2024 pterosaur palate study focuses again on Dsungaripterus

Chen et al 2024 wrote:
“Among the least studied portion of the pterosaur skeleton is the palate,”

Actually an 8-part study of pterosaur palates (Fig 1) was published here in 2012.
This study revisited the same Dsungaripterus palate described by Chen et al 2020, plus several others for comparison this time around.

Actually there are many parts of the pterosaur skeleton that have received too little attention. e.g. The tip of the Dsungaripterus snout is a single overlooked tooth (Fig 2).

“To shed new light on this region, we employed advanced X-ray imaging techniques on the non-pterodactyloid Kunpengopterus (Wukongopteridae), and the pterodactyloids Dsungaripterus (Dsungaripteridae), Hongshanopterus (Istiodactylidae), and Hamipterus (Hamipteridae).”

Pterodactyloidea is paraphyletic and has been since Peters 2007. So this study is out-of-dating itself by continuing the myth of the clade ‘Pterodactyloidea’.

Figure 1. The palates of several Tapejaridae and Pteranodontidae, both evolving from Germananodactylus. ” data-image-caption=”

Figure 1. The palates of several Tapejaridae and Pteranodontidae, both evolving from Germananodactylus.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=227″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=584″ class=”size-full wp-image-13028″ src=”https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=584&h=773″ alt=”Figure 1. The palates of several Tapejaridae and Pteranodontidae, both evolving from Germananodactylus.” width=”584″ height=”773″ srcset=”https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=584&h=773 584w, https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=113&h=150 113w, https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg?w=227&h=300 227w, https://pterosaurheresies.files.wordpress.com/2012/03/tapejarid-palates5882.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 1. The palates of several Tapejaridae and Pteranodontidae, both evolving from Germananodactylus from 2012. Note the variation while keeping certain patterns.

Chen et al continue,
“We also point out that the presence of a maxillopalatine fenestra (previously identified as postpalatine fenestra), is unique within Diapsida.”

1. The clade Diapsida has been paraphyletic for over a decade.
2. The ‘maxillopalatine fenestra’ identified by Chen et al is actually one of two internal nares (= choana) split apart by the medial migration of the palatal process of the maxilla.

Pterosaurs and their nonvolant fenestrasaur kin
have two internal nares (Figs 2–4). That’s the tricky part understood only after accepting the fenestrasaur-tanystropheid-lepidosaur origin of pterosaurs. Unfortunately for paleontology that hypothesis remains unacceptable to current workers more than two decades after publication (Peters 2000, 2007).

Peters 2000 was cited in Chen et al 2024, but not otherwise mentioned.

Figure 2. The palates of Macrocnemus, Rhamphorhynchus and Dsungaripterus, plus a drawing of the open mouth of Iguana showning the lateral internal nares (choanae). Note the evolution of the dual internal nares. ” data-image-caption=”

Figure 2. The palates of Macrocnemus, Rhamphorhynchus and Dsungaripterus, plus a drawing of the open mouth of Iguana showning the lateral internal nares (choanae). Note the evolution of the dual internal nares.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=300″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=584″ class=”size-full wp-image-85346″ src=”https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=584&h=569″ alt=”Figure 2. The palates of Macrocnemus, Rhamphorhynchus and Dsungaripterus, plus a drawing of the open mouth of Iguana showning the lateral internal nares (choanae). Note the evolution of the dual internal nares.” width=”584″ height=”569″ srcset=”https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=584&h=569 584w, https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=150&h=146 150w, https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif?w=300&h=292 300w, https://pterosaurheresies.files.wordpress.com/2024/04/macrocnemus_palate_kuhn588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 2. The palates of Macrocnemus, Rhamphorhynchus and Dsungaripterus, plus a drawing of the open mouth of Iguana showning the lateral internal nares (choanae). Note the evolution of the dual internal nares. Frame 2 restores the missing vomer and pterygoid only to enhance understanding the palate morphology. Note the fusion of the ectopterygoid and pterygoid in Rhamp;horhynchus.

On this point, Chen et al wrote,
“Although an extensive comparison of the pterosaur palate with other diapsids is beyond the scope of this paper, there are a few comments and  differences that can be highlighted. The most significant is the presence of a maxillopalatine fenestra, which is absent in other diapsids and might turn out to be a synapomorphy of Pterosauria.

As mentioned earlier, that ‘fenestra’ is the internal naris, lateral, as in lepidosaurs.

“In prolacertiformes and lepidosauromopha, the internal naris is positioned at the lateral side of the palate, followed by a postpalatine fenestra, and lacks any extra opening between them. But the postpalatine fenestra is similar to the one observed in pterosaurs, since it is formed anteriorly by the palatine and posteriorly by ectopterygoid.”

At least they mentioned lepidosaurormorphs.
In the LRT, after adding pertinent taxa, pterosaurs are lepidosaurs (Peters 2007).

