Arthropulmonians are the largest and most diverse class in the invertebrate phylum Stegospondyla, and they include both non-modular insect-like organisms and the much more advanced, modular Panathyrmatheria. Though most Athyrmatherians are restricted to Borea and Comedia, the non-Athyrmatherian varieties are present in nearly every single terrestrial ecosystem on the planet, filling incredibly important niches in lower and middle trophic levels that are equivalent to those of insects. Some are obnoxious pests that feed on other animals and ravage vegetation, while others are important pollinators and serve as an important food source for animals. They resemble insects quite a bit as well, sharing a similar segmented external skeleton composed of chitin, and a muscular system that connects on the inside rather than outside. To refer to them as insects would of course be a misnomer, as these organisms are of totally alien origin.
One thing that sets Arthropulmonians apart from insects is their age. Athyrmagaia is about 8.043 billion years old, and it has been harboring life for a much greater span of geological time than Earth has. The earliest fossils of what appear to be Arthropulmonians originate from rock layers that date back to the Archaeocarbonic period (Athyrmagaia's equivalent of the Carboniferous), roughly 635 MYA; At this exact point in time, Earth was only just beginning to enter the Ediacaran period, and insects wouldn't evolve until 250 million years later. Additionally, Arthropulmonians have oddly more developed and centralized nervous systems compared to most of our own insects, and their brains superficially resemble those of agnathan fish. Their bodies are also organized differently. They have four main tagmata; the head, thorax, abdomen, and tail. The thorax and abdomen are less distinct from one another than those of insects, and both play equal parts in housing the vital organs. The segmented tail only contains the organism's anus and outer reproductive tract, and almost universally ends in a telson that extends past the cloaca. Their exoskeletons, much like those of crustaceans, contain layers of chromatophores that are capable of slowly changing color. Though nearly all species possess this ability to some degree, the effectiveness tends to vary greatly, with some species even forgoing them entirely in exchange for chemical-based or structural pigments. They have eight limbs, each of which are biramous and end in bifurcated tarsi that function somewhat like toes. The second, third and fourth pairs are used for locomotion (locomotor legs), whereas in the absence of antenna, the first pair of limbs act as antenna-like sensory appendages (sensory legs) used for tactile senses, self-grooming, and grasping. Like insects, Arthropulmonians also come in flight-capable varieties, but since the class ancestrally lacks endites or exites, they evolved their wings by modifying entire limbs. These limb-derived wings are called "pteropods". Arthropulmonians possess a three-part jaw structure, with two double-jointed lower mandibles, an upper mandible-like labrum or premaxilla, and a claw-tipped lower labium that serves as both a swallowing aid and a sheathe for a retractable hypopharynx. The hypopharynx acts as a chemosensory organ analogous to a snake's tongue, and is used for both taste and smell. The muscles of both the labium and the hypopharynx anchor to an anatomical structure in their upper thorax and throat that resembles a hyoid complex, a very unusual trait for an exoskeletal invertebrate. The upper portions of the mandibles form the kinetic lower portion of their cephalon, and allow for mouth that is semi-expandable. Combined with the efforts of the claw-bearing labium, this allows many Arthropulmonians to swallow larger pieces of food than most similarly sized insects. Rather than solid-state lenses or compound eyes, Arthropulmonians possess pinhole or camera-style eyes comprised of soft tissue, which bear structural and anatomical similarities to those of mollusks and vertebrates. These eyes lack eyelids and are protected by a clear, glass-like dome of transparent chitin, and since they are fixed within their sockets, each eye bears multiple pupils and corneas that allow them to see in multiple angles simultaneously. An additional eye used to regulate circadian rhythms often lies between them, functioning somewhat like the parietal eye of a tetrapod. Arthropulmonians sport a rather insect-like tracheal system for breathing atmospheric air, but lack the multitudinous abdominal spiracles associated with their Earthly analogues. Instead, they bear just a singular pair of facial spiracles that are located near their mouths, and their thorax houses six muscular lung-like chambers that are used to facilitate active, cyclic respiration during periods of high activity. A nearly identical system is also present in closely related marine Stegospondyls, albeit with gills in place of a tracheal system. In Athyrmatherians these have evolved into complex lungs, but among lesser Arthropulmonians these "lungs" act merely as bellows to more quickly pump oxygenated air across the trachea. Distribution of this oxygen, on the other hand, is more like that of arachnids than insects, and is accomplished via a hemolymphatic open circulatory system that uses copper-based hemocyanin as an oxygen carrier. While this does allow them to reach slightly larger maximum sizes than most of Earth's insects, it still limits their size in much the same way, as their size constraints are in large part proportionate to their planet's gravity and the abundance of atmospheric oxygen (both of which just so happen to be 99% similar to that of Cenozoic Era Earth).
