Lindwyrmus nychognathus

Lindworms, also know as "Chinese dragons" and "lungs" are medium to large-sized creatures with short, powerful forearms but lack hindlimbs entirely. Although secretive and rarely seen, it's clear that lindworms are totally flightless, having no wings to speak of. At first glance, that fact may imply their relationship with dragons, rather than wyverns, as dragons have both forelimbs and wings. However, preliminary genetic comparisons between a dead lindworm recovered from China (in press) and a living wyvern and dragon show that, in fact, lindworms are genetically closer to wyverns than dragons. The obvious taxonomic implications are that lindworms are stem wyverns, sitting somewhere below true wyverns on the archosaur family tree, branching off before wyverns developed wings.

Given their rarity, lindworms are rarely studied. In fact, the first named wyvern is Lindwyrmus nychognathus from Argentina, and it was described only recently (Farke, 2005). Although it features plenty of its own peculiar anatomical features, Lindwyrmus is generally viewed as diagnostic of the entire group. Most impressively, Lindwyrmus is semi-arboreal, rather than semi-aquatic as lindworms are usually thought to be. Farke noted that the bright-green lindworm curls its serpentine body on a branch as would an anaconda or constrictor. The arms are used to help pull the animal up through the trees and clamber across the canopy. While the body is bright green, Lindwyrmus has bright orange flaps of skin on the arms and along the spine. These structures are usually draped across the body or arms, but when aggravated, the lindworm can "flare them up" and create a striking threat display. In fact, the flaps are structured by thin muscular rods akin to the actinofibrils of pterosaur wings.

The skull of Lindwyrmus is very distinctive. There are two pairs of cranial horns which point posteriorally behind the eyes. Two small nasal horns rise from the tip of the snout. The skull is dorsoventrally flattened. Unlike most dragons and wyverns, there is no "lip" around the teeth. Instead, the teeth erupt from the underlying skin as in crocodilians. Most interestingly, however, is the pair of laterally-placed, spike-like mandibles that boarder the mouth. These mandibles are mobile in two places: at the point of articulation with the skull itself and immediately behind the spike-like process. This novel structure seems to serve two purposes: first, it greatly increases the effect of the animal's threat display, and second, the mandibles actually impale prey while the mouth grips it. Bizarrely, the main articulation of the mandibles is a cup-like depression in the jaw just above the mandibular joint, and the proximal surface of the mandible is a ball joint. However, in life, the mandibles are limited to fore and aft movement. Exactly what environmental forces combined to create the need for such a strange structure is not understood. Incidentally, while the mouth is closed, the mandibles are held forward, overlying the exposed teeth and framing the face.

Lindwyrmus subsists on small vertebrates almost exclusively, though Harrison (2007), on a trip to Argentina's jungles, witnessed a pair of lindworms feasting on a dead hog. He was unable to determine whether the lindworms actually killed the hog or were simply scavenging on it. Both he and Farke were unable to tell males from females, and Harrison commented that, generally, lindworms seemed to be solitary animals. Neither man saw juvenile animals, which is unfortunate. It would interesting to see whether juveniles and subadults also posess the strange mandibular array that adults do. Farke estimated the total length of an adult Lindwyrmus at "about twenty feet," though he left open the possibility that larger individuals existed.

Farke said a recent draconology conference that he intends to travel back to Argentina to further observe Lindwyrmus and hopefully find some juvenile animals. Other workers, perhaps inspired by Farke's work with Lindwyrmus, are traveling to Asia to track down other species of lindworm. Hopefully, this little-known branch of the wyvern family tree will soon be better understood!

Farke, A. (2005). The first description of a "lindworm," from the jungles of Argentina. European Journal of Draconology 106(2): 245-258.

Harrison, M. A. (2007). Observations on Lindwyrmus nychognathus (Farke 2005). Draconium 48(3): 402-405.

Jugoceras horriblus

The largest and most ferocious modern wyvern, Jugoceras horriblus (Northrop, 1985) lives alongside two dragons, the basal dracolympian Spinodracos dysonii and the bizarre cat mimic, Felimimus paradoxus. By all accounts, Jugoceras does not relish the company, and chance encounters between it and Spinodracos are violent and brutal (Naish, 2005). This great wyvern can grow up to forty feet long, although a large portion of that length is made up of neck and tail, both of which are surprisingly thick and muscular. Aside from its immense size, Jugoceras is notable for several reasons. First, it has a full compliment of five toes, and one is fully reversed. The individual toes are fairly short and terminate in large black claws. The wyvern is heavily armored, with black plates running down its entire ventral side and light brown, keeled plates running from between its shoulderblades to the end of the tail. Both ventral and dorsal plates overlap to provide some flexibility. The keels on the dorsal tail plates are taller than those on the body. The animal's neck is decorated differently on its dorsal surface: The plates are small, retangular, and do not overlap. Their keels are more like spines. Black, conical spines erupt from the skin just to each side of the dorsal cervical plates.

