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This is taken from the 2000 book Walking with Dinosaurs the Evidence, written by Darren Naish and David Martill, a tie-in to 1999’s Walking with Dinosaurs, which explains some of the thought processes behind the (now very dated) portrayals of the animals included in that series. I’ve posted this for the sake of offering some easily accessible information regarding some of the most controversial creative choices taken in the series.

So the WW series certainly suffered from the law of diminishing returns, with 1999’s Walking with Dinosaurs attracting 18.91-15.69 million viewers (depending on the episode), 2001’s Walking with Beasts only 13.99-5.87 million (a more notable dip as it went on compared to WWD), 2002’s “The Giant Claw” about 6 million and “Land of Giants” 5.76 million, 2003’s Sea Monsters 7.59-6.52 million, and lastly, and 2005’s Walking with Monsters a meager 4.57-4.23 million, and likewise, the budget kept getting slashed from 6 million for WWD to 4 million for WWB, and finally only 3 million for Sea Monsters and WWM, explaining why the latter instalments were shorter, and likely also why they got less supplementary material (WWM didn’t even get its own companion book), and why there were no equivalents to 2000’s Walking with Dinosaurs: The Evidence, which was much more informative than any later book tied to the WW franchise.

Which is really annoying, because frankly, I’m dying to know just what their justification for describing Pteranodon as a near-cosmopolitan genus that lasted almost the entirety of the Cretaceous was , because I can’t find any other source that describes it as anything other than a Late Cretaceous North American genus, maybe offhandedly referencing possible contemporary material from Japan and Europe, but that’s about it. Pteranodon isn’t alone in this though, as the 2005 book describes Ornithocheirus in much the same vein, as a near-cosmopolitan genus that lasted the entirety of the Cretaceous , and “Land of Giants” shows the same gigantic species appearing 27 million years after its debut in “Giant of the Skies”, and Sea Monsters, likewise, has the exact same Pteranodon show up 25 million years after its debut in “Land of Giants”.

Again, it’s important to stress that endothermic, highly active tetrapods such as pterosaurs simply aren’t made to remain stagnant in an ever-changing world for more than 5 million years at best, being too specialized to survive unless they subtly change over millennia and inevitably evolve into new species, new genera, and eventually wholly new types, and 25 million years is a whole lot of time for them to evolve into something completely unrecognizable compared to its progenitor. While wastebasket taxonomy is evident here, in this case of Ornithocheirus, it’s easy to infer why they described it as such, because Ornithocheirus was a notorious wastebasket taxon, the pterosaur equivalent of Iguanodon, with both originally getting described from very fragmentary material found in Britain during the 19th century.

As is well known (but criminally underdiscussed), “Land of Giants” put Pteranodon in Argentina 100 million years ago (very early Cenomanian), and it actually crosses paths with the giant Ornithocheirus (Tropeognathus) cameo, as we briefly see them sharing the sky, and later, in the final segment of Sea Monsters, we see Pteranodon in North America 75 mya (upper Campanian), which is still anachronistic (the youngest fossils are about 5 million years older) but at least they put it in its proper biome (the Niobrara Chalk) with animals that it would have actually encountered in life, though according to the 2005 book, Ornithocheirus could just as well have made a cameo here.

As I said, the latter is easier to explain, as Ornithocheirus simus (the type species) was first described from very fragmentary remains from the upper Albian Cambridge Greensand in England and subsequently, the genus became a dumping ground for various ornithocheirid fossils (the group would more accurately be called anhanguerians today) from all across the globe, and as is usually the case with wastebasket taxa, which pterosaur fossils counted as Ornithocheirus would depend on who you asked, though like with Iguanodon, the bulk of the assigned fossils came from the Lower Cretaceous of England. The 2005 book seems to take a very generous definition, and gives Ornithocheirus a fossil range of 140-70 mya, and says that it inhabited Europe, South America, Africa and...Australia?

