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INTRODUCTION & HISTORY

Besides Azhdarchidae, Spec also harbours another diverse clade of toothless pterosaurs. Pteranodonts are perhaps the most iconic clade of pterosaurs, well established in the pop cultural imaginarium, and while utilmately losing the spotlight in the HE, Spec’s pteranodonts are perhaps the most inventive and bizarre of the world’s pterosaurs.

Having possibly appeared as early as the Albian – if not deeper into the Lower Cretaceous -, pteranodontians gradually replaced their cousins in the Late Cretaceous, their domain beginning right after the systematic disappearence of their cousins, the ornithocheirids. The Santonian and Campanian periods are of particular relevance as the most iconic pteranodonts, †Pteranodon logiceps, †Geosternbergia sternbergi, †Geosternbergia maysei and †Dawndraco kanzai, dominate the flying vertebrate guilds of the north american seaways. At least a thousand †Pteranodon longiceps fossils are known, and many are so well preserved that for the longest time they proposed an important role in the understanding of pterodactyloids in general. An unique lineage of pteranodonts, the bizarre, nyctosaurids, also date back to this era, though they are much less common. Pteranodont remains before and after this span of time in the Santonian-Campanian are rare and often fragmentary, but they are found on a cosmopolitian scale, with remains occuring in not only North America, but South America, Europe and Japan as well (as well as a possible australian foot), and they span all the way to the Maastrichtian, both in the form of nyctosaurid humeri as well as well as specimens assigned to Pteranodontidae.

Since the late Cretaceous that pterodactyloid pterosaur diversification has been focused on two main linages. The first is Azhdarchoidea, a clade of terrestrial pterosaurs that spawned stork, hornbill, terror bird and condor like forms, which are probably Spec's most famous pterosaurs. The other one, Pteranodontia, diversified instead on aquatic environments, giving a special touch to Spec's shores that HE's lack. By the end of the Mesozoic, it seems, larger pteranodontians were in turn replaced by the smaller, more adaptable nyctosaurids; the last known non-nyctosaurid genus was Antarctoptera, a genus which included the largest known pteranodontian ever, Antarctoptera hyperion, a massive flier with a wingspan of over 10 meters, competing directly with azhdarchids in terms of size. Said giant disappeared with the Paleogene/Eocene Thermal Maximum, which disrupted oceanic environments and caused the collapse of many food chains. During the Eocene nyctosaurids did better in freshwater environments, much like HE's early frigate birds such as Limnofregata, and with the changes imposed by the Oligocene they diversified once more, albeit with new competitors, the large radiation of icthyornithes.

In Spec, Pteranodontia enters the Cenozoic with a bang, diversifying in the Eocene. Here, they witness global success and diversity, being the most common pterosaur lineage of the epoch, and the main component of the “Cenozoic Pterosaur Renaissance”. Besides characteristic marine piscivores, a variety of inland forms also appear, a few like Messelosauravis having wingspans beneath 1.5 meters. Much like in earlier ornithocheiroids, a wide range of wing proportions can be seen: while all retain large wings, in many of these forms either the wing finger or the fourth metacarpal is dramatically shorter, showcasing the tendency for inland flyers to have shorter wings. In some forms, the clawed finger metacarpals are still long and connected to the wrist, which may either be an atavism or an ancestral feature, since the phylogeny of many of these pterosaurs is only now starting to be understood. Most retain the “normal” stork like beak, though some have serrations and strange jaw curvatures.

Pteranodontid fossils decrease drastically in number by the Oligocene, but this appears to be an artifact of preservation rather than a genuine feature, as the diversity achieved in the Eocene still occurs well into the Miocene; Pteranodontidae itself, however, appears to have disappeared in the Oligocene. This period witnesses a dominance of nyctosaurs in particular, which become some of the most common pterosaur finds in contrast to their Maastrichtian rarity. Other pteranodontians are mostly found on terrestrial/freshwater settings, a situation that goes on the present day. Unlike azhdarchids, which witness an evident shift in diversity by the Plio-Pleistocene, it’s difficult to evaluate how pteranodont diversity coped with the glaciations, given the rarity of non-nyctosaur remains. These seem largely unaffected, and middle-Pleistocene New Zealand sites seem to imply a sudden diversity increase at least during this era.

