The Snapscrub

Trees and Pseudotrees

The snapscrub is an open, semiarid biome found in the interiors of all three former Panapterran continents, with its largest extent in central Abeli. This page explores the central Abeli snapscrub, but many of these species can be found elsewhere. Some evolved from the inhabitants of the Middle Arthrocene's Seamless Savanna, which occupied the same location. Since then, further waves of migrating plants and animals have arrived from every direction, including some whose ancestors lived in very different climate zones. The region gets its name from its dominant tree group, the palmsnaps, of which three genera are prevalent here. They have become further adapted for xeric life, having shed inefficient features found in their recent ancestors. For example, the Provider Parasol (Umbeloconfractus magnificus) is descended from the Eartly Arthrocene parasol palmsnap, which used a large sacrificial leaf-whorl to provide shade for several smaller low-growing stems. This strategy is no longer necessary, for the Umbeloconfractus genus has since evolved to use dark purple pigments to protect its blades from sunburn, allowing it to grow as an isolated stalk that reaches a maximum of five meters high. It has an exceptionally robust stem for a palmsnap, only breaking under the force of rare cyclonal storms that make their way into the continental interior once a decade or so. It gets its common name from its abundant seeds, which it produces year-round, even during the dry months when few other plant-based foods are available. In these harsh times, herbivore species that normally partition along specialized niche boundaries gather together, feeding at every height level, for the tree grows inflorescences at each of its scarred nodes.

The jumping palmsnap genus (Spinoconfractus) has two representatives on the snapscrub. The first, known as the Shadestickle (S. umbrensis) essentially occupies the niche once held by small parasol shoots: it grows in the shade of the provider, away from the harsh desert sun. It spreads easily from one host to another, hitching a ride on the pelt or plumage of any animal that brushes by in search of parasol seeds. Its close cousin the Crown-of-Bugs (S. coronatus) is more at home in the direct sunlight. In fact, it fares best on patches of bare sand or rock where few other plants can be found. It sprouts in recently disturbed areas, using arboreal isopod seed-carriers to ensure it arrives before any other plants. These carriers are usually pregnant female Sucker-Sheriffs (various members of the genus Suctucaudus), who quickly get to work tending their host seedling and founding the colony that will protect it. For its first two years, the crown-of-bugs grows straight up, reaching 1-1.5 meters in height. From its uppermost scar nodes, it then sends out many smaller segmented stems, each tipped with tiny barb-like leaves, creating the "crown" that gives the species its name. 

One species of snapstick is also found here, growing relatively slowly as a consequence of its leafless structure. However, because of its decreased water demands, it can survive even in spots overgrown with other plants, eventually poking above smaller shrubs and grass-like grasses. It relies on its neighbors to keep it stable, as it has a relatively small root system, which allows it to put more energy into its stems. If ever the surrounding foliage is destroyed or consumed by herbivores, this plant can sense that its days are numbered; the next decent-sized storm will surely be enough to uproot it. It thus focuses its efforts on producing thousands of small, spherical seeds, borne on dense inflorescences deep within its tangle of segments. The stems themselves curl inward as their stored water is redirected, forming a tight ball that finally breaks off from its base. The Tumblesnap (Aphylloconfractus mobilis) then uses the wind to roll across the snapscrub, depositing its seeds wherever it goes. It may be a week or more before it runs out, by which time it may have traveled as much as twenty-five kilometers from its starting point. 

Only one other true tree can be found here - a diminutive pocketwood called the Pocketpod (Brevicavernus ubiquitus), which has subspecies all across Abeli and Loxodia, from the snapscrub to the matplains to the tropical rainforest. All share a small, squat shape, with only a single large pocket making up the main body of the plant. The snapscrub variety has short branches that live for up to ten years, long enough for their bases to be subsumed into the growing trunk, making them look like they emerge from random points on the pocketpod. In reality, they originate from the upper apical node and from the scar nodes contained in the pocket. Both openings face up, allowing water to collect in the basin of the pocket. This is absorbed quickly into the tree's body to prevent evaporation, as the temperature in the pod may rise as high as 33 degrees on sunny days. The pocketpod's large chamber serves as a home for nearly half the snapscrub's insect and vertebrate species, and larger animals often eat its trunk during the hot summer months when other water sources are unavailable. 

The tallest snapscrub plant is not a tree but a pseudotree. The Brevifolious Basket-Bush (Ramidendrocanistrum brevifolium), which belongs to a branching-stemmed basket-bush genus commonly called Spikeballs, can sometimes exceed eight meters tall, though six is more typical. Most members of its genus have large, broad leaves arranged in multiple closely-packed rosettes, but this species has much smaller foliage that helps it retain water. Like most basket-grasses, its leaves grow quickly and can regenerate from their base if eaten, and like most basket-bushes, its stem grows much slower, usually just a few centimeters a year. However, it can live for up to two centuries, gaining height slowly the entire time. Though it has no secondary growth, its trunk is composed almost entirely of living vascular tissue, with only a thin dead outer layer and a narrow air-filled tube in the middle. This basket-grass is second only to the provider parasol in terms of sheer seed production, though it bears its inflorescences high above the ground, so only a handful of animals can easily access them.