Figure 3. Skull and palate of Cosesaurus, Oculudentavis and Longisquama. These are nonvolant pterosaur relatives. ” data-image-caption=”

Figure 3. Skull and palate of Cosesaurus, Oculudentavis and Longisquama. These are nonvolant pterosaur relatives.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=291″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=584″ class=”size-full wp-image-85347″ src=”https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=584&h=603″ alt=”Figure 3. Skull and palate of Cosesaurus, Oculudentavis and Longisquama. These are nonvolant pterosaur relatives.” width=”584″ height=”603″ srcset=”https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=584&h=603 584w, https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=145&h=150 145w, https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg?w=291&h=300 291w, https://pterosaurheresies.files.wordpress.com/2024/04/cosesaurus_longisquama_oculudentavis.palate588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 3. Skull and palate of Cosesaurus, Oculudentavis and Longisquama. These are nonvolant pterosaur relatives.

Peters 2000 was NOT the first to correctly identify
the anterior palate of pterosaurs was maxillary in origin. Recent prior workers (e.g. Wellnhofer 1978, 1991, Bennett 1991, 2001a,b) identified that plate as the palatine. It was correctly identified a century earlier by Newton 1888 and Seeley 1901. Unaware of those older citations, Peters 2000 compared Macrocnemus (Fig 1) to Cosesaurus (Fig 3) and Rhamphorhynchus (Fig 1). Ten years later, Osi, Frey and Pohl 2010 proclaimed a ‘new interpretation of the palate of pterosaurs’, confirming the interpretations of Newton 1888, Seeley 1901 and Peters 2000.

Figure 3. Germanodactylus, Phobetor and Dsungaripterus palates compared. ” data-image-caption=”

Figure 3. Germanodactylus, Phobetor and Dsungaripterus palates compared.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=123″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=419″ class=”size-full wp-image-85382″ src=”https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=584&h=1428″ alt=”Figure 3. Germanodactylus, Phobetor and Dsungaripterus palates compared.” width=”584″ height=”1428″ srcset=”https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=584&h=1428 584w, https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=61&h=150 61w, https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg?w=123&h=300 123w, https://pterosaurheresies.files.wordpress.com/2024/04/germanodactylus-dsungaripterids.skull-recon2-1.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 4. Germanodactylus, Phobetor and Dsungaripterus palates compared.

Chen et al continue,
“Controversy persists, particularly concerning the extent and shape of the palatines and whether a single, generalized palatal pattern configuration can be applied to Pterodactyloidea.”

This where a misunderstanding of pterosaur phylogeny starts to confuse these authors. A valid phylogeny must be the first step. Just add taxa to minimize taxon exclusion. Chen et al still do not accept the fenestrasaur, tanystropheid, lepidosaur origin of pterosaurs (Peters 2000, 2007), which hampers and hobbles their paper.

Chen et al continue,
“Regarding pterosaur phylogenies, there are several proposals/hypothesis published. Here we have followed mainly Kellner et al. 2019.”

The phylogenetic portion of Chen et al 2024 (Fig 5), derived from Kellner 2003, suffered greatly from cherry-picking = taxon exclusion. Unfortunately, adding pertinent taxa is not on the to-do list of any academics at this point. Perhaps this is due to the online presence of the large pterosaur tree (LPT, 269 taxa) and Peters 2007 which invalidated earlier studies by simply adding traditionally omitted taxa.

Unfortunately academic workers too often turn a blind eye to competing studies. No one wants to be wrong or shown the error of their methods and taxon lists. That’s human nature. The scientific method requires criticism from competing methods.

Figure 5. Chen et al 2024 cladogram. Note they don’t group the toothless pterosaurs together and lack about 260 taxa, also excluding all nonvolant outgroups. The LPT does not confirm the Monofenestrata and Pterodactyloidea. Rather the pterodactyloid grade was achieved five times by convergence and neotony. ” data-image-caption=”

Figure 5. Chen et al 2024 cladogram. Note they don’t group the toothless pterosaurs together and lack about 260 taxa, also excluding all nonvolant outgroups. The LPT does not confirm the Monofenestrata and Pterodactyloidea. Rather the pterodactyloid grade was achieved five times by convergence and neotony.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=126″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=429″ class=”size-full wp-image-85385″ src=”https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=584&h=1393″ alt=”Figure 5. Chen et al 2024 cladogram. Note they don’t group the toothless pterosaurs together and lack about 260 taxa, also excluding all nonvolant outgroups. The LPT does not confirm the Monofenestrata and Pterodactyloidea. Rather the pterodactyloid grade was achieved five times by convergence and neotony.” width=”584″ height=”1393″ srcset=”https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=584&h=1393 584w, https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=63&h=150 63w, https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif?w=126&h=300 126w, https://pterosaurheresies.files.wordpress.com/2024/04/chenetal2024cladogram588-1.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 5. Chen et al 2024 cladogram. Note they don’t group the toothless pterosaurs together and lack about 260 taxa, also excluding all nonvolant outgroups. The LPT does not confirm the Monofenestrata and Pterodactyloidea with 260+ taxa. Rather the pterodactyloid grade was achieved five times by convergence and neotony. Add taxa to your own pterosaur cladogram to see for yourself.

