Thursday 16 August 2018

Dinosauria Caspakensis: Pterodactylus tyleri

Above the trees there soared into my vision a huge thing on batlike wings - a creature large as a large whale, but fashioned more after the order of a lizard.
  - Chapter 4, "The Land That Time Forgot," Edgar Rice Burroughs, 1918

There are three general "groups" of animals which are not members of the Dinosauria, but due to their size, majesty, and the terror they instill, are included as "honourary Dinosaurs" in the public consciousness. There are the "pre-Mesozoic reptilimorphs" like Dimetrodon ("two measures of teeth"), Scutosaurus ("shield lizard"), and Gorgonops ("Gorgon face"), who may belong to wildly distinct groups, but are sufficiently morphologically similar that they are counted among them; there are the marine reptiles like the Plesiosaurs, Ichthyosaurs, and Mosasaurs; then there are the Pterosaurs, who are the closest related to the Dinosaurs as fellow members of the Ornithodira.

While Burroughs only granted a specific name to Plesiosaurus olsoni, I thought that the other fauna of Caspak deserved that honour. As Bradley coined P. olsoni in honour of the man who slew & subsequently cooked it, I figured that as the first person of the U-33 to see a Pterosaur should be its namesake. That being none other than Bowen Tyler himself, I introduce to you Pterodactylus tyleri, "Tyler's wing finger."

Discovery of the Pterodactylus

Mighty forms could be seen moving upon the ground in the thick forest, while the bosom of the river wriggled with living things, and above flapped the wings of gigantic creatures such as we are taught have been extinct throughout countless ages.
  - Chapter 4, "The Land That Time Forgot," Edgar Rice Burroughs, 1918

Much like the remains of Plesiosaurs, Pterosaur fossils have been stumbled upon throughout human prehistory. Fossils of Pterosaurs and marine reptiles are plentiful in the Niobrara Chalk, the strata preserving what was once the Great American Seaway: people dwelt in the region for thousands of years before Othniel Charles Marsh's 1870 expedition. Adrienne Mayor conjectures that ancient Cheyenne and Sioux may have come across fossils of Pterosaurs and marine reptiles in association, which then inspired tales of great birds and water monsters which battled one another, their bones petrified in the very earth.

It's easy to imagine early peoples throughout the world making similar mythological journeys. Herodotus' "Winged Serpents" of Arabia certainly sound familiar:

75. There is a region moreover in Arabia, situated nearly over against the city of Buto, to which place I came to inquire about the winged serpents: and when I came thither I saw bones of serpents and spines in quantity so great that it is impossible to make report of the number, and there were heaps of spines, some heaps large and others less large and others smaller still than these, and these heaps were many in number. This region in which the spines are scattered upon the ground is of the nature of an entrance from a narrow mountain pass to a great plain, which plain adjoins the plain of Egypt; and the story goes that at the beginning of spring winged serpents from Arabia fly towards Egypt, and the birds called ibises meet them at the entrance to this country and do not suffer the serpents to go by but kill them. On account of this deed it is (say the Arabians) that the ibis has come to be greatly honoured by the Egyptians, and the Egyptians also agree that it is for this reason that they honour these birds.
76. The outward form of the ibis is this:—it is a deep black all over, and has legs like those of a crane and a very curved beak, and in size it is about equal to a rail: this is the appearance of the black kind which fight with the serpents, but of those which most crowd round men's feet (for there are two several kinds of ibises) the head is bare and also the whole of the throat, and it is white in feathering except the head and neck and the extremities of the wings and the rump (in all these parts of which I have spoken it is a deep black), while in legs and in the form of the head it resembles the other. As for the serpent its form is like that of the watersnake; and it has wings not feathered but most nearly resembling the wings of the bat. Let so much suffice as has been said now concerning sacred animals.
 - The History of Herodotus, Part II, 75-76
While winged snakes can be easily explained as imaginary combinations of primal fears (snakes + wings = flying snakes!), and reconstructing a Pterosaur into a winged snake would require dissociating from the leg bones (articulated skeletons like P. antiquus, "ancient wing finger," are not as common as partial ones) it's worth noting that Pterosaur remains have been found on the other side of a mountainous area between Egypt and Arabia not too far away from Buto:

Then one considers the African Kongamoto - could it have been inspired by fossils from the Tendaguru Beds? Could the dread Balaur of Transylvanian folklore, a serpentine monster which grows wings when it has not sated its appetite, be the mythic incarnation of pre-Cuvier encounters with bones of the monstrous Azhdarchids of Hateg Island? Is Quetzalcoatlus ("feathered serpent") an example of "recursive" mythology - an animal named after a mythological being, which was the Aztecs' mythological rationalisation of that very animal's remains? It's fun to imagine...