Though the animals shown on this page are excellent model organisms for the class itself, they are but a small taste of the vast variety of forms these creatures take. Arthropulmonians are incredibly biodiverse, and are represented by a dense cast of different orders and over a hundred millions species, too many to be described in summation. Generally speaking, however, a majority of the insect-grade Arthropulmonians can be organized into two primary subclasses: Archaeopulmonia ("first lungs") and Cruraptera ("limb wing"). Archaeopulmonians are a comparatively more basal branch, and are represented by ametabolous flightless organisms that and are born as nymphs that resemble the adults. Only one extant infraclass, the Repenopoda ("crawling feet") is known from this group, with a number of other primitive species that currently remain unranked. Unlike Crurapterans, Archaeopulmonians retain multiple sets of small pleuropod-like gonopods under their tail, more archaic pinhole-style eyes or eyespots, and a namesake side-to-side body movement described as being similar to that of a reptile. The most primitive representatives of this subclass have three articulated scale-like tergites per segment as opposed to a singular piece per segment, a much more primitive condition exhibited by aquatic, non-Arthropulmonian Stegospondyls. Cruraptera is comprised of more derived, often flight-capable species that possess camera style eyes and lack gonopods, and is represented by two infraclasses that evolved flight independently of one another despite a shared common ancestry: Biramoptera ("two-branched wing") and Ambuloptera ("wandering wing"). Biramopterans walk on only four limbs, and possess multi-jointed biramous pteropods that are modified from the tarsi of the third pair of limbs. Some biramopterans are hemimetabolous, but the majority of the clade has evolved a more complex holometabolous life cycle where they undergo a larval, pupal, and eft stage prior to adulthood. Ambulopterans differ in that they are all hemimetabolous, and possess four semi-uniramous pteropods that are modified from the second and third pairs of legs. These novel appendages function both as wings for flight and as legs for walking, and when a Ambulopteran is grounded, the papery membranes of its wings neatly fold away with the aid of a tarsus-derived wingtip.
Order Floraptera ("flower wing"): Florapterans (floraflies and duskwings) are an incredibly speciose order of brightly colored Biramopterans belonging to the superorder Tunicoptera ("coated wing"). Easily one of the most stunning examples of interstellar convergent evolution currently known, the Florapterans are especially remarkable because they almost the spitting image of our own Lepidopteran insects, the moths and butterflies, even though they aren't even remotely related to them. These parallels are not only in their appearance, but also in their ecological niche. Florapterans are specialized nectarivores that feed on the nectar contained within the flowers of Angiodactyl Olekirkophytes, and come in a vast number of colorful and beautiful forms that are specialized to feed on specific species of flowering plants. They are represented by both diurnal (floraflies) and nocturnal (duskwings) forms, which often resemble butterflies and moths respectively. They are also important pollinators, and when they feed on nectar they often take some amounts of pollen with them on their small, furry bodies. They even boast a similar global distribution, made possible in large part by the evolution of flower-bearing Olekirkophytes.
