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INTRODUCTION

When you usually think about a pterosaur, you probably imagine it to be the first vertebrate that have to capability to fly tough the sky. Of course you might think of other species of Pterosaur as well. From the small Dimorphodon, to the bizzare Tupandactylus, to the iconic Pteranodon and all the way up to the terrifying giants like Quetzalcoatlus or Hatzegopteryx who ruled the skies during the final days of the Mesozoic. On recent expeditions back to Spec, we have made some new discoveries. Living Pterosaurs.

HISTORY

Azhdarchs first appeared in the Lower Cretaceous, quite possibly as far back in time as the Jurassic/Cretaceous boundary with forms like †Palaeocursornis showing unambiguous azhdarchid material from this era. Part of Azhdarchoidea, a clade of toothless, terrestrial omnivorous pterosaurs, azhdarchs were fairly low key during the Lower Cretaceous, even overshadowed by some of their relatives, such as the bizarrely-crested tapejarids or the macropredatorial thalassodromedids, increasing somewhat in diversity in the early Late Cretaceous as their relatives dwindled. However, it was after the Turonian where azhdarchs got their chance to shine: in an evolutionary blink of an eye, they dominate the world’s pterosaur faunas, far outnumbering all other forms in terms of preserved remains. More so, they also showcase some degree of diversity and specialisation, with different jaw shapes (most common being a dichotomy between forms with long, curved bills and forms with short, spear like beaks), different degrees of neck elongation, and most impressively bigger sizes: azhdarchids produced the largest flying animals ever known, with the titanic giraffe-sized †Quetzalcoatlus being the most iconic example. In HE, their story ends right in their golden age, as the KT event killed azhdarchs as they underwent their radiation, but in Spec it obviously progressed.

In Spec’s Palaeocene, the most studied pterosaur is the former “Spec’s last pterosaur”, †Gigantala. Remains of this pterosaur are best known from rock formations in western Canada, though the animal is found across North America, and possibly other Laurasian landmasses as well. Two species are recognised, †Gigantala cranitus and †Gigantala hastarostris, the former being by far the most common and well studied. †Gigantala cranitus is a mid-sized azhdarch, with a wingspan of 4.5 meters, while †Gigantala hastarostris was seemingly larger at an estimated 7 meters; the former occurs throught the Palaeocene, while the latter is restricted to Selandian deposits. †Gigantala is unique among azhdarchids for it’s short neck, having reversed the elongation and flatenning of the azhdarch cervical vertebrae to the condition seen in earlier azhdarchoids like chaoyangopterids. With long jaws, typical limb proportions for neoazhdarchians and their normal bias towards inland terrestrial settings, †Gigantala was most likely a generalistic carnivore, perhaps akin to thalassodromedis in habits. The skull doesn’t seem to show much deviation from normal azhdarchid skulls except for it’s abnormally small nasoantorbital; G. cranitus has a longer, stork-like bill, while G. hastarostris has a shorter beak with scizzor-like jaw edges. Gigantala disappears from the fossil reccord in the Eocene, the youngest fossils dating to two million years before the PTM. However, other pterosaurs explode in diversity in an event at times called “Cenozoic Pterosaur Renaissance” (CPR), and among these we see several types of azhdarchids, diversified as very unusual and bizarre forms that deviate from the “stork-like” norm. Among these are the limuazhdarchines, a lineage of small azhdarchids with long, wide webbed toes, seemingly converging ecologically with the long gone ctenochasmatoids; †Heusia, a diminutive azhdarchid with a wingspan beneath a meter that may show adaptations for aerial foraging; and †Verpacheirus, a lanky, sloth like azhdarchid. Giant azhdarchids are rare, presumably due to the spread of global forests, though remains belonging to animals with a presumed ten meter wingspan are known from China, Morocco and Australia; otherwise, the largest forms seem to be beneath a wingspan of 4 meters.

With the Oligocene, most of the more bizarre forms disappear from the fossil reccord – though this could be due to limitations in the availiable fossil reccord, as Spec palaeontology is still a new field -, returning to a normal dominance of longer legged plains-dwellers, some of which returning to giant sizes – one species, †Aetiogenoi eletherios, is quite possibly the largest flying animal ever known, with a wingspan of 15 meters; at an estimated weight of 500 kilos, it may very well had reached the limit of azhdarchid flight capacity. The Miocene remains most of the same, though with an increase in morphological diversity. The Pliocene and Pleistocene see a mild weeding out of their diversity, but the survivors quickly exploit the vast grassland world that has formed, and become an important component on grassland ecosystems, though sacrificing most of their inovation in favour of conservatism.

