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Draconology © VikasRao

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The reason this took so long is because I wanted to find a way to distinguish these dragons from the previous ones, but still retain some similarities so that their relatedness is obvious. In case anyone hasn't figured it out yet, the eudraconids from a previous post are based on the standard Middle Eastern/European dragon, which, although famous in pop culture, are often quite similar in terms of their morphology, behavior, & even their associated legends (evil dragon terrorizes village/hordes treasure/kidnaps maidens, is slain by a mighty hero...), with some cultural variations. But not only do dragon/flying serpent legends from many other cultures vary from simply being temperamental to entirely benevolent, they also show incredible morphological and cultural diversity. For example, although Eastern dragons are almost always depicted as the ever-popular water deity, there are plenty of examples of flying ones that are associated with the skies as well, and even within the same culture, every variant is associated with a completely unique legend. Of course, as with the last dragon post, most of these are not based on any singular dragon myth (except one), but rather common regional/cultural motifs or concepts, and I thought it fitting to make them just as diverse, and so here we are!

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With 26 species distributed across every continent except Antarctica, the Neodraconidae are the second-most diverse clade of true dragons (Eudraconia) after the Eudraconidae. As their name suggests, this is the most recent clade of dragons to have evolved, originating in the Oligocene roughly 27-28 Mya.

They are closely related to Eudraconidae, with both families being grouped together within the superfamily Eudraconoidea. Compared to eudraconids, neodraconids show a greater degree of morphological and ecological variation, with members of different subfamilies showing a high degree of dissimilarity to each other. This appears to have evolved as a means of minimizing competition with eudraconids, which were quite widespread around the time these dragons first appeared.

Anatomically, they are quite similar to their eudraconid cousins, with some species being so similar that it is nearly impossible to determine their true affinities without a more detailed study. However, there are some subtle differences. Most notably, neodraconids have a larger uropatagium, a more gracile build, denser flocculus (the region of the brain responsible for fine motor control & thus flight control), and teeth that are more conical than blade-like. Most species are also more proficient fliers than their eudraconid cousins, and smaller species therefore spend considerably more time in the air than on the ground, and hunt smaller, aerial prey unlike most eudraconids of similar size, although there are exceptions in both families.

One rather peculiar point of difference between the two is that neodraconids are in general less aggressive than eudraconids towards their own kind as well as towards many other animals. The latter aspect is likely due to the fact that they generally feed on smaller game; however, the reason for the former is unclear. It may be due to the fact that these dragons rely more on their agility & precision than brute strength compared to eudraconids, because of which even a minor limb injury or impaired vision will significantly impede their ability to hunt. As such, fights to the death are somewhat rarer among these dragons (though by no means unheard of). Although many species are more ornate than eudraconids, most still lack display structures made of soft tissue, as these can be burnt off or damaged rather easily in a combat situation involving fire. While many do possess horns or horn-like structures, unlike those of eudraconids (which are made of compacted keratin), these are mostly made of spongy bone with a thin keratinous covering. Many do however possess crests that, like those of eudraconids, start out as a keratinous sheath covering a bony structure, but eventually the underlying bone is reabsorbed leaving behind a solid, sturdy keratin crest.

The Neodraconidae are believed to have evolved either from an offshoot of the Eudraconidae or a common ancestor. Evidence from more recent studies leans more towards the latter. Both families evolved from a clade of basal eudraconoidean true dragons known as the Agniraptoridae, which first appeared in the late Cretaceous and inhabited the then insular Indian subcontinent and were among the few Mesozoic dragon lineages that escaped the K-Pg extinction due to their isolation, general small size, and unspecialized nature.

During the late Paleocene to early Eocene, as the insular subcontinent drifted northward towards Eurasia, it came in close proximity to Africa. During this time, some agniraptorids reached the continent alongside several other taxa that had until then been evolving in isolation. These African immigrants eventually gave rise to the Eudraconidae around 50 Mya. The other agniraptorids that remained isolated dispersed into Eurasia during the late Eocene to early Oligocene, once the collision between the Indian subcontinent and Eurasia had begun. This lineage eventually gave rise to the Neodraconidae around 28 Mya. But by the time these dragons reached even Asia, their eudraconid cousins were running riot across 3 continents (Africa, Europe, & Asia).

However, having evolved on a continent that had been witnessing the expansion of grasslands for millions of years, the eudraconids had become open habitat specialists. The neodraconids though had until then been evolving denser habitats such as forests and woodlands, and were thus better adapted to such habitats which at the time covered most continents, allowing them to catch up to their eudraconid cousins to some extent.

With their numerous similarities in terms of both anatomy and intelligence, competition between early representatives of both groups appears to have been rather intense. Indeed, it is only in the early Miocene that members of both groups begin to show enough differences to enable clear and accurate assignment to either clade, indicating that by this time, both had begun to develop specialized characteristics as a result of niche partitioning.

The 26 extant species of the Neodraconidae are classified into 3 subfamilies: the Longidraconinae, Neodraconinae, and Aquilasaurinae.

The Longidraconinae, also called serpentine dragons, are the smallest neodraconid subfamily, consisting of 3 species in 3 genera. They get their name from their long necks and short legs, which gives them a passing resemblance to a winged snake when in flight. These dragons often inhabit more closed habitats than most other dragons of similar size, such as dense woodland and jungles, and rather than fly, use their elongated bodies to "snake" their way through their environment. As with snakes, they are adept ambush hunters, hunting on the ground and using their long necks to strike prey from a distance. As much of their prey consists of animals much smaller than themselves, they are known to generally avoid using their fire breath unless targeting very agile prey. However, this does not mean that they rarely breath fire in general. Their long necks actually house similarly elongated fuel-producing glands, meaning that they can actually produce more flame than a dragon of similar size.

The plumed dragon (Amphipteryx kukulkanus) or amphiptere/amphithere is the only member of this clade in the Americas. With a 6-7 m wingspan and weighing roughly 55-75 kg, it is not the largest longidraconine, but is nonetheless a fearsome predator of forests and jungles throughout southern and southeastern North America, the Abya Yalan archipelago, and most of northern South America. Its elongated body allows it to easily navigate through dense woodland that most other dragons its size would avoid, and can even attack prey hidden underground.

Although they can breathe fire like all true dragons, plumed dragons almost never use their fire when hunting, likely due to their choice of small prey that can be easily subdued and which do not require a great deal of effort to digest. They will, however, burn the flesh of large carcasses when scavenging, especially because their conical, curved teeth are ill-equipped for tearing stiff meat. Such feeding habits may explain how they manage to coexist alongside other dragon species of similar, or larger, size.

Sexual dimorphism is minimal but noticeable, with males having bright red crests and females having less contrasting blue crests. Males have larger territories than females, sometimes spanning multiple islands, and mate with multiple females. Females too are known to mate with several males, especially when their own territories lie on the territorial borders of more than one male. During the breeding season that coincides with the wet season, male dragons engage in spectacular, vicious battles over the canopies of their jungle habitats. Being rather weak physically due to their lean builds, they avoid coming to blows, but do make full use of their prodigious fire-breathing capabilities, and on occasions when these fights occur shortly after a downpour, the heat from their fiery battles can cause the water accumulated on tree tops to evaporate, creating vast mists. These mists in turn create a paradise for lichens, as well as quicken cloud formation. In fact, the territorial boundaries of large males can often be identified simply by an unusual abundance of lichens.