Florapterans are liquivores, with reduced mandibles and a retractable proboscis derived from the hypopharynx. Feeding is achived with contractions of the mandibular joints and the labium, which sucks in liquid nourishment through the proboscis via a pump-like action. The shape of the proboscis also varies, ranging from the more typical retractible type to a permanently exposed beak-like structure. The most recognizable features of this clade are their enormous pteropods, which are typically many times larger than their own bodies and unable to fully fold their membranes. The wings are and covered in small, lepidopteran-like scales, but unlike the Lepidopteran equivalents these scales are not the source of the bright colors. The actual color of the wing comes from unique chromatophores called iridophores, which use contractable layers of light-scattering guanine crystals to change color. These chromatophores are much faster acting than those of most other Arthropulmonians, able to shift nearly instanteously with very little effort. Among diurnal floraflies, such as the blue spotwing (Maculopapilia caeruleapterus), these iridophores are fixed to form a specific pattern and set of colors that are usually aposematic in function, with patterns such as brightly colored stripes and spots that they are able to roll and ripple to confuse and deter predators. These flashy colors are not unwarrented either, and many of the more brightly colored floraflies tend to be either foul-tasting or even mildly poisonous. The duskwings more commonly utilize mimicry, their wings mimicking the bark of trees or the petals of their favorite flowers. If the environmental disguise doesn't work, they are able to terrify a to-be predator by flashing a pair of eyespots that are normally invisible. Some species utilize another form of mimicry, imitating the body and wing colorations of their venomous predatory cousins, the Raptoripterans.
Florapterans are members of the holometabolous branch of the Biramopteran phylogenetic tree, and go through a larval, inert pupal and ambulatory pupal stage before becoming flighted adults. Among Athropulmonians, Florapterans are very notable in the fact that photosynthesis plays a role in their life cycle. The average individual hatches from its egg in the form of an herbivorous, benign caterpillar-like larvae that spends most of its youth gorging on leaves. As it does so, it stores the chloroplasts from the food it eats in the tissue of its stomach lining, as well as the proteins needed for the production of eggs. Once the humble caterpillar has reached a large enough size, it sheds its skin and enters the inert phase of its pupal stage, which turns bright green as the gut chloroplasts migrate into the tissues of the chrysalis's outer shell. The muscular, digestive, nervous, respiratory and circulatory systems are the first anatomical features to reform, which triggers the pupa to awaken as it enters the eft stage, a mixotrophic animal that powers its metabolic processes partly through photosynthesis until the rest of its adult features finish developing. Once this is complete, the adult Florapteran emerges from its green-tinted pupal shell and flies away. In the same way as our own Lepidopterans, a majority of their life is spent in the larval stage, and their short lived alate adult forms only serve as a means of reproduction and dispersal. Some, like the blue spotwing, only live for about 13 days after its wings develop. They are almost exclusively R-strategists that put very little investment into parental care, leaving as soon as they finish laying their eggs.
Order Raptoriptera ("seizing wing"): Raptoripterans, or "eagle-wasps", are a group of regionally endothermic Tunicopterans that live in a variety of highly productive ecosystems, and represent a closely related sister group of the Florapterans. Though they share the same colorful, scale covered pteropods of their placid nectar-drinking cousins, they differ significantly from them in the fact that they are all active predators to some degree, fierce hunters with murderous mandibles instead of a graceful proboscis. Most often they will hunt flightless invertebrates, but the largest Raptoripterans, such as the tigris eagle-wasp (Raptoripodus tigris) shown here, are known to to hunt small Tagmatasaurians and Saurophysans.
Though they are referred to as "wasps", they are obviously in no way related to their earth namesakes, and do not possess a stinging ovipositor. They do, however, have a venomous bite for incapacitating prey, which causes itching and swelling for larger animals. One family, the pepperbees, possess the ability to spray liquid capaicin from glands in their cloaca, which causes a burning sensation in mucous membranes. True to the name given to them by researchers, Raptoripterans have evolved a morphology that is optimized for tracking down and apprehending prey via a "swoop and strike" method of prey capture, while also avoiding larger airborne insectivores. Their tremendous eyes can see in both nearly full color and in the ultraviolet spectrum, and bear three specialized pupils. One pupil is large and horizontally elongated for a wide field of vision, and two smaller more forward-facing pupils above and below it provide binocular vision at both upper and lower angles. Their ability to track prey is further enhanced by their sense of smell, as they possess a feathery hypopharynx designed to effectively pick up airborne scent particles. Their biramous wings, which are comprised of the lengthened toes of their middle locomotor legs, bear joints that are capable of stiffening and unstiffening to adjust the shape of the aerodynamic surface, allowing for various mid-air maneuvers when avoiding larger predators. Their primary weapons for prey apprehension are a pair of enlarged, muscular raptorial hind limbs, which possess prehensile tarsi and lethal, armor-piercing tibial spines. These formidable appendages are used to quite literally snatch their prey off the ground, grasping the unfortunate soul with their feet before skewering it between their spiked femur and tibia while delivering a venomous bite that turns the prey functionally brain-dead. Once this is accomplished, they begin to feast on their prey with their powerful mandibles, regardless if their catch is dead or alive following capture. They tend to be particularly ravenous, and may even swallow the prey whole without the need for venom if it is small enough for their highly kinetic cephalon to allow it.