BIOLOGY

Azhdarchids are among some of the most derived pterosaurs, having taken the “aberrant” pterodactyloid traits to their extreme, yet ironically also fairly conservative. Following the lophocratian tendency for terrestrially, azhdarchids have long, powerful hindlimbs, hatchet-shaped postacetabular processes and very long fourth metacarpals – the forelimb elements being proportionally comparable to those of fast running ungulates, and having the longest fourth metacarpals among pterosaurs. They are efficient walkers and runners, even being suberb gallopers, being some of the most terrestriality adapted pterosaurs ever known. Their feet are generally short and narrow, yet quite strong, being of poor use to swim or even to wade, but proper for galloping and, in some forms, for climbing, having developed strong claws. They have rather mammal-like footpads, covered with scales.

While terrestrial, azhdarchids in general also display several further adaptations for powered flight. Up to 50 kilos of robust flight musculature are anchored to their shoulders, attached to large, robust scapulae and coracoids of subequal length, large glenoids and anterior cervicals bound into a notarium, and attaching to the humeri in large, though fairly simple dectopectoral crests. The rather large, blocki humeri have a large pneumatic opening in their proximal anterior surface, to which air sacs are attached: like ornithocheiroids and other neoazhdarchians, azhdarchids have extended their pulmonary air sac system into the patagia, the propatagium being completly inflatable and the brachiopatagium being pneumatised along the forelimb, allowing the animal to save weight as well as to help the muscle fibers and the pteroid to manipulate the shape of the wing membranes.

While the fourth metacarpal is predictably very long and robust, the metacarpals supporting the smaller, clawed fingers are much smaller and locked entirely at the end of the fourth metacarpal, having lost their connection to the wrist bones, a feature also seen in other neoazhdarchians and some ornithocheiroids. The wing finger is the smallest by pterosaurian standards, usually 47% of the overall wing length. The first phallange is the longest, while the second and third phallanges have a T-shaped cross-section that reinforce the torsional motions of the dystal part of the wing during flight.

With the short, curved wing finger and long metacarpal, the wing has normally a rather characteristic low aspect ratio, as to be expected from flyers in terrestrial settings; however, much like other pterosaurs, azhdarchids can contract their wing membranes, increasing their wing loading. While large flying birds like rocs are soarers, azhdarchids are better described as flap-gliders, something facilitated by the very energy economic pterosaurian launching and the shape-changing wing membranes. With just a couple of wing strokes, azhdarchids can cover very large distances, and with the largest forms being able to cover as much as 16,000 km in a single, non-interrupted flight, geographical barriers mean very little for the larger animals.

Azhdarchids have proportionally the longest necks of all pterosaurs, and perhaps the longest in porportion to the body size of all known vertebrates and the longest necks outside of Sauropoda, period, achieved through the elongation of the cervicals three to eight, with the fifth characteristically stretched to a length eight times it’s width. Flattened and with reduced neural spines, their vertebrae have an unique tubular appearence, rather apropriate considering their pneumatisation; due to their sheer length, it was thought that azhdarchid necks were rather inflexible, and indeed the tubular central neck vertebrae allow little motion, but the vertebrae at each end of the neck allow a wide range of motion. Like most pterosaurs, azhdarchids have rather elastic neck tissues, and can expand their easophagi considerably, allowing them to contain proportionally very large prey. As the azhdarchid torso is rather small, the animals often just store their prey in the neck, not needing a gular pouch, where the trapped prey item will stay as it moves downwards, being digested and processed slowly.

Azhdarchid skulls tend to be proportionally very large, the extinct Aetiogenoi eletherios holding the reccord for the longest skull on a terrestrial animal at 6 meters from jaw tip to the back of the skull. The nasoantorbital, the fusion of the nostril and the antorbital fenestra so characteristic for pterodactyloids, is rather large, sometimes longer than the torso, and extends above the eye, a trait only seen in other azhdarchoids; uniquely, the premaxillary bar starts out thin and becomes rapidly thick more dystally, while in other azhdarchoids it runs in a subparallel fashion. As with all azhdarchoids, teeth have been lost altogether, being replaced by a rhamphotheca sheet that covers most of the jaws, connecting the beak to the crest in the skull. Azhdarchid crests follow the two pterosaur crest models: either a bony casque, or a mainly keratinous crest supported by fibrous bone.