The head is unique among wyverns in that it is short, squared-off, and fairly narrow. Northrop described the teeth as "oversized," comparing the dentition to Ceratosaurus nasicornus. Jugoceras has a very broad range of binocular vision. Its most striking cranial features are what its namesake: Each jugal bone is equipped with an impossibly long and rigid horn. Additionally, a small nasal horn is present just above the external nares. Like many wyverns, Jugoceras is capable of breathing fire. Yates (2001) noted that rather than using its fire breath for attacking prey, Jugoceras "spurted" its fire breath, and seemed to use it primarily for intimidation. Indeed, Naish never mentioned the giant wyvern using its breath weapon during encounters with Spinodracos, where such an offense would be an obvious advantage.

The oddities, however, do not end there. Jugoceras has short arms but long metacarpals and extremely long, but thin, phalanges. Unlike most wyverns, there are five fingers in the wing. The first fingers is short and flexible, and is used for terrestrial movement. The other four fingers end in similarly elongate claws. As they have no use in flight, Northrop concluded that the claws must have a display function. Not surprisingly given its bulk and unusual wing construction, Jugoceras is restricted to the lower, less snowy sections of the Alps and spends a lot of its time clambering about the rocks on all fours. It feeds mainly on ungulates. Juveniles have never been observed, though we can probably assume that they are not as weighed down as their parents and may live a very different lifestyle.

While Jugoceras obviously presents a great many unique derived characters, it also displays some obviously basal ones. The five-toed pes and manus, for instance, make it more primitive than even Eowyvern. It would appear that the wings were once useful for flight, but the creature's lifestyle and enormous bulk preclude such ability. A small, squarish head and overszied teeth may be primitive for the group, as Eowyvern also demonstrates these characters. Mantell (2001) placed Jugoceras in its own family, the Jugoceratidae, and placed it below Eowyvernidae. He cautioned, however, that Eowyvern could represent a basal wyvern that independantly lost many basal features, such as pentadactyly and quadrapedality. In that case, Mantell argued, Eowyvern would likely originate from a more basal clade than Jugoceras, as Jugoceras is more derived in its wing construction and ability to fly.

Northrop, A. (1985). Comments on a gigantic wyvern from the lower Alps. Natura Historia 392: 322-331.

Naish, D. (2005). Jugoceras is unquestionably aggressive toward Spinodracos. Draconium 46(1): 56-58.

Yates, A. (2001). Fire-breathing behaviors among wyverns. European Journal of Draconology 102(3): 294-297.

Mantell, G. (2001). A broad look at wyvern systematics. European Journal of Draconology 102(2): 305-311.

Eowyvern dorsetti

Known in South Africa for years as the "caped dragon," Nielson provided the first description of Eowyvern dorsetti in 1929. Given that so few wyverns were known at the time (only Fugaxotitan and Gracilopterus predate it in the literature), Nielson was at first unsure of how to categorize the creature. At a mere six feet long, Eowyvern is slightly larger than Gracilopterus but much heavier, and completely flightless. Its build it far more dinosaurian than wyvernian given its functionally tridactyl pes, reduced hallux, facultatively bipedal (and horizontal) posture. Nielson even considered a connection with Aves, suggesting that Eowyvern was, perhaps, an ancestral relic of birds, surviving to the present. Perhaps not surprisingly, this possibility was briefly revived and endorsed by Feduccia in 1996. However, Nielson eventually placed Eowyvern at the base of the Wyvernia, a location it has not moved from since. Although some workers believe that Eowyvern represents a more modern wyvern which has become secondarily flightless, the current concensus is that it represents an incredibly basal lineage.

The most obviously basal feature of the animal are its wings. More like a pterosaur than a wyvern, Eowyvern's patagium is attached to a single extended digit IV, and the patagium extends down to the end of the ribcage on the body. The wing is incredibly small and has no flight capabilities. Digits II and III are long and move more or less as a single unit. Digit I is opposable. The arms themselves are not particularly long, and a propagagium extends from the wrist to the shoulder, strengthened by a thick tendon. Eowyvern is surprisingly broad-chested, unlike the deep-chested flighted wyverns, and long-bodied. The legs are powerful and tridactyl, making Eowyvern the most cursorial of all wyverns (or dragons). The skull is fairly small given the animal's overall size, and is equipped with numerous dagger-like teeth. The snout is blunt and subrectangular, more like dragons than wyverns. The eyes posess a fair degree of binocular vision. Eowyvern has a stout, powerfully-muscled neck.

