The first specimen of the extinct pterosaur genus Campylognathoides, which dates to the early Toarcian period, was found in Germany's Württemberg Lias deposits. However, it only had portions of its wings. Felix Plieninger created a new genus based on additional, better-preserved specimens that were discovered in the Holzmaden shale.
Discovery
Friedrich August Quenstedt designated Pterodactylus liasicus as a new species in 1858. Its foundation was a fossil, holotype GPIT 9533, including a few wing bones that was discovered in early Toarcian strata close to Metzingen. The strata date back around 180 million years. The Lias were referred to by the particular name. Quenstedt concluded that the new species did not belong to more basic taxa, such as the long-tailed Rhamphorhynchus, since he believed he had found lengthy metacarpals in the wing. Near Holzmaden, in 1893, commercial fossil collector Bernhard Hauff Sr. found the skeleton of a big pterosaur. Felix Plieninger created the new genus Campylognathus in 1894 based on this specimen. Given the bent lower jaw, the genus name is derived from the Greek words kampylos, which means "bent," and gnathos, which means "jaw." Zitteli Campylognathus is the type species. The particular name is in honor of Alfred von Zittel. Holotype number is SMNS 9787. Hauff produced another specimen in 1897, which was subsequently purchased by Pittsburgh's Carnegie Museum of Natural History in 1903. The most comprehensive source of knowledge on the genus is this fossil, CM 11424, because of its completeness. When Plieninger examined P. liasicus for the first time in 1901, he found that Quenstedt had misidentified the species as a basic pterosaur when in fact the short metacarpal was really a coracoid. P. liasicus and the Pittsburgh specimen were assigned to Campylognathus by Plieninger in 1906, but the precise status of each of the three exemplars was not yet determined. However, Plieninger identified C. liasicus as a second species of Campylognathus in 1907, and CM 11424 was also attributed to this species. The African bug Campylognathus nigrensis, a genus of the Heteroptera described in 1890, was previously known by the name Campylognathus, as Norwegian entomologist Embrik Strand found in the 1920s. In 1928, he gave the pterosaur a new name, Campylognathoides, because the original name was taken. The total number of known specimens has increased to about twelve over the 20th century thanks to additional discoveries.
Species
The type species is Campylognathoides zitteli (Plieninger, 1894).
With a wingspan of 1.75 m (5 ft 9 in), the bigger Holzmaden Campylognathoides, C. zitteli (SMNS 9787), was seen.
With a wingspan of three feet, Campylognathoides liasicus (Quenstedt, 1858; formerly known as Pterodactylus liasicus) was smaller than its Holzmaden counterpart, C. zitteli.
Indicus "Campylognathoides" Jain, 1974
Based on a jaw fragment (ISI R. 48) found in the Chanda area of India, Sohan Lal Jain described C. indicus. Kevin Padian believes this to be a nomen dubium, maybe derived from a fossilized fish. Even if it were a pterosaur, the fact that the Kota Formation in which it was discovered has now been redated to the Middle Jurassic or later, seems to rule out any close relationship to Campylognathoides.
It is difficult to distinguish between C. liasicus and C. zitteli. Plieninger only identified the smaller species because he didn't think its fossil was good enough to act as a reference for other specimens. After examining specimen UUPM R157 in 1925, Swedish researcher Carl Wiman came to the conclusion that the two species could be distinguished by a basic morphological difference: C. zitteli has a wing that is proportionately considerably longer than the other. Padian (2008), however, noted that larger animals may have grown abnormally large wings to reduce the wing load as a result of ontogenetic development. There's a chance that other distinctions, such having more teeth in the lower jaw, having a longer snout and nares, having five sacrals rather than four, having perpendicular sacral ribs, and having a longer leg, are also connected to size. Only a continuous development series, as demonstrated in the cases of Rhamphorhynchus and Pterodactylus before, could provide conclusive evidence. Padian maintained his distinction between the two species for the time being, but due to their bigger size and physical similarities, SMNS 51100 and GPIT 24470 were transferred to C. zitteli.