Generally social animals, azhdarchids can often be seen in large flocks. Azhdarchids have proportionally small brains, but displaying an astonishing level of complexity, which translates to problem solving skills. For the moment, no detailed study on pterosaur intelligence has been made, but some researchers have compared azhdarchid social behaviours to those of elphabas, Spec’s weird flying primates.

HASTAZDARCHINAE (Azhdarchs)

Hastazdarchines are common denizens on the world's grasslands, and the most diverse and successful of modern azhdarchids, large flocks being observed across the world's open spaces. Hastazhdarchines follow the classic "short jawed model", with their rostrums being spear-like and often deep. They have keratinous crests, often without a bony crest underneath, in some species falling off after the breeding season. Their claws are small and hoof-like, being of little use for climbing. Ovovivipary is common in these animals, and seems to have evolved and possibly even been lost (and redeveloped) multiple times.

Hastazdarchines seem to have diverged from their closest relatives in the Miocene, and indeed the oldest remains known so far are from the earliest Pliocene. More so than other azhdarchids they seem to have enjoyed the global cooling, their ovoviviparous tendencies allowing them to cope better with colder environments than other pterosaurs, not having the restrictions laid out by having to bury their eggs, as well as ensuring a higher clutch survival rate without sacrificing mobility. This is particularly evident in relation to plains dwelling birds like the palaeognath bastards, which need to built nests and incubate their eggs, leaving them vulnerable. It does add detrimental weight to the animal, but this is lessened out by the animals' energy saving launching and relatively small young.

Purple Azhdarch (Hastazhdarcho purpurea)

The Purple Azhdarch occurs throught the plains of the Old World, frequenting the steppes of Eurasia from Central Europe to Siberia during Spring and Summer months, and wintering as far south as Tanzania during Winter, with many areas of it's range supporting sedentary animals, such as South Africa. With a wingspan of 5 meters, it overshadows the largest rocs and gorgeese it co-exists with, with only the Rukh competing with it in terms of size, though still leagues beneath the largest living pterosaurs. Like most azhdarchids, it spends most of it's time on the ground, where it forages and rests, though it still flies frequently even as sedentary animals, be it for move around quicker, seeking new feeding or resting spots, evading predators when physical violence or galloping doesn't work, or simply for the hell of it. The Purple Azhdarch is mostly coloured in light and golden shades of brown, with violet markings of various shapes along it's back, achieved thanks to unique porphyrins. The face is also violet, with the headcrest being of a deep indigo. The beak is of a normal brown colour, while the limbs have whiteish, unpigmented coats, and the dorsal part of the wing membrane has a dark colour.

The Purple Azhdarch is a rather generalistic omnivore. The bulk of it's diet is composed of mammals, squamates, dinosaurs (particularly birds, small oviraptors, jackalopes and mattiraptors), large insects, eggs, berries, fruits, soft plants and seeds. Food, animal or plant, is generally ingested alive, the typical thick sauropsid easophagus walls rendering most claws, teeth and beaks useless to fight with, though some prey like poisonous squamates or animals with horns may be grabbed by the beaked jaws and are either suffocated or have their heads beaten against a hard object for as long as it takes, an activity hastazhdarchines are particularly suited to do thanks to their shorter and deeper jaws. Subduing large prey is harder, but a Purple Azhdarch has been recorded to capture a cheetaur, managing to suffocate it long enough for it to be too weak to fight back. The pterosaurs may also attempt to capture birds and other flying animals on the wing, particularly during migrations. Purple Azhdarchs often forage in the company of ungulipedes and other large herbivores, whose tolerance for the pterosaurs may decrease when they have young.