Despite all of their differences, Raptoripterans are quite similar to Florapterans in that they have a holometabolous life cycle, starting out as a caterpillar and becoming a pupa before emerging as an alate adult. As a largely predatory clade, however, their version of this life style has some noteworthy differences. Raptoripteran caterpillars are ambush predators that disguise themselves as bare twigs, using their skin chromatophores to match their colors with the bark of the tree they inhabit. When prey flies near them, they strike with their clawed forelegs. Their eft stage is a cleverly disguised ambush predator as well, forgoing mixotrophy by simply waiting for prey to arrive so that it doesn't waste calories that are needed to complete its ongoing metamorphosis. Their adult alate stages are also much longer lived than those of their cousins, since they are able to ingest protein-rich food as both adults and larvae. The tigris eagle-wasp in particular can live to be a year old, with the adult stage taking up about 48% of its lifetime.
Most Raptoripterans are solitary, and the only time they ever willingly meet is to briefly mate before moving along with their lonesome lives. Other than that, a majority of meetings between Raptoripterans are rather hostile. There are exceptions to this, of course, as the species that belong to the "pepperbee" family often live as small congregations in arboreal nests made of a regurgitated bioplastic mixture. These congregations have hardly any social order to speak of, however, with the cohabitants of the nest being chummy roommates at most. Solitary Raptoripterans tend to be marginally more dedicated parents compared to their Florapteran relations, and will usually lay their eggs in soil, on leaves, or in microbially induced galls, where they will warily guard their unborn young until they hatch.
Order Cyclopoptera ("one-eyed wing"): Cyclopopterans, best known by researchers as "flyclopses" or even just "flies", are a very diverse clade of often very small Biramopterans that have evolved to occupy a fly-like niche in Athyrmagaia's biosphere. These pesky little creatures are just as ubiquitous as their Terran analogues, and are present on every single landmass on the planet as in the form of scavengers, decomposers, predators, pollinators, and parasites. This makes them a very significant part of the biosphere, as multiple species of this order can adopt the niches of both predator and prey within the same ecosystem. Quite a number of them, such as the rot fly (Cyclopomusca necrophageus) shown here, are cosmopolitan species that are able to live in any terrestrial environment where their preferred food source is readily available. They even play a role in population control, with disease-carrying bloodsuckers and sap sucking plant ravagers helping cull organisms that are less resistant to disease in favor of those who are more so. As a consequence in their place in the food chain, however, many species that belong to this order, particularly those with a taste for blood or rotting matter, have earned a somewhat mixed reputation as obnoxious, swarming pests.