Pterosaur fossil sites throughout the world - no wonder flying reptilian monsters abound in European, Asian, and American mythologies!
(Purple = Triassic; Blue = Jurassic; Green = Cretaceous)

The modern history of Pterosaur knowledge begins much later - in 1784, with Cosimo Alessandro Collini. Collini was a Florence-born historian who eventually ended up in the Electorate of Bavaria, where he became a member of the Palatine Academy of Sciences and director of the Mannheim Naturalienkabinett (Cabinet of Natural History) in 1763. Collini was thus the custodian to the collections held in the Elector Count's castle - which included a curious item.

Collini concluded that, rather than being the remains of a sinner killed in the Flood - a popular explanation for these bones at that point in history - it was an animal from a previous epoch of Earth's history. Since this creature was found in the Early Jurassic Solnhofen Plattenkalk, it was associated with various marine creatures - turtles, fish, belemnites - as well as creatures which would later be called Ichthyosaurs. Combined with a belief that the likeliest environment to host unknown animals is the ocean, Collini theorised that this was a marine reptile, with the long wing finger supporting a membranous "fin." As theories go, it wasn't that out there considering the state of research: in fact, Johann Georg Wagler would revisit the marine Pterosaur theory in 1830, and the notion occurred to Dougal Dixon when writing The New Dinosaurs.

The new species was not yet named, but it fascinated the scientific community. Johann/Jean Hermann was the first to argue that the long finger from which Pterodactylus derives its name supported a wing, and also drew the first recorded palaeontological reconstruction of the creature - albeit as a batlike mammal rather than a reptile:

George "Father of Modern Palaeontology" Cuvier disagreed with the mammalian conclusion, and stuck with the reptile diagnosis. Richard Owen concurred - but not everyone did. The Pterosaurs-as-bats meme reached its arguable zenith with Edward Newman's 1843 hypothesis, where he depicted Pterodactylus as looking like some sort of terrifying long-nosed vampire fiend:

Unfortunately, most of these reconstructions were shared in the scientific community alone: it wasn't until the middle of the 19th Century that popular science brought Pterosaurs into the public consciousness, from Henry de la Beche's Duria Antiquior to John Martin's Great Sea Dragons - the old American myths of sea-against-sky monsters echoed in nightmarish detail. The most enduring of popular depictions of Pterosaurs remains, once again, Benjamin Waterhouse Hawkins' monumental Crystal Palace statues:

Like their marine Mesozoic mates, Pterosaurs have also featured in Theosophy and cryptozoology. Madame Blavatsky supposed in The Secret Doctrine that they "served as the prototypes for the Seraph of Moses and his great Brazen Serpent." The aforementioned Kongamato is but one of many alleged sightings of extant Pterosaurs reported in living memory, with others including the Central African Olitiau , the South American Washoriwe, and the Papua New Guinean Ropen. The cultural impact of Pterodactylus casts a long shadow indeed.

While the discovery of Rhamphorhynchus ("beak snout") in 1825 and Pteranodon ("winged & toothless") in 1870 hinted at the extreme size and morphological variety of Pterosaurs, for the majority of its years in the limelight, the humble Pterodactylus has remained the "default" Pterosaur in the public eye.

The Pterodactylus in Literature
The "impudent" Pterodactyl as illustrated in John Mill's The Fossil Spirit; A Boy's Dream of Geology (1854)

We saw nothing of the wild men of the previous day, and only once were we menaced by any of the strange denizens of Caprona, when some frightful nightmare of the sky swooped down upon us, only to be driven off by a fusillade of bullets. The thing appeared to be some variety of pterodactyl, and what with its enormous size and ferocious aspect was most awe-inspiring.
 - Chapter 6, "The Land That Time Forgot," Edgar Rice Burroughs, 1918

Pterosaurs didn't have to wait long to be inducted into literature: again, one of the earliest was John Mill's The Fossil Spirit - A Boy's Dream of Geology (1854), which uses the fantastical storytelling method of One Thousand and One Nights (one chapter's even called "The Fakir's Story") and others to cast the reader's mind back through time, encouraging one to imagine themselves in the role of an Ichthyosaurus, a Cetiosaurus ("whale lizard") - or, indeed, a Pterodactyl.

Higher up still, the first monkey ever seen on the face of the globe clambered, gamboling and playing up the granite hills. Still farther away, ran the Pterodactyl, with the winged hand, gliding or rather sailing through the dense and compressed air like a huge bat.
 - Chapter 29: "On the Waters - A Raft Voyage," A Journey to the Centre of the Earth, Jules Verne (Griffith and Farran translation), 1871

Pterodactyls haunted the strange skies of the literary world throughout the 19th Century, frequently in the role of Primeval Winged Terror: along with Verne's subterranean odyssey, John Morgan's King Solomon's Treasures (1887), Phil Robinson's "The Last of the Vampires" (1893) and Mary Anderson's "A Son of Noah" (1893) all feature surviving Pterodactyls menacing the protagonists in modern (or in the latter's case, Holocene) times.

