Anhanguera

Credit: Matt Martyniuk.

*Anhanguera* is an extinct genus of anhanguerine pterosaur that lived in Brazil from the Early to Late Cretaceous.

## Description

*Anhanguera* was piscivorous. Rounded crests on the anterior of the jaws were present, as in all anhanguerids, with conical, curved teeth of many sizes and orientations. The jaws taper in width, expanding in a spoon-shaped rosette at the tip. Small differences in the crest and teeth. The crest of the upper jaw is set farther down the skull. Like pteranodonts, another crest sits at the back of the skull. However, it is small and blunt. A 2003 study finds *Anhanguera* held it's head downwards due to its inner ear, which it used to detect balance. Their eyes are supported by sclerotic rings. *A. araripensis*, a dubious species, is described as similar to other *Anhanguera* and possibly *Coloborhynchus*, of which it has a complicated history in both. The rostral tip is blunt and is adorned with 2 forwards-projecting teeth that sit higher in the jaw. It still has the jaw crests. Like *Coloborhynchus* and unlike the type Anhanguera sp., the crest emerges at the tip of the jaws. Unlike *Coloborhynchus*, however, it lacks a dent in the jaw tip, with the teeth appearing smaller and with a more uniform size.

*A. bittersdorfi* is known from a complete skull from the Santana Group, Romualdo Formation's calcareous concretions. A. piscator is known from a near-complete skeleton, and has been proposed to be *Coloborhynchus*, with Andres and Myers (2013) referring it to *Anhanguera. A. spielbergi* is known from a complete skeleton, originally assigned *Coloborhynchus*, but has been reassigned to *Maaradactylus*.

AMNH 22555 may represent a new *Anhanguera* species, being previously assigned to *A. "santanae"*. AMNH 22555 is known from remains persevered in 3D, this includes bones from all of the upper jaw, a complete braincase, a fragment of the rear bottom jaw, most of the vertebral column, The shoulder girdle, as well as fragmentary pieces of the hips, legs, wing, and pteroid. AMNH notes that the specimen was preserved in a hard concretion. It is likely a new species. AMNH 22555, along with invertebrates and fishes, were illegally exported from Brazil to the American Museum of Natural History by Herbert Axelrod in the 1980s. The publications mentioning this specimen neglect the date of acquiring, which is likely purposeful.

## Classification

Andres and Myers (2013) placed *Anhanguera* into pteranodontia like so:

## Species

* *A. araripensis*? (=Santanadactylus araripensis?, =Coloborhynchus?)
* *A. piscator*
* *A. robustus*? (=*Tropeognathus robustus*, =*Coloborhynchus*)
* A. santanae? (=*Araripesaurus*?)
* *A. spielbergi*?
* *A. n. sp.*?

### Reassigned Species

* *A. cuvieri* (=*Ornithocheirus cuvieri*, =*Pterodactylus cuvieri*, =*Cimoliopterus*)
* *A. fittoni*
* *A. ligabuei* (=*Cearadactylus ligabuei*, =*Coloborhynchus* or *Cearadactylus*)

## Synonyms

* "Pricesaurus"
* *Santanadactylus araripensis* (=*A. araripensis*)
* *Coloborhynchus araripensis* (=*A. araripensis*)
* *Coloborhynchus piscator* (=*A. piscator*)
* *Coloborhynchus robustus* (=*A. robustus*)
* *Tropeognathus robustus* (=*A. robustus*)
* *Coloborhynchus spielbergi* (=*A. spielbergi*)
* *Maaradactylus spielbergi*?

## Paleoecology

The Romualdo formation represents a very humid equatorial climate like the older Crato formation. Although no remains of soft plant matter or insects are found directly in the formation a concept of the flora can be made from the Crato formation. AMNH 22555 would have shared it's environment with the Spinosaurid *Irritator*, the Tyrannosauroid *Santanaraptor*, the basal Coelurosaur *Aratasaurus*, and the Compsognathid *Mirischia*. It would have also shared it's environment with pterosaurs like *Tapejara*, *Thalassodromeus*, *Tropeognathus*, *Tupuxuara*, *Unwindia*, *Santanadactylus*, *Maaradactylus*, *Cearadactylus*, *Barbosania*, the Notosuchid *Araripesuchus*, and the Turtles *Santanachelys*, *Cearachelys*, and *Araripemys*, as well as 19 species of fossil fishes.

## References

* https://en.wikipedia.org/wiki/Romualdo\_Formation
* https://en.wikipedia.org/wiki/Anhanguera\_(pterosaur)
* AMNH 22555 plaque.
* Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control? PeerJ 5:e3285, doi:10.7717/peerj.3285
* https://twitter.com/TylerGreenfieId/status/1437579603594383364