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The Speculative Dinosaur Project: Azhdarchidae

August 27, 2013

Once delegated to comments on their size and occasional mentions in
non-professional literature, azhdarchids have since become one of the most
iconic lineages of pterosaurs. Well adapted to terrestrial locomotion and being
the first pterosaurs where this trait was discovered, azhdarchids have become a
window to the true nature of pterosaur diversity, not as aberrant fish eating
gliders as once thought, but as generalistic and diverse animals not unlike
terrestrial birds, some of which possibly even competing with predatory
dinosaurs. Azhdarchidae was one of the most successful and long lived lineages
of pterosaurs: at a timespan of 80 million years, azhdarchids lived almost
through half of the entire Mesozoic, and were only killed off because of the
catastrophic KT event.

Naturally, in Spec, the 80 million years had to them added another 66 million
years, but this wasn’t appearent at first. For some inexplicable reason, early
spexplorations did not found pterosaurs, returning instead simply with a
Palaeocene age skull found in a Pacific Northwest shore; this skull, named
Gigantala cranitus, was for a while thought to be the last of Spec’s
Pterosauria. However, posterior investigations found flocks of azhdarchs across
the world’s plains, and even more careful and extensive research revealed
several other types of pterosaur. While pterosaur faunas are often more subtle
than dinosaur or even mammalian faunas, azhdarchs at least are an important
component in tetrapod biotas in most landmasses, in some cases even accounting
for as much as a third of the present megafaunal biomass, and being quite
visible. It has been suspected that, for some reason, the first spexplorers
ignored or ommited pterosaurs from their research, but for the moment Hanlon’s
Razor is applied.

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 terrestriality,
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.

DIVERSITY

HASTAZDARCHINAE

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 reccorded 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, speanding 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 prefference 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)

Preffering 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

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 reccord 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.

CARNOCURSORINAE

Carnocursorines are the apex of azhdarchid specialisation for terrestriality,
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* 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.

Curnocursorines have modified their body greatly for both flightlessness and
cursoriality. The wing membranes have been completly 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 hindlimbs, 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, preffer
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
preffers above all to feed on hard shelled molluscs 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.

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 seperate 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
fenestrae, reminiscient 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 ocuring in groups above three
individuals.

Hyakarra (Carnocursor novaehollandiae)

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 huntinger larger prey. It is usually white, to repell the intense
midday light.

Contrary to it’s marsh and forest dwelling relatives, and like other raptorial
carnocursorids, the Hyakarra’s hindfeet are quite narrow and not webbed, as
apropriate for acceleration based hunting. It hunts a wide variety of
Australia’s herbivorous dinosaurs, and even animals far larger than it are
targetted, 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 preffers 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)

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, reminiscient of phorusrhacid rostrums, capable of breaking bones.

Endemic to central and western Australia, the adults are the apex predators of
their ecosystem, rivalled only by the largest rhynchoraptors. They are less
specialised than their ornithopod competitors, however: while they are capable
of killing the largest herbivorous dinosaurs, they preffer 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 preffer 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. Preffering 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 wingfinger, serving
as a signalling 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 apropriate.
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 molluscs 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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