With a wide distribution spanning much of Asia, the Asiatic serpentine dragon (Longidrakon ornatus) is similar in size to its Central American counterpart, but differs in its more ornate features and greater adaptability. In contrast to the plumed dragon, which is a forest/jungle specialist, the serpentine dragon appears to have little difficulty surviving across a range of habitats, from open woodland to wetlands and montane forests and even sparse scrubland. Considering this wide distribution, it is perhaps unsurprising that the species shows a high degree of variation across populations, from the size and shape of their cranial ornamentation to coloration, with at least 5 color morphs being known, although red and golden are the most common. Both sexes are similar in size and coloration and therefore are not very easy to distinguish. Males are notable for having a pair of elongated bony crests, which are extensions of the parietal bone, as well as a "beard" of fine skethers, similar to the beard of bearded dragons (Pogona spp), however, even these can vary considerably between populations. Both sexes possess short, fleshy tendrils, which may help navigate dense vegetation at night, similar to mammalian whiskers. Such ornamentation is rare among true dragons in general, as their fire breath would make such structures obsolete in the event of a fight, however, this species is rather unusual in its relatively peaceful disposition towards others of its kind. Outside the breeding season, territorial boundaries, even between males, are loosely defined. The breeding season of this species generally coincides with spring or monsoon, depending on where they live. During this time, several males will gather together and begin bellowing to attract a mate, emitting a deep, booming call that can be mistaken for thunder. This form of lekking behavior is rather unusual, but may have evolved as a form of defense. The serpentine dragon shares its range with the Eurasian mountain devil, which breeds at the same time of year and is far more aggressive, and is more than capable of killing this smaller, weaker species. However, adult mountain devils are solitary, and while a single male dragon is vulnerable, a group of 5-6 or larger can drive one away with their combined "firepower". This is further corroborated by the fact that populations that live near mountains form larger leks, as female mountain devils preferentially breed in mountains.

Once the females arrive, each male will attempt to impress a single female. If the female reciprocates, both engage in an elaborate aerial mating dance, after which both begin building a suitable nest. While both parents look after the eggs, only the female will care for the offspring for a few months.

With a 9-10.5 m wingspan and weighing 185-200 kg, the denwen (Hajedrakon deinopyrus) is the largest longidraconine. This large dragon inhabits most of Africa and the Middle East, and is an occasional vagrant in the Mediterranean and western Asia. Genetic evidence indicates that this species is basal to the other 2, implying that the Longidraconinae may have evolved in Africa before dispersing to Asia and the Americas, roughly 12-15 Mya. A notable characteristic of this species is its "hood", formed by flaps of skin on the sides of its neck with elongated skethers that can be flared as part of an intimidating threat display, similar to the hood of a cobra and some wyrms such as nagas. Should this threat display fail, the denwen will quickly switch to a far more devastating strategy. The denwen is notable among dragons in that it produces the hottest fire of any dragon species known. Whereas the fire of most dragons burns at roughly 800-1200 °C (similar to charcoal), the denwen produces fire that burns at a scorching 2000 °C (comparable to an acetylene blowtorch). While a direct blast from most dragons would cause instant third or fourth degree burns and char skin and flesh, a direct blast from a denwen would instantly burn straight to the bone. This particular trait allows it to take down relatively large, fast prey comparable to the kind that many larger eudraconids hunt simply using its fire breath alone, and all but negates the need for a slashing, killing bite. In Africa, it is the only dragon, and one of the few animals, capable of holding its own against the African mountain devil (Magnadrakon regius). This frighteningly hot fire is due to a high concentration of nitrocarbons in the "fuel" that this species produces. Unlike regular hydrocarbons such as ordinary lipids (fats), when these pyrophoric substances burn, they produce little to negligible amounts of water vapor, which would reduce the temperature of the flame produced. However, such a trait does have its limitations. While the denwen does have elongated fuel-producing glands as with other serpentine dragons, as much 2/3rds of each of these glands is actually dedicated specifically to producing the fuel, and only the remaining 1/3rd stores it. This is because the process of converting lipids derived from body fat to the highly volatile nitrocarbons requires considerable energy, and nitrogen is difficult to handle biologically. While ordinary lipids can be obtained simply by feeding on the fat of prey and can be produced in abundance in the body, nitrogen can only be obtained from protein, and is necessary to build protein as well; plus, an overabundance of nitrogen can easily turn fatal. As such, the denwen cannot produce enough fuel to sustain prolonged use of its fire breath, although its lethality more than makes up for this.

Through most of their range, both eudraconids and neodraconids manage to coexist with surprisingly little competition. This is primarily due to niche partitioning. While most larger eudraconids are macropredators, many of their neodraconid counterparts are piscivores, which is somewhat surprising considering the fact that despite several members of the former practicing piscivory to some extent today (especially those of the Halodraconinae subfamily), none went on to become as specialized as members of the latter, despite their longer evolutionary history. This may be related to differences in dentition between the ancestors of the 2 groups: early eudraconids (as with most of their modern descendants) possessed curved, blade-like serrated teeth ideal for slicing flesh but less effective at catching slippery fish, whereas early neodraconids possessed straighter, semi-conical teeth that were much better suited for this task. This appears to have given the latter a major head start in dominating aquatic niches. Indeed, by the time highly piscivorous eudraconids (such as the halodraconines) evolved similar adaptations, the neodraconids had already taken over this niche across most of world, thus forcing most of them to become pelagic animals, and competition with seabirds kept their diversity rather low. Today, the 9 species of the Neodraconinae are prime examples of this. Distributed across eastern Asia, Australia, and the Americas, as well as Atlantis and Lemuria, several species are often called dracolisks on account of their ornate appearance and close association with water, which resembles to a degree those of basilisk lizards. Although the majority are simply generalists that often engage in piscivory, a few are specialist piscivores that spend considerable time in and around water.

The oriental dracolisk (Ripariodrakon orientalis) is the most widespread dracolisk, distributed across wetlands, lakes, and riparian ecosystems across eastern and southeast Asia. This species is by far the most specialized piscivorous neodraconine, and may spend anywhere from half to 2/3rds of its time in water as it forages for fish, amphibians, invertebrates, and occasionally small mammals, waterbirds, and reptiles. While its vision is good enough for it to hunt underwater prey by sight, in murkier waters, it relies on its sense of smell & sensitive barbels to locate prey by sensing underwater movements. Owing to its low density (courtesy of its air sac system, hollow bones, & large lungs), this species is able to float on the water surface much like a bird. Its skethers can also be pressed tightly against the body and have microserrations that trap air, acting as a sort of life jacket. This allows the dracolisk to swim in the surface with minimal effort using its webbed hind feet. However, this makes diving underwater to catch anything rather difficult. To do so, the dracolisk swallows several (sometimes large) stones and completely empties its lungs to increase its density to the point that it sinks.

Once underwater, the oxygen stored in its blood and muscles allow to stay submerged for 5-15 minutes depending on the level of activity. To rise back up to the surface, it regurgitates the swallowed stones. Small prey is generally eaten without the need to burn it first, however, larger prey is first swallowed, and then regurgitated near land, where it is burnt and eaten in a manner typical of eudraconians. During this process, it is not uncommon for certain stones that had been churning in its stomach unregurgitated for weeks, months, and in some cases years to be spat out as well, either intentionally or otherwise. These stones, having been ground smooth and blasted by fire to the point of vitrification under the right conditions, are often highly prized by the hominins that live alongside them, mostly in the northeastern parts of the Indian subcontinent and parts of western southeast Asia that represents the westernmost extent of its range. Sometimes termed 'dragonstones', these stones are mostly prized for their ease of working (a result of partial digestion in concentrated acid) as well as their smoothness, which allows them to be made into very sharp stone daggers that can produce smoother cuts and penetrate deeper, thereby making them excellent for hunting or butchering. White dragonstones, which are sometimes called 'dragonpearls', are considered especially valuable, and are usually reserved for high-ranking tribe members, though their properties are no different from those of other stones and this is simply a matter of aesthetic value.

Sexes are similar in size, with a 5.5-7.5 m wingspan, possessing a pair of antler-like pronged bony crests, and in coloration, with leafy green skethers that aid in camouflage. However, during the breeding season, males turn a brilliant golden-yellow, making them instantly distinguishable from females. An exceptionally rare blue color morph of this species also exists. As with most animals in general, the green coloration of this dragon actually results from a combination of yellow pigment and refractive structures in their skin that reflect blue light. Blue dracolisks have a low concentration of yellow pigment, which causes their blue skin and skethers to appear as it were unfiltered. This also means that males of this color morph do not appear golden yellow, but rather a striking teal or deep cyan, as the color change in breeding males is brought on by a hormonally triggered surge in yellow pigment production. It is unclear if this limits the breeding success of such males, but as blue females are just as successful as regular green females, the gene responsible for this color morph is still prevalent in the population.