Moving on to the 20th Century, Eden Phillpott's Fancy Free featured the Pterodactyl in "The Archdeacon and the Deinosaurs" in an elaborate dream sequence alongside several other prehistoric beasties. Robert Ames Bennet's Thyra: A Romance of the Polar Pit expands on the Theosophy-inspired sector of Lost Worlds with an adventure in a Polar realm populated by beastmen, dinosaurs, pterodactyls, and the civilisation founded by lost Viking explorers (anyone else thinking "Marchers of Valhalla" or War Eagles?) Cryptofiction continued to populated the present with shadows out of time in Samuel Hopkins Adams' "The Flying Death" (1903, later expanded to novel length under the same name in 1908) and Thomas Charles Sloane's "The Pterodactyl" (1907).

Some famous authors also mentioned the wing-fingers. Ambrose Bierce wrote of it a number of times: it is (not entirely correctly) listed as a contemporary of the Deinotherium in The Devil's Dictionary (1906), and features in the truly weird short tale "For the Ahkoond" (1909). Arthur Machen was also fond of evoking the creature, mentioning it in The Novel of the Black Seal (1895) and "The White People" (1904), and is the focus of "St. George and the Dragon," a delightful essay on the mythology of dragons.

But it is in Arthur Conan Doyle's The Lost World that the Pterodactyl truly gets a chance to shine: in fact, it features more prominently and more regularly in the story than any of the dinosaurs. A piece of its wing was part of the evidence which convinced Malone to embark on the expedition, then the sight of one allows Summerlee to express a final chance at scepticism, and it provides the unforgettable first encounter with a living, breathing, hunting dweller of Maple White Land:

What occurred was this. Lord John had shot an ajouti—which is a small, pig-like animal—and, half of it having been given to the Indians, we were cooking the other half upon our fire. There is a chill in the air after dark, and we had all drawn close to the blaze. The night was moonless, but there were some stars, and one could see for a little distance across the plain. Well, suddenly out of the darkness, out of the night, there swooped something with a swish like an aeroplane. The whole group of us were covered for an instant by a canopy of leathery wings, and I had a momentary vision of a long, snake-like neck, a fierce, red, greedy eye, and a great snapping beak, filled, to my amazement, with little, gleaming teeth. The next instant it was gone - and so was our dinner. A huge black shadow, twenty feet across, skimmed up into the air; for an instant the monster wings blotted out the stars, and then it vanished over the brow of the cliff above us. We all sat in amazed silence round the fire, like the heroes of Virgil when the Harpies came down upon them.
 - Chapter IX, "Who could have Foreseen it?" "The Lost World," Sir Arthur Conan Doyle

The Pterodactyl (Malone notes that they could not tell whether the creatures they met were Dimorphodon or Pterodactylus), along with the Megalosaurus ("great lizard"), proves to be the most terrible reptilian foe of Challenger's expedition, and it's difficult not to imagine John Martin going wild with a scene like this:

 If, as Lord John said, the glade of the iguanodons will remain with us as a dream, then surely the swamp of the pterodactyls will forever be our nightmare...
The place into which we gazed was a pit, and may, in the early days, have been one of the smaller volcanic blow-holes of the plateau. It was bowl-shaped and at the bottom, some hundreds of yards from where we lay, were pools of green-scummed, stagnant water, fringed with bullrushes. It was a weird place in itself, but its occupants made it seem like a scene from the Seven Circles of Dante. The place was a rookery of pterodactyls. There were hundreds of them congregated within view. All the bottom area round the water-edge was alive with their young ones, and with hideous mothers brooding upon their leathery, yellowish eggs. From this crawling flapping mass of obscene reptilian life came the shocking clamor which filled the air and the mephitic, horrible, musty odor which turned us sick. But above, perched each upon its own stone, tall, gray, and withered, more like dead and dried specimens than actual living creatures, sat the horrible males, absolutely motionless save for the rolling of their red eyes or an occasional snap of their rat-trap beaks as a dragon-fly went past them. Their huge, membranous wings were closed by folding their fore-arms, so that they sat like gigantic old women, wrapped in hideous web-colored shawls, and with their ferocious heads protruding above them. Large and small, not less than a thousand of these filthy creatures lay in the hollow before us.
 - Chapter X, "The most Wonderful Things have Happened," "The Lost World," Arthur Conan Doyle
And then we have one of - if not the - first examples of a prehistoric creature escaping into a city:

All sound had hushed in the audience and everyone was absorbed in the spectacle before them. Professor Challenger drew off the top of the case, which formed a sliding lid. Peering down into the box he snapped his fingers several times and was heard from the Press seat to say, 'Come, then, pretty, pretty!' in a coaxing voice. An instant later, with a scratching, rattling sound, a most horrible and loathsome creature appeared from below and perched itself upon the side of the case. Even the unexpected fall of the Duke of Durham into the orchestra, which occurred at this moment, could not distract the petrified attention of the vast audience. The face of the creature was like the wildest gargoyle that the imagination of a mad medieval builder could have conceived. It was malicious, horrible, with two small red eyes as bright as points of burning coal. Its long, savage mouth, which was held half-open, was full of a double row of shark-like teeth. Its shoulders were humped, and round them were draped what appeared to be a faded gray shawl. It was the devil of our childhood in person. There was a turmoil in the audience—someone screamed, two ladies in the front row fell senseless from their chairs, and there was a general movement upon the platform to follow their chairman into the orchestra. For a moment there was danger of a general panic. Professor Challenger threw up his hands to still the commotion, but the movement alarmed the creature beside him. Its strange shawl suddenly unfurled, spread, and fluttered as a pair of leathery wings. Its owner grabbed at its legs, but too late to hold it. It had sprung from the perch and was circling slowly round the Queen's Hall with a dry, leathery flapping of its ten-foot wings, while a putrid and insidious odor pervaded the room. The cries of the people in the galleries, who were alarmed at the near approach of those glowing eyes and that murderous beak, excited the creature to a frenzy. Faster and faster it flew, beating against walls and chandeliers in a blind frenzy of alarm. 'The window! For heaven's sake shut that window!' roared the Professor from the platform, dancing and wringing his hands in an agony of apprehension. Alas, his warning was too late! In a moment the creature, beating and bumping along the wall like a huge moth within a gas-shade, came upon the opening, squeezed its hideous bulk through it, and was gone.
 -  Chapter XVI, "A Procession! A Procession!" "The Lost World," Arthur Conan Doyle

Barely two years after Malone's spectacular publication, Edgar Rice Burroughs published David Innes' account of a similar prehistoric realm in At the Earth's Core (1914) - which introduced the Thipdar, the most immediate literary ancestor of Pterodactylus tyleri.

And at the first glance there broke upon my horrified vision the most frightful thing I had seen even within Pellucidar. It was a giant dragon such as is pictured in the legends and fairy tales of earth folk. Its huge body must have measured forty feet in length, while the batlike wings that supported it in midair had a spread of fully thirty. Its gaping jaws were armed with long, sharp teeth, and its claw equipped with horrible talons...
... So this was a thipdar. I might have known it. The cruel bloodhound of the Mahars. The long-extinct pterodactyl of the outer world. But this time I met it with a weapon it never had faced before. I had selected my longest arrow, and with all my strength had bent the bow until the very tip of the shaft rested upon the thumb of my left hand, and then as the great creature darted toward us I let drive straight for that tough breast.  Hissing like the escape valve of a steam engine, the mighty creature fell turning and twisting into the sea below, my arrow buried completely in its carcass.
 - Chapter XIV, "The Garden of Eden," At The Earth's Core, Edgar Rice Burroughs (1914)
The Pellucidarian Pterodactyl - Pterodactylus thipdar, perhaps? - is distinctive from P. tyleri in a few ways which will become evident. Both are different again from the other major Burroughsian pterosaur, the Mahars (Pteranthropoides?), but that's a whole other article. Suffice to say, the notion of a connected Burroughs universe is one of the most beloved flashpoints in popular culture - if Pellucidar could exist in the same reality as Tarzan's Africa, why not Caspak?

Pterodactylus tyleri

Near the summit I came upon a huge cavern. It is the abode of some mighty winged creature of the Triassic--or rather it was. Now it is mine. I slew the thing and took its abode.
 - Chapter 10, "The Land That Time Forgot," Edgar Rice Burroughs, 1918
Plesiosaurus olsoni was notable in that it accurately reflected contemporaneous palaeontological descriptions of its Mesozoic counterparts. Pterodactylus tyleri, on the other hand, is a different beast in one important respect - its size.

The largest Pterosaur known at the time "The Land That Time Forgot" was written was the 20ft/6m Pteranodon longiceps ("long-headed winged & toothless," originally Pterodactylus ingens, "enormous wing finger"), though some specimens were claimed to achieve 25ft/7.6m) and would remain so until the discovery of the monstrous 40ft/9m Quetzalcoatlus northropi ("Northrop's feathered serpent") in 1976. Few other Pterosaurs known to science at the time approached either in wingspan: the largest Pterodactylus specimens suggest a maximum wingspan of 5ft/1.5m.

P. tyleri has an observed length and wingspan of 80ft/24m.

Rough illustrative comparison using Pterodactylus antiquus as a basis.

Would you look at the size of that thing!

This is larger than the higher estimates for the current King of the Skies, Arambourgiana philadelphiae ("Arambourg's creature from Philadelphia," 43ft/13m), the wildest previous estimates for Quetzalcoatlus northropi (52ft/15.9m) and even the widely & prematurely reported Terror of Israel (60ft/20m) whose wing bone turned out to be a piece of fossilised wood. (Not to mention the giant Thipdar at the end of At The Earth's Core.) There were WW1 bombers with smaller wingspans than P. tyleri. There is nothing on the face of this planet, living or extinct - no bird of prey, no teratorn, no seabird - with remotely comparable dimensions of P. tyleri. At least, nothing that we know of...