The 2000 book does offer some context to it. A number of isolated pterosaur bones had been recovered from the Eromanga Basin of Queensland (Aptian-Cenomanian) by the turn of the millennium. The piece of jaw in the book seems to be reffering to Aussiedraco (known from a partial mandibular symphysis found in 1980, and named in 2011), but it does not mention Mythunga (known from an incomplete skull , found in 1991 and described in 2008), though some of the other finds have also been likened to Ornithocheirus , so I can see where WW would get the idea to include Australia as part of Ornithocheirus’s range (as there we no named Aussie pterosaurs by 2005), and the 2000 book also brings up a shoulder girdle likened to Pteranodon , though also notes similarities with Ornithocheirus. When you only have an isolated bone to work with, determining the identity of the pterosaur who left it is inherently difficult, though it was apparently good enough for WW to lump at least some of the Eromanga material into Ornithocheirus. I also recall some papers likening other Australian fossils to pteranodontids, including a partial pelvis (more on that later). However, differentiating isolated pteranodontian material from anhanguerians or azhdarchoids has proven time and time again to be difficult, due to convergences in certain aspects of their anatomy.

The rest is more clear-cut; the South American specimen is Tropeognathus (who also accounts for the large size), the “recent African material ” refers to (still unnamed) anhanguerid fossils discovered in the Elrhaz Formation in Nigeria alongside the new Sarcosuchus material at the turn of the millennium (potentially also finds from the Chenini Formation, 2000 ), though fragmentary fossils of pterosaurs from the Lualaba Group in Central Africa have also been classified as cf. Ornithocheirus back in 1948, and as I said, a lot of the other material comes from Europe, which included taxa from the very Early Cretaceous; Serrodraco (Valanginian) and Targaryendraco (Hauterivian), as well as the Campanian "Ornithocheirus" buenzeli", known only from a partial jaw and humerus from Austria, and was previously hailed as evidence of toothed pterosaurs surviving into the Late Cretaceous, while current evidence points towards them being one of many victims of the Turonian extinction, with only toothless types soaring the Late Cretaceous skies. There is also Piksi, known from fragmentary remains from the upper Campanian Two Medicine Formation, initially described as a bird but later suggested to be a dwarf pterosaur, specifically an ornithocheiroid of some sort (though others still argue bird), but extraordinary claims require extraordinary evidence so I'm very much inclined to the former theory; that toothed pterosaurs simply weren’t a thing in the Late Cretaceous, much like allosaurs and diplodocoids or pliosaurs and ichthyosaurs.

So yes, the 2005 book was quite generous when judging what counted as an Ornithocheirus (basically anything ever likened to the genus) but it’s easy enough to guess what they were citing. Ironically, the two other named Australian pterosaurs (discovered after WW wrapped up) turned out to be keel-crested tropeognathines, those being the giant Thapunngaka, known from a partial jaw with a crest found in 2011 (named in 2021), which was almost as large as its Brazilian cousin, and the smaller but more complete Ferrodraco (2019), Australia’s most complete pterosaur to date . Worth noting is that David Unwin, the main consultant for “Giant of the Skies”, was of the opinion that the keel-crested Tropeognathus mesembrinus was synonymous with Ornithocheirus simus, and that the former represented the male of the species , as depicted in the episode (with females being crestless), and this was also the main inspiration for the migratory behavior . Conversly, others like Peter Wellnhofer (who named Tropeognathus) were critical and already citing Ornithocheirus as a wastebasket taxon, as mentioned in the 2000 book, and in Wellnhofer's own 1992 The Illustrated Encyclopedia of Pterosaurs, he's very critical and elaborates by stating the same issues modern pterosaur researchers have with the genus (quite ahead of his time).