In contrast to many dragon species, mating hierarchies are mostly prevalent in females rather than males. Essentially, the females engage in generally non-lethal fights to establish dominance, and then choose the best males in the territory to mate with. Although they don't mate for life, both sexes only mate with a single partner for the duration of a breeding season. The hatching of the eggs usually coincides with the onset of the monsoon, however, if the monsoon arrives early, or if the eggs fail to hatch quickly enough that the nest is at risk of flooding, the parents will carry the eggs in their jaws as they travel to higher ground. Typically, either or both parents will look after the young for roughly 1-1.5 months.

While their dentition did grant them an edge over their eudraconid cousins, it did have its drawbacks. Most notably, it left many neodraconids unable to compete with their cousins for the role of macropredator across most of their shared range. However, in regions where eudraconid diversity is low, room for specialized niches exists, or due to a quirk of paleobiogeography, certain species have arisen to fill similar roles, often with surprising solutions. One of the best examples of this is the South American Andean dragon (Neodrakon amaroca).

This neodraconine inhabits forests and jungles though much of South America, from the foothills of the Andes to the Amazon basin as well as the southernmost Central American islands. Although it shares these habitats with a few species of forest-dwelling eudraconine dragons, it is the largest forest-dwelling dragon and indeed the largest neodraconid in South America, with a 6.5-7.5 m wingspan and weighing upto 95 kg, making it comparable in size and niche to the Asian helmeted forest dragon (Pyrolophosaurus indicus).

But while its teeth are somewhat more flattened and recurved than those of most of its relatives, they aren't quite the serrated knives of eudraconids, making them less efficient at dispatching prey in the same manner. To compensate for this, the Andean dragon has evolved 4 pairs of elongated fang-like teeth, 2 at the front of its upper jaw and 2 at the front of the lower jaw. Being an ambush hunter, the dragon typically incapacitates larger prey with a blast of fire or rushes smaller prey and, restraining it with its powerful wings and large wing talons, bites down on the neck, belly, or even head, and then pulls its head back with sufficient force to remove a significant chunk of its victim's flesh clean off (including bones or even the entire head in some cases). The fangs essentially act as ice picks when dealing with smaller prey (perforating internal organs) or aid in gripping flesh when dealing with larger prey.

The Andean dragon's technique is brutally efficient compared to even most eudraconids. A eudraconid with a similar niche and hunting strategy may take 15-30 seconds to kill its prey by slashing open its throat/flank and letting it die of blood loss; the Andean dragon however can kill its prey in 5-10 seconds. One explanation for this may be competition with other predators, particularly jaguars, which are both similar in size and occupy a broadly similar niche. Jaguars are also the among the only predators capable of killing an adult dragon, their powerful jaws and unique skull-puncturing technique enabling them to kill one should they manage to execute a successful ambush. In fact, such predation and competition, as well as factors such as intraspecific combat, may be the reason for the Andean dragon's distinctive horns. While many true dragons possess horns or other forms of cranial ornamentation, the Andean dragon is rather unusual in that its horns jut upwards rather than backwards, giving them a more than passing resemblance to mammalian ears. Most dragons though have backward-pointing horns to protect the backs of their heads from attacks by predators or other members of their own kind. Andean dragons, especially those near the Andes, often lair in or near caves. The region is volcanically active, and it is not unheard of for several nesting pairs to fly out of their lairs en masse when an earthquake or increase in volcanic activity occurs, making for a remarkable if not intimidating sight.

The largest neodraconine, as well as the largest neodraconid and one of the largest extant true dragons is the Lemurian dracolisk (Limnarchon grandis). With a 13-14 m wingspan and weighing 300-320 kg, this species is similar in size to the Eurasian and Western mountain devils, but occupies a very different ecological niche. Endemic to the highly productive aquatic ecosystems of Lemuria, this dragon has a lifestyle similar to that of the oriental dracolisk, but one taken to an extreme. The sole extant member of its genus, genetic studies indicate that it is related to, and may possibly be a descendant of the Ripariodrakon genus. It is believed that either a common ancestor or perhaps an early representative of the Ripariodrakon genus may have made its way to Lemuria sometime in the late Miocene to early Pliocene (7-5 Mya), where an abundance of prey, suitable habitats, and low competition resulted in insular gigantism. Fossil evidence suggests that the genus Limnarchon was more diverse in the past, and that a species potentially even larger than L. grandis may have existed in the past, though it is possible that this was merely a subspecies.

Although a top predator of Lemuria's aquatic ecosystems, the Lemurian dracolisk is not as specialized for an aquatic lifestyle as its Asian relatives. It lacks whisker-like barbels and instead relies on smell and sight. While it does have crocodilian-like pressure sensors on its head, these do not appear to be quite as sensitive as the barbels, though this isn't an issue as the waters it inhabits generally have greater visibility than the ones inhabited by its relatives, and as it hunts much larger prey. As with many neodraconines, the Lemurian dracolisk has teeth of unequal size, with 2 pairs of enlarged teeth located at the front of the jaws. These teeth are actually used to not just grip, but also dispatch prey by stabbing them. They're particularly effective for rapidly dispatching armored prey such as small crocodilians and large fish.

Unlike the oriental dracolisk, the Lemurian dracolisk is monogamous. Mated pairs generally share a territory, but rarely hunt together outside the breeding season. Interestingly, the male appears to share the bulk of parenting duties, from looking after the eggs to protecting flaplings, likely because the male's larger size allows it to better defend against threats such as other dragons.

As with many large dragons, this species shows ontogenic shifts, with juveniles filling different ecological niches than adults. Young flaplings generally hunt invertebrates and other small prey near the water's edge rather than in the water; being no bigger than cats, they are highly vulnerable to predation by large fish and reptiles such as crocodilians & snakes. Once they're 5 years old, they gain significant mass and begin swimming in the water. While heavy enough to sink with minimal effort, they are lightweight enough to be agile swimmers, using their grebe-like lobed feet to propel themselves underwater and their wings for bursts of great speed. Around the time they turn 15-17, their size increases rapidly disproportionate to their mass, meaning that while they continue to grow larger, they do not gain as much mass. This makes them lighter than the water and makes diving and swimming underwater more difficult, and it is around this age that they are similar in size and ecology to the oriental dracolisk. By the time they reach full size, around 25-28 years of age, their skin and some of their larger bones become thicker, which reduces their buoyancy but not to the point that they sink, though it does make the task easier. Around this age, they move to hunting in much deeper waters, as they are large enough to be able to comfortably walk along the riverbed like a gigantic wading bird, and their feet lose their hydrodynamic lobes in exchange for larger, recurved claws, which provide better traction on the riverbed. While they still need to swallow stones (sometimes hundreds if not thousands of them) to further reduce their buoyancy, their greater size and mass means that they do not need to empty their lungs and air sacs to dive, and instead can use them as oxygen reserves to dive deeper and longer, sometimes upto 1 hour as they lay still in deep water to ambush large prey, such as crocodilians, turtles, taraihonu (Barysuchus cheloniformis), and most often, large fish.

Because of their ecology and generally rare instances of combat, Lemurian dracolisks are the longest lived of all dragons. As they age, it is not unheard of for certain individuals, under the right set of conditions, to grow so large that they become flightless. At this stage however, their exclusive reliance on aquatic prey means that this does not mean a death sentence, as it does to some other dragons. Should a flapling live to adulthood, it will easily live for over a century. The oldest known individual, and indeed the oldest known dragon ever, was a dracolisk that lived to the incredible age of 230.