The creature is undeniably predatory, often chasing down medium-sized mammalian herbivores such as boars and young or sick antelope. Eowyvern has been known to share territory with Tauropesa ungulatus, although Brimely (1953) noted that, given the latter's larger size and more aggressive attitude, Eowyvern always backs down from a direct confrontation. Eowyvern attacks prey with its hands extended, grabbing prey and tearing away a chunk of meat with its teeth before letting go. The target often dies of shock and bloodloss within a short time. The wyvern is not above scavenging carcasses, either.

Switek (1978) was lucky enough to observe several instances of wing use in Eowyvern. These observations are very powerful, in that they may allow draconologists to study how the wing structure evolved in more derived wyverns. In one instance, a large male Eowyvern had extended its wings, which had become brightly colored, to a female who he was courting. The male stood fully upright, or at least as upright as it could get, and "shivered" its wings in an attempt to show off the bright breeding stripes which had lit up the patagium's underside. The normal-colored female seemed unimpressed, and eventually ran away. In another instance, Switek watched as a female shaded her brood with her wings. The hatchling wyverns were small but looked much like their mother aside from larger relative eyes and proportionately longer hind limbs. However, at no time did Switek observe a wyvern flapping its wings or making use of the wings in any way that would suggest a precursor for flight. Switek reasoned that perhaps over the millenia, Eowyvern, in finding no use for flight-related activities, had lost such behaviors entirely, and that its own uses for its wings had developed apart arboreal needs. Switek's paper is interesting in that he only discusses Eowyvern for about two-thirds of the paper, and spends the last third complaining about the his institution (wink!).

Eowyvern continues to be a point of fascination today. In an upcoming publication on wyvern genetics, Milnar & Fries hope to accurately pinpoint the "first wyvern" on an extent family tree.

References:

Nielson, L. (1929). Comments on the "cape dragon" of South Africa. Brevia (February): 45-52.

Feduccia, A. (1996). Eowyvern dorsetti: evidence for the ancestry of birds. Bulletin of Zoological Curiosities 47: 468-473.

Brimley, W. (1953). Co-occurance of Eowyvern and Tauropesa. Brevia (June): 34-37.

Switek, B. (1978). Observations on wing use in Eowyvern. European Journal of Draconology 68(4): 533-539.

A Persistant Misunderstanding

At least two of my readers (Sheryl and Glendon) currently think or have thought (respectively) that the dragons and wyverns and lindworms I write about are sculptures, sculptures that I have made. Nothing could be further from the truth. Anybody can buy these creatures at toy stores or comic shops. Or online.

What I do is make up scientific descriptions for these dragons and wyverns as if they were living animals. The McFarlane dragons have their own epic fantasy storyline, something about a hydra destroying the dragon kingdom and then humans gaining control of some magical stone and learning how to talk and...it's wierd. I'd prefer to think of these dragons as real animals, and I'm interested in how they might be related to each other. Incidentally, there are several McFarlane dragons I have not included (or bought) based on my preference for what dragons and wyverns are supposed to be. So I'm not sculpting these dragons, although whoever did is a great sculptor. I just try to give them life!

Lindworm!

Lindworms are flightless dragons which have further lost their hind limbs. At left is Lindwyrmus nychognathus, a large arboreal lindworm from Argentina. We'll be covering it soon, after we finish the wyverns and a recently-discovered dragon (a new member of the Dracolympidae). The last third of the dragons/wyverns series will be problematic taxa, which Lindwyrmus most certainly is.

Harryhausina marinus

This giant wyvern, the only living marine species known, was named in 1999 by Heathcote when she thought it looked like a beast from one of the famed special-effects guru's films. "Harryhausia marinus would not look out of place in some Greek adventure like Clash of the Titans," she wrote in the wyvern's initial description. And like Ray Harryhausen's creations, Harryhausia is a fearsome, destructive beast, greatly feared by the captains of smaller fishing vessels in the Pacific Ocean. Although principally found near Indonesia, Harryhausia has been sighted much farther south, in the waters west of Australia and between that continent and New Zealand. Although Heathcote's description was based largely on beached carcasses, authors since then (notably Garcia & Garcia, 2002) have published accounts of the wyvern's activities in the wild.

Harryhausia is a whopping sixty feet long, with a twenty-foot "wingspan." Its wings are short, and the patagium is thick and fairly inflexible. The forelimbs are quadradactyl, in an odd configuration. The thumb and index finger are free from the patagium, while the middle finger and ring fingers anchor the patagium to the arm, while bony spines erupt from the bottom of the wrist and elbow to broaden the wing surface. Draconologists are unsure exactly what purpose the vestigal wings serve, as Harryhausia's main method of propulsion is to undulate its body from side to side through the water, using its sizeable caudal fins to provide thrust. Heathcote suggested that perhaps the wings were used for threat displays, or aided in mating rituals, but to this day, their exact function remains mysterious.

