Eudimorphodon

Eudimorphodon was a pterosaur that was discovered in 1973 by Mario Pandolfi near Bergamo, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic (mid to late Norian stage), making Eudimorphodon the oldest pterosaur then known. It had a wingspan of about 100 cm and at the end of its long bony tail may have been a diamond-shaped flap like in the later Rhamphorhynchus. If so, the flap may have helped it steer while maneuvering in the air. Eudimorphodon is known from several skeletons, including juvenile specimens.

Description
Eudimorphodon was a small pterosaur, especially when compared to the later giant Quetzalcoatlus. Compared to it, Eudimorphodon was only 1 m in length, and only weighed up to 10 kg. Clearly a pterosaur, its fourth finger had a very large size, and attached to the membrane making up the wing.

Dentition and diet
Eudimorphodon showed a strong differentiation of the teeth, hence its name, which is derived from ancient Greek for "true dimorphic tooth". It also possessed a large number of these teeth, a total of 110 of them densely packed into a jaw only six centimeters long. The front of the jaw was filled with fangs, per side four in the upper jaw, two in the lower jaw, that rather abruptly gave way to a line of smaller multipointed teeth, 25 in the upper jaw, 26 in the lower jaw, most of which had five cusps

The morphology of the teeth are suggestive of a piscivorous diet, which has been confirmed by preserved stomach contents containing the remains of fish of the genus Parapholidophorus. Eudimorphodon had slightly differing dentition with fewer teeth and may have had a more insectivorous diet. The top and bottom teeth of Eudimorphodon came into direct contact with each other when the jaws were closed, especially at the back of the jaw. This degree of dental occlusion is the strongest known among pterosaurs. The teeth were multi-cusped, and tooth wear shows that Eudimorphodon was able to crush or chew its food to some degree. Wear along the sides of these teeth suggests that Eudimorphodon also fed on hard-shelled invertebrates. The teeth distinguish Eudimorphodon, because almost all other pterosaurs either had simple teeth, or lacked them altogether. Benson et al. (2012) noticed that the teeth would have been perfect for grabbing and crushing fish.

Phylogeny and classification
Despite its great age, Eudimorphodon has few primitive characteristics making the taxon of little use in attempting to ascertain where pterosaurs fit in the reptile family tree. Basal traits though, are the retention of pterygoid teeth and the flexibility of the tail, which lacks the very long stiffening vertebral extensions other long-tailed pterosaurs possess. Paucity of early pterosaur remains has ensured that their evolutionary origins continues to be a persiseant mystery with some experts suggesting affinities to dinosaurs, archosauriformes, or prolacertiformes.

Within the standard hypothesis that the Dinosauromorpha are the pterosaurs' close relatives within an overarching Ornithodira, Eudimorphodon is also unhelpful in establishing relationships within Pterosauria between early and later forms because then its multicusped teeth should be considered highly derived, compared to the simpler single-cusped teeth of Jurassic pterosaurs, and a strong indicator that Eudimorphodon is not closely related to the ancestor of later pterosaurs. Instead it is believed to be a member of a specialized off branch from the main "line" of pterosaur evolution, the Campylognathoididae.

Discovery and species
Eudimorphodon currently includes two species. The type species, E. ranzii, was first described by Zambelli in 1973. It is based on holotype MCSNB 2888. The specific name honours Professor Silvio Ranzi. A second species, E. rosenfeldi, was named by Dalla Vecchia in 1995 for two specimens found in Italy. However, further study by Dalla Vecchia found that these actually represented a distinct genus, which he named Carniadactylus in 2009. One additional species is still currently recognized: E. cromptonellus, described by Jenkins and colleagues in 2001. It is based on a juvenile specimen with a wing span of just 24 centimeters, MGUH VP 3393, found in the early nineties in Greenland. Its specific name honors Professor Alfred W. Crompton; the name is a diminutive because the exemplar is so small.

In 1986 fossil jaw fragments containing multicusped teeth were found in Dockum Group rocks in western Texas. One fragment, apparently from a lower jaw, contained two teeth, each with five cusps. Another fragment, from an upper jaw, also contained several multi-cusped teeth. These finds are very similar to Eudimorphodon and may be attributable to this genus, although without better fossil remains it is impossible to be sure.

Many fossils have been found, making Eudimorphodon represent one of the most abundant pterosaurs from Italy.