BIOLOGY

Pteranodontia is the remaining branch of a prosperous Mesozoic branch of the pterosaur dynasty known as Ornithoehiroidea. Ornithocheiroids have quite possibly became the vertebrates most specialised for an aerial existence, having taken the already efficient pterosaur design and redefining it for life on the air: the shoulders ae higher than in other pterosaurs, the sternum keel is deeper – and, as predicated, the muscles associated with the downstroke are proportionally massive -, the humerus has developed a block-like form with large, warped dectopectoral crests, more robust and broad scapulocoracoids – in form forms no longer articulating with the notaria -, well developed notaria, an extensive rearrangeent of the flight musculature, and of course proportionally massive wings and small hindlimbs. In pteranodontians, these traits were taken to their logical extreme, with pneumatisation becoming so extensive that even the hindlimbs are pneumatised, and rendering them the most pneumatised vertebrates to have ever existed. The toothed jaws present in other ornithocheiroids were instead replaced by a toothless, keratinous beak, forming a continuous sheet with their casque-like supraoccipital crests. Like in azhdarchids, the metacarpals I-III lost contact with the wrist bones, being now anchored at the end of the fourth metacarpal, smaller than the clawed digits themselves. The fourth metacarpal, on the other hand, has enlarged further, extending the already proportionally enormous wing, becoming some of the proportionally largest fourth metacarpals in any pterosaur and indeed any vertebrate. Some forms have taken these specialisations even further, developing inflexible scapulocoracoids, hatchet-shaped dectopectoral crests and a complete loss of the clawed digits of the hand, leaving only the wing finger.

Having specialised to a different ecological niche than azhdarchids, pteranodonts can in some ways be described as the opposite of them. Their shoulder blades, much like in other ornithocheiroids, are located higher than in other pterosaurs, an adaptation for flight. Their fingers (aside from the wing finger, that is) are pretty much reduced to the point that modern forms, the nyctosaurids, lack them, again an indicator of their specialisation towards flight. Their anteorbital frenestra as quite small in porportion to the skull than those of many other pterosaurs, including their ornithocheiroid ancestors, in some species being about as big as a large bird nostril. Their crests vary in shape and size, from small stumps of the back of the head to quite long ones; some species pretty much have their crest[s] on the beak instead, much like long gone ornithocheiroids such as Ornithocheirus and Anhanguera. Pteranodonts, much like azhdarchoids, gave up their teeth, having instead long rostrums; interestingly their jaws are usually actually more stork like than those of azhdarchids. Unlike seabirds, which often land inn the water surface to feed, nyctosaurids are generally more specialised to hunt prey on the wing, due to the fact they would have tremendous difficulties to take off from the water and because they could loose a lot of body heat from the wings in cold waters, though some species do land on the water. Small nyctosaurids are able to skim on the water surface, but larger ones cannot engage in this behvaiour, so they grasp prey from the water surface with the jaws; some species engage in a hovering behaviour, which is either followed by diving or by spiralling flights above the water surface, snatching the prey from the water surface.

Besides the adaptations discussed above, pteranodontians are also weird among pterosaurs for other reasons. They lack a “gizzard”, relying exclusively on their stomachal acids for processing prey. Some pteranodonts have atypically long tails for pterodactyloids, with biforcated terminal caudals, in some cases supporting small structures thought to be similar to the diamond-like tail “fins” of early pterosaurs like †Rhamphorhynchus. Their nasonantorbital fenestrae are rather small for pterosaurs, providing them with iconic shallow, stork-like beaks. Their crests do not have keratin extensions, being entirely composed of bone sans the keratinous integrument envelope, as opposed to the elaborate keratin crests of other pterosaurs, including some azhdarchids; nonetheless, in some animals these bony crests can grow to be proportionally quite large and bizarrely shaped. All pteranodontians have rather wide pelvic cannals in females, and indeed some forms have become fully viviparous with true placentas.

PTERANODONTIDAE

Impundulu (Keranopterus scopus)

The skies of Africa, southern Europe, Arabia and Madagascar are crossed by a rather curious pteranodont. With white pelage and black wing membranes, beak and crest – which is rather small, barely noticeable unless under a closer inspection -, the Impundulu has an average wingspan of 2.5-3 meters, males generally being larger than females, though otherwise not much different beyond a slightly larger crest. The Impundulu, like all ornithocheiroids, has proportionally large wings, but they are smaller than those of the more characteristic pteranodonts, with metacarpal and wing finger proportions closer to that of the long gone istiodactylids and boreopterids, perfect for flight in inland settings. More curiously, the Impundulu still has it’s first metacarpal connected to the wrist, a trait unique among pteranodonts, as well as non-pneumatic hindlimbs, which might imply that this is a rather basal animal. Indeed, genetic analysis place it outside of the rest of Pteranodontia, implicating that it is in fact he most basal member of this clade. Similar animals are found in the Eocene and Oligocene in Asia and North America, and extinct forms of Keranopterus are known from the late Miocene of Europe.

The Impundulu favours generally dry open areas, particularly steppes, though it can also be found on wetter environments on occasion. Spending most of it’s time in the air, it nonetheless forages on the ground, feeding on squamates, beetles, small mammals, small dinosaurs and many invertebrates from snails to scorpions are also predated upon. This similarity in diet to HE’s Geronticus has labelled these pterosaurs as “Otherworld Bald Ibis”, though unlike the birds the Impundulu does not probe the ground with it’s beak outside of already made holes and crevices, preffering to capture prey on the surface substrates. With typical ornithocheiroid hindlimb proportions, it is a less efficient terrestrial walker than the azhdarchids it shares it’s habitat with, though like them it also has small, compact feet, as opposed to the larger and spread-toed pes of the average pteranodont. It roosts in cliffs, either near or miles away from it’s foraging grounds, gathering in large flocks. Occasionally, these flocks are targetted by large avisaurs, but generally only terrestrial predators attack these animals when foraging, and life expectancy can be quite high. The Impundulu is serially monogamous, with couples forming every breeding season to protect the eggs and then disbanding when the flaplings hatch. The “nest” is a pile of decaying vegetation in the cliffs, which is tended by both parents as to control the temperature within the eggs. After the flaplings are born, they generally remain in the cliffs for a while, foraging on insects and lizards that live there, before moving on to lowland areas.