The wyvern's body is fairly wide in the middle and tapers towards the head and tail. The body is green and smooth, though the belly is covered in wide, overlapping white scales. A series of cartiliginous spines runs down the back, anchoring a short dorsal sail. The spines are tallest above the arms and become more tightly packed toward the end of the tail. The sail is made of the same tough dermal material as the patagium and is resistant to bending.

Harryhausia marinus lacks any trace of hindlimbs. Were it to lose its wings, the monster would resemble an overweight sea snake more than a wyvern. The head is flat and roughly triangular in dorsal view. The dentition is extremely reduced, with only four large teeth in the upper jaw, and two in the lower jaw. The jaw tips are covered by a horny beak, and Harryhausia has a series of recurved palatal teeth. The wyvern also has two bulbous fleshy pads on its head, one over either eye, which are bioluminescent. Garcia & Garcia reported that Harryhausia will often dive to great depths, and that the pads may aid in prey capture or intraspecies signaling in darker waters. Despite its size, Harryhausia feeds mainly on prey much smaller than itself, mostly fish and young marine mammals like dolphins and pinnipeds.

Young marine wyverns are at risk from shark attacks, but juveniles will often swim together in groups of up to ten individuals (Cope, 2001) for their first few years, leaving the safety of the creche only when they approach thirty feet in length. Whether adults lay eggs or give bith to live young is unknown at this time, although Harryhausia is perfectly able to beach itself. Individuals are often seen sunning themselves on islands and beaches across the sea, although they do not venture farther than the beach on any landmass. Cope wrote that babies "must be hatched from eggs, as groups of juveniles are almost always the same age. Unless adults females were giving birth to up to ten, and possibly more, brothers and sisters at a time, one must conclude that these babies hatched at the same time and, from birth, swam together as a group." Munster (2002) dealt a blow to Cope's assumption, noting that many living animals, especially reptiles, give birth to multiple live young at once, and that the babies often stay together for a short time.

Harryhausia marinus is unique from a phylogenetic perspective in that it seems to have no living or extinct relatives. A swath of wyvern fossils are known from marine deposits, but most are from arboreal forms. Even the most fragmentary marine-buried wyvern fossils are no indication of a marine wyvern. In his brief review of wyvern systematics, Mantell lamented: "The fact that wyverns and dragons are primarily arboreal means that they, like seagulls, could die while over water, and thus fall into the sea and be buried and fossilized, only later to be unearthed by paleontologists who would be remiss in theorizing that they lived in a marine environment. As such, even fragmentary wyverns discovered in marine deposits cannot be readily shown to be relatives of Harryhausia marinus . . . without specific features linking it to the extent marine wyvern. As of this writing, there are no such specimens." This situation may change in the coming years. A team from Australia recovered a partial skull of what they consider a relative of Harryhausia in 2005, but alas, the specimen remains unpublished.

References:

Heathcote, J. (1999). A giant sea-serpent wyvern from the Indian Ocean. European Journal of Draconology 100(3): 450-461.

Garcia, M. J. & Garcia, N. G. (2002). Notes on the behavior of Harryhausina marinus (Heathcote, 1999). Natura Historia 409: 677-689.

Cope, E. D. (2001). Incidental observations of Harryhausina marinus. Brevia (July): 56-61.

Mantell, G. (2001). A broad look at wyvern systematics. European Journal of Draconology 102(2): 305-311.

Caprawyvern mencotti

As far as wyverns go, none of them are quite as strange as Caprawyvern mencotti (Dawson, 1976). Large and ferocious, the "goat wyvern" is also one of the most dangerous. Caprawyvern is a fairly rare wyvern that lives in the American southwest, feeding primarily on domestic cattle. The creature is not above stalking smaller vertebrates, however, and in fact young wyverns are fond of jackrabbits. At twenty-two feet long, with a wingspan of between fifty-six and sixty feet, Caprawyvern is a sight to behold. The wyvern's most obvious feature is its namesake--a pair of curved, ram-like horns emerging from the head, one behind each eye. The horns are used in intraspecific territorial and dominance disputes. Only males seem to have curved horns. Like mammalian rams, females have fairly small, back-swept horns. The large horns of the males are in stark contrast to the relatively small skull and narrow mouth.