With 14 known species, the Aquilasaurinae are the most diverse neodraconid subfamily, and are descended from a more basal lineage of neodraconids, which split from the other subfamilies very early in the clade's evolution, roughly 22 Mya. The group gets its name from their hind feet, in which the first digit (which is highly reduced or even absent in most other dragons) is modified into a hallux similar to those of eagles and other raptors. They also have shorter tails and a particularly large uropatagium, which in many species is connected to the tail to form a deltawing structure, giving them a passing resemblance to a flying eagle. In some other species however, the uropatagium, along with the legs, form a second pair of wings, which, although cannot generate much lift, do vastly improve maneuverability.

Their most distinctive feature however, is their single, retractable sickle-like wing talon. This bears a resemblance to the sickle claws of certain dinosaurs, particularly seriemas, dromaeosaurs, & troodontids.

Members of this group are widely distributed across the tropics, from the Americas to Africa all the way to Australia, but are noticeably rare in northern latitudes, probably due to increased competition with their namesakes as well as other dragons, particularly eudraconine eudraconids in North America and varanopterine eudraconids in Asia.

Aquilasaurines occupy several ecological niches throughout their range, with size being a good indication of their niche. The smallest species are agile forest predators, medium-sized species are speedy open air hunters and the largest are terrestrial macropredators. However, the vast majority of these dragons are rather small, with most not growing much larger than a medium-sized eagle.

In stark contrast to their namesakes however, most aquilasaurines are either strictly nocturnal or cathemeral, with no known species being predominantly diurnal. This may be a means of avoiding competition with eagles and other diurnal raptors.

The smallest member of this group and the smallest neodraconid is the black dragonet (Parvidraco nigriscens). Endemic to the Andaman and Nicobar Islands, this small dragon has a wingspan of just under 90 cm, making it smaller than many of the birds and bats it lives alongside. The short, broad wings and expanded uropatagium of this dragon allow it to navigate the dense jungles of its native range as it hunts insects, small birds and bats. Being a crepuscular predator in a dense habitat, this species relies on both large eyes as well as an acute sense of hearing to navigate. A pair of large crests, extensions of the lacrimal bones, act almost like ear pinnae to focus incoming sound waves to its ears. The ear holes themselves are asymmetrically placed on the head, much like those of owls. Because of this, sound reaches one ear a fraction of a second before reaching the other, allowing the dragonet to accurately pinpoint the source of a sound.

Males and females are similar in size, and, unlike many dragon species, mate for life. Mated pairs also live together outside the breeding season, likely because their small size means that finding enough prey isn't an issue, or perhaps because it offers protection against predators such as raptors.

The spiral dragon (Aquilasaurus platycaudatus) is a typical member of this clade that is native to Australia, with a distinct subspecies in New Guinea. Its common name is derived from its highly acrobatic flying style, particularly during mating flights, during which it repeatedly flies in increasingly tight and sometimes highly erratic spirals. With a 1.5-2 m wingspan, this species is equally at home in forests and brush, but avoids open habitats with sparse tree cover such as deserts. This species is one of the fastest dragons over short distances, able to fly at an impressive speed of 160-180 km/h. However, it cannot maintain this speed for long, and therefore must rely either on ambush tactics or attack prey in confined environments where it can outmaneuver it. Its legs and large uropatagium are often held outstretched in flight, which aids in maneuverability and appear almost like a second pair of wings, though they cannot be flapped and as such do not contribute much to lift. Its tail is characterized by large, flattened skin flaps, which further aid in maneuverability much like that of a bird, compared to most other dragons whose tails act merely to counterbalance their heavy chests and heads when in flight.

Spiral dragons are predominantly crepuscular or cathemeral, although in brush and open woodlands are generally more active at night. As with most aquilasaurines, they specialize in hunting flying prey, such as birds, bats, wyverns, and even smaller dragons. Powerful muscles force flaming fuel from their mouths at high velocity, because of which, a blast of fire from these dragons is more like a flaming liquid bullet than flaming gas or liquid shot from a flamethrower. This allows them to strike a target quickly even when pursuing prey at high speed in the air.

Although typically solitary, it is not unheard of for these dragons to occasionally hunt in pairs or small groups. These generally tend to be mated pairs or related individuals, respectively, and appear to be more common in areas with a high abundance of competitors and/or predators. Hunting cooperatively allows these pint-sized dragons to better defend their kills against predators such as large varanids, land crocodiles, mammalian predators such as marsupial lions & thylacines as well as dragons, though it does necessitate more frequent hunts.

Sociality is much more common in the New Guinean subspecies (Aquilasaurus platycaudatus novaeguineae), which is roughly 30% smaller than its Australian cousin. Mated pairs generally live and together throughout the year rather than just during the breeding season, and siblings will often live together for years rather than months, until they reach adulthood around 10 years of age. This may be due to predation pressure from raptors such as Papuan eagles as well as the New Guinean giant wyvern (Haplorhynchus pennocristatus), which is often over twice the size of this dragon subspecies.

With a 5.5-7 m wingspan and weighing 55-60 kg, the spectral dragon (Spectropyrus infernaetos) is the largest aquilasaurine. This dragon inhabits southern Asia from the Indian subcontinent to the islands of southeast Asia, and is notable for its rather peculiar, if not dangerous, feeding habits. This species preys almost exclusively on other dragons, including other true dragons capable of breathing fire, in addition to wyverns, wyrms, and other large reptiles such as large snakes and small crocodilians. Mammal predation, while not unheard of, is exceptionally rare and is generally only practiced by starving individuals. A cathemeral predator active at intermittent periods during the day and night, the spectral dragon hunts in grasslands, open woodland, and mixed forest, avoiding dense forests. It's leathery wings, as with those of other dragons, wyverns, and bats, allow it to fly much more silently than birds, however, the wings of this species in particular are notable for having a sparse covering of hair-like coelofibers on the patagia. The patagia also have fine, frayed edges, and these traits are thought to further dampen any sound emitted by forceful wingbeats, thereby affording it near silent flight despite its size. This is a crucial trait considering that it hunts primarily by ambushing its oftentimes dangerous prey. Oddly enough, this dragon frequently hunts at noon, exploiting a major weakness of diurnal dragons: their inability to see directly above their heads. As with diurnal raptors, many dragons that are active during the day possess eyes that are shielded from the sun by bony ridges or are sunken into their skulls. The spectral dragon uses this to its advantage by divebombing its prey directly from above and blasting it with fire before physically impacting it with its legs. At night, the dragon typically strikes from a tree or closer to ground level, as most dragons can see rather well in moonlight but aren't blinded by it as they are by the sun. Regardless of the time of day, the end technique is always the same: prey is pinned down using the legs, the massive sickle claw is used to disable the victim via stabbing, and a fatal bite is delivered directly to the head. An enlarged pair of fang-like teeth in the upper jaw are used to stab the victim directly in the eyes, thereby puncturing them and the brain directly behind them in the process. Smaller or weakly armored prey is often killed with fire sickle claws alone or with a bite to the neck.

The slender, concave crests of this species bear a passing resemblance to mammalian ear pinnae, and appear to function in a similar way: channeling incoming sound waves to the ears. As with other aquilasaurines, spectral dragons have asymmetrical ears, and these crests, combined with an expandable hood/ruff, direct sound waves to the ears and allow them to pinpoint the source of a sound from significant distances, an ability that becomes especially useful at night.

Although not social, both parents have been known to raise their offspring together. Spectral dragons typically breed with a single partner year after year, but they aren't strictly monogamous. If one member of the pair cannot be found during the breeding season, the other will breed with another male/female. It appears that their associations are based more on territorial proximity than anything else. Although both parents share territory when looking after their offspring (a period of roughly 7-8 months), they will go their separate ways not long afterwards, back to their own individual territories. As with many medium to large dragons, this species does not appear to breed every year, which is understandable considering that prey availability may vary depending from year to year.