Purple Azhdarchs usually gather in large flocks, which thin out when the animals forage during the day, the animals walking away from the congregation as they wander around in search of food. During the breeding season, both sexes try to impress mates by raising their heads and emitting characteristic roars that may echo for miles. There is little sexual dimorphism, as interspecific competition for mates occurs in both males and females, an elaborate set of "mating groups" forming, that may either be polygynous, polyandrous or somewhere in between. These groups remain together for most of the breeding season, sleeping and foraging together, and seem to be a measure of protection for gravid female/s. Pregnancy lasts for about 2 months, the female/s then giving birth. They soon hatch, and forage around the adults, often eating insects disturbed when the adults forage. They grow rapidly for the first two years of their lives, reaching 53% of the adult size by sexual maturity, spending another five growing slowly until they reach their maximum size.

Hesper Azhdarch (Hastazhdarcho zephyrus)

A smaller cousin of the Purple Azhdarch, the Hesper Azhdarch has an average wingspan of 3.5 meters. It occurs across North America's open habitats, being some of the most common continental flyers in the landmass at flocks of thousands. It's pelage is large composed of various tones of gray, leaning towards white, with a black head and a red headcrest. While typically opportunistic, it has a preference for grasshopers and similar large insects, their breeding seasons often coinciding with the arthropods'. While the massive flocks may branch out during foraging, it is very rare to see an animal alone, being vulnerable to larger azhdarchids and avisaurs. Juveniles often perch on the backs of hmungos and other large plains dwellers, though they scarcely behave like oxpeckers, only paying attention to the largest ticks.

Yhi (Hastazhdarcho deserti)

One of the only two hastazhdarchines endemic to Australia, the Yhi sticks out like a sore thumb in a land of more bizarre pterosaurs. With a bright gold pelage, orange wing membranes and a white headcrest, the Yhi strides the outback, unbothered by the hot australian Sun. It feeds on more plant matter than most other hastazhdarchines, being an important distributor of several fruits and seeds in the scrubland; however, like all azhdarchids, the Yhi is a hazard to animals as large as the average tingamarroid. The Yhi frequently rests atop the outback's strange mountain rocks like the Uluru, wandering across the crags like alien mountain goats, though it is more than happy to rest in the lowlands. Unlike the rest of Hastazhdarcho, it is fully oviparous, the female laying eggs in a secluded mound, tended by her and her mate for 3 months, before the flaplings emerge and leave, a trait that seems to have evolved secondarily as it's closest relatives are ovoviviparous.

Partridge Azhdarch (Hastazhdarcho xenoperdix)

The Partridge Azhdarch is one of the smallest living azhdarchids at a wingspan of just 80 centimeters. Residing across the Levant and the Mediterranean, this animal bears a cryptic colouration of various browns and grays, as opposed to the more vibrant colouration of it's larger relatives. During migration, it gathers in large flocks, but it is otherwise solitary, only gathering in pairs during the breeding season. Although it has a similar omnivorous diet, it co-exists with several species of streks and bastards, often feeding on them as well.

Condor Azhdarch (Vulturazhdarcho sudamericanum)

Proffering alpine grasslands and other open highland biomes, the Condor Azhdarch is nonetheless happy to occur in lowlands with frequency: it ranges across most of western America, from the middle of the Rockies' range well into the Chilean Andes, soaring over the mountains with a four meter wingspan. It has a rather short neck with block-like neck vertebrae, similar to those of thalassodromedids except with the typical azhdarchid lack of neural spines, rendering the neck more flexible than that of most other azhdarchids. Combined with it's jaw tips being compressed into sharpened edges, the Condor Azhdarch is better suited to deal with proportionally large prey, wrestling commonly with viriosaurs and other notosuchians, bastardsloths and juvenile hadrosaurs and therizinosaurs, using the beak to bite off chunks of flesh and to bite the throat to suffocate the victim. Regardless, most of it's prey are still small animals, and it also frequently scavenges, often intimidating all but the largest harpies away from corpses.

Coloured mostly in black, with silver wing membranes and a gray-purpleish head - still covered with pycnofibrils, having no thermoregulatory use for a naked head thanks to the wing membranes as with all pterosaurs -, this pterosaur showcases perfect sexual dimorphism, with the male having a large headcrest, coloured in red alongside the beak, while the female has no crest and has the beak coloured in black. Males are more sedentary than females, keeping large territories while females fly around. Males may tolerate other males within their territory, but a clear pecking order is formed, with the dominant male always feeding first, either at normal carcasses or at kills performed by other pterosaurs within the territory, with other males and females feeding afterwards; only small prey are not regulated in this way, as they are instantly swallowed anyway. Females give birth to live young, the flaplings following her around, often feeding from her kills or on carcasses she feeds at. Juveniles between birth and two years of age seem to be exempt from the feeding hierarchy, though they sheldom aproach a non-occupied dominant male, lest he eat them instead.