From a taxonomic perspective, however, Caprawyvern's most striking feature is the structure of its wing, which is totally unlike any other known living wyvern. The arms are short but dominated by an elongate humerus. The radius has been lost--only a rod-like ulna remains. The radius' job is supplemented by a strong tendon which seems to help flex the tiny hand bones. Two fingers, most likely digits I and II, are free from the patagium and assist in prey capture and terrestrial movement. The other fingers have degenerated completely, replaced by a series of nine rigid cartiliginous structures. Three of these "spines" eminate from where one would normally find digits III-V, two laterally from the wrist, two more from the elbow, and two more from the middle of the humerus. In addition to these "main" spines, a smaller spine erupts from between each pair of "main" spines but does not reach the end of the patagium. Rather, Benton (1994) believes they serve simply to strengthen the patagium.

The spines are rigid and inflexible. The groups of spines are so distinctly placed along the arm that they have each been given name. The "manual group" is the first group and erupts from from the distal end of digit II. Unlike the others, the manual group is composed of four spines--two long spines, a short spine, and a final long one. The "carpal group" originates from the wrist and is positioned perpendicular to the ulna. The "elbow group" originates, as the name would imply, from the elbow. Finally, the "humeral group" erupts towards the proximal end of the humerus. The longest spine in all the groups is CG-III, or "carpal group III," the second long spine in the carpal group. Benton hypothesized that the wing spines were a sort of hypertrophied version of pterosaur actinofibrils, though they obviously evolved independantly. Interestingly, Caprawyvern hatchlings only have a single spine per adult group (except the manual group, where hatchlings have two). Growth studies carried out by Delphi (1997) suggest that these single rods correspond to the "short" spines in adult growth stages, except for MG-I, which is the distalmost spine in both adults and hatchlings. The main spines grow out as the animal matures, eventually surpassing their predecessors.

As noted above, Caprawyvern is extremely hostile to territorial invaders. Five human deaths have been attributed to the goat wyvern, and there have been dozens of reported attacks. The wyvern apparently attacks from above with its talons, scratching and kicking while flapping its wings and screeching. Males are far more aggressive than females, though females are, themselves, surprisingly confrontational.

Without fossil forms, Caprawyvern would be impossible to place among other wyvern clades. The strange wing configuration alone makes classification difficult. Thankfully, a few extinct forms have shed light on the process by which Caprawyvern lost the more traditional wyvern wing. Zhouwyvern haduken (Ryu, 1995) is a small wyvern from China that only has four wing fingers, including a free thumb. What makes Zhouwyvern unique is that the humerus, like that of Caprawyvern, is unusually long while the radius/ulna is proportionatelly short. The structure of the wing fingers suggests that the wing was narrow. Ryu suggested that Zhouwyvern may have had a spine (or spines) erupting from the elbow, as such a structure would have deepened the wing surface. Another caprawyvernine, Protocaprawyvern tusconi (Elyard, 2003), only has three wing fingers, just one of which is connected to the wing surface. Its wing strongly resembles that of Harenadracos. Elyard theorized that, given the resemblence of the wrist and elbow joints to those of Caprawyvern, that Protocaprawyvern must have had cartiliginous spines from at least these two points.

As far as where Caprawyvern, Protocaprawyvern, and Zhouwyvern (the Ceratowyvernidae), we'll leave that to Mantell (2001), who suggested that, based on the unique curved horns and reduced number of wing fingers (four in Zhouwyvern), that the Ceratowyvernidae is a sister group to the Gigantowyvernidae.

References:

Dawson, G. M. (1976). The first comprehensive description of Caprawyvern mencotti, the well-known "goat wyvern" from the American southwest. American Journal of Zoology 23(4): 67-89.

Benton, C. S. (1994). On the unique wing structure of Caprawyvern. Draconium 35(1): 88-94.

Delphi, J. (1997). Ontogeny and Caprawyvern. Wellnhoferia 15(2): 105-111.

Ryu, K. (1995). A primitive fossil wyvern with similarities to Caprawyvern (Dawson 1976). European Journal of Draconology 86(3): 361-366.

Elyard, S. (2003). A new spine-winged wyvern from Arizona. Draconium 44(2): 235-241.

Mantell, G. (2001). A broad look at wyvern systematics. European Journal of Draconology 102(2): 305-311.

Coming Up: Harryhausina marinus, a marine wyvern named after the famed stop-motion animator!