Herbaceous Plants

Due to the near-desert conditions of the snapscrub, its most abundant herbaceous grasses are of the wellstalk and basket-grass clades, both of which have maintained a strong foothold in dry areas for millions of years. In particular, the well-whips have found extraordinary success here. Originating in the now extinct ten-year desert, this group (now treated as a subfamily, Monophylloideae) conserves water by growing only a single leaf continuously over the course of its life. At the base of the leaf is a semi-wooden caudex that grows amorphously and, in some cases, calls into question well-whips' classification as herbaceous grasses. The Ephemeral Well-Whip (Ephemeromonophyllum magnigramenus) is one such example, relying on a large aboveground caudex to store energy through its dormant period, which lasts through the entirety of the harsh snapscrub summer. Because this nodeless, lumpy stem sits atop the ground, it has developed a hard exterior, deterring herbivores. The leaf, on the other hand, is fast-growing and rather nutritious compared to other wellstalks, often finding itself grazed upon upwards of a dozen times over the course of its fall-through-spring growing season. This is no trouble to the well-whip, which can regrow its blade at a rate of more than a meter a week. By the time summer arrives, the plant will have gathered enough solar energy to send out a shoot of large, nutlike caryopses, packed full of calories to allow its offspring to start life at a similarly feverish growth rate a few months later. 

The Tunneling Well-Whip (Ambulomonophyllum persistens), on the other hand, grows slowly all year round, packing its leaf full of poisons to deter animals from feeding on it. The underground caudex is edible but difficult to dig out; adhesive roots anchor it to rocks below the surface. Soon after sprouting at ground level, it quickly produces deeper roots that use fine hairs to cling to sediment. After a storm, as the ground dries and shrivels, these roots are placed under tension, pulling the plant's upper parts down into the cracked soil. This continues over the course of several years and dozens of dry-wet cycles until the nutritious caudex is securely out of reach of most animal threats. The meristem, which is more UV-sensitive than the blade of the leaf, also finds itself protected from the sun, so it doesn't have to go into dormancy during the summer to protect itself.

Sharing a genus and a subterranean lifestyle with the tunneling species, the Walking Well-Whip (A. copiogramenus) has an elongated caudex that grows sideways just below the surface, seeking rare pockets of damp shade beneath rocks or other plants. If it's lucky, it will find its way into a parasol grove, pocketpod chamber, or cavity in a rotting log, all of which grant access to more water than can be found beneath bare ground. This is another method of facilitating year-round growth, and it also puts the walking well-whip in contact with higher-than-average soil nutrients, in turn allowing it to flower and set seed many times per year, though rarely during the heat of summer. 

The only non-well-whip wellstalk found in the snapscrub is the Windwell (Venticalorhiza delicata), which has traded its ancestors' small, generic-looking grains for fluffy, wind-dispersed ones that, at the beginning of autumn, blanket low-lying areas in a layer several centimeters thick. Individual plants can cover hundreds of square meters and consist of thousands of individual stems, all connected by a winding, maze-like rhizomatous network. In the low-elevation basins where it grows, groundwater is mostly stagnant and many meters below the surface. The windwell easily accesses this hydrathion with its ten-meter-long fibrous roots. Forming similarly massive colonies on open hillsides is the Rock Loop-Grass (Torqueodolius xericus), whose clade's typical growth habit is that of a circularly-expanding rhizome system that eventually exhausts the nutrients in its center, resulting in a ring or spiral shape in older individuals. By contrast, this species grows in a line or gentle arc of small, scraggly rosettes, aligning itself perpendicular to the slope of its hill. This lets it catch streams of surface-level runoff, which it absorbs through its comparatively shallow roots. Multiple individuals can grow along different topographic contours on a single hill, forming terrace-like layers that catch debris over time. Each arc slowly moves uphill, feeding on the nutrients collected above itself. There is thus a continuous cycle of new individuals sprouting at the base of every snapscrub hill, while the oldest ones eventually reach the hilltop and die.

Of the snapscrub's three sweetstalk species, Scrubjade (Xericlarisaccharus vulgaris) is the most widespread, relying on its woodlouse mutualists to pile up decaying organic matter around its base. This mound holds moisture for up to two weeks after it rains, allowing the scrubjade to grow when other skystalks can't. However, population density is limited by the availability of leaf litter, which is only readily available at the beginning of summer, when many plants drop their leaves to conserve water during their dormant phase. During the rest of the year, scrubjade struggles to grow, except around stands of provider parasols, which serve as an abundant source of leaf fibers and discarded seed hulls. During wintertime, each parasol finds itself surrounded by a ring of scrubjade just beyond its drip line; these individuals' offspring then go on to recolonize the rest of the snapscrub once leaf litter is more widely available a few months later.