Tohil (Tohilus aviserpens)

The Tohil is the second largest flying animal alive and the largest of the hastazhdarchines at a wingspan of 9 meters, and towering above four meters on the ground. True to the capacities of giant azhdarchids, it is found on nearly every landmass except remote oceanic islands and Antarctica - though it may occur as a vagrant in either -, occuring in grasslands from as far away as the american midwest as to New Zealand's highland moors; it is an efficient flying animal, managing to remain an efficient flapper at large sizes, and only impeeded by wide oceanic expanses, where the absence of thermals make the animals reluctant to go much further. Predictably, the Tohil have a diverse diet, feeding on a variety of animal species as well as browsing frequently, swallowing fruits and leaves like a pterosaurian giraffe. The male bears a deep black pelt, with gray and white bands around the shoulders and extending midway into the throat, a bright yellow beak and a bright crimson crest with an eye-like dot in it, while the female is of a simple gray, with a brown beak. Tohils, being cosmopolitian animals, breed year round, often flying large distances in search of a viable partner. Gravid females can still fly and are not significantly impaired, though they spend more time on the ground when not foraging. Tohils reach their sexual maturity at around 7 years of age.

Rhea Azhdarch (Tohilus minor)

In contrast with it's closest relative, the Rhea Azhdarch has a wingspan of just two meters. It occurs widely across South America's open environments, north and south of the Amazon. Coloured mostly in various shades of gray, with black bands in the shoulders and neck and a black beak, the Rhea Azhdarch is a frequent companion of the flocks of nandrakes that roam the continent's grasslands. Both animals have similar diets, though the Rhea Azhdarch is more carnivorous, feeding on snakes and small notosuchians with frequency. It is a rather crepuscular animal, and it is occaisonally active at night.

BUCEROSAURINAE (Calahaos)

Bucerosaurines are in some ways basically the opposite of hastazhdarchines: their jaws are long, slender and often surved, and their crests are bony casques without keratin extensions, earning them designations such as "hornbill pterosaurs". Their claws also tend to be larger and quite curved, allowing them to climb trees, hanging upside down like sloths (like other pterosaurs, the ungrasping feet don't allow for bat like suspension). Bucerosaurines first appear in the fossil record in the Eocene, and seem to have had a lot of global success during the Oligocene and Miocene, bing seemingly the dominant azhdarchids during the latter, before being largely replaced by hastazhdarchines in the Pliocene, reduced to the genus Bucerosaurus. Nonetheless, a number of species still remain, one of which Spec's largest flying vertebrate.

Golden Calahao (Bucerosaurus magnificens)

The Golden Calahao is a common denizen of the tropical rainforests of Asia. At a mere wingspan of 3 meters, it is nonetheless one of the largest rainforest residents, overshadowing all but the largest birds and elphabas. The Calahao retains the usual azhdarchid limb proportions, but while it frequently descends from the canopy to feed on the forest foor, it spends most of it's time hanging upside down like a sloth, using it's large and powerful claws to remain suspended. It is faster than a sloth, however, and certainly more active than the languours it shares the forest with: it is an omnivore, using the long azhdarchid neck and jaws to snatch fruits or small animals while otherwise standing still, allowing the Golden Calahao to forage cryptically, without being detected by predator or prey. With it's short and broad wings, it glides effordlessly across the open canopies of the asian rainforests.

The male is larger, has a more radiant pelage and a larger casque. During the breeding season, he builds a mound of vegetation amidst the branches or in a large tree cavity, where females lay their eggs. He tends to this mound for three months, removing or adding plant matter as to keep a constant temperature, before the flaplings leave.

Kongamato (Bucerosaurus africanus)

A denizen of Africa's tropical rainforests and swamps, the Kongamato is smaller at a wingspan of 2.5 meters, and bears a dark gray pelage and wing membranes, only the bright yellow or orange beak and casque standing out. It often descends to forage at the water's edge, having a prefference for aquatic snails and crustaceans, usually grabbing them from the shore rather than wading. Nonetheless, it is still mostly a frugivore, and through most of the time it is hard to detect as it forages in the deep forest.