It doesn't really take much more than a solid glance to figure out what gives this curious group its systematic name. Cyclopopterans generally possess a huge, singular eye that accounts for anywhere between 30% to 50% of their head's mass. This monocular eye originally evolved from two separate eyes, which fused together as the ancestors of this clade got progressively smaller. The result is a singular combined ocular organ with a single covering, The primary purpose of this cyclopticism is the regulation of visual information; A complex lens-based eye gathers much larger quantities of visual information than compound eyes, and such a large amount of information could be overwhelming for a tiny creature with an even tinier brain. By reducing the number of cones in their eyes and merging both eyes into one, Cyclopopterans reduce the amount of visual information being captured, but increase the rate by which it is processed by their tiny brains, allowing them to essentially see the world in slow motion. They don't even need depth perception; The single lens of the combined eye is multifocal, and the two focal lengths are processed by two separate, albeit simplified retinas. Though their vision only extends a couple of yards at most, this forms the closest approximation of depth perception they can achieve, allowing them to easily estimate the distance of an object of interest or an approaching threat. A few other flying Biramopterans have convergently evolved partial or full cyclopticism as well, but Cyclopopterans are the only order of Arthropulmonians that is represented by exclusively cycloptic organisms. Even the biggest species, which can be as large as a hornet, possess this large, singular eye. Aside from that, many of their other adaptations are quite similar to those of Earth's dipterans. Their mouthparts are usually adapted for sucking, slurping and/or piercing, with reduced mandibles and a proboscis modified from the labium and hypopharynx. Other species may possess a stylet, with internalized mandibles used to bore into skin or chitin. They often search for their food by scent, picking up traces of ammonia using a tetained pair of tines under their proboscis. Unlike any other Biramopterans, the pteropods of Cyclopopterans are uniramous and lack a tarsal hindwing, and the forewing only has a single joint at its base to allow it to fold back at rest. Replacing the hindwings are a modified pair of sensory legs called halteres, vibrating club-like organs that are used for balance. This gives Cyclopopterans an insane level of stability and agility in flight, and though the structures involved are quite different, these halteres serve the same exact purpose as the wing-derived halteres of dipterans and strepsipterans. Their high maneuverability on the wing combined with their ability to react faster than we can blink has put them very high on the list of "most annoying animals in the universe", right next to our very own flies.
Cyclopopterans are holometabolous, and have a fairly nuanced life cycle where they go through a larval, pupal and eft stage before becoming an adult. Their larvae can vary quite a bit in appearance, ranging from eyeless and limbless tubular blobs to spiny aquatic critters. When these larvae feed and live on decaying matter, however, they are often referred to as "maggots". In the latter case, the mother will lay her eggs on or inside of decaying organic material, such as carrion or rotting fruit. When the egg hatches, the maggot will feed on the surrounding matter until it grows large enough and pupates. Upon emerging as an eft, it continues to feed until it finally sprouts the wings and halteres of the adult stage.
Order Lepidotergita ("scaled back"): Lepidotergitans are an order of small Repenopodans that thrive in humid environments. Loosely resembling silverfish or firebrats in appearance, these creatures are similarly primitive, and represent one of the oldest groups of Arthropulmonians still living. Indeed, many Archaeopulmonians look more similar to their aquatic ancestors than other members of their class, and are similarly prone to desiccation under direct sunlight exposure. Lepidotergitans are no different, and they are only capable of living in areas that are either sheltered from sunlight or humid enough to mitigate desiccation. There are multiple species that belong to this order on both the Borea-Comedian landmasses and Austrus. The most notable of these ancient creatures is the pangobug (Lepidotergitus griseosquamus), a cosmopolitan creature that lives in warm, high humidity or low-light areas and feeds mainly on plant detritus.
Lepidotergitans are obviously a very archaic group and bear a variety of characteristics that are lost entirely in most other Arthropulmonians. In addition to their two sensory legs and six locomotor legs, their elongated tails are rimmed with fourteen pleopods, an ancient holdover from their aquatic ancestors. Having long lost their use as swimming appendages, these redundant limbs function as gonopods for holding eggs. Their eyes, while fairly well developed, exhibit a more basal pinhole-styled condition, and lack a solid, enclosed lens. Due to this, their vision is fairly poor, and they are unable to perceive color. To compensate, Lepidotergitans rely mainly on their sense of touch, smell and taste, and have an exceptionally large hypopharynx with exaggerated tines. As opposed to a single tergite per segment, Lepidotergitans have three per segment, with dorsal and lateral tergites that overlap like the scales of a fish or a lizard. This much more kinetic exoskeleton is much less rigid than the fortified singular tergites of most other Arthropulmonians, and leaves these creatures prone to desiccation when directly exposed to the sun. It does, however, allow them to easily squeeze between tight cracks. Male Lepidotergitans do not possess penises. Mating is instead accomplished through the passing a spermatophore to the female. The Lepidotergitan life cycle is very simple and ametabolous, and the nymphs are born physically identical to adults apart from size. Throughout their lives, they molt continuously even after reaching full maturity.