Deinowyvern enigmatus

You would think that the world's largest living wyvern would be easier to find, but it's not. It was "discovered" quite by accident, flying over the Panama Canal in 1925. Since then, sightings have been brief and rare. Draconologists are unsure where, exactly, such a beast would hide, but most accounts would have you believe that it makes its home in the mountains of the Sierra Madre. Indeed, most sightings are from Guatemala and Honduras. Calvin (1945) attempted the first scientific description of Deinowyvern enigmatus based on a bevvy of interviews. He characterized the monster as having "a wingspan exceeding 100 feet," and "massive hindlimb talons for tearing prey assunder." Most (but not all) interviewees made mention of large bull horns on the head. The exact number of wing fingers varied between interviewees, from three to five. Calvin's necessarily short description concluded that Deinowyvern enigmatus was a "holdover from some mythical time in the past when giants ruled the Earth, perhaps making meals of humanity's greatest natural predators including saber-tooth cats, mammoths, and bears." If nothing else, Calvin seems to have had a flair for the dramatic.


It would be another 25 years before science was presented with a more factual description, based on a corpse, by Reed (1970). The specimen in question had been found on a beach in Costa Rica and immediately quarantined. And while not quite as awe-inspiring as Calvin's monster, Deinowyvern has proven to be every bit as fascinating. Reed's Deinowyvern had an eighty-five-foot wingspan (larger than Europe's massive Megalodracos), and a nose-to-tail length of forty-four feet. The wings, aside from being long, were surprisingly broad and supported by three massive fingers. Like most wyverns and dragons, the thumb was free of the patagium and featured an oversized claw. The animal's tail is long and fairly inflexible. It is topped by three rows of keratinous spines. Deinowyvern's torso is narrow but extremely deep, and the chest is covered in large overlapping scales. Reed found the beast's head to be most peculiar. He noted that it seems to share features with Africa's Tauropesa ungulatus including a squared muzzle, external fleshy nares, and the anteriorly-directed, curved "bull" horns. Also like Tauropesa, Deinowyvern has a sizeable "chin."

Odd for a wyvern, Deinowyvern exhibits a true tridactyl foot and a hallux which is only partially reversed. The two forward-pointing toes are large and broad, while the hallux toes point inward, toward each other. Reed was struck with the contradictory nature of the foot: While the reduction in toe number is a derived trait for the Wyvernia, the lack of a fully-reversed hallux is quite primitive, and in fact missing entirely from central Africa's Eowyvern. "Whether at some point the perching toe was lost, or it had simply not formed yet," he wrote, "I am unable to say." He also noticed that, with just four wing fingers, Deinowyvern was more derived in the structure of the hand than most other wyverns, which retain all five wing-fingers.

Deinowyvern's range has been nominally estimated to run the distance of southern Mexico to Costa Rica by Mandelbaum (1992) in his review of extent wyvern species' ranges. His estimate was based on reported sightings and physical evidence in the form of shed teeth and scales. A few sightings in Jamaica since then suggests that Deinowyvern is able to frequent great distances overseas, perhaps in an effort to find food. Despite its enormous range, the enormous wyvern's main population seems concentrated in the Sierra Madre mountains, as the vast majority of sightings still occur in and around them.

Although frustratingly secretive, Reed's corpse was able to provide Mantell (2001) with some idea as to the creature's taxonomic significance. Given the monster's size, he placed it in a family previously occupied by the large North American fossil form Gigantowyvern dawsonii (Brahe, 1987), the Gigantowyvernidae. Frustratingly, the foot skeleton of G. dawsonii is unknown, so Reed's musings about the animal's derivity are still unknown. However, Mantell suggested that the Deinowyvernidae is a derived family, and that its closest living relative may be Caprawyvern mencotti from the American southwest. Mantell cautioned that the two groups probably split sometime in the Miocene, and that Caprawyvern is the odd duck in its family. The suggestion that the Gigantowyvernidae and the Ceratowyvernidae share a common ancestor is based mostly on fossil forms from the latter group.

References:

Calvin, R. R. (1945). Reports on a new, massive wyvern from Central America. European Journal of Draconology 35(2): 340-346.

Reed, R. (1970). Description of a corpse of the giant Central American wyvern, Deinowyvern enigmatus Calvin, 1945. Draconium 11(3): 413-456.

Mandelbaum, I. J. (1992). A review of known wyvern home ranges. In Wyverns: A Global View (Bordel & Barnes, eds.). University of Liverpool Press: 108-122.

Mantel, G. (2001). A broad look at wyvern systematics. European Journal of Draconology 102(2): 305-311.

Brahe, T. E. (1987). A new fossil wyvern from Texas and a new upper size limit for Wyvernia. Natura Historia 394: 1252-1256.

Coming Up: The most aggressive wyvern on record just happens to be suspected of erroneously starting the myth of the European dragon!