Wellnhofer was definitely on the right track, though Unwin’s sexual dimorphism theory was certainly novel and forward-thinking, complementing what had already confirmed to be the case in Pteranodon and Nyctosaurus during the 90s, with females of the latter being smaller than the males and lacking as extravagant of a crest (which is especially pronounced in Pteranodon), though evidence of such has not been confidently confirmed in other pterosaurs. Perhaps unsurprisingly, as they hail from Niobrara, Pteranodon and Nyctosaurus have a much more extensive fossil record than the vast majority of pterosaur taxa, since most fossil formations that otherwise preserve dinosaurs and marine reptiles well simply don’t have the right conditions to preserve delicate-boned pterosaurs, a major fossilization bias that is unfortunately quite prevalent across the Triassic, Jurassic and Cretaceous. Really, just name any famous Mesozoic formation, like Chinle, Morrison, Oxford Clay, Djadochta, Dinosaur Park, Nemget or Hell Creek, and their pterosaur fossil record only comes in two varieties; very scrappy or nonexistent. By contrast, fossil formation that are rich in pterosaur material, like Niobrara/Peirre Shale, the Santana Group or the Solnhofen Limestone tend to have few if any good fossils of non-avian dinosaurs. In a nutshell, this means that the chances of us finding evidence of sexual dimorphism or migratory behavior in animals like Tropeognathus or Quetzalcoatlus is pretty slim (Caiuajara, known from a bonebed, implies sexual dimorphism wasn't a thing in azhdarchoids), leaving it in the realm of speculation. Ironically, in contrast to the nuance of "Giant of the Skies", “Land of Giants” and Sea Monsters fail to showcase the strong sexual dimorphism in Pteranodon, even though it would have been very easy to pull off (them repurposing the design for Azhdarcho has effectively created female Pteranodon).

Of course, back in the early 2000s, they would have faced the issue of not having any pterosaur material from Mid Cretaceous Argentina to cite, other than the filter-feeding, upper Albian ctenochasmatid Pterodaustro...and (given that they were starting from scratch) it's weird that they didn’t just go with it. It isn’t a very big pterosaur, but it certainly is weird and intriguing. On the upside, we have a diverse, if fragmentary fossil record of Cenomanian pterosaurs from the contemporary Kem Kem Beds in Morrocco, which (biogeographically and biochronologically) is very consistent with the earlier taxa from the Santana Group, housing fossils of azhdarchoids but also anhanguerians like Siroccopteryx (also considered a tropeognathine) and Nicorhynchus (a coloborhynchine), the latter also being known from slightly older rocks in Britain (Cambridge Greensand again), which include the jaw tip of a large specimens estimated to have had a 7-meter wingspan, and Kem Kem also has jaw fragments tentatively attributed to the Brazilian Anhanguera and also Ornithocheirus (though take those with a giant grain of salt).

Notably, Africa and South America had very low faunal endemism during the Mid Cretaceous, as evident from its native dinosaurs (titanosaurs, rebbachisaurids, carcharodontosaurids, spinosaurids, and both large and small abelisaurids), and the flying pterosaurs were likely no different. Another giant from this time period is the coloborhynchine Uktenadactylus from the upper Albian Paw Paw Formation in Texas, who is estimated to have reached similar proportions to its cousin Nicorhynchus (though it’s only known from a snout tip ). Tropeognathus, Thapunngaka, Nicorhynchus and Uktenadactylus show that huge anhanguerids were not uncommon in the Mid Cretaceous, and ironically, despite Australia’s lackluster fossils record, Thapunngaka and Ferrodraco, in a roundabout way, validate the inclusion of Tropeognathus in “Land of Giants”. Ferrodraco proves that these keel-crested tropeognathines made it to the upper Cenomanian, coming from the Winton Formation, which is contemporary to the Huincul Formation, home of Argentinosaurus (and after Giganotosaurus), and gigantism is a very common occurrence in evolution, which, coupled with the biogeography of anhanguerids, makes it’s pretty plausible that giant anhanguerids soared across the skies of Cenomanian South America, but fossils of pterosaur from that time and place are basically nonexistent at the moment, with only isolated anhanguerian teeth from the Cenomanian of Brazil being known. Suffice to say, finding any pterosaur fossils from the Rio Limay Group, giant or not, will be a very important find, and anhanguerians are the most likely type to pop up there (so far we only have undescribed footprints).

But what about Pteranodon and pteranodontians? Well...this is where things get very baffling ...