Southern Ropen (Saurolarus ominosus)

Saurolarus is a cosmopolitian genus of nyctosaurids that can be best described as analogous to oversized terns. A typical example of this clade is the Southern Ropen, a species whose distribution extends along Australasia, from southern New Guinea to Tasmania and New Zealand, occuring as east as New Caledonia and as west as the western australian coast, and which is about of the same dimensions as HE's caspian tern. Much like other ropens, these are generalistic piscivores, occuring in both coastoal environments and on lakes and rivers well inland. It is main grey coloured, with a white underside and bright red beak and crest, which is small and disc shaped. It preys mostly on fish and crustaceans, which captures via skimming and/or kinglerfisher like hovering and diving. Occasionally it also preys on amphibians, small reptiles and bird chicks and eggs. Spending most of their time on the air, they can occasionally be seen resting on river banks or in large rocks and trunks; to compensate for the lack of digits they have small keratinous hooks on the front limbs should they need to climb. During the breeding season they gather in huge flocks in large waterways; here, males compete with each other for the right to mate, displaying impressive aerial acrobacy. Females then proceed to bury their eggs on the sand, and after a few months the young dig their way upwards to leave. Like in all pterosaurs, they are independent from the moment they are born, resulting in them occupying several ecological niches. However, nyctosaurids in general grow quickly for pterosaur standards, reaching full size and thus sexual maturity on an year.

Royal Privateer (Archipirata magnificiens)

If ropens are Spec's terns/skimmers, then these are the frigate birds. Pterosaurs have stereotypically been seen as seabird like for a long time in the scientific community, with frigate birds as being considered their closest analogue; its perhaps ironic that, in all their vast diversity, these winged reptiles reproduced a Fregata analogue. Privateers are divided in more or less four species, which occur in the oceans worldwide, the biggest being the Royal Privateer, a flying sauropsid with a wingspan of five meters, more or less the maximum size of the other main linage of large oceanic soarers, the icthyornithe diomedeodontornids. Unlike them, which land on the water surface, these catch prey from the air, and much like frigate birds they don't hesitate to harass smaller fliers, either eating their catch or eating them. The largest of the privateers is a black coloured animal, with a white underside and an yellow bill, with dark green naked skin pacthes around the eyes and white spots along the neck and back. Pelagic nomads as they are, during the summer their ranges can extend as far north as Iceland and as far south as Patagonia. Nonetheless, each year or so they gather in specific breeding spots, always islands spread across the tropics in the open ocean. Males develop keratinous crests along the upper jaw, similar to those of the long gone Nyctosaurus bonneri.

White Cherere (Lacusoptera cardiocrista)

The biggest of modern nyctosaurids are the chereres, large pterosaurs that occupy roughly the same ecological niche as HE's pelicans. Occuring in coastoal or inland waters across tropical and temperate latitudes (including South America), these pterosaurs land on the water to feed, capturing their prey with their large pouches on the lower jaw, and using their powerfull wings to take off from the water. This particular species, a relatively small one with a wingspan of 3 meters, occurs in the coastoal and freshwater environments of southern Europe, North Africa and Asi all the way to India. Mostly white coloured with black uppersides of the patagia and a grey beak, the males develop keratinous crests on the tip of the jaws, resembling the shape of a heart when the animal has the beak closed. Other species have crests along the upper jaw or even at the back of the head. They are usually calm animals, but many specexplorers have been pecked to death by the males when protecting the eggs. Males form harems during the breeding season, and every single female lays her eggs on his nest; this strategy is used by several dinosaurs, and is perfected by the chereres because they often form homosexual pairs with other males. Needless to say that the young are independent.

ANATOCHASMIDAE

Anatochasmids are a lineage of pteranodontians of about 10 species occuring across the world’s freshwater bodies, known from as far back as the Paleocene – though genetic data seems to place their origin much earlier into the Cretaceous. It’s been suggested that these are perhaps living members of Pteranodontidae, though their nyctosaurid like dectopectoral crests and rather more generalised habits seem to imply another placement in Pteranodontia. With similar wing proportions to those of the Impundulu, these animals appear to occupy a similar ecological niche to that of their long gone boreopterid relatives, trapping small aquatic animals in ridges in their beaks while they swim in rivers, lakes and wetlands. However, unlike their dead cousins, these animals are actual filter feeders, opening and closing their jaws to allow water to flow in and other of their beaks in order to trap multiple prey at once. In a way, they are basically like large dabbling ducks, and indeed Spec does not have the same diversity of dabbling ducks that developed in HE’s Miocene, presumably due to competition with these pterosaurs.