Draconology: True dragons pt 1

Draconology: The Eudraconia
Draconology: The Draconimorpha

Description

Draconology © VikasRao

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The reason this took so long is because I wanted to find a way to distinguish these dragons from the previous ones, but still retain some similarities so that their relatedness is obvious. In case anyone hasn't figured it out yet, the eudraconids from a previous post are based on the standard Middle Eastern/European dragon, which, although famous in pop culture, are often quite similar in terms of their morphology, behavior, & even their associated legends (evil dragon terrorizes village/hordes treasure/kidnaps maidens, is slain by a mighty hero...), with some cultural variations. But not only do dragon/flying serpent legends from many other cultures vary from simply being temperamental to entirely benevolent, they also show incredible morphological and cultural diversity. For example, although Eastern dragons are almost always depicted as the ever-popular water deity, there are plenty of examples of flying ones that are associated with the skies as well, and even within the same culture, every variant is associated with a completely unique legend. Of course, as with the last dragon post, most of these are not based on any singular dragon myth (except one), but rather common regional/cultural motifs or concepts, and I thought it fitting to make them just as diverse, and so here we are!

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With 26 species distributed across every continent except Antarctica, the Neodraconidae are the second-most diverse clade of true dragons (Eudraconia) after the Eudraconidae. As their name suggests, this is the most recent clade of dragons to have evolved, originating in the Oligocene roughly 27-28 Mya.

They are closely related to Eudraconidae, with both families being grouped together within the superfamily Eudraconoidea. Compared to eudraconids, neodraconids show a greater degree of morphological and ecological variation, with members of different subfamilies showing a high degree of dissimilarity to each other. This appears to have evolved as a means of minimizing competition with eudraconids, which were quite widespread around the time these dragons first appeared.

Anatomically, they are quite similar to their eudraconid cousins, with some species being so similar that it is nearly impossible to determine their true affinities without a more detailed study. However, there are some subtle differences. Most notably, neodraconids have a larger uropatagium, a more gracile build, denser flocculus (the region of the brain responsible for fine motor control & thus flight control), and teeth that are more conical than blade-like. Most species are also more proficient fliers than their eudraconid cousins, and smaller species therefore spend considerably more time in the air than on the ground, and hunt smaller, aerial prey unlike most eudraconids of similar size, although there are exceptions in both families.

One rather peculiar point of difference between the two is that neodraconids are in general less aggressive than eudraconids towards their own kind as well as towards many other animals. The latter aspect is likely due to the fact that they generally feed on smaller game; however, the reason for the former is unclear. It may be due to the fact that these dragons rely more on their agility & precision than brute strength compared to eudraconids, because of which even a minor limb injury or impaired vision will significantly impede their ability to hunt. As such, fights to the death are somewhat rarer among these dragons (though by no means unheard of). Although many species are more ornate than eudraconids, most still lack display structures made of soft tissue, as these can be burnt off or damaged rather easily in a combat situation involving fire. While many do possess horns or horn-like structures, unlike those of eudraconids (which are made of compacted keratin), these are mostly made of spongy bone with a thin keratinous covering. Many do however possess crests that, like those of eudraconids, start out as a keratinous sheath covering a bony structure, but eventually the underlying bone is reabsorbed leaving behind a solid, sturdy keratin crest.

The Neodraconidae are believed to have evolved either from an offshoot of the Eudraconidae or a common ancestor. Evidence from more recent studies leans more towards the latter. Both families evolved from a clade of basal eudraconoidean true dragons known as the Agniraptoridae, which first appeared in the late Cretaceous and inhabited the then insular Indian subcontinent and were among the few Mesozoic dragon lineages that escaped the K-Pg extinction due to their isolation, general small size, and unspecialized nature.

During the late Paleocene to early Eocene, as the insular subcontinent drifted northward towards Eurasia, it came in close proximity to Africa. During this time, some agniraptorids reached the continent alongside several other taxa that had until then been evolving in isolation. These African immigrants eventually gave rise to the Eudraconidae around 50 Mya. The other agniraptorids that remained isolated dispersed into Eurasia during the late Eocene to early Oligocene, once the collision between the Indian subcontinent and Eurasia had begun. This lineage eventually gave rise to the Neodraconidae around 28 Mya. But by the time these dragons reached even Asia, their eudraconid cousins were running riot across 3 continents (Africa, Europe, & Asia).

However, having evolved on a continent that had been witnessing the expansion of grasslands for millions of years, the eudraconids had become open habitat specialists. The neodraconids though had until then been evolving denser habitats such as forests and woodlands, and were thus better adapted to such habitats which at the time covered most continents, allowing them to catch up to their eudraconid cousins to some extent.

With their numerous similarities in terms of both anatomy and intelligence, competition between early representatives of both groups appears to have been rather intense. Indeed, it is only in the early Miocene that members of both groups begin to show enough differences to enable clear and accurate assignment to either clade, indicating that by this time, both had begun to develop specialized characteristics as a result of niche partitioning.

The 26 extant species of the Neodraconidae are classified into 3 subfamilies: the Longidraconinae, Neodraconinae, and Aquilasaurinae.

The Longidraconinae, also called serpentine dragons, are the smallest neodraconid subfamily, consisting of 3 species in 3 genera. They get their name from their long necks and short legs, which gives them a passing resemblance to a winged snake when in flight. These dragons often inhabit more closed habitats than most other dragons of similar size, such as dense woodland and jungles, and rather than fly, use their elongated bodies to "snake" their way through their environment. As with snakes, they are adept ambush hunters, hunting on the ground and using their long necks to strike prey from a distance. As much of their prey consists of animals much smaller than themselves, they are known to generally avoid using their fire breath unless targeting very agile prey. However, this does not mean that they rarely breath fire in general. Their long necks actually house similarly elongated fuel-producing glands, meaning that they can actually produce more flame than a dragon of similar size.

The plumed dragon (Amphipteryx kukulkanus) or amphiptere/amphithere is the only member of this clade in the Americas. With a 6-7 m wingspan and weighing roughly 55-75 kg, it is not the largest longidraconine, but is nonetheless a fearsome predator of forests and jungles throughout southern and southeastern North America, the Abya Yalan archipelago, and most of northern South America. Its elongated body allows it to easily navigate through dense woodland that most other dragons its size would avoid, and can even attack prey hidden underground.

Although they can breathe fire like all true dragons, plumed dragons almost never use their fire when hunting, likely due to their choice of small prey that can be easily subdued and which do not require a great deal of effort to digest. They will, however, burn the flesh of large carcasses when scavenging, especially because their conical, curved teeth are ill-equipped for tearing stiff meat. Such feeding habits may explain how they manage to coexist alongside other dragon species of similar, or larger, size.

Sexual dimorphism is minimal but noticeable, with males having bright red crests and females having less contrasting blue crests. Males have larger territories than females, sometimes spanning multiple islands, and mate with multiple females. Females too are known to mate with several males, especially when their own territories lie on the territorial borders of more than one male. During the breeding season that coincides with the wet season, male dragons engage in spectacular, vicious battles over the canopies of their jungle habitats. Being rather weak physically due to their lean builds, they avoid coming to blows, but do make full use of their prodigious fire-breathing capabilities, and on occasions when these fights occur shortly after a downpour, the heat from their fiery battles can cause the water accumulated on tree tops to evaporate, creating vast mists. These mists in turn create a paradise for lichens, as well as quicken cloud formation. In fact, the territorial boundaries of large males can often be identified simply by an unusual abundance of lichens.