Gracilipterus pyros

This small wyvern, one of the only members of its order found in South America, was described in 1927 by Winston Kramer. The creature lives in the lowlands of Brazil, although unconfirmed sightings have been reported as far south as Chile. Gracilipterus, meaning "slender wing," is an apt name, given the wyvern's overall small, slender build but especially in reference to the wings themselves. The arms are very thing, and the widely-spaced fingers are long and splint-like. Gracilipterus lacks a propatagium, but it makes up for this absence by way of a very flexible wrist which allows the overall plane of the hand to be nearly perpendicular to that the lower arm. At a mere five feet long, with a nine-foot wingspan, Gracilipterus is one of the smallest living wyverns. It certainly stands out in its forest home with a grey-to-black body and red spots on the neck leading to a red chest. Uniquely, this wyvern's neck is adorned with dorsal spines, although the spines do not have bony cores. Instead, they seem to be made of flexible cartilage and dermal tissue. They can be folded back against each other or raised straight up. Adams (1967) suggested that the spines were used in courtship or threat displays.

Aside from its miniscule size, Gracilipterus is not far removed from the typical wyvern bauplan. Its head is triangular and the jaws end in a small rostrum. Small cranial horns sweep back behind the orbits. The feet are functionally tridactyl, but a prominant reversed hallux is present and is used, like most wyverns, to get a better grip on the substrate. In Gracilipterus' case, that substrate is tree branches. It is surprisingly capable in the jungle canopies, leaping and parachuting from branch to branch, and dive-bombing small vertebrates, toes outstretched--ready to grab the target in a second. In fact, Kramer rarely saw any Gracilipterus individuals on or near ground-level. Hicks (1982) would later relay that, in fact, Gracilipterus spend a large amount of time near the ground, where larger-bodied prey items could be caught. In one instance, Hicks witnessed three wyverns consuming a large boar, which he assumed had been killed by the group. While grounded, Gracilipterus lopes about awkwardly, in what Hicks described as "knuckle-walking, but more like a bat than any gorilla." He determined that, given their slow movement on the ground, the wyverns must have attacked the boar from above.

While not the first wyvern to display fire-breathing, Gracilipterus was the first wyvern studied in hopes of explaining this odd phenomenon (thus the species name). Arsenault (1944) described the "fire-breathing apparatus" of a dead wyvern, determining that large chemical glands existed on either side of the wyvern's tongue, wedged in a depression of the mandible. The glands, when compressed, would each squirt a chemical compound diagonally at such an angle that the two unique compounds would collide in mid-air about six inches in front of the dragon's mouth, at which point the chemicals would ignite. As neither compound was itself flammable, the resulting fire could not travel back down either stream, thus the wyvern did not risk burning its mouth. Bizarrely, the chemical makeup of either stream has not been analyzed to this day.

Gracilipterus pyros is placed in its own family, the Gracilipteridae, which does not have any other living members. However, the fossil record of the family is quite robust. A partial skeleton of Graciliwyvern lewins (Dawson, 1955) from the Miocene of Paraguay. Another fossil form, Lagopterus isopus (Dawson & Clarke, 1973), hails from Panama and may be from the Oligocene, suggesting that the Gracilipteridae is older than most other living clades. Only the Gigantowyvernidae may be more ancient. The unique manual anatomy of the group separates it from all others, but pedal morphology suggests a kinship with the Wyvernidae.

References:

Kramer, W. (1927). A small Brazilian wyvern. There be Dragons 12: 213-225.

Hicks, A. (1982). Feeding strategies of Gracilipterus pyros (Kramer 1927). Natura Historia 391: 333-41.

Arsenault, J. J. (1944). A chemical component for the fire-breath of Wyvernia. European Journal of Draconology 34(1): 92-104.

Dawson, R. L. (1955). Graciliwyvern lewins, a new Miocene relative of the modern Gracilipterus pyros. Draconium 19: 33-45.

Dawson, R. L. & Clarke, A. C. (1973). A fossil wyvern from Panama. Draconium 37: 245-251.

Next Up: Deinowyvern, a giant monster of a wyvern from Central America!

Fugaxotitan elegans

Fugaxotitan elegans was the first wyvern to be described, by Gilbert, in 1854. This large Australian wyvern was at first erroneously ascribed to the Draconia, but further inspection by Green (1872) warranted the erected of a new group of flying reptiles, the Wyvernia. Green's approach to the differentiation was as follows: "Wyvernia is separated from the Draconia by the lack of a third pair of limbs. Wyverns have opted to take to the skies by the usual method for vertebrates--by modifying the existing forelimbs into large wings." In addition, Green noted many other anatomical differences from Eudracos. Among them, Fugaxotitan has a roughly triangular head, with a thin snout. It also has a reversed hallux. As draconologists would see over the next century, these features are fairly diagnostic of wyverns in general.