Order Repoadacta ("crawling biter"): Repoadactans, also known as "beetiles", are a highly varied order of flightless, almost exclusively carnivorous Arthropulmonians that sport a mixture of both basal and derived traits. Unencumbered by any sort of wing to speak of, they rely on speed and brute strength to kill prey. These creatures are found in a variety of environments worldwide, such as forests, deserts, grasslands, and even in or near the ocean.
Repoadactans were initially classified as derived Archaeopulmonians related to Repenopodans, due to their possession of a similar side-to-side style of locomotion and lack of pteropods. Upon further analysis of their anatomy and comparisons of DNA, it has been found that Repoadactans are actually basal Crurapterans closely related to both Ambulopterans and basal Biramopterans. Though they lack the limb-derived pteropods that are almost ubiquitous among their flighted cousins, the other similarities they do share with them outweigh their similarities to ancient Repenopodans. Rather than fleshy nautiloid-like pinholes, they possess proper camera eyes with lenses and retinas, and their abdomens lack gonopods. They also have a single tergite per body segment as opposed to three, an adaptation that evolved to add more rigidity and muscular support to the body and limbs on land. It is strongly believed that the common ancestor of both the Biramopterans and Ambulopterans would have been a creature quite similar-looking to a Repenopodan.
Repoadactans come in multiple varieties, many of which are fast-moving terrestrial carnivores that forage on the ground or in the trees. Many of the forms representing this order might resemble wingless beetles or cockroaches, with their fluttering, darting hypopharynx and peculiar side-to-side locomotion also calling to mind squamates such as lizards. Many species also retain the ability to change color by expanding and contracting exoskeletal chromatophores, adjusting the brightness and saturation of their patterns to easily blend in while hunting prey or avoiding predators. Their jaws are very formidable tools, ranging from piercing hooks to crushing vices, and the tarsi of their sensory legs act like prehensile ropes as they grab hold of prey. Not only do they have excellent vision and an effective sense of smell, but they also have fairly good hearing to boot, possessing rudimentary "ears" on their legs that can pick up sound in multiple directions, and the setae that cover their bodies can pick up even the slightest vibrations. With the retention of six locomotor legs used exclusively for terrestrial movement and no modifications for flight, the land speeds of Repoadactans are almost unrivaled, making them some of the fastest-running terrestrial invertebrates the planet has to offer. Similar to beetles on earth, their order also includes some of the largest species of non-Panathyrmatherian Arthropulmonians, with the largest known species species being slightly longer than our own goliath beetles. To defend themselves from predators, Repenopodans often have a variety of physical or chemical defenses, such as powerful jaws, sharp spines, or even icky tasting slimes or sprays secreted from their joints or anus. The species pictured, the giant emerald beetile (Macroadacta macroadacta), scares off predators with a harsh, rasping hiss that it emits from its spiracles.
Repenopodans are hemimetabolous, and go through an incomplete metamorphosis that lacks a larval or pupal stage. Unusually for a less derived clade, some species are known to engage in parental care, tending to their underdeveloped children until they are large enough to live by themselves. Others, still, may simply lay their eggs in a safe enough place and leave. Regardless of the strategy, their nymphs undergo visible ontogenetic niche partitioning as they mature, avoiding competition with their adult forms by seeking out different food sources. Tropical ground-dwelling species such as the emerald beetile lay their eggs near the bottoms of trees, and have arboreal nymphs that hunt prey in the treetops. Coastal species, such as various beach-combing sea beetiles, lay their eggs near rocky tide pools, where their amphibious larvae eat algae off of rocks, somewhat like an herbivorous version of a dragonfly naiad.