Quite why Burroughs chose to depict a Pterosaur more than thrice the wingspan of the largest recorded species yet discovered by the 1910s is something of a mystery. It would be easy to assume that he just didn't know or care, and just put 80 feet because it sounded big and scary - but then, why would he have his P. olsoni a "mere" 16-18 feet, when the likes of Verne vastly inflated Plesiosaur size to 100 feet? Burroughs was clearly familiar with contemporaneous palaeontology, with plenty of esoteric references that only someone prehistorically-minded would recognise: in "The Land That Time Forgot" alone he namedrops palaeontologists and individual specimens. I'm not convinced Burroughs would have introduced such a monster through plain ignorance - especially when he later explains "how infinitely different was the true, live thing from the crude restorations."

More likely to me is that P. tyleri's incredible size was deliberate on Burrough's part. But for what reason? Perhaps it is representative of a common theme in animal horror - gigantism. Reptiles keep growing throughout their lives, and they can live for a very long time: the popular mythology that crocodiles and other reptiles would keep growing indefinitely if not killed by disease or injury persists to this day. So perhaps the sun-blocking terrors were the apex predators of Caspak: those individuals which survived their encounters with the other creatures of the isle eventually grew to a size where none could feasibly prey upon them.
 "I suppose I'm a fool," remarked the assistant secretary; "but by George, I can't help believing it, and I can see that girl now, with the big Airedale at her side protecting her from the terrors of a million years ago. I can visualize the entire scene--the apelike Grimaldi men huddled in their filthy caves; the huge pterodactyls soaring through the heavy air upon their bat-like wings; the mighty dinosaurs moving their clumsy hulks beneath the dark shadows of preglacial forests--the dragons which we considered myths until science taught us that they were the true recollections of the first man, handed down through countless ages by word of mouth from father to son out of the unrecorded dawn of humanity."
 - Chapter 1, "The People That Time Forgot," Edgar Rice Burroughs, 1918
The mention of Caspak's heavy air is also a nod to the question of just how such an enormous creature could fly in the first place. The giant Azhdarchids already appear to stretch the absolute limits of biological earthly flight, with only the mighty Argentavis magnificens ("magnificent silver bird") coming close to the Pterosaur league: how could a creature twice the wingspan and more than three times the length even get off the ground? The clue is in the air itself.

While Burroughs wouldn't write his first Amtor series until the 1930s, Venus as a tropical, thunderstorm-riven landscape was well explored in fiction for decades. Venus was regularly compared to Earth's Carboniferous period, a time where the planet was covered in forests and populated by gigantic insects - and, crucially, the atmosphere was very different. Since we know Caspak was also home to "huge insects" at the southernmost regions, it follows that in at least this area in Caspak, atmospheric conditions would have been more akin to the Carboniferous too - the land of the Galus was "considerably colder" than even the Alus' country. Quite how even a continent-sized Caspak could support several different atmospheres along with its strictly territorial flora & fauna is another facet of the Mystery of Caspak, and one noted in the text.

There's also a thematic resonance to the idea of a 100-foot-wingspanned monster terrorising ground-dwellers from the skies - indeed, it might even have proven darkly prophetic. Burroughs was clearly very invested in the Great War raging across Europe: he would surely have heard the stories of Operation Türkenkreuz, the Gotha bombing raids bringing the war to England:

On 25 May the Gothas first appeared in force over England. They bombed Shorncliffe Camp; 110 Canadian soldiers were among the casualties. On 13 June Kampfgeschwader 1 made its first raid on London, causing 600 casualties including 66 children killed and injured when a bomb struck an East End school. On 7 July the Gothas made their most spectacular raid on the capital. Appearing over London in mid-morning, 21 bombers dropped explosives over a wide area. Though casualties and damage were not heavy, the raid, viewed by millions of Londoners, vividly disclosed the impotence of a defense system designed to ward off airships, not airplanes. 
Through the periodic transfer of front line squadrons from France to England and the stiffening of home defense squadrons by allocations of first class fighter aircraft and experienced combat pilots, the British home defense organization forced the Germans to go over to night raiding by late 1917. These night raids, in which Germany employed giant aeroplanes (Riesenflugzeuge) as well as Gothas, were never seriously impeded by the defenses until the final raid of the war in May 1918. At that time the Germans gave up the campaign of their own volition, deciding that the bombers could more directly serve the needs of the army in the field by operations against the Allied back areas on the Western Front.
 - Christon Archer, Men at War: Politics, Technology, and Innovation in the Twentieth Century

At the time "The Land That Time Forgot" was published, aerial bombing was an unfamiliar, speculative idea, a scientifiction. Many people simply walked outside to see the plane, totally oblivious to this new age of warfare and the horrors to come - Guernica, Barcelona, Nanking, Berlin, the Blitz, Tokyo. The primal hiss of the Pterodactyl became the scream of the Stuka; the Pterosaur's swoop became the Kamikaze; the deadly bite & rend of ancient teeth & claws became terror bombing.