Description

This is taken from the 2000 book Walking with Dinosaurs the Evidence, written by Darren Naish and David Martill, a tie-in to 1999’s Walking with Dinosaurs, which explains some of the thought processes behind the (now very dated) portrayals of the animals included in that series. I’ve posted this for the sake of offering some easily accessible information regarding some of the most controversial creative choices taken in the series.

So the WW series certainly suffered from the law of diminishing returns, with 1999’s Walking with Dinosaurs attracting 18.91-15.69 million viewers (depending on the episode), 2001’s Walking with Beasts only 13.99-5.87 million (a more notable dip as it went on compared to WWD), 2002’s “The Giant Claw” about 6 million and “Land of Giants” 5.76 million, 2003’s Sea Monsters 7.59-6.52 million, and lastly, and 2005’s Walking with Monsters a meager 4.57-4.23 million, and likewise, the budget kept getting slashed from 6 million for WWD to 4 million for WWB, and finally only 3 million for Sea Monsters and WWM, explaining why the latter instalments were shorter, and likely also why they got less supplementary material (WWM didn’t even get its own companion book), and why there were no equivalents to 2000’s Walking with Dinosaurs: The Evidence, which was much more informative than any later book tied to the WW franchise.

Which is really annoying, because frankly, I’m dying to know just what their justification for describing Pteranodon as a near-cosmopolitan genus that lasted almost the entirety of the Cretaceous was , because I can’t find any other source that describes it as anything other than a Late Cretaceous North American genus, maybe offhandedly referencing possible contemporary material from Japan and Europe, but that’s about it. Pteranodon isn’t alone in this though, as the 2005 book describes Ornithocheirus in much the same vein, as a near-cosmopolitan genus that lasted the entirety of the Cretaceous , and “Land of Giants” shows the same gigantic species appearing 27 million years after its debut in “Giant of the Skies”, and Sea Monsters, likewise, has the exact same Pteranodon show up 25 million years after its debut in “Land of Giants”.

Again, it’s important to stress that endothermic, highly active tetrapods such as pterosaurs simply aren’t made to remain stagnant in an ever-changing world for more than 5 million years at best, being too specialized to survive unless they subtly change over millennia and inevitably evolve into new species, new genera, and eventually wholly new types, and 25 million years is a whole lot of time for them to evolve into something completely unrecognizable compared to its progenitor. While wastebasket taxonomy is evident here, in this case of Ornithocheirus, it’s easy to infer why they described it as such, because Ornithocheirus was a notorious wastebasket taxon, the pterosaur equivalent of Iguanodon, with both originally getting described from very fragmentary material found in Britain during the 19th century.

As is well known (but criminally underdiscussed), “Land of Giants” put Pteranodon in Argentina 100 million years ago (very early Cenomanian), and it actually crosses paths with the giant Ornithocheirus (Tropeognathus) cameo, as we briefly see them sharing the sky, and later, in the final segment of Sea Monsters, we see Pteranodon in North America 75 mya (upper Campanian), which is still anachronistic (the youngest fossils are about 5 million years older) but at least they put it in its proper biome (the Niobrara Chalk) with animals that it would have actually encountered in life, though according to the 2005 book, Ornithocheirus could just as well have made a cameo here.

As I said, the latter is easier to explain, as Ornithocheirus simus (the type species) was first described from very fragmentary remains from the upper Albian Cambridge Greensand in England and subsequently, the genus became a dumping ground for various ornithocheirid fossils (the group would more accurately be called anhanguerians today) from all across the globe, and as is usually the case with wastebasket taxa, which pterosaur fossils counted as Ornithocheirus would depend on who you asked, though like with Iguanodon, the bulk of the assigned fossils came from the Lower Cretaceous of England. The 2005 book seems to take a very generous definition, and gives Ornithocheirus a fossil range of 140-70 mya, and says that it inhabited Europe, South America, Africa and...Australia?