Granduke (Anatochasma fritzli)

The Granduke is a presence on the wetlands of most of Africa and southern and western Eurasia, occuring from the Mediterranean to the Ganges and into the Caucasus’ inland seas, with occasional breeding populations making a living as far East as the Baikal and as far west as Brittain; most of the extremes in it’s range usually occur in breeding and wintering seasons, when these pterosaurs migrate huge distances and are taken to explore new territories. With a 4.5 wingspan and standing at 1.7 meters of height, the Granduke may not be huge by pterosaur standards, but it is the largest living anatochasmid, and certainly a very impressive animal, on the air or on the ground. Occuring in a variety of freshwater habitats, from estuaries to swamps to highland lakes, it tends to preffer large lakes, where it can gather to quite large flocks. With a wide beak, it displays very well the anatochasmid filter feeding method, pumping water in and out of it’s snout in biting motions, trapping crusctaceans, insects, tadpoles, small fish and other small prey in the pecten-like ridges.

The Granduke has a white pelage, gray wing membranes and a pink bill. It’s crest, barely distinguishable in colour from the beak, is rather oddly positioned for a pteranodontian, forming a disc in the jaw tip, not too dissimilar from the crests of the typical ornithocheirid. In males, this crest is predictably bigger, but it is also wider and has a more robust internal support, instead of the usual spongy casque inner bone. More bizarrely yet is how these crests are used; contrary to what one may think, there is very little animousity between adult males, which are content with foraging alongside each other. They instead have a much more sinister purpose: males strike at females with these crests, ramming their robust jaw tips against them, bruising them until they are too weak to fight against anatid-style violations. At times, groups of males may gang up, isolating females from groups and beating them with the crests in all directions, causing severe bruises. In the breeding season, females may even be rounded up in groups, guarded by several males that prevent escaping and help subdue the rape victims.

This seemingly barbaric method is observed occasionally in HE dolphins, but the Granduke seems to have taken it to it’s logical extreme, the tendency for forced intercourse having evolved into a grotesque herding strategy. How and why these pterosaurs switched from male rivalry in lek systems to cooperation in herding and gang-raping the opposite sex is unknown, though the presence of similar crests in some extinct relatives may suggest it has began as far back as the Eocene. It has been suggested that this is simply the logical conclusion of the sexual selection expressed in rape strategies, where an arms race of male and female strength determines if the male’s genes will be passed on to the next generation.

Although adult males do not act violently against each other, as sexual competition is detrimental in a scenario where cooperation is vital, younger males that haven’t yet reached sexual maturity are fair game, often raped and rounded up alongside females. After they reach sexual maturity, they may join males of a specific group, or move elsewhere. The Granduke is also not limited to animals of it’s same species: several animals, from ducks to juvenile mokeles, may also be gang-raped and often die of internal organ fracture.

Thankfully, these strategies only occur in the first month or so of the breeding season: afterwards, males stop herding females, which are free to go away and move to the safety of larger flocks, if they aren’t too wounded. They gather in large numbers in sandy bars and river banks, burying their eggs, often alongside crocodiles and turtles. At least a few females remain nearby, helping to fend off predators. After an incubation period that may take up to three months or so, the juveniles are born, enough nests incubating at the same time to ensure predators only get a small fraction of the flaplings. They stay near the adults, even migrating alongside them, foraging on aquatic insects and small crustaceans. Half an year later, they’re about a quarter of the adult size, growing slowly for about three years or so until they reach sexual maturity, and continuing to grow for about three more years.

Trausduke (Anatochasma moorei)

A smaller cousin of the infamous Granduke, the Trausduke usually doesn’t grow above a two meter wingspan. It is certainly a large animal among the freshwater flyers of the continent, however, and occupies an almost identical ecological niche across both Americas. With a black and orange pelage, the Trausduke also bears a similar crest, and engages in the same violent sexual behaviours, generally occuring in Spring in either hemisphere. Compared to it’s Old World relative, the Trausduke is more migratory: part of the year, the population has a more northern range towards North America and is rarer in South America south of Pantanal, while during another part of the year the focus becomes reversed, disappearing from most of North America north of Florida and extending it’s range southwards into Patagonia. Such shifts are seemingly a recent phenomena, as the glaciations presented a more narrow band for these shifts, and indeed it appears that it was largely sedentary during the glaciations, occuring mostly in subtropical America.