Still, there is one animal not native to Caspak which could challenge their supremacy of the skies - and sometimes the only thing to beat a flying monster is another one:

The first intimation I had of it was the sudden blotting out of the sunlight from above, and as I glanced quickly up, I saw a most terrific creature swooping down upon me. It must have been fully eighty feet long from the end of its long, hideous beak to the tip of its thick, short tail, with an equal spread of wings. It was coming straight for me and hissing frightfully - I could hear it above the whir of the propeller. It was coming straight down toward the muzzle of the machine-gun and I let it have it right in the breast; but still it came for me, so that I had to dive and turn, though I was dangerously close to earth.
 - Chapter 2, "The People That Time Forgot," Edgar Rice Burroughs, 1918

While the creature Tyler sighted was "large as a large whale," Tom Billings provides the reader with the 80ft figure in a rather more up-close-and-personal encounter. If the notion of aerial bombing was still fairly new to the world, then aerial battles must have seemed like fantasy - H.G. Wells' visionary The War in the Air was ten years old, and the first genuine dogfight only took place four years before "The People That Time Forgot" appeared in The Blue Book. Taking Conan Doyle's "The Horror of the Heights" to its logical martial conclusion, Burroughs' scrap between Monster and Man-in-Machine feels deeply authentic (at least from the machine's standpoint!):

The thing didn't miss me by a dozen feet, and when I rose, it wheeled and followed me, but only to the cooler air close to the level of the cliff-tops; there it turned again and dropped. Something - man's natural love of battle and the chase, I presume - impelled me to pursue it, and so I too circled and dived. The moment I came down into the warm atmosphere of Caspak, the creature came for me again, rising above me so that it might swoop down upon me. Nothing could better have suited my armament, since my machine-gun was pointed upward at an angle of about degrees and could not be either depressed or elevated by the pilot. If I had brought someone along with me, we could have raked the great reptile from almost any position, but as the creature's mode of attack was always from above, he always found me ready with a hail of bullets. The battle must have lasted a minute or more before the thing suddenly turned completely over in the air and fell to the ground.
Bowen and I roomed together at college, and I learned a lot from him outside my regular course. He was a pretty good scholar despite his love of fun, and his particular hobby was paleontology. He used to tell me about the various forms of animal and vegetable life which had covered the globe during former eras, and so I was pretty well acquainted with the fishes, amphibians, reptiles, and mammals of paleolithic times. I knew that the thing that had attacked me was some sort of pterodactyl which should have been extinct millions of years ago. It was all that I needed to realize that Bowen had exaggerated nothing in his manuscript.
 - Chapter 2, "The People That Time Forgot," Edgar Rice Burroughs, 1918
As with P. olsoni, the reptilian nature of P. tyleri makes it tremendously difficult to kill - even with a machine gun.

P. tyleri is the largest carnivorous predator in Caspak by a substantial margin: while the local Tyrannosaurus is undoubtedly heavier, it is half the length, and a fraction the span. But not all the Pterosaurs of Caspak are gigantic:

I knew how anxious they would be for word from me, and I was equally anxious to relieve their minds and also to get them and our supplies well within Caspak, so that we might set off about our business of finding and rescuing Bowen Tyler; but the pterodactyl's carcass had scarcely fallen before I was surrounded by at least a dozen of the hideous things, some large, some small, but all bent upon my destruction. I could not cope with them all, and so I rose rapidly from among them to the cooler strata wherein they dared not follow; and then I recalled that Bowen's narrative distinctly indicated that the farther north one traveled in Caspak, the fewer were the terrible reptiles which rendered human life impossible at the southern end of the island.
 - Chapter 2, "The People That Time Forgot," Edgar Rice Burroughs, 1918

Prehistoric Pterosaurs - that is, one which we know of from the fossil record - have a great variety of shapes and sizes, from the bluejay-sized Nemicolopterus to the aforementioned draconic Azdharchids, the long-tailed Rhamphorynchus to the comb-toothed Pterodaustro, with many in between. It's possible the "hideous things" of varying sizes are all P. tyleri at different ages, as the many P.olsoni which swarmed the U-33 appeared to be of the same species, but it's just as possible they represent the great variety of Pterosauria in the fossil record.

One curious aspect of P. tyleri is their apparent defiance of the land's unspoken law:

Almost instantaneously a huge bulk swooped down upon me, and as I looked up, I realized that there were flying reptiles even in this part of Caspak. The creature dived for my right wing so quickly that nothing but a sheer drop could have saved me. I was already close to the ground, so that my maneuver was extremely dangerous; but I was in a fair way of making it successfully when I saw that I was too closely approaching a large tree. My effort to dodge the tree and the pterodactyl at the same time resulted disastrously. One wing touched an upper branch; the plane tipped and swung around, and then, out of control, dashed into the branches of the tree, where it came to rest, battered and torn, forty feet above the ground.  Hissing loudly, the huge reptile swept close above the tree in which my plane had lodged, circled twice over me and then flapped away toward the south. As I guessed then and was to learn later, forests are the surest sanctuary from these hideous creatures, which, with their enormous spread of wing and their great weight, are as much out of place among trees as is a seaplane.
 - Chapter 2, "The People That Time Forgot," Edgar Rice Burroughs, 1918

Billings crashed somewhere between Bo-Lu ("Club-Men," Neanderthals) and Sto-Lu ("Hatchet-Men," between Neanderthal and Grimaldi Man) territory: roughly analogous of the time period from the Late Pleistocene to the Upper Palaeolithic.