Floodflame (Pluvipratisaccharus ripariensis) is a species of meadowflame that grows on the shady sides of dry streams and lakebeds. In late fall, its flower stalk pokes just above the riverbank, appearing like a small red flame when viewed across the landscape from ground level. Each spikelet retains this coloration as its seed grows within, advertising the presence of bitter alkaloids. Once the seed is ripe, these toxins are removed and glucose is directed into the "petals", which turn yellow to signal that they are now edible. Animals eat the fruit-like flowers but discard the pebble-like seed, often at a different location along the bank. Since floodflame, like all sweetstalks, must die after flowering, its stem and leaves dry out as the seeds grow, rerouting water into the grains themselves. This allows them to immediately grow a wide system of wiry roots after being deposited on the ground. Leaves are not produced at this stage; they won't grow until a major flood provides a flush of water and nutrients. Until then, the floodflame goes dormant, waiting up to two years for ideal conditions.

Found in the canopies of pocketpods, parasols, and especially basket-bushes is the Basketbloom (Canistrophilosaccharus elasticus). Instead of latching onto tree stems like most epiphytic trichopileads, this species wraps its fine roots around large leaves, so it can live at the very top of tree and pseudotree canopies. It may span between several blades, forcing it to contend with twisting and pulling forces whenever a breeze shakes its host. To cope with this, the basketbloom's roots have an exterior made of elastic fibers that can stretch by a factor of three, then return to their original length. The soft core of the root is ripped apart in the process, but as a trichopilead, the basketbloom's roots are used only for anchoring, so no functionality is lost. Water is taken up through hairs on the blades, which also capture dust in the wind; this decomposes and serves as the basketbloom's main source of nutrients. 

Though trees on the snapscrub are too small to support aerial networks of creeping shadeblades, one species of basal epiphytic bladevine is commonly found on the trunks of local palm-grasses. The Starleaf Bladevine (Astrovinea pliabilis) protects itself from herbivores with spiky, digitate leaves, though in some cases this backfires and makes it even more of a target; climbing mammals have learned to grasp its petioles and pluck off these defensive structures to use as part of their own nests. Another issue the starleaf bladevine faces is the ever-expanding nature of its host. Because it is an epiphyte, it can't put roots into the tissue of the tree it grows upon; it simply clings to the trunk's outer surface. As the palm-grass's diameter grows, the roots get stretched outward, but unlike the basketbloom, they will never need to return to their original position. Instead, the branching points on each root are made of soft tissue that can expand freely throughout the vine's lifespan.

The Scrub Bloatseed (Volafructus compactus) is the snapscrub's only fruitgrass. It is a close relative of floatseeds, with both belonging to a subfamily known as Void Fruits (Cavitofructoideae). Like the floatseed, much of the bloatseed's fruit is actually a hollow, air-filled chamber. Uniquely among non-drunkfruit fruitgrasses, this genus allows its fruit to ferment as it grows, but in this case the goal isn't to attract frugivores hoping to get a buzz from consuming it. Rather, as yeast grows beneath the fruit's skin, carbon dioxide fills its chamber. When an animal or a strong gust of wind disturbs the plant, ripe bloatseed fruits violently tear themselves from their stems, using the built-up gas pressure to propel each fruit's single seed up to four meters away. If a frugivore eats the berry before it ferments, the seed can survive a trip through most digestive systems, giving this species two viable dispersal routes. 

Abelitheres

The Squamasnout (Communabelitherium fossorialis) is a descendant of Notoabelitherium communalis, which made its home in ground-level pocketwood chambers eight million years ago. This trait is still shared by all communal Notoabelitheres, of which the squamasnout is just one of about two dozen representatives. The genus is found all across the continent for which the Abelitheriid family gets its name, having spread and diversified alongside its pocketwood companions, and like many Abelitheres it has retained an insectivorous diet ancestral to swattermice. However, while many swattermice of a similar size continue to specialize on mosquitoes in flight, sqaumasnouts and several other communal Notoabelitherian species have pivoted to focus on insects and isopods on or in the soil, from the surface down to about ten centimeters underground. Compared to the more wooded environment from which its genus emerged, the squamasnout has to contend with a habitat full of heavily armed insects and crustaceans, from Dermestemimids to dipterachnids to stagmice to the toughest challenge of all, castlebugs, the latter of which have little presence in forests. In response to increasingly fierce counterattacks from its prey, this species has evolved folds of facial armor. This is made from a tougher version of the scaly-textured skin of typical rat tails. While on the tail this integument is rather thin and mainly used for thermoregulation, the modified "scales" of the nose and forehead are keratinized, and their texture is that of rough, hard leather. However, like the tail skin, the snout covering can deglove if pulled on strongly by a predator or an unexpectedly resistant prey item, regrowing in about two weeks. The rest of the squamasnout's face is also adapted to avoid serious damage from struggling victims; the ears are small, point backward, and can be tucked flat over the back of the neck during combat. The eyes are tiny and sink into their sockets when closed, and the whiskers are short and stiff, making them perfect for feeling the movements of squirming prey while the eyes are shut.