Australian Calahao (Bucerosaurus dorfi)

The Australian Calahao is larger than it's eurasian cousins, reaching a five meter wingspan. While capable of climbing trees - and outback rocks -, it spends most of it's time on the ground, foraging in a more traditional azhdarchid manner, both on terrestrial prey and on fruits it browses from the trees; it feeds on less hard plant matter than the local hastazhdarchines, and on less animal prey than other local pterosaurs. It bears a distinctive white pelage with pink wing membranes and a metallic black beak and casque. Uniquely among calahaos, the male doesn't take part in protecting the nest, females simply laying their eggs on termite nests.

Giant Calahao (Bucerosaurus giganteus)

The largest living flying animal, the Giant Calahao reaches a wingspan of 13 meters and stands at six meters high; combined with the normal bucerosaurine slender and long jaws, it almost seems like a time shifted Quetzalcoatlus, albeit enlarged. Once thought to represent an older line of bucerosaurines, genetic evidence lists it amidst the bucerosaurines, closest to the Australian Calahao; the oldest remains date from the mid-Pleistocene, belong to a possible palaeosubspecies that was smaller by a third.

With shorter claws than it's smaller relatives, the Giant Calahao is a very aerial animal, spending most of it's non-roosting or non-foraging time flying. It can be found anywhere on Earth, appearently unfettered to geographical barriers: an individual lunching on monotremate pups in the Antarctic Peninsula may tomorrow be flying over Tanzania. Not even the open sea deters it from trasversing it, crossing the waters by flapping powerfully, covering miles in a matter of minutes. Only colder temperatures discourage the Giant Calahao, and even then the poles are rarely safe during Summer months.

While it feeds on fruits, the Giant Calahao is mostly a carnivore, using it's long jaws to snatch up animals as big as a human being from the ground. In some areas it is the apex predator, by virtue of colonising remote islands where the other large carnivores are either azhdarchids or avisaurs, and even in the mainland it only fears the larger tyrannosaurs and abelisaurs, though draks, rhynchoraptors and carnocursorines may very well bring down weak unhealthy pterosaurs.

HUIKARAE (Terrorsaurs aka Carnocursors)

Carnocursorines are the apex of azhdarchid specialization for terrestrially, being the only pterosaurs to have lost flight altogether. Genetic analysis indicate that these animals diverged from their relatives back in the Cretaceous, and the unusual Eocene form known as Murgonocheirus australis is occasionally suspected to be closely related to these animals. The first unambiguous carnocursorines appear in the Oligocene, already flightless, and the Miocene sees the expansion of this group in Oz, from small omnivores to large dromornithid-like herbivores, as well as the terrifying predators that still stalk the continent. In the case of Murgonocherius, it was an animal very similar to a "normal" chaoyangopterid, but with more thalassodromedid-like limb proportions, with longer legs and metacarpals and shorter wing fingers, as well as a longer, thicker stork-like rostrum and smaller nasantorbital. Fossils of this animal have been found in Australia and Seymour Island, dating to the Eocene. It was probably a rather azhdarchid-like animal ecologically, except more suited to wade in wetlands with it's longer, more splayed toes, and perhaps even more terrestrial, with it's shorter wing proportions.)

Curnocursorines have modified their body greatly for both flightlessness and cursoriality. The wing membranes have been completely lost, though the pneumacy of these animals remains, the forelimbs still filled with air by pulmonary airsacs. The hindlimbs have become digitigrade, an unique trait among pterosaurs, allowing the animals to gallop very efficiently: their efficient lungs, combined with the typical pterosaurian anaerobic power, allows them to gain speed very quickly, and maintaining it for very long distances, being some of the most efficient cursorial predators to have ever evolved. Some of the muscle complexes that powered their ancestors' wings still remain, allow the forelimbs to provide extra power during their strides. The wing finger has been further reduced, and has develop a keratin coating akin to a claw, functioning as a dagger either for defense against larger predators or to help subdue prey. The hand fingers have enlarged, though the metacarpals are still small and located at the end of the fourth metacarpal.

