The “first Quetzacoatlus”
Pterodactyloids are the pterosaur version of the theropods: they were the spotlight stealing squad. And frankly, it’s no wonder: nearly all of the most spectacular flying animals of all time are pterodactyloids. Which pterosaurs reached the biggest sizes? Ornithocheiroids and azhdarchoids. Which pterosaurs competed directly with theropod dinosaurs to the point of almost prevalence in some cases? Lophocratines (dsungaripteroids and azhdarchoids FYI). Which pterosaurs rivalled modern birds in terms of ecological niches? Pterodactyloidea as a whole, from vulture-like istiodactylids to eagle-like ornithocheirids to petrel-like nyctosaurids to pelican-like boreopterids to duck-like ctenochasmatids to flamingo-like Pterodaustro to woodcock-like Pterodactylus to boar/bear-like dsungaripteroids to terror bird-like thalassodromedids to antelope/ratite-like tapejarids and many more. Which pterosaurs were the last (and therefore the most attention grabbing, as most people only care about the terminal Cretaceous)? Pterodactyloids, of course, though anurognathids might had survived unpreserved.
However, just as very interesting things have been noted amidst non-theropod dinosaurs, the same can be said for non-pterodactyloid pterosaurs. The so called “rhamphorhynchoids” (a purely vernacular term, unless you include pterodactyloids as “rhamphorhynchoids”), while far less morphologically aberrant than the “it’s hard to believe these things are sauropsids” pterodactyloids, and certainly nowhere as impressive size wise, were certainly spectacular animals on their own right. They were the first vertebrates ever to fly, and pioneered many adaptations and niches long before pterodactyloids, birds and bats evolved, and in some ways they still did them better than their successors.
Among the most impressive of all of the non-pterodactyloid pterosaurs were the campylognathoidids. The exact relationship these pterosaurs had with other early flyers is unknown; it has been suggested synonimy with Eudimorphodontidae (which would make them among the oldest known pterosaurs), though most modern studies seem to ally them with the Rhamphorhynchidae + Monofenestrata clade, known as Novialoidea. At any rate, this clade appears to have been a short lived success; unless eudimorphodontids are early campylognathoidids, which would make theirs the first pterosaur dynasty, dominating the Triassic skies (and, most incredibly, making their reign uphased by the Triassic extinctions), these pterosaurs dominated pterosaur faunas during the Early Jurassic, being the most common pterosaur fossils from this time period, but they rapidly declined in the Middle Jurassic and were quickly replaced by rhamphorhynchids as the dominant pterosaurs. Some species might have survived as recently as the Late Jurassic or even the Cretaceous, but their golden days were long over, and were henceforth non-chalantly replaced by the then newly emerged ornithocheirids.
When they were alive, however, they were the most magnificent flying animals to have ever existed, and their imediate successors, the rhamphorhynchids, paled in comparision. Campylognathoides itself was the largest non-pterodactyloid pterosaur ever to fly; on average, adult specimens had a wingspan of 2.7 meters, and exceptionally large specimens had wingspans up to 3.5 meters. Compared to posterior pterosaurs, this is outrightly miniscule, but by the standards of modern birds and bats, Campylognathoides was a pretty much a giant, comparable in size to modern albatrosses and condors. The closest contenders in size were the very largest rhamphorhynchids, with wingspans of up to 2.3 meters.
Equally interesting is it’s ecological niche. Like other giant pterosaurs, Campylognathoides was historically assumed to have been nothing more than a giant bulldog-bat, cruising the warm seas of it’s era in search on fish it caught on the wing. However, recent studies of it’s jaw morphology show that hunting fish from the air was at best very difficult, and more likely borderline impossible. On the other hand, it’s short, conical jaws and piercing teeth were quite apropriate for a terrestrial predator, for delivering bone crushing bites to small lepidosaurs, mammals and perhaps the unlucky young dinosaur. This is futher supported by it’s rarity on marine deposits and more common finds on inland areas, while the reverse occurs with piscivorous pterosaurs like Dorygnathus. Furthermore, much like pterodactyloid pterosaurs, Campylognathoides had an erect gait, allowing it to walk and run efficiently on the ground, much like a mammal.
In short, with it’s large size, efficient terrestrial locomotion and raptorial habits, Campylognathoides was the azhdarchid prototype, haunting the Mesozoic plains long before pterodactyloids evolved. So early in the evolution of flying vertebrates, and pterosaurs already produced the archetypical King of the Skies.