The Whiskerwig (Paniculabelitherium procerotus) is from a non-communal genus in the Notoabelithere tribe commonly called Tufted Abelitheres. This name comes from the whisker hairs that grow from the top of its head and the tips of its ears. The whiskerwig is of about average size for its genus: in other words, exceptionally large among southern Abelitheres in general, with a typical shoulder height between fifteen and eighteen centimeters. Though the tufts on its head have become important display features, they originally evolved to facilitate burrowing; while sqaumasnouts live aboveground but forage in the dirt, whiskerwigs and other tufted Abelitheres make their nests in large tunnels and graze on the surface. The burrow may extend as far as eight meters horizontally, and it is usually built into the face of a hill. This allows the whiskerwig to ascend slightly as it digs, ensuring that water can't flood in when it rains. While digging with its paws, the rat constantly uses its tuft to feel the consistency of the roof above its head; snapscrub soils are a patchwork of loose gravel and semi-concreted sediment, the former of which must be avoided to prevent a deadly cave-in. The ears, which are exceptionally large even within its genus, are used to scope for danger before the whiskerwig emerges. Keeping its head and body safely within the rim of the burrow, the rat sticks its ears out and swivels them around in every direction, listening for any unexpected noises. Older individuals build up an encyclopedic auditory memory of all the normal, non-threatening noises that occur near its home, reducing false alarms and increasing the likelihood that a genuine threat will be recognized. Once it determines the coast is clear, it wanders a short distance away before beginning to forage, as it must avoid feeding on the grasses that hide its burrow. Because it's the smallest grazing vertebrate in its habitat, it has the advantage of being able to pick through the scraggly desert grasses and choose only the softest, freshest growth, leaving the tough older plants for bigger herbivores with less discerning tastes.

Capiodocids

The tiptoe swattermice have a long history of thriving in desert environments, so it comes as no surprise that several of them reside in the comparatively mild snapscrub. The arid-adapted subfamily, which comprises most of Capiodocid diversity worldwide, is known as Docidipodinae. Common features of docidipods include triangular, often oversized ears, bare limbs, and powerful, grasping hands with only three digits. Some of the toes may also be reduced; cursorial species usually have just two, while their fossorial cousins have three functional ones and a tiny dewclaw, usually buried in the flesh of the heel. A common omnivorous species from the latter group is the Cracklemouse (Atrocidipus minimus), which can be found across Abeli's plains, savannas, snapscrub, and hot deserts alike. It eats hard-shelled food, primarily seeds and armored arthropods. Its common name references the sound that emanates from its burrow when it gets to work disassembling its victims, which include many families of isopods and flies. It hunts at night and can tackle prey up to 150% its own size, dragging them underground before removing the dangerous legs, mouthparts, and pointy pieces of armor. It is prone to surplus-hunting, even when plant-based foods are also available. It builds a stockpile of protein-rich food, with each piece of meat protected from spoilage by its owner's exoskeleton and immune system. During the scorching months when most plants aren't producing and most prey is hunkered down beneath the surface, the cracklemouse sates its high metabolic demands by finally ending the suffering of its defeated targets, eating a few each day until other food sources return.

The Quilleye (Macrodipus hystricotrichus)'s common name refers not to any features around its own eyes, but rather to what is likely to happen to any predator that gets too close. Its tail hairs are sharp and covered in microscopic barb-shaped scales, and they pop out of their follicles as soon as they stop growing. They are then held in place solely by overlapping layers of scaly tail skin. When threatened, the quilleye can swivel its tail in typical swattermouse fashion, often crouching down and holding its tail over its body. If a predator chooses to take a swipe anyway, the pain of the quills may force it to halt, giving the rat time to flee at speeds of up to 30 km/h. Unfortunately, owing to the quilleye's small size, it is just as likely to be killed outright in such an attack. The predator, of course, learns not to mess with quilleyes in the future, but this is no advantage to the deceased individual. There is thus a selective pressure for this species to aposematically advertise the danger it poses and to use its quills only as a last resort. To this end, its skin and hair are packed with phaeomelanin, giving the quilleye a school-bus-yellow coloration that contrasts against the grey-beige earth of the snapscrub.