With a wide distribution spanning much of Asia, the Asiatic serpentine dragon (Longidrakon ornatus) is similar in size to its Central American counterpart, but differs in its more ornate features and greater adaptability. In contrast to the plumed dragon, which is a forest/jungle specialist, the serpentine dragon appears to have little difficulty surviving across a range of habitats, from open woodland to wetlands and montane forests and even sparse scrubland. Considering this wide distribution, it is perhaps unsurprising that the species shows a high degree of variation across populations, from the size and shape of their cranial ornamentation to coloration, with at least 5 color morphs being known, although red and golden are the most common. Both sexes are similar in size and coloration and therefore are not very easy to distinguish. Males are notable for having a pair of elongated bony crests, which are extensions of the parietal bone, as well as a "beard" of fine skethers, similar to the beard of bearded dragons (Pogona spp), however, even these can vary considerably between populations. Both sexes possess short, fleshy tendrils, which may help navigate dense vegetation at night, similar to mammalian whiskers. Such ornamentation is rare among true dragons in general, as their fire breath would make such structures obsolete in the event of a fight, however, this species is rather unusual in its relatively peaceful disposition towards others of its kind. Outside the breeding season, territorial boundaries, even between males, are loosely defined. The breeding season of this species generally coincides with spring or monsoon, depending on where they live. During this time, several males will gather together and begin bellowing to attract a mate, emitting a deep, booming call that can be mistaken for thunder. This form of lekking behavior is rather unusual, but may have evolved as a form of defense. The serpentine dragon shares its range with the Eurasian mountain devil, which breeds at the same time of year and is far more aggressive, and is more than capable of killing this smaller, weaker species. However, adult mountain devils are solitary, and while a single male dragon is vulnerable, a group of 5-6 or larger can drive one away with their combined "firepower". This is further corroborated by the fact that populations that live near mountains form larger leks, as female mountain devils preferentially breed in mountains.

Once the females arrive, each male will attempt to impress a single female. If the female reciprocates, both engage in an elaborate aerial mating dance, after which both begin building a suitable nest. While both parents look after the eggs, only the female will care for the offspring for a few months.

With a 9-10.5 m wingspan and weighing 185-200 kg, the denwen (Hajedrakon deinopyrus) is the largest longidraconine. This large dragon inhabits most of Africa and the Middle East, and is an occasional vagrant in the Mediterranean and western Asia. Genetic evidence indicates that this species is basal to the other 2, implying that the Longidraconinae may have evolved in Africa before dispersing to Asia and the Americas, roughly 12-15 Mya. A notable characteristic of this species is its "hood", formed by flaps of skin on the sides of its neck with elongated skethers that can be flared as part of an intimidating threat display, similar to the hood of a cobra and some wyrms such as nagas. Should this threat display fail, the denwen will quickly switch to a far more devastating strategy. The denwen is notable among dragons in that it produces the hottest fire of any dragon species known. Whereas the fire of most dragons burns at roughly 800-1200 °C (similar to charcoal), the denwen produces fire that burns at a scorching 2000 °C (comparable to an acetylene blowtorch). While a direct blast from most dragons would cause instant third or fourth degree burns and char skin and flesh, a direct blast from a denwen would instantly burn straight to the bone. This particular trait allows it to take down relatively large, fast prey comparable to the kind that many larger eudraconids hunt simply using its fire breath alone, and all but negates the need for a slashing, killing bite. In Africa, it is the only dragon, and one of the few animals, capable of holding its own against the African mountain devil (Magnadrakon regius). This frighteningly hot fire is due to a high concentration of nitrocarbons in the "fuel" that this species produces. Unlike regular hydrocarbons such as ordinary lipids (fats), when these pyrophoric substances burn, they produce little to negligible amounts of water vapor, which would reduce the temperature of the flame produced. However, such a trait does have its limitations. While the denwen does have elongated fuel-producing glands as with other serpentine dragons, as much 2/3rds of each of these glands is actually dedicated specifically to producing the fuel, and only the remaining 1/3rd stores it. This is because the process of converting lipids derived from body fat to the highly volatile nitrocarbons requires considerable energy, and nitrogen is difficult to handle biologically. While ordinary lipids can be obtained simply by feeding on the fat of prey and can be produced in abundance in the body, nitrogen can only be obtained from protein, and is necessary to build protein as well; plus, an overabundance of nitrogen can easily turn fatal. As such, the denwen cannot produce enough fuel to sustain prolonged use of its fire breath, although its lethality more than makes up for this.

Through most of their range, both eudraconids and neodraconids manage to coexist with surprisingly little competition. This is primarily due to niche partitioning. While most larger eudraconids are macropredators, many of their neodraconid counterparts are piscivores, which is somewhat surprising considering the fact that despite several members of the former practicing piscivory to some extent today (especially those of the Halodraconinae subfamily), none went on to become as specialized as members of the latter, despite their longer evolutionary history. This may be related to differences in dentition between the ancestors of the 2 groups: early eudraconids (as with most of their modern descendants) possessed curved, blade-like serrated teeth ideal for slicing flesh but less effective at catching slippery fish, whereas early neodraconids possessed straighter, semi-conical teeth that were much better suited for this task. This appears to have given the latter a major head start in dominating aquatic niches. Indeed, by the time highly piscivorous eudraconids (such as the halodraconines) evolved similar adaptations, the neodraconids had already taken over this niche across most of world, thus forcing most of them to become pelagic animals, and competition with seabirds kept their diversity rather low. Today, the 9 species of the Neodraconinae are prime examples of this. Distributed across eastern Asia, Australia, and the Americas, as well as Atlantis and Lemuria, several species are often called dracolisks on account of their ornate appearance and close association with water, which resembles to a degree those of basilisk lizards. Although the majority are simply generalists that often engage in piscivory, a few are specialist piscivores that spend considerable time in and around water.

The oriental dracolisk (Ripariodrakon orientalis) is the most widespread dracolisk, distributed across wetlands, lakes, and riparian ecosystems across eastern and southeast Asia. This species is by far the most specialized piscivorous neodraconine, and may spend anywhere from half to 2/3rds of its time in water as it forages for fish, amphibians, invertebrates, and occasionally small mammals, waterbirds, and reptiles. While its vision is good enough for it to hunt underwater prey by sight, in murkier waters, it relies on its sense of smell & sensitive barbels to locate prey by sensing underwater movements. Owing to its low density (courtesy of its air sac system, hollow bones, & large lungs), this species is able to float on the water surface much like a bird. Its skethers can also be pressed tightly against the body and have microserrations that trap air, acting as a sort of life jacket. This allows the dracolisk to swim in the surface with minimal effort using its webbed hind feet. However, this makes diving underwater to catch anything rather difficult. To do so, the dracolisk swallows several (sometimes large) stones and completely empties its lungs to increase its density to the point that it sinks.

Once underwater, the oxygen stored in its blood and muscles allow to stay submerged for 5-15 minutes depending on the level of activity. To rise back up to the surface, it regurgitates the swallowed stones. Small prey is generally eaten without the need to burn it first, however, larger prey is first swallowed, and then regurgitated near land, where it is burnt and eaten in a manner typical of eudraconians. During this process, it is not uncommon for certain stones that had been churning in its stomach unregurgitated for weeks, months, and in some cases years to be spat out as well, either intentionally or otherwise. These stones, having been ground smooth and blasted by fire to the point of vitrification under the right conditions, are often highly prized by the hominins that live alongside them, mostly in the northeastern parts of the Indian subcontinent and parts of western southeast Asia that represents the westernmost extent of its range. Sometimes termed 'dragonstones', these stones are mostly prized for their ease of working (a result of partial digestion in concentrated acid) as well as their smoothness, which allows them to be made into very sharp stone daggers that can produce smoother cuts and penetrate deeper, thereby making them excellent for hunting or butchering. White dragonstones, which are sometimes called 'dragonpearls', are considered especially valuable, and are usually reserved for high-ranking tribe members, though their properties are no different from those of other stones and this is simply a matter of aesthetic value.

Sexes are similar in size, with a 5.5-7.5 m wingspan, possessing a pair of antler-like pronged bony crests, and in coloration, with leafy green skethers that aid in camouflage. However, during the breeding season, males turn a brilliant golden-yellow, making them instantly distinguishable from females. An exceptionally rare blue color morph of this species also exists. As with most animals in general, the green coloration of this dragon actually results from a combination of yellow pigment and refractive structures in their skin that reflect blue light. Blue dracolisks have a low concentration of yellow pigment, which causes their blue skin and skethers to appear as it were unfiltered. This also means that males of this color morph do not appear golden yellow, but rather a striking teal or deep cyan, as the color change in breeding males is brought on by a hormonally triggered surge in yellow pigment production. It is unclear if this limits the breeding success of such males, but as blue females are just as successful as regular green females, the gene responsible for this color morph is still prevalent in the population.