In case you hadn't guessed, Pterodactyls vs. Aircraft is quite a popular theme.

Although P. tyleri's specific name is my own appellation, Burroughs bestowed upon the creature an honour that P. olsoni does not possess - a Caspakian name.

"The dim outlines of Oo-oh were unfolding below us when there came from above a loud whirring of giant wings. The Wieroo and I glanced up simultaneously, to see a pair of huge jo-oos" (flying reptiles - pterodactyls) "swooping down upon us.
 - Chapter 5, "The People That Time Forgot," Edgar Rice Burroughs, 1918
Many Caspakian words have translations, direct or approximate, in English provided: -lu for "man" under its original definition, cor-sva-jo for "from the beginning," and so on. As for jo-oo, there could be a number of possibilities. The first is the plainest, taking Billings' parentheses into account: jo = "flying"; oo = "reptile." However, this does not account for the presence of both compounds in other Caspakian words. Oo-oh is the land in which the Wieroos dwell: as the Wieroos are also winged, perhaps the oo element of jo-oo, Ooh-oh, and Wieroo is something to do with wings, flight, or the sky. As for jo, if we take the aforementioned cor-sva-jo as 1=1 word to translation, it could be cor = "from"; sva = "the"; jo = "beginning." In that sense, jo-oo could mean something along the lines of "beginning wing" = a reference to pterosaurs being the first true rulers of the skies, as well as their general territory on Caspak being far closer to the original source.

The Wieroos may consider themselves the masters of Caspak, but in an aerial between the Wieroo and the P. tyleri, there is no question of who rules the skies of Caprona:

"The Wieroo wheeled and dropped almost to sea-level, and then raced southward in an effort to outdistance our pursuers. The great creatures, notwithstanding their enormous weight, are swift on their wings; but the Wieroo are swifter. Even with my added weight, the creature that bore me maintained his lead, though he could not increase it. Faster than the fastest wind we raced through the night, southward along the coast. Sometimes we rose to great heights, where the air was chill and the world below but a blur of dim outlines; but always the jo-oos stuck behind us. 
"I knew that we had covered a great distance, for the rush of the wind by my face attested the speed of our progress, but I had no idea where we were when at last I realized that the Wieroo was weakening. One of the jo-oos gained on us and succeeded in heading us, so that my captor had to turn in toward the coast. Further and further they forced him to the left; lower and lower he sank. More labored was his breathing, and weaker the stroke of his once powerful wings. We were not ten feet above the ground when they overtook us, and at the edge of a forest. One of them seized the Wieroo by his right wing, and in an effort to free himself, he loosed his grasp upon me, dropping me to earth. Like a frightened ecca I leaped to my feet and raced for the sheltering sanctuary of the forest, where I knew neither could follow or seize me. Then I turned and looked back to see two great reptiles tear my abductor asunder and devour him on the spot.
 - Chapter 5, "The People That Time Forgot," Edgar Rice Burroughs, 1918

It is notable that the Wieroo's literary ancestors and subterranean counterparts, the more Pterodactylian Mahars, have domesticated - or at least trained - their Pterosaurs, yet the great Wieroo have not. Indeed, it is arguable that they cannot tame P. tyleri: their size alone would make them formidable to break, but a greater barrier is the Mystery of Caspak itself. The pan-genetic biological imperatives which keep species in each of their chrono-biomes, their interminable and self-destructive battling, and the Mystery of Caspak itself all prevent us from the terrifying sight of a Flight of Wieroos riding Jo-oos in the night sky. It is just as well for the world that the awesome terrors of Caspak are contained - one can only imagine what would happen if they could get out.

Depictions & Successors

As Plesiosaurs were to the prehistoric ocean, so Pterosaurs were to the antediluvian skies. In the decades after "The Land That Time Forgot," the winged wonders appeared everywhere. The Weird Tales Triumvirate all made some mention of them. Lovecraft's Elder Things constructed their cities with huge stone blocks lifeted "by vast-winged pterodactyls of a species heretofore unknown to paleontology" in "At the Mountains of Madness" (1936) - perhaps the same trenemdous stock from which P. tyleri derives descent? Similarly, Clark Ashton Smith's “Ubbo-Sathla” (1933) features a rather cosmic-horrific riff on the Mystery of Caspak - Pterosaur included - while he compares the depredators of "The Flower-Women" (1935) to Pterodactyls. Perhaps surprisingly, Howard is a bit more sparing on the wing-fingers than his compatriots: one could argue the "flying dragon"/"winged dinosaur" of "The Shadow Kingdom" (1929) was intended to be a Pterosaur, though I'm of the opinion Howard used that phrase over the more familiar Pterodactyl for a reason. The same with the thing "that was neither bat nor bird" in "The Scarlet Citadel" (1933), which Howard referred to as a dragon in his synopsis.