Minkaduke (Purpurorhamphus eyrensis)

The Minkaduke is Australia’s dominant anatochasmid, found all over the continent’s wetlands, but most often seen in the southern great lakes, which in Spec are still sizeable, permanent waterbodies, unaffected by the extreme drying in HE; it often also frequents coastoal bays and lagoons. The Minkaduke is the last of it’s genus, of which several species are known in Australia since the Oligocene, bearing a rounded, thick supraoccipital crest, present only in the males. The pelage is mostly of a cream colour, of a darker yellow in the head, with a black line on the torso and black shoulders; the wing membranes are of a gray tone, and the beak and headcrest bear a distinctive violet colour, achieved through porphyrins not too dissimilar to those present in the Hastazhdarcho azhdarchids. Reaching a wingspan of three meters, it is among the largest non-azhdarchid flyers in the continent, and the massive congregations of these animals darken the skies above wetlands. Breeding occurs year round, males forming temporary territories that may cover entire waterbodies, though usually restricted to a patch of shoreline in large waterbodies. Passing females lay their eggs on a mound of decaying vegetation the male has built, which is tended by him for around three months until the flaplings hatch. During this period the male guards the “nest” viciously against even large rhynchoraptors and carnocursorines, all the while checking the mound’s temperature and removing and adding vegetation in accordance to the eggs’ ideal incubation temperatures, only occasionally taking breaks to filter feed.

Clamduke (Tachyeromimus sudamericum)

Occuring in southern South America, the Clamduke frequents both large lakes as well as coastoal waters. It has deviated from the usual filter feeding norm of it’s relatives by becoming a specialised molluscivore, it’s pecten like ridges having become larger and thicker, less useful as sieves but helping in cracking bivalve shells open. Unlike dsungaripterids and the Dupap, and like most pteranodontians, the Clamduke can swim very well and thus has easier access to molluscs, foraging either by dabbling or even shallow diving, depending on the water’s depth. While occuring in lakes all over South America south of Pantanal, it only occurs in the Atlantic shores, the chilean coastline being instead occupied by the oviraptorian Waldo, which occupies a similar niche; neither animal occurs in southernmost South America. Coloured in a rich black and white pelt and an orange beak, the Clamduke has a small supraoccipital crest as well as a disc like crest in the tip of it’s bill, present only in the female; in a rare case among pterosaurs, the female is the one who forms harems, defending a territory where the smaller, less ornamented males tend the eggs, buried by the female on sandbanks and bars. Generally, all it takes is the male’s loud screams to drive the attention of the larger and more powerful female, which can reach a wingspan of 3.6 meters, fighting off most predators. 

NYCTOSAURIDAE

The most morphologically derived clade of Pteranodontia, and indeed Ornithocheiroidea as a while, nyctosaurs have transformed their bodies further. Besides losing the fingers and metacarpals aside from the wing digit. the fourth metacarpal has also elongated further, being the proportionally longest of all non-azhdarchid pterosaurs, which combined with the wing finger it often renders their wings not only the longest in proportion to the body size of any animal, but also allowing very high aspect ratios, making nyctosaurs potentially the best soarers that have ever lived. The wing finger itself now only has three phalanges and is further stiffed, while the dectopectoral crests have become unwarped but rather hatchet like, offering a wider surface of attachment for the wing muscles. Tendons in the arm have become mineralised, an unique feature among pterosaurs, allowing the wing to remain locked in a fixed position as the animal soars continuously, and decreasing the danger of arthritis so common among old aged pterosaurs. The scapulocoracoids do not articulate with the notarium, forming a stable support for flight muscles, though at the expense of water based launching.

Nyctosaurids are for the most part pelagic animals, spending most of their lives flying above the sea, never touching land unless under rare circumstances. Spec’s first living nyctosaur specimens come from carcasses washed ashore in Tasmania, and since then reccords of living animals have increased sporadically, but they are still incredibly cryptic. Nyctosaurids can most often be seen in large congregations of other marine animals, be them schools of prey driven to the surface by marine predators, or the nocturnal migrations of deep sea organisms to surface waters. Unlike other pteranodontids, nyctosaurs are poorly suited for water based take off, and thus they are forced to forage aerially, much like HE frigatebirds and the contemporary enantiornithes known as jarilos. Various techniques have been observed in nyctosaurids, from typical dip feeding to impressive feats of hoovering, remaining more or less stationary on the air and simply picking up prey from the water surface, to angled strikes at high speed. Nyctosaurid diets are well documented in their carcasses, a wide variety of small animals having been found on the stomachal contents; while usually generalised, nyctosaurid diets appear to be largely focused on crustaceans and cephalopods. Kleptoparasitism has also been documented, with the pterosaurs targetting seabirds of various sizes, pecking them until they vomit their cargo. A few carcasses have also been found with petrels and other small seabirds inside the esophagus or stomach, implying that occasional predation on these birds occurs.