Loxoditheres

Sphynxes (Insilloxoditherium) are a genus of fully carnivorous rats descended from the ailurat. Having adopted a raptorial ambush style of hunting, they are now able to occupy a much wider range of habitats - essentially, anywhere with small prey species and sufficient cover to set up their attacks. The Scrub Sphynx (I. triangularis) is a bit smaller than average for its genus, but no less effective a predator. It hunts at ground level and in the trees, and its typical food items include whiskerwigs, squamasnouts, cracklemice, stashers, and the chicks and eggs of larger birds. It lies in wait, concealing itself below a thicket of vegetation, behind a rock, or within a pocketpod. The moment a small animal wanders by, the sphynx leaps, powered by muscular hindlimbs and a flexible spine. Its forepaws are small and not very dextrous, but they can be turned inward to grasp victims between them. The hand claws, sharp and highly curved, emerge at a slightly upward angle from the fingers, allowing their tips to hover just a couple millimeters above the abrasive soil. This keeps them sharp for clinging onto prey, which is killed quickly with a single bite to the neck.

Clutchbirds

Stashers (Ophiothylacavis) are a genus of arboreal pouchwing clutchbirds descended from the small-bodied tree-dwellers of the previous age's Bayouland flood forests. They have a large, fluff-covered pouch like all members of their subfamily, but they have ceased to use it for incubating their eggs. Instead, all species in this genus are cavity or cup nesters, only rarely carrying their eggs if the nest is damaged or threatened by predators. In day-to-day life, the pouch is used primarily for storing and carrying food, serving as a second crop with the added benefit of keeping its contents dry. There are over a hundred species of stashers found across all three major continents, and they range in size from about a hundred grams to over three kilos. All stashers have zygodactyl feet, but those that forage on the ground can lean forward and put weight solely on the front two, allowing them to walk efficiently. Only one stasher species lives on the snapscrub: the Desert Flystash (O. dipterachnophilus), which as its name suggests is more at home in environments even drier than can be found here. It lives in pocketpods and feeds on tough venomous insects, especially dipterachnids. It dispatches its prey by grabbing an extremity and flinging it back and forth with frantic movements of its snake-like neck, a feature that evolved in the first stashers to make it easier to reach into the pouch. It finds greater difficulty when dealing with armored isopods, which tend to be resilient against its mode of attack, simply curling up to avoid damage. Since woodlice are more common on the snapscrub than dipterachnids, the desert flystash can only exist here in small numbers. However, in harsher deserts, dessication-resistant dipterachnids compose a greater fraction of arthropod biomass, so this species can reach greater population densities and body sizes. Regardless of habitat, flystashes also eat a significant number of flying flies, mostly Pugilopsids, and they are happy to take advantage of easy prey in the form of dustflies. Their long neck comes in handy again when catching flighted prey, as it can coil backward before suddenly springing towards unsuspecting insects.

The other tribe of the pouchwing subfamily are called Ouchwings (Unguithylacavini) because they retain a claw on each arm, allowing them to fend off would-be egg thieves. Ouchwings are the combined descendants of woodland and plains pouchwings, which continued interbreeding until just three million years ago, while the arboreal tribe branched off a further three million before that. The Apteroos (Macropodornis) are the widest-ranging ouchwing genus, distributed across open areas of Abeli, Loxodia, and Ailuropia. This is because their highly efficient locomotion requires relatively clear ground. The Bluntbeaked Apteroo (M. brevirostris), though on the small side at only about 120 centimeters tall, exemplifies its genus's method of movement perfectly. It has an elastic tendon running all the way from the back of each knee to the base of the middle toe. When it lands from a hop, this tendon is stretched as the angle of the heel joint gets sharper. It then springs back to its neutral position, straightening the leg and propelling the apteroo into its next bound. It can hop for many kilometers across the snapscrub, hardly ever returning to the same place twice. Family groups consisting of one male, one to four females, and as many as thirty juveniles travel together, occasionally joining together in migratory grazing herds of a thousand or more during the height of the spring growing season.

Wallabirds (Frugivornis) are more sedentary and generalistic, though their greatest affinity is to sweet foods like bloatseed fruits and floodflame flowers. They also eat soft-shelled seeds, high-quality foliage, exposed rhizomes and tubers, medium-to-large arthropods with modest armor, and occasionally small vertebrates. The Woolly Wallabird (F. tomentosus), whose feathers are white and reflect the heat of the sun, is a common sight in and around parasol groves. It avoids venturing too far away, as it can't hop and doesn't run particularly fast, relying instead on the trees for cover. It is among the largest animals still capable of squeezing into an unoccupied pocketpod, and it is also a decent climber, often perching for hours atop a parasol or basket-bush until it feels safe.