Other Weird Tales stalwarts joined in the Ptero-scramble: Edmund Hamilton brought the Inner Earth raging into the Outer World borne on Pterosaur wings in the bombastic "The Abysmal Invaders" (1929); E. Hoffman Price imbued the occult detective yarn with Mesozoic menace in "The Devil's Crypt" (1934); Manly Wade Wellman's boisterous Hok the Mighty encounters one in "Hok Visits the Land of Legends" (1942). A personal favourite post-Burroughs tale is Geoff St. Reynard's "The Plaid Pterodactyl" (1949) which provides an unexpected environ for the winged one - 1940s Scotland!

The Pterosaur really arrived in cinema in The Lost World (1925). While referred to as a "Pterodactyl" in the film itself, the creature is quite clearly a Pteranodon rather than any genus of Pterodactylus: it bears the distinctive crest, and appears to be toothless. Since Pterodactyl has been used synonymously with Pterosaur, we can presume that the choice was made to use the then-largest known species to represent "P. acernalbum" in the film. Willis O'Brien's groundbreaking stop-motion creature changed the face of cinema, ushering in the likes of the human-snatching P. craninsularis (King Kong, 1933) and the gigantic as-yet-undiscovered pterosauriform relative of Yi qi (One Million Years BC, 1966)

As for P. tyleri, it of course followed P. olsoni into cinematic cult status in 1975's The Land That Time Forgot. Here, it is has the opposite problem of P. olsoni: rather than being vastly oversized, P. tyleri is arguably undersized compared to its literary counterpart.

According to the Internet Movie Database, Bobby Parr (who played Ahm) is 5' 4"/1.63m. Therefore, using highly sophistimacated measuraming, we can calculate the 1977 P. tyleri to be roughly in the region of 37-40 feet or 11-12 metres. This places it well into the limits of known Pterosaur sizes - interestingly, Quetzalcoatlus northropi was publicly announced the same year The Land That Time Forgot was released - but barely half the size of the original iteration. I may be an absolute stickler for accuracy in my fantastical vintage retro-prehistoric creatures, but given that the only size comparison happens in a scene that isn't actually in the book (Ahm's fate in the original story is rather more mysterious) we can perhaps just presume that it's one of the smaller Pterodactyls Caspak is known for.

P. tyleri snatching away Ahm like a demon kingfisher plucking a frog from a pond may have been a consequence of the prop's lack of mobility, but it's much more feasible than the ubiquitous "people-snatching Pterosaur" meme common to palaeofiction. Due to Pterosaurs' light body weight relative to their size, most of them would struggle mightily with carrying away a human, and none of them had the muscles or the ligaments to grasp them with their talons. In fact, a book-accurate P. tyleri with an 80ft wingspan would be a more accurate way of depicting a Pterosaur carrying off a human, thanks to the square cube law.

Burroughs himself played with this idea with his Mahar, but given they are clearly a distinct species with unnatural joukerie-pawkerie in their genetic ancestry... Well, we can presume that the biological constraints which would prevent known prehistoric Pterosaurs from carrying humans away would not apply to them.

The sizing aside, the film's P. tyleri is about as accurate both to novel and contemporary science as can be expected: leathery skin, membranous wings that stretched to the ankles, narrow conical teeth. In fact, I'd even say it's a pretty close cousin to the beasties of palaeoartist extraordinaires Zdeněk  Burian and Neave Parker:

As with the books, P. tyleri gets a more substantial role in the sequel, 1977's The People That Time Forgot. Of course, in the adaptation it's still quite a bit smaller than the book (and the sweet poster), and rather than Billings' thrilling dogfight chase, the aerial battle is a bit more... strategic:

The film, being a bit lighter in tone than its predecessor, dilutes the menace of the dinosaurs: in the first film, the dinosaurs are a constant threat to the battle-hardened crew of a World War 2 submarine, with only heavy concentrated gunfire, ship-mounted artillery, or natural disaster stopping them. The Pterosaurs in the sequel are threatening to a troupe of explorers for exactly one scene, before they become little more than a sight gag for the rest of the adventure:

How to get 8-year-old Aly to hate your character, exhibit A
Sadly - or thankfully, depending on your appraisal of the films - Kevin Connor & Amicus never got around to the final third of the Caspak trilogy, Out of Time's Abyss, and thus depriving the world of a mid-air chase scene of two P. tyleri hunting our hero as he is carried away by a Wieroo.

Pterodactyls have been fixtures of popular culture for as long as the dinosaurs have - perhaps even longer - for their dominance of the skies. Birds and bats have conquered the air since the great calamity that ended the Mesozoic and began the Cenozoic - but in their time, it was the Pterosaur whose wings reigned supreme over the dreaming earth. They were the dragons of reality long before any kind of man stepped from the mists of apedom into humanity. It's little wonder that any works which brings them back from extinction places them right back at the top of the sky.

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