Chen et al 2024 concluded:
“1. In some non-pterodactyloids (e.g., Rhamphorhynchus and Cacibupteryx),
the anterior process of the pterygoids is not as developed as it is in pterodactyloids.”

Actually just the opposite is true. The anterior pterygoids are lost of reduced to nubbins at the back of the palate in derived pterosaurs.

“2. The angle between the two rami of the palatine in some nonpterodactyloids are nearly 90°, in Kunpengopterus is 45°, and in pterodactyloids are <60° (e.g., ~30° in Dsungaripterus, ~60° in Hamipterus).”

According to their figures 4 and 6, the palatine angle is similar-to-identical in Rhamphorhynchus, Gnathosaurus and Hamipterus. The authors don’t realize that palatine angle shrinks to 0º in Pteranodon, as we learned yesterday.

“3. In pterodactyloids the jugal process of the maxilla extends posteriorly until the ectopterygoid, but not in the non-pterodactyloids (Rhamphorhynchus and Cacibupteryx).”

Actually the pterodactyloid-grade was achieved 5x in pterosaurs. The authors are not aware of this due to taxon exclusion in their cladograms. So the extent of this process has to recalibrated to reflect this reality.

“4. The last tooth is posterior to the anterior margin of maxillopalatine fenestrae in non-pterodactyloids (e.g., Dorygnathus, Cacibupteryx, Rhamphorhynchus, and Kunpengopterus), but is anterior to the maxillopalatine fenestrae in toothed pterodactyloids
(e.g., Hongshanopterus, Anhanguera, Tropeognathus, Liaodactylus, Gnathosaurus, Aurorazhdarcho, Hamipterus, and Dsungaripterus).”

Actually the pterodactyloid-grade was achieved 5x in pterosaurs. Revisit this assertion.

“5. Pterodactyloids have a pair of pterygoid openings (foramen or fenestra) bordered by the lateral ramus of the palatine and the median process of pterygoid. This opening is present in the nonpterodactyloid Kunpengopterus and Cacibupteryx, but not in
Rhamphorhynchus.”

Actually the pterodactyloid-grade was achieved 5x in pterosaurs. Revisit this assertion.

“6. Along with the great variation in the sizes of the palatal openings such as the pterygoid foramen or fenestra, the posterior margin of the choanae moved posteriorly to the maxillopalatine fenestra in Kunpengopterus and pterodactyloids and even posterior to the
postpalatine fenestra in some pterodactyloids (e.g., Caupedactylus, Tupuxuara, and Dsungaripterus).”

Actually the pterodactyloid-grade was achieved 5x in pterosaurs. Revisit this assertion.

Figure 6. The palate of the CM1143 specimen of Rhamphorhynchus. ” data-image-caption=”

Figure 6. The palate of the CM1143 specimen of Rhamphorhynchus.

” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=182″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=584″ class=”size-full wp-image-85390″ src=”https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=584&h=964″ alt=”Figure 6. The palate of the CM1143 specimen of Rhamphorhynchus. ” width=”584″ height=”964″ srcset=”https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=584&h=964 584w, https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=91&h=150 91w, https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif?w=182&h=300 182w, https://pterosaurheresies.files.wordpress.com/2024/04/rhamphorhynchus.cm11434-588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 6. The palate of the CM1143 specimen of Rhamphorhynchus.Those openings borderd by the blue palatines are lateral portions of the dual choanae = internal nares. See Macrocnemus in figure 1. Note the anterior placement of the anterior pterygoid tips. Dsungargipterus lacks these processes.

The Chen et al 2024 paper provided a great opportunity
to review pterosaur and fenestrasaur palates, always an intriguing subject. On the other hand, I hate to see pterosaur workers wading through a dark age of their own making.

References
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Chen et al. (7 co-authors) 2020. New anatomical information on Dsungaripterus weii Young, 1964 with focus on the palatal region. PeerJ 8:e8741 DOI 10.7717/peerj.8741
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Peters D 2007. The origin and radiation of the Pterosauria. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27.
Pinheiro FL, Schultz CL 2012. An Unusual Pterosaur Specimen (Pterodactyloidea, ?Azhdarchoidea) from the Early Cretaceous Romualdo Formation of Brazil, and the Evolution of the Pterodactyloid Palate. PLoS ONE 7(11): e50088. doi:10.1371/journal.pone.0050088
Seeley HG 1901. Dragons of the air. An account of extinct flying reptiles. – London, Methuen: 1-240.
Wellnhofer P 1978. Pterosauria. Handbuch der Paläoherpetologie, Teil 19.– Stuttgart, Gustav Fischer Verlag: 1-82.
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Reconstructing the palate of Pteranodon in 1910, 1993 and 2024

Dsungaripterus palate news: Chen et al. 2020

The Pterosaur Palate: More workers coming on board.

Evolution of the Pterosaur Palate – part 8: Tapejaridae and Pteranodontidae