So well specialised for powered flight and soaring, nyctosaurids usually spend most if not all of their lives in the air, sleeping while flying like some of the most aerial birds, and uniquely they are among the few vertebrates that mate in the air. Landing is very well thought to almost never happen in most species, and should it happen the animals are virtually helpless on the ground, the immense wings being too long to coordinate with the hindlimbs when walking, though still competent swimmers even if water-based launching is very hard at best. Indeed, unlike most seabirds, nyctosaurs have completly cut off all ties to land, being viviparous, having developed a quite sophisticated placenta. This has the side effect of having much shorter “litters” than most pterosaurs, somewhere between one and four flaplings, but surprising at least some species seem to have reasonably high population densities; so far, it’s unknown if nyctosaurs engage in parental care, though no rookeries or similar nesting grounds have been identified, and many specimens are rather young animals, though all thought to be above six months in age; no significant deviations from the usual pterodactyloid growth rates have been observed thus far. Like all ornithocheiroids, their casque-like crests do not bear keratin extensions, but since the Cretaceous that these animals have provided rather impressive headgear.

Being the dominant marine pterosaurs in the Cenozoic, nyctosaurs had a rather impressive presence in Oligocene and Miocene marine fossil sites, being far more common than the few Cretaceous specimens. The Plio-Pleistocene seems to have done wonders to this clade, as the productivity of krill and other prey has increased in th globe’s oceans – though genetic studies imply a genetic bottleneck around the late Pliocene -, and Pleistocene subfossil beach sites in New Zealand’s Southern Island have provided thousands of specimens, mostly belonging to several species of Monochiropterus and Kerguelenorhamphus, presumably coinciding with an especially common presence in the southern oceans. As of this, at least 26 species have been recognised, and more seem to be on the way.

George’s Nycto (Monochiropterus georgi)

The first nyctosaur specimens discovered on Spec were three washed up carcasses of these animals in Tasmania. Since then, wild animals have been found across the world’s southern oceans, with occasional sightings as far north as Scottland, being one of the most well documented nyctosaurs. Largely of a light, silvery grey with a predictable bright white underside, the George’s Nycto displays a yellow and black beak, and males have an impressive headcrest of similar colours, albeit arranged in stripes and strange, axe-like dots. Roughly as long as the headcrest of the iconic †Nyctosaurus, it is rather distinct in shape: rather than a branched, antler-like structure, it is rather laterally flat and with a very large surface area, forming a structure similar to a much enlarged, reversed †Pteranodon longiceps crest. Hollow and filled with delicate, spongy internal support, it is occasionally seen broken in some specimens, and being reserved to the males it clearly bears no sigificant aerodynamic function. The wingspan usually ranges around 2.3 meters, with males being larger: the largest reccorded specimen had a wingspan of 3 meters. As always, the wings are massive in proportion to the rather small body.

The George’s Nycto is seemingly nocturnal, very rarely seen during the day and uncommon even in twilight hours, being most often encountered well after the Sun has set. Stomach contents show a variety of cephalopods, including baleen-whale larvae, as well as krill, with only occasional fish remains, mostly small or soft bodied like eel larvae. The reccords of the animal foraging in the wild show that it uses dip-feeding, flying over the ocean surface and quickly grabbing prey with it’s jaws. These reccords show a wide variety of strategies, from quick, angled attacks to hovering over the ocean surface, from the usual grabbing with neck flexion to angled strikes that “spear” the prey in the lower jaw, initially mistaken for skim-feeding. These attacks tend to have impressively high success rates, something poorly understood as the only other known nocturnal aerial dip-feeders are [spec]bats, and they use ecolocation to find prey. No ecolocation has been reccorded in nyctosaurs, which may suggest either their eyesight is so sophisticated that they can make out prey in dark water, or they have some sense currently undetected. Like most pterosaurs, it seems to have a poor sense of smell, and it doesn’t respond to oil baits like petrels, aside from following seabirds. Kleptoparasitism has been observed, with the pterosaurs grabbing small seabirds or other nyctosaurids by the tail or hindlimbs and shaking them, or pecking them in the flanks or throat, until they regurgitate their load. No actual predation has been so far observed, though one carcasse of a male was found with a storm-petrel inside of the esophagus.

Carcasses of gravid females suggest that these animals give birth to two or four flaplings. The juveniles’ wings are already well developed like those of most flaplings, but such a small brood size is atypical for supposedly superprecocial sauropsids. It is possible that nyctosaurs raise their young, or that this small offpsring production is a consequence of the needs of powered flight – though, since nyctosaurids seemingly almost never land, the main weight-saving pressure in flying vertebrates is absent. For a while, it was thought that this small brood size kept nyctosaur populations small, allowing the high diversity of species with so little competition, though studies on the population dynamics of the George’s Nycto and other species has shown that they have expected population sizes for pelagic foragers, at times even more common than some seabirds. Juvenile animals have been found in the wild, but they’re generally no younger than 6 months, where they’re more or less a third of the adult size.