The Kiweasel (Mustelapteryx vulgaris) is a species of microcarnivorn that descends from the slinkstepper. It has relatively long legs for a fossorial animal, but it can tuck its knees above its shoulders when underground, relying only on its feet to push it along. Across Abeli and Loxodia, this species has largely outcompeted small ratweasels, as its beak allows it to more easily pluck wedged-in prey from their burrows. It sometimes then steals these burrows to raise its own young, especially when it finds a sturdy whiskerwig tunnel, but it is also capable of building its own if necessary. It isn't purely a subterranean predator; it also goes after small rodents like squamasnouts and cracklemice if it catches them out in the open, and it is a rather efficient egg thief. Relying on its small size to go undetected, it can sneak into nests on the ground or in pocketpods. If it feels daring or desperate enough, it may even use its tweezer-like beak to quietly pluck an egg from the pouch of a sleeping ouchbird.

Packslashes (Macrocarnivornis socialis) are descended from the slashstepper and live all over the Abelian continent. They come in two morphs: a robust form has its greatests numbers in forests, a gracile one is most common in rough habitats like mountainsides and the snapscrub, and their ranges overlap on the open temperate plains. In some cases, their genetics get messy due to interbreeding, but the snapscrub's gracile packslash has no genetic contribution from robust conspecifics, so it can easily be defined as a distinct subspecies, M. s. megageococcyx. It is not purely carnivorous, sometimes subsisting on fruits and tubers, but it is a speedy and formidable predator - the largest found here. It has many features that help it pursue prey: its "free" caudal vertebrae are held together as a single stiff rod by overlapping bony struts (though the pygostyle can lift up and swivel around for display purposes); its legs are long and lean, with most of their force coming from the neocaudofemoralis that connects to the tail; and its neck is held high when running, allowing it to swing downward with force to land a powerful first bite as it catches up to its target. It then begins inflicting cuts with the side of its beak in the standard slashstepper way, and feeding commences as soon as its prey isn't moving, though it may remain alive for quite some time, as the packslash first goes for the protein-packed muscles of the limbs, only later moving on to eat the internals. The lack of massive herbivores on the snapscrub means packslashes here rarely hunt together, but they still help each other in other ways. Because the availability of small game in a particular area fluctuates with boom-and-bust population cycles, individuals in neighboring territories may see wildly different success rates. A packslash that experiences a surplus often offers some of its catch to another that wasn't so lucky, for it may only be a matter of months before the roles are reversed.

The Common Struttio (Ornatocaudornis velox vulgaris) belongs to the sole species of a third clutchbird subfamily, one whose ancestor was the greater steppestepper. Its recently-connected populations across all three major continents are genetically too closely related to merit even subspecies status, but they display a tremendous range of sizes; the snapscrub struttio (itself not the smallest type) is under a meter high at the shoulder, while its savanna counterpart may stand thrice as tall and weigh 300 kilograms. The struttio is in a sense a successor to its Early Muricene ancestor Apteryx grandis, sharing its niche of "large-bodied 'normal' ratite". In fact, its long tail is one of the only traits that would make it stand out in a lineup of ostriches, emus, and rheas from Holocene Earth. Since those three families each evolved this lifestyle independently, it should come as no surprise that a kiwi found a place for itself doing the same thing. This was the case for A. grandis, but it was driven off the plains by the rise of early rat predators. With its large clutch size and impressive speed, the steppestepper gave its lineage a chance to retread old ground, the result of which is the struttio.

O. velox is all around the best runner on the snapscrub. Apteroos can travel longer distances without getting exhausted, and packslashes can reach higher top speeds when running in a straight line, but the struttio has a powerful combination of endurance, speed, and most importantly a tight turning radius, made possible by its flexible tail. Though this sacrifices some of the strength of the neocaudofemoralis muscle, this species can still reach a speed of 60 km/h, powering its legs primarily with muscles that attach to the core of the body and secondarily with large muscles on the legs themselves. This speed can be maintained for ten minutes or more, and it drops by only a few kilometers per hour even when making tight turns; at full tilt, the struttio can switch directions by ninety degrees in just two and a half body lengths. It also doesn't require flat ground; it easily navigates rocky terrain where packslashes must slow down for fear of tripping. Though unmated males may spend several years wandering the snapscrub and avoiding predation using these athletic abilities, breeding adults live the remainder of their lives in large territories, typically marked by natural barriers like rivers, rock faces, or dense parasol groves. Only in featureless areas are disputes likely to occur, and these are settled using an escalating series of displays. First, the two belligerents show off their pygostylic feathers, trying to appear large and intimidating. Next, they coil their necks around one another. This gives each the chance to truly assess its opponent's strength before an all-out fight begins, and the weaker of the two usually shies away at this stage. However, if the two are evenly matched, or if the weaker one refuses to back off, they begin kicking each other with their spike-like claws, aiming to bruise and bloody their enemy's ribs, belly, and legs. This may prove fatal, in which case the victor doesn't always claim the entirety of its defeated competitor's old territory, leaving space for roaming juveniles to recruit into the local area.