Description
While the snout of this species is very short compared to its contemporary from the same strata, Dorygnathus, the skull is still generally elongated, but considerably lighter constructed. Given that the enormous eye sockets are positioned low in the skull over a small jugal, some experts suggest that Campylognathoides may have had very keen vision or maybe lived a nocturnal existence. The skull's back is flat and comparatively high, with a sharp dip just in front of the eyes. At its final end, the nose curves slightly upward in a narrow point. Long, bony nares cover a significant portion of the nose. The fenestra antorbitalis, a little triangular skull aperture, is located underneath them. In line with the shallower snout, Campylognathoides' teeth are likewise small and devoid of any laniaries or fangs, unlike those of the distinctly heterodont Dorygnathus. Their large base and conical, recurved point are beveled off from the inside, creating a robust and sharp cutting surface. The praemaxilla of the upper jaw contains four very widely spread teeth that gradually get bigger from front to rear; the biggest pair is the fourth pair. Ten smaller teeth in the maxilla, which gradually get smaller posteriorly, are located behind them. In C. liasicus, the lower jaw contains twelve to fourteen teeth, whereas C. zitteli has sixteen to nineteen teeth. Thus, 66 is the maximum total number. Eight cervical vertebrae, fourteen dorsals, four or five sacrals, and up to 38 caudal vertebrae are known to exist, according to a research by Kevin Padian. With around six short vertebrae making up the flexible tail base, the caudals behind them lengthen and become stiffer from extremely long extensions, which enables the tail to act as a rudder. The sternum of Campylognathoides was a relatively big bone plate that was rectangular in shape and had a small crest called a cristospina that faced forward. With a square deltopectoral crest, the upper arm is strong despite its modest length. The hand, which has exceptionally long wing fingers for a basic pterosaur but small metacarpals, with the longest phalanx being the second, accounts for the substantial wing length in the lower arm as well. Pteroids are strong and short. The pelvis is not well understood. In 1986, a fossil collector in a Braunschweig shale quarry discovered a well-preserved Campylognathoides hip. Because the hip socket, according to Peter Wellnhofer, was in an upward lateral position, prohibiting the animal from being able to arrange its legs erectly, as in the case of dinosaurs, birds, and mammals, this pelvis, BSP 1985 I 87, proved to be scientifically noteworthy. This would demonstrate that Campylognathoides was incapable of walking on its hind legs and that it was more likely to have been a quadruped. It has also been established that other "rhamphorhynchoids" (basic pterosaurs) exhibited this gait stance. In contrast, Padian (2009) came to the opposite conclusion, contending that while a quadrupedal gait was feasible, a bipedal manner of movement was a prerequisite for a rapid gait and that an elevated stance was required to place the feet on the ground. This is still a very contentious topic. The feet are little and the leg is fairly short. The fifth toe is very small for a basic pterosaur and is frequently thought to be transporting a membrane between the legs.
Classification
In his subsequent works, Plieninger included Campylognathus within the "Rhamphorhynchoidea" taxon. This suborder only indicates that it was not a pterodactyloid, as it is a paraphyletic group of basal pterosaurs that are not very related. Baron Franz Nopcsa attempted the first definite diagnosis in 1928, placing the genus in the Rhamphorhynchinae subfamily of the Rhamphorhynchidae family. Following an era of minimal research on pterosaur systematics, Oskar Kuhn classified Campylognathoides in 1967 as a member of the Campylognathoidinae subfamily of the Rhamphorhynchidae. Peter Wellnhofer came to the conclusion in 1974 that it belonged below the Rhamphorhynchidae in the evolutionary tree, at a more basic location. The first comprehensive precise cladistic studies were conducted in the early 21st century, which corroborated this. The clade Campylognathoididae was established in 2003 by both David Unwin and Alexander Kellner. According to Unwin, this clade is the sister clade to the Breviquartossa within the Lonchognatha, while according to Kellner, it is the most primitive branch within the Novialoidea. The two places are not significantly different from one another. Based on the findings, it appears that Campylognathoides is closely linked to Eudimorphodon, with which it has similarities in terms of humerus morphology, sternum, and skull. Padian (2009) corroborated this, but he also noted a few primitive traits that are unique to Eudimorphodon and absent from Campylognathoides. In 2010, Brian Andres presented a study demonstrating that Eudimorphodon and Austriadactylus constituted a relatively primitive clade, leaving Campylognathoides as the sole member of the Campylognathoididae that is currently recognized. These findings are supported by more recent phylogenetic studies, which also indicate that Campylognathoides is more derived than all Triassic-aged pterosaurs, the Early Jurassic Dimorphodon, and Parapsicephalus. As a node-based taxon that includes Quetzalcoatlus, the last common ancestor of Campylognathoides, and all of its progeny, Campylognathoides is the most basal member of the Novialoidea.
Paleobiology
Like other pterosaurs, Campylognathoides is traditionally thought to have had a piscivorous lifestyle, which is supported in this instance by the fact that the bones were found in maritime sediments and by the length of its wings. However, pterosaurologist Kevin Padian has hypothesized that the form may have instead been a predator of tiny terrestrial animals because to the strong short teeth, which are perfect for delivering a piercing bite. On the other hand, Mark Witton hypothesizes that the remarkably strong forelimbs of Campylognathoides, which have proportionately large wing fingers, may represent a specialization for a swift aerial lifestyle similar to that of mastiff bats and falcons.
C. liasicus cast.
Campylognathoides Reconstruction.
C. liasicus skull.
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