Order Dracoptera ("dragon wing"): Dracopterans are an order of atavistic, predatory Ambulopterans that live in tropical and subtropical regions across Athyrmagaia. They are a very primitive yet successful grouping of flying predators, and have been around since Heteropods inherited the land. Despite being older by many millions of years, they are actually fairly comparable to odonates in terms of niche and flight adaptations, but unlike their Earthly imitations their life cycle isn't tied to the water, and they are able to inhabit a wider variety of habitats. Their fairly large size and superior ability to catch flying prey while on the wing has given them a major advantage over other flying Arthropulmonian predators such as contemporary Raptoripterans, and has allowed them to avoid extinction through niche partitioning. With their slender bodies and long, grasshopper-like hind legs, these aerial hunters are often called "dragonhoppers" or "jaguar flies".
Dracopterans are both very archaic and superbly specialized, and are some of the most well-adapted flying predators of all Arthropulmonians. Part of their success comes from the anatomy of their pteropods. Biramopterans evolved their biramous pteropods from their middle pair of walking legs, with each limb bifurcating into tarsal forewings and hindwings. This leaves most Biramopterans with only four functional legs to walk with, and while they do possess direct flight musles, their four tarsal wings are unable able to flap independantly because they are derived from a single branched limb pair, and therefore only possess a single set of flight muscles. As Ambulopterans, however, Dracopterans do not suffer from such limitations. Rather than evolving a singular bifurcating pair, they evolved four entirely separate pteropods from their first and second pairs of walking legs, with a modified tarsus acting as a foldable wingtip. Not only do these pteropods double as fully functional legs, but since they have their own separate sets of direct flight muscles, Dracopterans are able to flap them independently from one another. Most Ambulopterans can do this to some degree as well, but the Dracopterans are the unrivaled masters of this ability, and they are able to fly in all directions like a VTOL aircraft. They have a variety of different wing strokes that make this possible, using counter-stroking for hovering and precise movement, phased-stroking for fast forward movement, and synchronised stroking to perform fast turns. The jointed structure of their pteropods also allows them to change the shape of the wing while in mid-air, partially folding them to enhance such maneuvers. They can even mate while in mid-air, and they have evolved an elongated, clasper-tipped tail to make this action possible. Taking off can be a bit cumbersome when you have wings for legs, so Dracopterans have evolved two different methods of takeoff to mitigate this issue. When grounded, they launch themselves into the air using their muscular, grasshopper-like hind limbs. When perched on a stem, they use their sensory legs to grab hold to their perch as their winged limbs release their grip and unfold, allowing them to take to the air almost instantaneously. These aren't the only adaptations that make these flying predators so effective, however. The eyes of Dracopterans are enormous, so much so that their chitinous coverings are fused at the middle to form an aviator-like visor. These eyes have a visual field that wraps around the head, with a total of eight curved, rorsach-like pupils that can contract and dilate independently. To further enhance this wide visual radius, Dracopterans have a highly mobile head and upper thorax, and even while in flight they are able to track their prey and seemingly predict their trajectory. Due to this, Dracopterans have some of the greatest hunting success rates of any animal on the planet.
These animals are proud predators not only as adults, but also as young. Like all Ambulopterans, Dracopterans are hemimetabolous animals that go through an incomplete metamorphosis, a trait they coincidentally share with their Earthly odonate counterparts. Of course, the flightless nymph of a Dracopteran is not an aquatic beast with extendable jaws. Instead, it is an air-breathing, tree-dwelling ambush predator that resembles both a mantis and a katydid in appearance, armed with barbed claws that are used to seize its prey. While it is not yet capable of flight, it is quite an effective jumper in all stages of development. The buds for the wing membranes start growing during the second half of the nymph's growth cycle, which causes the first and second walking limb pairs to flatten out. At this point in development, the Dracopteran nymph already starts to develop the instincts for flight, enhancing its jumps by fluttering its flattened limbs. Dracopterans are not very sociable animals, but they will gather on occasion to mate, wooing one another by doing aerial dances. Once this flamboyant display of airborne acrobatics is finished, they mate while in flight, and the female lays her eggs on the branch of a tree, coating them in a foamy, protective ootheca.