Other Birds

The Boxerbird (Pugilavis continentalis) is a mid-sized gannetgrouse species and the second snapscrub animal to have evolved sharp quilly integument. Unlike the quilleye, its needle-like feathers do not detach. They have a smooth exterior because their function is to inflict multiple puncture wounds as the bird repeatedly backs its rear end towards any attacking carnivore. Intraspecific conflicts are resolved in a very different fashion. Seacocks have long used their wings for display purposes, beginning with the green-winged seacock, which lived around the year 2,500,000 Post-Abandoment and fluttered its colorful plumes to attract mates. These feather tufts emerged from the tip of the stubby wing, and the shoulder and elbow joints gained strong muscles to move them in complex patterns. This was further developed in earlier gannetgrouse species, some of which began boxing for mating rights. It was at this stage when the bones of the shoulder began to realign, resulting in a forward-backward range of motion instead of typical up-down flapping. This boxing behavior is, as the name suggests, one of the defining features of the modern boxerbird. The hand tufts have shrunk away, with the feathers turning into scaly keratinous bumps that increase the damage of each punch. However, the arms are not used for fighting purposes only. When foraging for plant matter on the ground, the boxerbird can drop to all fours, bringing its face close to its food without setting itself off balance. It is specialized for eating roughage, with its cheek pouches serving as a repository for powerful digestive bacteria that let it extract energy from dry leaves, twigs, bark, roots, and seeds. It gorges itself on windwell fluff during the autumn bonanza, gaining so much weight in the process that it becomes more comfortable to walk quadrupedally for up to a month afterward.

The Barbelbird (Nudapteryx subxericus) is a large, short-beaked mudslider groundgoose that builds its burrows into floodflame patches on steep, low-lying slopes near ephemeral water sources. Starting near the top of the bank, mated pairs build a tunnel that curves down and back up again, terminating in a large central chamber where one egg is laid by the female per year. The top of this room is sculpted into a dome, then coated by mucus-laden saliva, which hardens into a glue-like coating. This makes the roof water-repellent, so very little drips down when rain batters the ground above. When the water level rises and floods the entryway, the dome holds a large air pocket, so its inhabitants never need to flee to higher ground. During dry times, both adults tunnel extensively, building a labyrinth of narrow passages impassible to all predators but the smallest kiweasels. This species eats any small subterranean animals it can find that don't have extensive armor: squashroaches, airgills, and basal Dermestemimids are among its favorite prey. However, it isn't totally restricted to the underground world. When the snapscrub's rivers and lakes flood, a short-lived aquatic ecosystem of algae, copepods, mosquito larvae, gillywogs, and small aquatic isopods takes shape. Of these, only the larval mosquitoes originate from parents that make their living on the snapscrub during dry times; the rest must remain dormant as eggs or cysts for many months in between floods. The barbelbird is the apex predator of this community. Though its digging feet are not webbed, they are broad enough to push a large volume of water, making this species the fastest underwater swimmer in its habitat. Just as it does beneath the soil, it uses its facial whisker-feathers to feel for prey in the turbid water. It can be seen churning the water just beneath the waves, sometimes even propelling itself above the surface in pursuit of prey. It then disappears into the murky green-brown soup again before its own predators can get a good look at it.

Scansoriarthriform Isopods

Woodlice of the order Scansoriarthriformes have an easier time than others when expanding into dry environments, for in many cases they can get all the water they need from their host plants, drinking nectar or wetting their gills with pure water, which pocketpods secrete in the tops of their chambers. The tree does this not only to promote the health of its inhabitants, but also to give herbivores an easier alternative than getting the water by eating the plant itself. It is up to the pocketpod's mutualists to ensure this water is protected from organisms that don't benefit the pocketwood in return, but thieves slip past these defenses relatively often. During the everyday life of the tree, the primary caretaker of its internal cavity is the Pod Scrubsnatcher (Gracilibrachiarthrus cavernensis). This species spends almost its entire life in the dimly-lit chamber, leaving only in mid-spring to reproduce with scrubsnatchers from neighboring colonies. About a hundred of them can live in a full-grown pocketpod, and they are content to share their space with the myriad other species that use the short tree for shelter. They have sensitive antennae and can quickly detect if another animal is inflicting damage on their home, in which case they will attack together, crowding around the central columnar stem in hopes of driving the intruder out through one of the chamber exits. More aggressive attackers, however, are not allowed to leave. The scrubsnatchers instead line up around the exit holes, then slowly close in around their target, pinning it down with their long, piercing arms. This genus isn't designed to rip and pull its enemies apart like the first snatchers did. Instead, it punctures flesh with a direct jab, then tears a long gash by pulling its arms back along their serrated edges. This causes death by blood loss within a few minutes when many of them work together.