Fugaxotitan exhibits several unique features all its own. Although quite large (twenty-five feet long, fifty-two foot wingspan), Fugaxotitan is by no means the largest wyvern. But unlike its bigger cousins, Fugaxotitan is bipedal while on the ground, walking along in the manner of birds, but with the upper body held more or less vertically. This ability is achieved thanks to a unique development in the lower back: Fugaxotitan has a mammal-like spine, with distinct dorsal and lumbar regions. The lumbar vertebrae are far more flexible than the dorsal series, and permits the wyvern's unique posture. Like Eudracos and many other dragons, Fugaxotitan has developed large, caudally-directed cranial horns. Males and females have different horn shapes--while male horns are thick at the base and are directed straight back, females have thinner horns which curve toward each other toward their distal ends.

Females are about 30% smaller than males, and tend to be a lighter green color. Hune (1945) detailed the animal's nesting behavior. Females build a nest atop some canyon ledge or rocky structure with large sticks and tree trunks. Very often, the animals will beachcomb for suitable nesting materials. The nest is relatively circular and is supplemented with vegetation. Females lay between two and four eggs, depending on their age, and the chicks stay in the nest for their first four months of life. Hatchling wyverns are very small and bright green. Their skin is smooth but their wings are large.

The chicks grow very quickly, and by the time they are six months old, they are half the size of their mother. By that time, they are capable of flight. However, chicks do not stray far from their nest site for the next year. Because Fugaxotitan nests in such hard-to-reach places, mothers will actually abandon the brood to find food for herself while the eggs are still in the next. She will also find food for her hatchlings until they can feed themselves. Fugaxotitan mothers are not terribly aggressive, but will squawk and make threat displays toward unwelcome guests. Actual physical confrontations are rare and short-lived. The only dangers facing the eggs (or chicks) are young, hungry dragons. Even so, raids on nests are extremely rare, and present mothers always scare away would-be raiders.

Fugaxotitan, while tolerant of humans, does not actively pursue their company. The animals do not live near large human settlements, and indeed seem to prefer the badlands. These large wyverns have also been sighted in New Zealand, indicating that they are able to fly across the sea. Despite their size, the wyverns are perfectly happy to hunt any small game they can find, although they have a preference for pigs. There have been several sightings of Fugaxotitan taking down kangaroos. Stowers (1987) reported finding a tasmanian wolf skull in an abandoned wyvern nest!

As the original wyvern named, Fugaxotitan occupies the type family, Wyvernidae. Its features are seen as generally diagnostic of the group, although the Wyvernidae itself is not a terribly diverse family. In fact, as draconologists would soon discover, the living Wyvernia is just as fragmented as the extent Draconia. Luckily, the fossil record for wyverns is much better, so not only do we know their origins (Archosauria), we are also far better aware of their ancient relationships.

References:

Gilbert, W. E. (1854). A large new dragon from Australia with a brief comparison to Eudracos. Journal of Zoology 14(3): 213-238.

Green, L. (1872). Fugaxotitan elegans (Gilbert, 1854) and a new category of flying vertebrates, wholly separate from the European Draconia. Royal Journal of the Natural Sciences 73(1): 45-72.

Hune, B. (1945). The nesting habits of Fugaxotitan elegans. Draconium 9(3): 313-352.

Stowers, N. A. (1987). Contents of an abandoned wyvern nest in Australia. Science Notes 64: 432-438.

Next up: A small South American form, known by most as the "spitfire" wyvern.

Wyverns are not Dragons

After a long hiatus from writing about dragons, I decided it was probably time to continue my discussion about large flying modern reptiles. But we're finished with the Draconia (for now), so what else is there to write about? How about wyverns? Although often mistaken for dragons, wyverns are an entirely different group of reptiles. Their poor fossil record suggests that the group evolved fairly late, perhaps originating as recently as the Miocene. Wyverns are generally larger than dragons, have more triangular skulls, can "breath" fire, but most importantly, wyverns only have two pairs of limbs.

Dragons are unique, of course, in having three functional pairs of limbs: Two sets of forelimbs and one set of hindlimbs. The upper pair of forelimbs have been modified into large wings, but they retain (in most cases) the same general structure as the unmodified forelimb. Wyverns, however, took the bat route: they modified their only pair of forelimbs into enormous wings by way of elongating (and strengthening) the individual fingers. Wyverns are more widespread than dragons, occupying the North and South America. They also have a strong presence in Africa and Australia. The largest and heaviest flying vertebrates to ever live are wyverns, and exactly how they manage to take flight while being so bulky is one of the great mysteries of evolution. Wyverns do not have the tendancy to abandon flight that dragons do, although this probably has to do with the inability to depend on a second pair of arms. Wyverns are a fascinating group, and I look forward to discussing them in the coming days (and weeks).