The “pterosaur dark ages” are not over

Professor David Martill (PhD) and his student, Roy E Smith (PhD),
pre-published an accepted manuscript that included the term “pterosaur dark ages.” The authors were lamenting the 1930s through 60s when AS Romer was publishing his university-level textbook ‘Vertebrate Paleontology’ in three editions (1933, 1945, 1966). Martill and Smith 2024 continued, “How times have changed! Pterosaurs, volant reptiles of the Mesozoic, are now among the best studied of any Mesozoic vertebrate group, including the Dinosauria.”

This statement is false. The “pterosaur dark ages” continue in 2024.

Pterosaur misinformation and myth continues
in academia largely due to taxon omission, a curious lack of curiosity, vague reconstructions and a gentlemans’ agreement not to discuss certain topics. So decades after Romer pterosaur workers are still wearing blinders, a topic discussed here in 2018. Wing shape, launch technique, giant azhdarchid flight, the origin and closest relatives of pterosaurs, what to do with pedal digit 5, what to do with the pteroid and its origin, reconstructing stance, phylogenetic interrelationships, phylogenetic miniaturiztion, gender, crests, clades, eggs… all these remain stuck in the mire of Wellnhofer’s coffee table book ‘The illustrated Encyclopedia of Pterosaurs’ (1991), praised by Martill and Smith.

Martill and Smith wrote,
“Pterosaurology’s true reawakening did not occur until a young German worker named Peter Wellnhofer (who) went on to become the world’s leading expert on pterosaurs for around four decades.”

In fact, Wellnhofer cataloged and measured specimens without understanding that pterosaurs were lepidosaurs, had a narrow wing chord, arose from flapping sprinting bipeds that left matching footprints, and phylogenetically achieved the pterodactyloid grade five times by convergence.

Martill and Smith also wrote,
“Following Wellnhofer, four young students, Christopher Bennett (USA), Alex Kellner (Brazil), Kevin Padian (USA) and David Unwin (England) studied pterosaurs for the PhDs. Hot on their heels, Dino Frey (Germany) switched allegiances from crocodiles to pterosaurs and entered the ‘game’.”

For the record, these were not Wellnhofer’s students. That means these four aging professors studied pterosaurs in their youth pretty much on their own, without an expert to guide them.

For the record, that means you, too, can do what they did.
You have the same access to libraries, collections, symposia and PDFs.

Now let’s review
the individual contributions to pterosaurs made by these five workers.

Bennett separated Pteranodon specimens into genders
based on statistics, not realizing the small ones were phylogenetically more primitive, closer to smaller germanodactylids with smaller crests. Bennett did not realize the big-opening pelvis he assigned to a female Pteranodon actually belonged to a large nyctosaurid. More recently Bennett mistook an anurgnathid maxilla for a giant eyeball (= scleral ring) in the anterior of the crushed skull. These are both widely accepted hypotheses in the literature now. When conducting his second phylogenetic analysis of pterosaurs Bennett omitted Cosesaurus, Longisquama and Sharovipteryx years after they were shown to be pterosaur outgroups. Instead Bennett restricted his taxon list to archosauromorphs because he knew he could get away with it under the current ‘gentlemans’ agreement’.

Bennett’s infamous curse “You will not be published, and if you are published you will not be cited,” was adhered to in Martill and Smith 2024. Pterosaur workers used to commonly cite works by yours truly, but they don’t do that anymore.

Padian correctly considered Dimorphodon a biped,
and so refused to accept that quadrupedal pterosaur tracks were made by pterosaurs. That is, until the famous Crayssac tracks changed his mind – but non-French tracks remained non-pterosaurian in Padian’s hypothesis.

Kellner benefited from his location in Brazil
as 3D Early Cretaceous pterosaurs began to pop out of Brazilian matrices. He is still hoping for an archosaur origin for pterosaurs, starting with a long wing finger, hopefully arising from his 2003 outgroup taxa: Lagerpeton, Marasuchus, Scleromochlus, Herrerasaurus and Ornithosuchus. After several more-inclusive phylogenetic analyses we can demonstrate that’s now how it happened. The wings came last, as in birds.

Unwin championed the false deep-chord pterosaur wing membrane
hypothesis by misinterpreting Sordes, a Russian pterosaur brought to his attention by his Russian co-author and wife. Similarly Unwin had access to and wrote about Sharovipteryx and Longisquama, but failed to see their interrelationship with pterosaurs. Instead his phyogenetic analysis favored an all zero outgroup. Unwin also imagined pterosaurs buried their eggs in some sort of matrix, never considering that as lepidosaurs pterosaur mothers could carry their eggs in utero to full term.

Frey championed the false anteriorly-directed pteroid
hypothesis and co-authored papers that championed the false deep chord pterosaur wing membrane attached to the ankles. There is no evidence for that, but plenty for the narrow chord design. To his credit, Frey noticed most pterosaurs had a more dorsal shoulder joint, but a few had a more ventral shoulder joint aligned with the sternum.

Oddly missing from this list of influential pterosaur workers
is Rupert Wild, a contemporary of Peter Wellnhofer. Wild described all the Triassic pterosaurs and elucidated the three-part sternal complex, still unrecognized and appreciated by today’s pterosaur workers.

Oddly, ironically, while R Wild is missing from the Martill and Smith citation list,
fiction writer, Jules Verne, is included.

In summary,
Martill and Smith 2024 is a cheery review of cherry-picked literaturegoing back to the 1700s. The co-authors attempted to confine their review to Cretaceous pterosaurs. Their understanding of phylogeny perpetuates old myths based on taxon exclusion. Citation omissions also weaken this report. There is nothing new here. Most importantly, there are no challenges to any old myths still simmering in the current ‘pterosaur dark ages’.

Instead, Martill and Smith continue their gentlemans’ agreement
not to challenge each other, not to create or confront challenging hypotheses,
and not to include challenging authors in their citations.

References
Martill D and Smith RE 2024 (accepted ms). Cretaceous pterosaur history, diversity and extinction. Geological Society, London, Special Publications https://doi.org/10.1144/SP544-2023-126