Order Polyformica ("many-ant") - If you are on Athyrmagaia and see what looks like an enormous centipede having its orifices invaded by ants, it is not at all what it looks like. These are the Polyformicans, a very broad order of eusocial Biramopterans that are most similar to ants. They have also been referred to as "legiants" due to their peculiar social structure; while still tremendously distant cousins at best, Polyformicans are some the closest living relatives of the great Athyrmatherians. The current intepretation of the phylogenetic tree places the Polyformica at the very base of a unique clade of zooid-based Arthropulmonians called the Panathyrmatheria, which are descended from ant/termite-like eusocial organisms and include a number of other transitional clades, both extant and extinct. While the Polyformicans are certainly the less derived than the Athyrmatheria and its sister groups (by a long shot), they share very similar adaptations for body modularity and polymorphism, which gives exceptionally clear insight into how such an unusual biology came to be. In the Polyformicans of course, their version is much more in line with a eusocial society, albeit a still very odd one.
Polyformican colonies come in a multitude of strange and unique forms, and have managed to colonize a variety of ecosystems. Regardless of the type of colony, however, all Polyformican societies are based on a caste system with two main divisions: the royalty (the matriarch and her suitors) and the laborers. The matriarch is the heart and soul of the colony, a tremendous centipede or worm-like superorganism that functions similarly to a queen. Much like in true Athyrmatherians, the matriarch's body is comprised of multiple embryonically separate "zooids" that connect in a single lane, each of them being anatomically specialized for different purposes. Though unlike the former, the zooids of Polyformicans are succeeding generations of clonal daughters and granddaughters rather than siblings, and each subsequent zooid that trails behind the matriarch's main body is created via asexual embyronic budding from the cloaca of the preceding unit. When a new segment finishes growing, it detaches from the preceding segment and reattaches using a basal form of the jaw-and-telson coupling mechanism. This process occurs continuously until the matriarch reaches a particular length, and the aformentioned length of the her body is often an indicator of her age. Her suitors are exceptionally large, non-modular reproductive males that accompany her wherever she goes. Rather than competing for the privelage to mate with the matriarch, the multiple suitors cooperate with one another to ensure that they stay in her good favor. The laborers are much smaller, ant-like non-modular creatures that act as the matriarch's caretakers and aids, and are produced by the more conventional production of eggs. The laborers are often highly polymorphic, with a variety of different caste-specific morphs such as soldiers, minor and major workers, and alates (fertile males and females) that are determined by a combination of nutrient quantity and genetics. The laborers are highly cooperative, communicating using a syntax-based "language" that utilizes gestures and emissions of pheromone mixtures. Due to the peculiar nature of the Matriarch's body, most Polyformicans have an arrhenotokous sex determination system: Exclusively female workers are hatched from parthenogenetic eggs, while reproductive males are produced by fertilized eggs. Polyformicans are univoline, and once a year the matriarch will mate with one of her suitors and produce reproductive male alates that disperse to become suitors for another colony's matriarch. Within a similar time frame, she will also produce an alate queen from one of her laborer larvae by feeding it a unique secretion from her crop, something akin to the royal jelly of bees. Once this new queen meets a group of suitors of her own, she will begin her transformation into a matriarch in a temporary "coronation nest", with her multiple spouses helping to fuel her growth by finding her food.
Although the colonies of Polyformicans can take a multitude of forms, nearly all of them assume one of two different colony "types". Stationary colonies are the most common variety, and are more like traditional ant nests in structure, albeit with the elongated body of a stationary matriarch serving as the foundation for a subterranean complex. In this case, tasks such as foraging are performed solely by the laborer caste. A main tunnel is dug along the length of the matriarch's body, and various rooms such as the nursery, food and waste storage are dug around her. Nomadic colonies, on the other hand, are constantly on the move, and the matriarch acts as both a queen and a mobile nest. The spiracles of the nomadic matriarch's zooids house specialized tracheal chambers that serve as living quarters for her laborers, and both her and her laborers work together in the task of foraging. Armed with powerful claws and mandibles, the matriarchs of nomadic Polyformican colonies are often the top invertebrate predators of their respective ecosystems, and have unparalleled tracking abilities among Arthropulmonians thanks to their use of subservient laborers as a means of mapping out their surroundings.
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