Less mysterious, however, is what leads to the flaplings. At specific times of the year, these pterosaurs gather in large numbers at locales rich in prey stock, such as emerging blooms of krill. These concentrations generally are represented by a thousand or so individuals, some of the largest observed concentrations of nyctosaurids in the world – most impressively, these massive flocks almost vanish during the day, the animals dispersed at higher altitudes as they sleep, quickly descending once night settles in. Males fly well above the rest of the flock, honking loudly and performing incredible aerobatics. During this period, they forage very little, and indeed many males work and starve themselves to death, falling into the sea and never rising again. About a third of the amount of carcasses of these animals appear to be males that died this way, and it’s not uncommon to see mosarks, kronosharks and other large marine predators attending to these congregations, feeding on the fallen males. Should a female be impressed, she flies in the male’s direction and both rise up in the air. They mate in the air, the male displaying a long penis about a third of the creature’s wingspan and much longer than the torso, only as much as a fifth of it entering as the animals hover. Mating itself lasts only a few seconds, and the animals may mate continuously for two hours or so, before the female flies away and the male quickly refuels, before returning to aerobatics. The female may mate at most two more times, before she flies off, away from the massive flock and into the usual loneliness of the open sea.

White Nycto (Monochiropterus alba)

A smaller relative of the Goerge’s Nycto, the White Nycto has an average wingspan of around 1.4 meters as a full grown adult. One of the few polar pterosaurs, the White Nycto forages primarily in antarctic waters in Summer months, being commonly sighted on Antarctica’s shores, and even occasionally roosting on flat clifftops and icebergs, a rare behaviour for a nyctosaur; it is usually cathemeral, and during Antarctica’s Summer the White Nycto is obviously entirely diurnal. As the name implies, it has a thick, white pelage, which extends into the wing membranes; it’s beak is of a greenish tone, with yellow beak tips. Feeding primarily on krill, it can hoover excellently above the sea, picking the crustaceans from the water surface; it may also swoop down quickly. Mating during on Antarctica during the Summer, the males develop a rather small headcrest, of a vivid crimson and yellow colour, gathering in large flocks where the males exhibit themselves to exhaustion and mate in the air like it’s relatives. Mating on Antarctica during the Summer, the White Nycto winters in the open waters of the South Atlantic, Pacific and Indian Oceans, on average reaching as far north as the Galapagos. Despiste their Summer coastoal habits, the mystery to how nyctosaurids spend the first 6 months of their lives hasn’t been solved, and it is thought they give birth during Winter months on yet unidentified locations.

Tristan Nycto (Monochiropterus sudatlanticum)

At an average wingspan of two meters, this pterosaur occurs mostly in the waters of the Atlantic south of Ascension and north of the Falklands. White with a gray wing upperside and pink beaks, it displays little sexual dimorphism compared to it’s cousins, the males bearing only a small, round crest; nevertheless, the engages in similar mating gatherings. More piscivorous than it’s cousins, it has a prefference for pachycormid young, though it will happily consume most fish, cephalopods and crustaceans that it can swallow, and it frequently harasses seabirds and other pterosaurs for easy meals.

California Nycto (Monochiropterus septentrionalis)

Monochiropterus as a whole is more widely distributed in the southern hemisphere, the California Nycto being an exception to this rule. While it indeed occurs with large frequency in the western north american coast, it is also a frequent forager in Hawaiian and Asian waters, and it of course forages all over the pelagic Pacific between them, far away from shore; it seemingly preffers eastern Pacific waters during the Summer, generally moving westerwards during the Winter. With a wingspan of 2.4-2.9 meters, it bears a black pelage with a bright yellow beak and white wing membranes, white spots lining it’s flanks; the male develops a large white crest similar to that of the George’s Nycto in shape and size. Stomachal contents show that it feeds primarily on cephalopods, foraging primarily at nighttime. It Hawaii it has also been reccorded preying on bird chicks, swooping them from unguarded nests on the wing, usually during the day, picking also newborn seaturtles in this way; indeed Hawaiian individuals may very well predate on small insular animals during the day and forage on surface-venturing deep sea squid at night. Conversely, juveniles of this species seem to be a common target for jarilos and other acrobatic large seabirds. Mating gatherings occur both in Spring and Autumn, the former in the eastern Pacific, the latter in western waters.

Risso Nyctos (Kerguelenorhamphus spp.)

Once thought to represent a single species, Kerguelenorhamphus griseus, Risso Nyctos migght actually be a more complex group of species, with four main genetically distinct populations being identified. The original, first found as a moribund specimen in Île de Castries, has a western Indian Ocean distribution, found from western India to South Africa, being most common in waters to the east of Madagascar. Another population, informally assigned as K. cooki, occurs in waters around Australia, New Zealand, Melanesia, Micronesia and Indonesia east of Borneo; populations in the Bay of Bengal, eastern Indian Ocean and the central indonesian islands are thought to represent a mosaic of both populations, with interbreeding strongly suspected, mudding the waters fruther. A third population occurs in the Mediterranean, the Atlantic around Europe – including the North Sea – as far north as Iceland, as westwards as the Azores and as southernly as the Canaries, with infrequent occurences in the Black, Baltic and Caspian seas as well as the Atlantic north american coast. A third population occurs in the Pacific shores of the Americas, from Antarctic waters to the Pacific Northwest; populations occur in the Caribbean, which may either represent a mosaic between the Pacific and European populations, or an independent population of their own. Another mosaic/independent population scenario occurs in the waters around Japan and the Phillipines. Subfossil remains occur all over the world, but they first occur and are most common in New Zealand, particularly in South Island, and it has been thought that Kerguelenorhamphus is part of a southern radiation of nyctosaurids alongside Monochiropterus, though genetic testing seems to place these two genera at opposite sides of the Nyctosauridae phylogenetic tree. At any rate, while Monochiropterus has dominated southern oceans, Kerguelenopterus has spread across the world.