The 2000 book does offer some context to it. A number of isolated pterosaur bones had been recovered from the Eromanga Basin of Queensland (Aptian-Cenomanian) by the turn of the millennium. The piece of jaw in the book seems to be reffering to Aussiedraco (known from a partial mandibular symphysis found in 1980, and named in 2011), but it does not mention Mythunga (known from an incomplete skull , found in 1991 and described in 2008), though some of the other finds have also been likened to Ornithocheirus , so I can see where WW would get the idea to include Australia as part of Ornithocheirus’s range (as there we no named Aussie pterosaurs by 2005), and the 2000 book also brings up a shoulder girdle likened to Pteranodon , though also notes similarities with Ornithocheirus. When you only have an isolated bone to work with, determining the identity of the pterosaur who left it is inherently difficult, though it was apparently good enough for WW to lump at least some of the Eromanga material into Ornithocheirus. I also recall some papers likening other Australian fossils to pteranodontids, including a partial pelvis (more on that later). However, differentiating isolated pteranodontian material from anhanguerians or azhdarchoids has proven time and time again to be difficult, due to convergences in certain aspects of their anatomy.

The rest is more clear-cut; the South American specimen is Tropeognathus (who also accounts for the large size), the “recent African material ” refers to (still unnamed) anhanguerid fossils discovered in the Elrhaz Formation in Nigeria alongside the new Sarcosuchus material at the turn of the millennium (potentially also finds from the Chenini Formation, 2000 ), though fragmentary fossils of pterosaurs from the Lualaba Group in Central Africa have also been classified as cf. Ornithocheirus back in 1948, and as I said, a lot of the other material comes from Europe, which included taxa from the very Early Cretaceous; Serrodraco (Valanginian) and Targaryendraco (Hauterivian), as well as the Campanian "Ornithocheirus" buenzeli", known only from a partial jaw and humerus from Austria, and was previously hailed as evidence of toothed pterosaurs surviving into the Late Cretaceous, while current evidence points towards them being one of many victims of the Turonian extinction, with only toothless types soaring the Late Cretaceous skies. There is also Piksi, known from fragmentary remains from the upper Campanian Two Medicine Formation, initially described as a bird but later suggested to be a dwarf pterosaur, specifically an ornithocheiroid of some sort (though others still argue bird), but extraordinary claims require extraordinary evidence so I'm very much inclined to the former theory; that toothed pterosaurs simply weren’t a thing in the Late Cretaceous, much like allosaurs and diplodocoids or pliosaurs and ichthyosaurs.

So yes, the 2005 book was quite generous when judging what counted as an Ornithocheirus (basically anything ever likened to the genus) but it’s easy enough to guess what they were citing. Ironically, the two other named Australian pterosaurs (discovered after WW wrapped up) turned out to be keel-crested tropeognathines, those being the giant Thapunngaka, known from a partial jaw with a crest found in 2011 (named in 2021), which was almost as large as its Brazilian cousin, and the smaller but more complete Ferrodraco (2019), Australia’s most complete pterosaur to date . Worth noting is that David Unwin, the main consultant for “Giant of the Skies”, was of the opinion that the keel-crested Tropeognathus mesembrinus was synonymous with Ornithocheirus simus, and that the former represented the male of the species , as depicted in the episode (with females being crestless), and this was also the main inspiration for the migratory behavior . Conversly, others like Peter Wellnhofer (who named Tropeognathus) were critical and already citing Ornithocheirus as a wastebasket taxon, as mentioned in the 2000 book, and in Wellnhofer's own 1992 The Illustrated Encyclopedia of Pterosaurs, he's very critical and elaborates by stating the same issues modern pterosaur researchers have with the genus (quite ahead of his time).