In contrast to many dragon species, mating hierarchies are mostly prevalent in females rather than males. Essentially, the females engage in generally non-lethal fights to establish dominance, and then choose the best males in the territory to mate with. Although they don't mate for life, both sexes only mate with a single partner for the duration of a breeding season. The hatching of the eggs usually coincides with the onset of the monsoon, however, if the monsoon arrives early, or if the eggs fail to hatch quickly enough that the nest is at risk of flooding, the parents will carry the eggs in their jaws as they travel to higher ground. Typically, either or both parents will look after the young for roughly 1-1.5 months.

While their dentition did grant them an edge over their eudraconid cousins, it did have its drawbacks. Most notably, it left many neodraconids unable to compete with their cousins for the role of macropredator across most of their shared range. However, in regions where eudraconid diversity is low, room for specialized niches exists, or due to a quirk of paleobiogeography, certain species have arisen to fill similar roles, often with surprising solutions. One of the best examples of this is the South American Andean dragon (Neodrakon amaroca).

This neodraconine inhabits forests and jungles though much of South America, from the foothills of the Andes to the Amazon basin as well as the southernmost Central American islands. Although it shares these habitats with a few species of forest-dwelling eudraconine dragons, it is the largest forest-dwelling dragon and indeed the largest neodraconid in South America, with a 6.5-7.5 m wingspan and weighing upto 95 kg, making it comparable in size and niche to the Asian helmeted forest dragon (Pyrolophosaurus indicus).

But while its teeth are somewhat more flattened and recurved than those of most of its relatives, they aren't quite the serrated knives of eudraconids, making them less efficient at dispatching prey in the same manner. To compensate for this, the Andean dragon has evolved 4 pairs of elongated fang-like teeth, 2 at the front of its upper jaw and 2 at the front of the lower jaw. Being an ambush hunter, the dragon typically incapacitates larger prey with a blast of fire or rushes smaller prey and, restraining it with its powerful wings and large wing talons, bites down on the neck, belly, or even head, and then pulls its head back with sufficient force to remove a significant chunk of its victim's flesh clean off (including bones or even the entire head in some cases). The fangs essentially act as ice picks when dealing with smaller prey (perforating internal organs) or aid in gripping flesh when dealing with larger prey.

The Andean dragon's technique is brutally efficient compared to even most eudraconids. A eudraconid with a similar niche and hunting strategy may take 15-30 seconds to kill its prey by slashing open its throat/flank and letting it die of blood loss; the Andean dragon however can kill its prey in 5-10 seconds. One explanation for this may be competition with other predators, particularly jaguars, which are both similar in size and occupy a broadly similar niche. Jaguars are also the among the only predators capable of killing an adult dragon, their powerful jaws and unique skull-puncturing technique enabling them to kill one should they manage to execute a successful ambush. In fact, such predation and competition, as well as factors such as intraspecific combat, may be the reason for the Andean dragon's distinctive horns. While many true dragons possess horns or other forms of cranial ornamentation, the Andean dragon is rather unusual in that its horns jut upwards rather than backwards, giving them a more than passing resemblance to mammalian ears. Most dragons though have backward-pointing horns to protect the backs of their heads from attacks by predators or other members of their own kind. Andean dragons, especially those near the Andes, often lair in or near caves. The region is volcanically active, and it is not unheard of for several nesting pairs to fly out of their lairs en masse when an earthquake or increase in volcanic activity occurs, making for a remarkable if not intimidating sight.

The largest neodraconine, as well as the largest neodraconid and one of the largest extant true dragons is the Lemurian dracolisk (Limnarchon grandis). With a 13-14 m wingspan and weighing 300-320 kg, this species is similar in size to the Eurasian and Western mountain devils, but occupies a very different ecological niche. Endemic to the highly productive aquatic ecosystems of Lemuria, this dragon has a lifestyle similar to that of the oriental dracolisk, but one taken to an extreme. The sole extant member of its genus, genetic studies indicate that it is related to, and may possibly be a descendant of the Ripariodrakon genus. It is believed that either a common ancestor or perhaps an early representative of the Ripariodrakon genus may have made its way to Lemuria sometime in the late Miocene to early Pliocene (7-5 Mya), where an abundance of prey, suitable habitats, and low competition resulted in insular gigantism. Fossil evidence suggests that the genus Limnarchon was more diverse in the past, and that a species potentially even larger than L. grandis may have existed in the past, though it is possible that this was merely a subspecies.

Although a top predator of Lemuria's aquatic ecosystems, the Lemurian dracolisk is not as specialized for an aquatic lifestyle as its Asian relatives. It lacks whisker-like barbels and instead relies on smell and sight. While it does have crocodilian-like pressure sensors on its head, these do not appear to be quite as sensitive as the barbels, though this isn't an issue as the waters it inhabits generally have greater visibility than the ones inhabited by its relatives, and as it hunts much larger prey. As with many neodraconines, the Lemurian dracolisk has teeth of unequal size, with 2 pairs of enlarged teeth located at the front of the jaws. These teeth are actually used to not just grip, but also dispatch prey by stabbing them. They're particularly effective for rapidly dispatching armored prey such as small crocodilians and large fish.

Unlike the oriental dracolisk, the Lemurian dracolisk is monogamous. Mated pairs generally share a territory, but rarely hunt together outside the breeding season. Interestingly, the male appears to share the bulk of parenting duties, from looking after the eggs to protecting flaplings, likely because the male's larger size allows it to better defend against threats such as other dragons.

As with many large dragons, this species shows ontogenic shifts, with juveniles filling different ecological niches than adults. Young flaplings generally hunt invertebrates and other small prey near the water's edge rather than in the water; being no bigger than cats, they are highly vulnerable to predation by large fish and reptiles such as crocodilians & snakes. Once they're 5 years old, they gain significant mass and begin swimming in the water. While heavy enough to sink with minimal effort, they are lightweight enough to be agile swimmers, using their grebe-like lobed feet to propel themselves underwater and their wings for bursts of great speed. Around the time they turn 15-17, their size increases rapidly disproportionate to their mass, meaning that while they continue to grow larger, they do not gain as much mass. This makes them lighter than the water and makes diving and swimming underwater more difficult, and it is around this age that they are similar in size and ecology to the oriental dracolisk. By the time they reach full size, around 25-28 years of age, their skin and some of their larger bones become thicker, which reduces their buoyancy but not to the point that they sink, though it does make the task easier. Around this age, they move to hunting in much deeper waters, as they are large enough to be able to comfortably walk along the riverbed like a gigantic wading bird, and their feet lose their hydrodynamic lobes in exchange for larger, recurved claws, which provide better traction on the riverbed. While they still need to swallow stones (sometimes hundreds if not thousands of them) to further reduce their buoyancy, their greater size and mass means that they do not need to empty their lungs and air sacs to dive, and instead can use them as oxygen reserves to dive deeper and longer, sometimes upto 1 hour as they lay still in deep water to ambush large prey, such as crocodilians, turtles, taraihonu (Barysuchus cheloniformis), and most often, large fish.

Because of their ecology and generally rare instances of combat, Lemurian dracolisks are the longest lived of all dragons. As they age, it is not unheard of for certain individuals, under the right set of conditions, to grow so large that they become flightless. At this stage however, their exclusive reliance on aquatic prey means that this does not mean a death sentence, as it does to some other dragons. Should a flapling live to adulthood, it will easily live for over a century. The oldest known individual, and indeed the oldest known dragon ever, was a dracolisk that lived to the incredible age of 230.