Risso Nyctos are rather variable in their habits, but they generally preffer to forage in waters off the continental shelf, in basically “shallow pelagic” environments; they forage cathemerally, predating on cephalopods and fish. Sometimes they forage on more coastoal waters, even occuring sporadically on lagoons and estuaries or even inland, but they generally remain at least a mile away from the shore. They are largely of a light gray with a white underside, bright orange beaks and rectancular headcrests that extend from the middle of the beak to the forehead. Variations on this design occur: K. griseus has a completly gray head pelage and black beak tips, while the face is white on K. cooki with gray extending from the neck to the pelage on the base of the lower jaw, having no black beak tips, and the headcrest extending into the supreoccipital, albeit lowly. The “european” population has a gray “mask”, a black upperjaw, and the crest restricted to the beak, while the eastern Pacific population is very similar to the european population except for having both jaw tips black, with the black area of the lower jaw extending well into the middle mandible. The “japanese” population has a white head like K. cooki,, but no supraoccipital crest and an entirely black lower jaw. No significant sexual dimorphism has been reccorded in any of the populations but the european one, where the head crest is absent in the females.

Breeding concides with Spring months in either hemisphere. Unlike Monochiropterus nyctosaurids, Risso Nyctos don’t form large congregations. The exact means of mate selection are unknown, but pairings are common during breeding months, implying serial monogamy for brief periods of time.

Ropen (Eucryptovolans novaeguineaea)

The seaways of Micronesia and Indonesia are frequented by nyctosaurid that is cryptic even for the standards of it’s clade, being very rarely sighted alive. With a wingspan of two meters, the Ropen is a nocturnal animal, almost never encountered during even twilight hours, foraging in both open waters and sheltered bays and estuaries for mostly invertebrate prey. It is coated in a light brown pelt, with similarly dull wings and and black beak; it has no headcrest, and indeed males look almost identical to females. However, this pterosaur has mastered a trick only two other groups of amniotes, the very unrelated baskervilles and tuataras, have managed: bioluminescence. At night time, the wings of the males glow with an eerie blue light, radiated from most of the membrane, and it is thought that these animals use this spectacular method instead of headcrests to determine worthy partners. Most spectacularly, unlike the baskervilles, the Ropen does not rely on a symbiotic relationship with fungi, producing it’s own luciferase reactions, though it is thought that they may use chemicals present in their prey, like marine glowing worms. This technique is not without some liability, however, as the blue light seems to affect the male’s melatonin production, and may endanger him to cancer. It seems, this dangerous mate selection mechanism favours indeed the strongest and healthiest males.

Besides it’s bioluminescence, the Ropen is also weird for a nyctosaur in regards to it’s still articulating scapulocoracoid. It has been suggested that it may represent a very basal member of it’s clade, and indeed it doesn’t appear to be closely related to the other nyctosaurids.

Giant Nycto (Giganokeanopterus magnificens)

The North Atlantic has a rather poor diversity of nyctosaurs, but it does compensate in having the largest species. With a wingspan of 6.7 meters, the Giant Nycto is the largest living non-azhdarchid pterosaur, and the marine flyer in the world, far overshadowing the seabirds it coexists with. White with brown patterns and a black head, the Giant Nycto has a rather unimpressive headgear, with it’s crests being a mere supraoccipital spike, golden in colour alongside it’s beak. It feeds primarily on fish, following other marine animals to schoals pressed against the surface, grabbing prey as large as small ichthyodectids. It also easily attacks seabirds, and indeed the Giant Nycto is a common menace in Summer rockeries, attacking adult birds leaving or returning to their nests, and occasionally snatching up chicks on the wing. On rarer occasions, it may swoop down to capture terretrial prey on the beaches and clifftops, and it may also scavenge, though doing this is rather tricky and energy taxing as the animal forages always on the wing.

Like Monochiropterus nyctos, it gathers in large open sea congrgations in the breeding season. However, these affairs are more violent, with males attacking each other on the wing, spearing and striking wih their bills. Many are fatally wounded and fall into the sea, and even triumphant males generally are too weak to mate more than once or twice at a time, though usually effectively as the penis has barbs, which remove the sperm of competitors. As usual per nyctosaurids, no animals younger than six months are known, and these are already large enough to be protected from most aerial animals besides adult Giant Nyctos. The Giant Nycto is the last member of a genus that has been prevalent in the world’s oceans during the late Miocene and Pliocene, with it’s closest relatives being the much smaller Prionopterus nyctos, distributed across the world’s oceans.

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