Wellnhofer was definitely on the right track, though Unwin’s sexual dimorphism theory was certainly novel and forward-thinking, complementing what had already confirmed to be the case in Pteranodon and Nyctosaurus during the 90s, with females of the latter being smaller than the males and lacking as extravagant of a crest (which is especially pronounced in Pteranodon), though evidence of such has not been confidently confirmed in other pterosaurs. Perhaps unsurprisingly, as they hail from Niobrara, Pteranodon and Nyctosaurus have a much more extensive fossil record than the vast majority of pterosaur taxa, since most fossil formations that otherwise preserve dinosaurs and marine reptiles well simply don’t have the right conditions to preserve delicate-boned pterosaurs, a major fossilization bias that is unfortunately quite prevalent across the Triassic, Jurassic and Cretaceous. Really, just name any famous Mesozoic formation, like Chinle, Morrison, Oxford Clay, Djadochta, Dinosaur Park, Nemget or Hell Creek, and their pterosaur fossil record only comes in two varieties; very scrappy or nonexistent. By contrast, fossil formation that are rich in pterosaur material, like Niobrara/Peirre Shale, the Santana Group or the Solnhofen Limestone tend to have few if any good fossils of non-avian dinosaurs. In a nutshell, this means that the chances of us finding evidence of sexual dimorphism or migratory behavior in animals like Tropeognathus or Quetzalcoatlus is pretty slim (Caiuajara, known from a bonebed, implies sexual dimorphism wasn't a thing in azhdarchoids), leaving it in the realm of speculation. Ironically, in contrast to the nuance of "Giant of the Skies", “Land of Giants” and Sea Monsters fail to showcase the strong sexual dimorphism in Pteranodon, even though it would have been very easy to pull off (them repurposing the design for Azhdarcho has effectively created female Pteranodon).

Of course, back in the early 2000s, they would have faced the issue of not having any pterosaur material from Mid Cretaceous Argentina to cite, other than the filter-feeding, upper Albian ctenochasmatid Pterodaustro...and (given that they were starting from scratch) it's weird that they didn’t just go with it. It isn’t a very big pterosaur, but it certainly is weird and intriguing. On the upside, we have a diverse, if fragmentary fossil record of Cenomanian pterosaurs from the contemporary Kem Kem Beds in Morrocco, which (biogeographically and biochronologically) is very consistent with the earlier taxa from the Santana Group, housing fossils of azhdarchoids but also anhanguerians like Siroccopteryx (also considered a tropeognathine) and Nicorhynchus (a coloborhynchine), the latter also being known from slightly older rocks in Britain (Cambridge Greensand again), which include the jaw tip of a large specimens estimated to have had a 7-meter wingspan, and Kem Kem also has jaw fragments tentatively attributed to the Brazilian Anhanguera and also Ornithocheirus (though take those with a giant grain of salt).

Notably, Africa and South America had very low faunal endemism during the Mid Cretaceous, as evident from its native dinosaurs (titanosaurs, rebbachisaurids, carcharodontosaurids, spinosaurids, and both large and small abelisaurids), and the flying pterosaurs were likely no different. Another giant from this time period is the coloborhynchine Uktenadactylus from the upper Albian Paw Paw Formation in Texas, who is estimated to have reached similar proportions to its cousin Nicorhynchus (though it’s only known from a snout tip ). Tropeognathus, Thapunngaka, Nicorhynchus and Uktenadactylus show that huge anhanguerids were not uncommon in the Mid Cretaceous, and ironically, despite Australia’s lackluster fossils record, Thapunngaka and Ferrodraco, in a roundabout way, validate the inclusion of Tropeognathus in “Land of Giants”. Ferrodraco proves that these keel-crested tropeognathines made it to the upper Cenomanian, coming from the Winton Formation, which is contemporary to the Huincul Formation, home of Argentinosaurus (and after Giganotosaurus), and gigantism is a very common occurrence in evolution, which, coupled with the biogeography of anhanguerids, makes it’s pretty plausible that giant anhanguerids soared across the skies of Cenomanian South America, but fossils of pterosaur from that time and place are basically nonexistent at the moment, with only isolated anhanguerian teeth from the Cenomanian of Brazil being known. Suffice to say, finding any pterosaur fossils from the Rio Limay Group, giant or not, will be a very important find, and anhanguerians are the most likely type to pop up there (so far we only have undescribed footprints).

But what about Pteranodon and pteranodontians? Well...this is where things get very baffling ...