Lepidosaurian epipterygoids in basal pterosaurs

In 1998 lepidosaurian epipterygoids
were found in the basal lepidosaur tritosaur, Huehuecuetzpalli (Fig. 1, Reynoso 1998; slender magenta bones inside the cheek area).

Figure 2. Huehuecuetzpalli has a tall, narrow epipterygoid, as in other lepidosaurs, and just a pore of an antorbital fenestra in the maxilla.

Figure 1. Huehuecuetzpalli has a tall, narrow epipterygoid, as in other lepidosaurs, and just a pore of an antorbital fenestra in the maxilla.

About two years ago
previously overlooked lepidosaurian epipterygoids were identified here in a more derived lepiodaur tritosaur, Macrocnmeus (Fig. 2, slender green bones in the orbit area) for the first time.

Figure 1. Macrocnemus fuyuanensis (GMPKU-P-3001) in situ and as traced by the original authors, (middle) flipped with colors applied to bones, and (above) bone colors moved about to form a reconstruction. Darker yellow and darker green are medial views of premaxilla and maxilla. Note the long ascending process of the premaxilla and the palatal elements seen through the various openings all overlooked by those with firsthand access to the fossil. Epipterygoids are lepidosaur synapomorphies not present in protorosaurs.

Figure 2. Macrocnemus fuyuanensis (GMPKU-P-3001) in situ and as traced by the original authors, (middle) flipped with colors applied to bones, and (above) bone colors moved about to form a reconstruction. Darker yellow and darker green are medial views of premaxilla and maxilla. Note the long ascending process of the premaxilla and the palatal elements seen through the various openings all overlooked by those with firsthand access to the fossil. Epipterygoids are lepidosaur synapomorphies not present in protorosaurs.

Until now,
no one has ever positively identified lepidosaurian (slender strut-like) epipterygoids in a pterosaur. In the large reptile tree (LRT, 1737+ taxa) and the large pterosaur tree (LPT, 251 taxa) Bergamodactylus (MPUM 6009) nests as the basalmost pterosaur. Here is the skull in situ with DGS colors applied, as traced by Wild 1978 (above), and reconstructed in lateral and palatal views (below) based on the DGS tracings.

Figure 3. Bergamodactylus skull in situ and reconstructed. Wild 1978 tracing above.

Figure 3. Bergamodactylus skull in situ and reconstructed. Wild 1978 tracing above. Note the break-up of the jugal. Note the fusion of the ectopterygoids with the palatines producing ectopalaatines.

The lepidosaurian epipterygoids of Bergamodactylus
(slender bright green struts in the cheek/orbit area in figure 3), or any pterosaur over the last 200 years, are identified here for the first time, further confirming the lepidosaurian status of pterosaurs (Peters 2007, the LRT). Sorry I missed these little struts earlier. When you don’t think to look for them, you can overlook them.

Figure 5. Eudimorphodon epipterygoids (slender green struts).

Figure 4. Eudimorphodon epipterygoids (slender green struts).

Now you may wonder how many other pterosaurs
have overlooked epipterygoids? A quick look at Eudimorphodon reveals epipterygoids (Fig. 4, bright green struts). Other Triassic pterosaurs include:

  1. Austriadactylus SMNS 56342: slender strut present
  2. Austriadactuylus SC 332466: slender strut present
  3. Raeticodactylus : slender strut is present (identified on link as a stapes)
  4. Preondactylus: slender strut present
  5. Dimorphodon: amber strut over squamosal (Fig. 5 in situ image), 
  6. Seazzadactylus MFSN 21545: slender struts present, tentatively identified by Dalla Vecchia 2019, but as more than the slender struts they are) (Fig. 6).
The skull of Dimorphodon macronyx BMNH 41212.

Figure 5. The skull of Dimorphodon macronyx BMNH 41212. Above: in situ. Middle: Restored. Below: Palatal view. The slender yellow strut on top of the red squamosal in situ is a likely epipterygoid.

Figure 6. Seazzadactylus from Dalla Vecchia 2019. Here the epipterygoid struts are more correctly and less tentatively identified.

Figure 6. Seazzadactylus from Dalla Vecchia 2019. Here the epipterygoid struts are more correctly and less tentatively identified.

Hard to tell in anurognathids
where everything is crushed and strut-like. Hard to tell in other pterosaurs because the hyoids look just like epipterygoids. Given more time perhaps more examples will be documented that are obvious and irrefutable.

Added a few days later:

Added Figure. Here's the Triebold specimen of Pteranodon (NMC41-358) with epipterygoid splinters in bright green.

Added Figure. Here’s the Triebold specimen of Pteranodon (NMC41-358) with epipterygoid splinters in bright green.

Here’s the Triebold specimen of Pteranodon
(NMC41-358, added figure) with epipterygoid splinters in bright green. So start looking for the epipterygoid in every pterosaur. We’ll see if it is universal when more pterosaur specimens of all sorts are presented.


References
Dalla Vecchia FM 2019. Seazzadactylus venieri gen. et sp. nov., a new pterosaur (Diapsida: Pterosauria) from the Upper Triassic (Norian) of northeastern Italy. PeerJ 7:e7363 DOI 10.7717/peerj.7363
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.
Wild R 1978. Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen Trias von Cene bei Bergamo, Italien. Bolletino della Societa Paleontologica Italiana 17(2): 176–256.

wiki/Bergamodactylus
wiki/Huehuecuetzpalli
wiki/Homoeosaurus
wiki/Bavarisaurus

Leave a Reply

Fill in your details below or click an icon to log in:

WordPress.com Logo

You are commenting using your WordPress.com account. Log Out /  Change )

Google photo

You are commenting using your Google account. Log Out /  Change )

Twitter picture

You are commenting using your Twitter account. Log Out /  Change )

Facebook photo

You are commenting using your Facebook account. Log Out /  Change )

Connecting to %s

This site uses Akismet to reduce spam. Learn how your comment data is processed.