Without the need to remain relatively lightweight, the torso has been allowed to expand, have body/proportions that are less skewed than in other azhdarchids, most comparable to those of the extinct ctenochasmatoid pterosaurs. The tail has also elongated, and in the more raptorial species it has become quite robust, allowing the animals manouverabiity when chasing their prey.

All carnocursorines are viviparous, giving birth to live young through placentas. The young animals still have membranes running along their forelimbs and hind limbs, using them to glide and in at least some cases flutter, being latter absorbed by the animal as it grows. Specscientists have speculated that these glides and flutters may give an idea about how pterosaurs got into the air in the first place, and further study has been issued.

Marabou-Beak (Leptoptilorhamphus mollusciphagus)

A denizen of the wetlands of northern Australia and southern New Guinea, the Marabou-Beak wades while it's distant flying relatives, the azhdarchids, prefer the drier terrains. It's beak is, as the name implies, straight, long and deep, in terms of shape similar to that of Leptoptilos storks, although with the obvious large nasanteorbital so characteristic of pterodactyloids. The very base of the beak right above the eyes produces a small pointed bony casque, of equal size in both genders. It is about 2.5 meters tall, and it is usually coloured in soft gold with a black line along the back and a black stripe branching from it that goes across the ear opening and beneath the eye to the beak's base.

With long webbed toes, this pterosaur spends most of it's time wading in the shallows, swimming very competently if it must. It feeds on pretty much any unfortunate creature it can catch, as well as fruits and mushrooms, but it prefers above all to feed on hard shelled mollusks such as bivalves and snails. Here, the dagger-like wing finger comes in handy, prying open shells. It also allows it to defend itself from the local rhynchoraptors and crocodilians of the region.
Picture

Left: Marabou-Beak, Leptoptilorhamphus mollusciphagus (Northern Australia and Southern Papua New Guinea) Right: Bill’s Bill, Cervociconia orodromaius (Mountainous regions of Australia and Tasmania)

Generally shy, these animals live alone or in small groups even during the breeding season, though they are tolerant of other members of it's species if there is enough food. During the mating season, both genders produce long, deep calls, often accompanied by infrasound vibrations similar to those of cassowaries. Mating is a brief, poorly disputed affair, and the female is gravid for three months or so, giving birth to seven to ten young that go separate ways afterwards.

Bill’s Bill (Cervociconia orodromaius)

The Bill’s Bill is endemic to the mountainous environments of Australia and Tasmania. Isolated for several thousand years, these populations might be significantly genetically distinct, though there are few anatomical differences, if any. It is about 2 meters tall, and coloured in grayish-brown with dark brown stripes on the neck and face.

This terrorsaur has a long, slender bill, with a very large nasanteorbital fenestrate, reminiscent of those from it’s chaoyangopterid ancestors. It has a very omnivorous diet, stopping short at complex plant tissue, but it has a predilection for fungi and worms, using it’s sensitive beak to search for either in the mud. Like it’s northern relative, it also has webbed toes, though nowhere as long.

Having lost it’s wing finger, it relies exclusively on evasion to survive attacks, often diving in deep water, which is usually near it’s foraging sites anyways. It too is usually shy, rarely occurring in groups above three individuals.

Hyakarra (Carnocursor novaehollandiae)

Terrorsaur concept sketches by osmatar-d6x2vp6

Top: Hyakarra (Carnocursor novaehollandiae) (Southern New Guinea, Tasmania and Eastern Australia)Bottom: Hyakarra sprinting at lightning fast speed. Credit for the artwork belongs to Mette Aumala.

The Hyakarra is the image most associated with terrorsaurs, being the most common species on the continent. Standing at about 3 meters tall, it possess a hooked tip on it’s beak, used to rip flesh as in predatory birds, while the neck is proportionally short, with blocky neck vertebrae, as suited to deal with the stresses of hunting larger prey. It is usually white, to repel the intense midday light.

Contrary to it’s marsh and forest dwelling relatives, and like other raptorial carnocursorids, the Hyakarra’s hind-feet are quite narrow and not webbed, as appropriate for acceleration based hunting. It hunts a wide variety of Australia’s herbivorous dinosaurs, and even animals far larger than it are targeted, the pterosaur ripping off chunks of flesh with the beak and/or daggers, though obviously such feeding style is risky. It is not a pack hunter per se, but it frequently overwhelms prey in loosely organised gangs, as other predatory archosaurs do. Most of the time, however, it prefers to eat smaller animals like tingamarroids.