With 14 known species, the Aquilasaurinae are the most diverse neodraconid subfamily, and are descended from a more basal lineage of neodraconids, which split from the other subfamilies very early in the clade's evolution, roughly 22 Mya. The group gets its name from their hind feet, in which the first digit (which is highly reduced or even absent in most other dragons) is modified into a hallux similar to those of eagles and other raptors. They also have shorter tails and a particularly large uropatagium, which in many species is connected to the tail to form a deltawing structure, giving them a passing resemblance to a flying eagle. In some other species however, the uropatagium, along with the legs, form a second pair of wings, which, although cannot generate much lift, do vastly improve maneuverability.

Their most distinctive feature however, is their single, retractable sickle-like wing talon. This bears a resemblance to the sickle claws of certain dinosaurs, particularly seriemas, dromaeosaurs, & troodontids.

Members of this group are widely distributed across the tropics, from the Americas to Africa all the way to Australia, but are noticeably rare in northern latitudes, probably due to increased competition with their namesakes as well as other dragons, particularly eudraconine eudraconids in North America and varanopterine eudraconids in Asia.

Aquilasaurines occupy several ecological niches throughout their range, with size being a good indication of their niche. The smallest species are agile forest predators, medium-sized species are speedy open air hunters and the largest are terrestrial macropredators. However, the vast majority of these dragons are rather small, with most not growing much larger than a medium-sized eagle.

In stark contrast to their namesakes however, most aquilasaurines are either strictly nocturnal or cathemeral, with no known species being predominantly diurnal. This may be a means of avoiding competition with eagles and other diurnal raptors.

The smallest member of this group and the smallest neodraconid is the black dragonet (Parvidraco nigriscens). Endemic to the Andaman and Nicobar Islands, this small dragon has a wingspan of just under 90 cm, making it smaller than many of the birds and bats it lives alongside. The short, broad wings and expanded uropatagium of this dragon allow it to navigate the dense jungles of its native range as it hunts insects, small birds and bats. Being a crepuscular predator in a dense habitat, this species relies on both large eyes as well as an acute sense of hearing to navigate. A pair of large crests, extensions of the lacrimal bones, act almost like ear pinnae to focus incoming sound waves to its ears. The ear holes themselves are asymmetrically placed on the head, much like those of owls. Because of this, sound reaches one ear a fraction of a second before reaching the other, allowing the dragonet to accurately pinpoint the source of a sound.

Males and females are similar in size, and, unlike many dragon species, mate for life. Mated pairs also live together outside the breeding season, likely because their small size means that finding enough prey isn't an issue, or perhaps because it offers protection against predators such as raptors.

The spiral dragon (Aquilasaurus platycaudatus) is a typical member of this clade that is native to Australia, with a distinct subspecies in New Guinea. Its common name is derived from its highly acrobatic flying style, particularly during mating flights, during which it repeatedly flies in increasingly tight and sometimes highly erratic spirals. With a 1.5-2 m wingspan, this species is equally at home in forests and brush, but avoids open habitats with sparse tree cover such as deserts. This species is one of the fastest dragons over short distances, able to fly at an impressive speed of 160-180 km/h. However, it cannot maintain this speed for long, and therefore must rely either on ambush tactics or attack prey in confined environments where it can outmaneuver it. Its legs and large uropatagium are often held outstretched in flight, which aids in maneuverability and appear almost like a second pair of wings, though they cannot be flapped and as such do not contribute much to lift. Its tail is characterized by large, flattened skin flaps, which further aid in maneuverability much like that of a bird, compared to most other dragons whose tails act merely to counterbalance their heavy chests and heads when in flight.

Spiral dragons are predominantly crepuscular or cathemeral, although in brush and open woodlands are generally more active at night. As with most aquilasaurines, they specialize in hunting flying prey, such as birds, bats, wyverns, and even smaller dragons. Powerful muscles force flaming fuel from their mouths at high velocity, because of which, a blast of fire from these dragons is more like a flaming liquid bullet than flaming gas or liquid shot from a flamethrower. This allows them to strike a target quickly even when pursuing prey at high speed in the air.

Although typically solitary, it is not unheard of for these dragons to occasionally hunt in pairs or small groups. These generally tend to be mated pairs or related individuals, respectively, and appear to be more common in areas with a high abundance of competitors and/or predators. Hunting cooperatively allows these pint-sized dragons to better defend their kills against predators such as large varanids, land crocodiles, mammalian predators such as marsupial lions & thylacines as well as dragons, though it does necessitate more frequent hunts.

Sociality is much more common in the New Guinean subspecies (Aquilasaurus platycaudatus novaeguineae), which is roughly 30% smaller than its Australian cousin. Mated pairs generally live and together throughout the year rather than just during the breeding season, and siblings will often live together for years rather than months, until they reach adulthood around 10 years of age. This may be due to predation pressure from raptors such as Papuan eagles as well as the New Guinean giant wyvern (Haplorhynchus pennocristatus), which is often over twice the size of this dragon subspecies.

With a 5.5-7 m wingspan and weighing 55-60 kg, the spectral dragon (Spectropyrus infernaetos) is the largest aquilasaurine. This dragon inhabits southern Asia from the Indian subcontinent to the islands of southeast Asia, and is notable for its rather peculiar, if not dangerous, feeding habits. This species preys almost exclusively on other dragons, including other true dragons capable of breathing fire, in addition to wyverns, wyrms, and other large reptiles such as large snakes and small crocodilians. Mammal predation, while not unheard of, is exceptionally rare and is generally only practiced by starving individuals. A cathemeral predator active at intermittent periods during the day and night, the spectral dragon hunts in grasslands, open woodland, and mixed forest, avoiding dense forests. It's leathery wings, as with those of other dragons, wyverns, and bats, allow it to fly much more silently than birds, however, the wings of this species in particular are notable for having a sparse covering of hair-like coelofibers on the patagia. The patagia also have fine, frayed edges, and these traits are thought to further dampen any sound emitted by forceful wingbeats, thereby affording it near silent flight despite its size. This is a crucial trait considering that it hunts primarily by ambushing its oftentimes dangerous prey. Oddly enough, this dragon frequently hunts at noon, exploiting a major weakness of diurnal dragons: their inability to see directly above their heads. As with diurnal raptors, many dragons that are active during the day possess eyes that are shielded from the sun by bony ridges or are sunken into their skulls. The spectral dragon uses this to its advantage by divebombing its prey directly from above and blasting it with fire before physically impacting it with its legs. At night, the dragon typically strikes from a tree or closer to ground level, as most dragons can see rather well in moonlight but aren't blinded by it as they are by the sun. Regardless of the time of day, the end technique is always the same: prey is pinned down using the legs, the massive sickle claw is used to disable the victim via stabbing, and a fatal bite is delivered directly to the head. An enlarged pair of fang-like teeth in the upper jaw are used to stab the victim directly in the eyes, thereby puncturing them and the brain directly behind them in the process. Smaller or weakly armored prey is often killed with fire sickle claws alone or with a bite to the neck.

The slender, concave crests of this species bear a passing resemblance to mammalian ear pinnae, and appear to function in a similar way: channeling incoming sound waves to the ears. As with other aquilasaurines, spectral dragons have asymmetrical ears, and these crests, combined with an expandable hood/ruff, direct sound waves to the ears and allow them to pinpoint the source of a sound from significant distances, an ability that becomes especially useful at night.

Although not social, both parents have been known to raise their offspring together. Spectral dragons typically breed with a single partner year after year, but they aren't strictly monogamous. If one member of the pair cannot be found during the breeding season, the other will breed with another male/female. It appears that their associations are based more on territorial proximity than anything else. Although both parents share territory when looking after their offspring (a period of roughly 7-8 months), they will go their separate ways not long afterwards, back to their own individual territories. As with many medium to large dragons, this species does not appear to breed every year, which is understandable considering that prey availability may vary depending from year to year.

Draconology: True dragons pt 1

Draconology: The Eudraconia
Draconology: The Draconimorpha