescue, enjoy this preview!

## Facts

***Campylognathoides*** ("curved jaw", Strand 1928) is an extinct genus of pterosaur discovered in the Württemberg Lias deposits (dated to the early Toarcian age) of Germany; this first specimen however, consisted only of wing fragments. Further better preserved specimens were found in the Holzmaden shale; based on these specimens, Felix Plieninger erected a new genus.

### Species

Three species of *Campylognathoides* have been named:

* *Campylognathoides zitteli* (Plieninger, 1894) is the type species.

The larger Holzmaden *Campylognathoides*, *C. zitteli*, had a six foot (1.825 metres for SMNS 9787) wingspan.
* *Campylognathoides liasicus* (Quenstedt, 1858 [originally *Pterodactylus liasicus*])

*C. liasicus* had a three foot wingspan, making it smaller than its Holzmaden contemporary, *C. zitteli*.
* *Parapsicephalus*"Campylognathoides" *indicus* Jain, 1974

*C. indicus* was described by Sohan Lal Jain on the basis of a fragment of jaw, ISI R. 48, recovered from Chanda district, India. Kevin Padian considers this a nomen dubium, possibly based on a fish fossil. That the Kota Formation in which it was found, has since been redated to the Middle Jurassic or later, seems to preclude any close connection to *Campylognathoides*, even if it were a pterosaur.
The distinction between *C. liasicus* and *C. zitteli* is problematical. Plieninger merely recognised the smaller species because he considered its fossil too poor in quality to refer other specimens to. However, in 1925 Swedish researcher Carl Wiman, studying specimen UUPM R157, concluded that a fundamental morphological difference could distinguish the two species: *C. zitteli* has a proportionally much longer wing. In 2008 however, Padian pointed out that this might well have been a matter of ontogenetic development, larger individuals growing extra large wings to limit the wing load. Other differences, such as the larger number of teeth in the lower jaw, a longer snout and nares, five instead of four sacrals, perpendicular sacral ribs and a longer leg, might conceivably also be size-related. Final proof could only be given by a continuous growth series, as previously has been done in the case of *Rhamphorhynchus* and *Pterodactylus*. Provisionally Padian kept distinguishing two species, but moved two specimens to *C. zitteli*: SMNS 51100 and GPIT 24470, because of their larger size and morphological similarities.

## Classification

Plieninger in his later publications assigned *Campylognathus* to the "Rhamphorhynchoidea". As this suborder is a paraphyletic assemblage of not specially related basal pterosaurs, this classification merely states the negative fact that it was not a pterodactyloid. A positive determination was first attempted by Baron Franz Nopcsa who in 1928 assigned the genus to the subfamily Rhamphorhynchinae within the family Rhamphorhynchidae. After a period in which very little work was done on pterosaur systematics, in 1967 Oskar Kuhn placed *Campylognathoides* in its own subfamily within the Rhamphorhynchidae, the Campylognathoidinae. However, in 1974 Peter Wellnhofer concluded that it was placed in a more basal position in the phylogenetic tree, below the Rhamphorhynchidae. In the early twenty-first century this was confirmed by the first extensive exact cladistic analyses. In 2003 both David Unwin and Alexander Kellner introduced a clade Campylognathoididae; within Unwin's terminology this clade is the sister clade of the Breviquartossa within the Lonchognatha; applying Kellner's terminology it is the most basal offshoot within the Novialoidea. There is no material difference between the two positions.

According to the analyses *Campylognathoides* would be closely related to *Eudimorphodon*, to which it is similar in skull, sternum and humerus form. This was confirmed by Padian in 2009, though Padian also pointed out several basal features present in *Eudimorphodon* but lacking in *Campylognathoides*. In 2010 an analysis was published by Brian Andres showing that *Eudimorphodon* together with *Austriadactylus* formed a very basal clade, leaving *Campylognathoides* as the only known member of the Campylognathoididae. Other recent phylogenetic analyses confirm these results, and suggest that *Campylognathoides* is more derived than all Triassic-aged pterosaurs, as well as the Early Jurassic *Dimorphodon* and *Parapsicephalus*. *Campylognathoides* is the basalmost member of the Novialoidea which is defined as a node-based taxon consisting of the last common ancestor of *Campylognathoides*, *Quetzalcoatlus* and all its descendants.

## Paleobiology

Traditionally a piscivorous lifestyle is attributed to *Campylognathoides*, as to most pterosaurs; in this case supported by the provenance of the finds from marine sediments and the very long wings. Pterosaurologist Kevin Padian, however, has suggested that, in view of the stout short teeth, ideal for delivering a piercing bite, the form might have been a predator of small terrestrial animals instead. Conversely, Mark Witton suggests the construction of *Campylognathoides'* extremely robust forelimbs, with proportionally long wing fingers, could be a specialization for a fast aerial lifestyle comparable to those of falcons and mastiff bats.