It is among the most widespread of the terrorsaurs, ranging from southern New Guinea to Tasmania. The only exception to their range is most of central and western Australia, however.

Greater Terrorsaur (Carnocursor giganteus)

Flightless pterosaur of doom by osmatar-d4dmb7r

Greater Terrorsaur, Carnocursor giganteus (Central and Western Australia) Credit for the artwork belongs to Mette Aumala.

The largest of Australia’s predators is the massive Greater Terrorsaur, standing at 4 meters tall. Like a larger version of the Hyakarra, it has a shorter and deeper beak, reminiscent of phorusrhacid rostrums, capable of breaking bones.

Endemic to central and western Australia, the adults are the apex predators of their ecosystem, rivaled only by the largest rhynchoraptors, most notably the strumtiger. They are less specialized than their ornithopod competitors, however: while they are capable of killing the largest herbivorous dinosaurs, they prefer to hunt smaller herbivores. Most interesting is the niche partitioning between animals of the same species: juveniles, being far more social animals, occupy a niche similar to that of the Hyakarra, while the solitary adults prefer to scavenge the kills of other predators. As such, the Hyakarra is less common in western and southern Australia, where the Greater Terrorsaur is most dominant.

Pardic Terrorsaur (Carnocursor pardus)

A denizen of the forests of Papua New Guinea, the Pardic Terrorssaur is similar in size to the Hyakarra, but bears a deeper and shorter beak like the Greater Terrorsaur. Preferring the denser rainforests, little is known about this animal's more specific habits, though it appears to select proportionally large prey.

AZHDARCHIDAE INCERTAE SEDIS

Dudu (Apteroazhdarcho novacaledonensis)

A rather strange animal, the Dudu is something of a mystery among pterosaur researchers. Endemic to the islands of Melanesia, the Dudu is a seemingly flightless azhdarchid, the adults bearing nothing but a thin brachiopatagium running along the forelimbs, only significantly large in the wing-finger, serving as a signaling device. However, juveniles not only have well developed membranes, but can fly quite well, being able to traverse easy the distance between islands, and begin to lose their ability to fly around the age of sexual maturity, which may be surprisingly delayed if conditions aren’t appropriate. Thus, the Dudu is not only well adapted to the volatile nature of island biomes, but also unique among flying vertebrates for being volant in younger stages of it’s life and flightless as an adult, a feature otherwise only observed in some deinonychosaurs. The exact placement of the Dudu in the azhdarchid tree of life is controversial, something not helped by odd atavisms like a long first metacarpal that still connects to the wrist bones, though it is thought to be closer to bucerosaurines.

The adult Dudu is a rather heavy, robust and overweight animal, a stark departure from the gracile azhdarchid bodyplan, especially that of the juveniles. With a large, deep beak, the Dudu is an omnivore, feeding primarily on fruits, fungi and small animals of all types, even stealing eggs from the dinosaurs it shares it’s habitat with like the Rath. It can best be described as Melanesia’s answer to a hogbird or a vulgure.

Dupap (Melanoazhdarcho molluscivora)

The Dupap is a black and white azhdarch endemic to the Old World. With a three meter wingspan, this animal is an unique specialist among azhdarchids, feeding on mollusks and crustaceans it finds on the shoreline: like most azhdarchids, it isn’t a good wader or swimmer due to it’s small and compact feet, though it will do so occasionally. It’s rostrum is filled with sensitive tissue, allowing it to detect borrowed prey and dig it out. It’s jaws are more robust than those of the average azhdarchid, and it does have stronger jaw muscles, allowing it to break smaller shells. Larger ones are delt with by opening them with the beak, the lower jaw curved upwards for this function.

Once thought to be an hastazhdarchine, the Dupap’s true place within Azhdarchidae is something of a mystery, though it does share with the Dudu the atavistic first metacarpal. During the mating season, both sexes bear a small red keratinous crest in the lower jaw. Females bury their eggs in sand, particularly alongside riverbanks; both parents protect the nest, distracting or harassing predators away. The young that bury their way out of the nests are left to fend for themselves.

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