The Parasol Scrubsnatcher (G. umbelensis) is bigger and stronger than its congeneric, as it must spend most of its time out in the open. It patrols the length of the provider's blades, deterring seed-eaters from opportunistically snacking on the plant's foliage as well. It does not, however, defend the infructescences in any way, which in some cases means sap-sucking parasites can gain a foothold there. On the smooth internode sections can be found the Ring Barkflap (Ambulosuber annulus), which belongs to a genus of unusually active barkflaps, who typically don't move much as adults. Because of the harsh environment of the snapscrub, provider parasols can only afford to secrete a relatively tiny supply of nectar along their trunks, meaning the ring barkflaps cannot sustain a population that covers the entire tree's surface. Instead, only a few dozen live on the parasol at any given time, usually hiding up in the canopy. Here the tree produces ample nectar to feed its snatchers, who largely dominate the nectaries, but the barkflaps can still secure a drop or two here and there. In the event of an attack by Pestilarthrid woodlice, the barkflaps line up in a single ring near the tree's base. This allows them to defend against Plague isopods and other herbivores up to a kilogram in size, and larger enemies can at least be slowed down enough to give the scrubsnatchers time to join the fight.

While parasols can protect themselves with only small numbers of two Scansoriarthriform species, pocketpods need a much larger army to stay safe. In addition to their scrubsnatchers, they also employ a species of sucker-sheriff, a mid-sized Altiperiid genus with highly derived tail anatomy. The distal segments are poorly sclerotized and overlap to create a rounded cup shape, allowing them to form an airtight seal when downward force is applied. Though the sucker-sheriff can attach itself to a flat spot on the pocketpod's broad trunk, it prefers to let its enemies do the work. When approached, it feigns injury, allowing several Pestilarthrids to pin it down. It folds in half, resting its back atop its sucker, and allows itself to be pressed into the stem. Its gills are hidden under a protective plate derived from the first segment of the hindlegs, a feature that originally evolved for water retention. The fused plate now also serves to shield the softest parts of the underbelly, repelling Plague while the sucker-sheriff's fourteen grasping legs search for a target. Once one is singled out, the sheriff's tarsal claws latch onto it, pulling it towards the mouth. The crushing mouthparts are short and non-dexterous, but their conical shape allows them to punch through arthropod armor, usually aiming for the middle of the face when dispatching Pestilarthrids.

The nectarleech subfamily Floricisorinae has two representatives here; the first is a member of a large radiation that diversified alongside the sweetstalks. The Scrubjade Nectarleech (Microfloricisor xericus) is an avid nectar thief, though it does not live exclusively on the scrubjade plant as an adult. Juveniles, however, are so small (under a third of a millimeter) that they risk drowning in the nectar of large flowers. Scrubjade, with its numerous tiny florets, provides a perfect nursery; during the warm months, mothers climb slowly from one flower to another, depositing a single egg on each. The young nestle into the base of the floret, squeeze their heads between the protective palea and lemma, and begin drinking without harming the nectaries at the tips of the lodicules. If the plant senses damage to a floret, it will self-prune, cutting off the supply of nutrients. If the mancae are successful, though, they'll be free to feed for several weeks, undetected by their host and its defenders. As they grow up, they may continue feeding on scrubjade, but such individuals are eventually forced to start eating florets directly, for their heads are no longer big enough to fit inside. If few jade skystalks are available, nectarleeches will start seeking out other snapscrub sweetstalks, especially the large flowers of floodflame. There they can return to a gentler feeding style, one that still allows the flowers to get pollinated by dustflies and develop into seeds.

The False Flipbug (Pseudosquamulus) is an early-branching genus in the leechbug family, one that feeds on trees instead of sweetstalks. It can parasitize all the different trees in the snapscrub; in fact, it's one of the few population controls on the highly prolific tumblesnaps and crowns-of-bugs. When it infests parasols, it displays a unique behavior to avoid being ousted by their barkflaps: it joins the ranks and pretends to be one of them. Though the resemblance is far from perfect, false flipbugs are a relatively new species in this biome, so the barkflap bugs haven't yet evolved to recognize them consistently. The nectarleeches can thus be bold, walking alongside their hostile relatives with impunity. They can't do this on any other snapscrub tree, for parasols are the only species here inhabited by barkflaps, but they evolved in forests further south where the latter are the primary defense mechanism for many trees. Since the Scrub False Flipbug (P. dimorphicus) arrived here less than a quarter of a million years ago, it has begun to diverge into two lineages; one is trending towards sessility and inconspicuousness, while the other is adopting scent markers and characteristic behaviors that help it blend in with barkflaps. 

The secondarily solitary Cursoriarthrids also have a two-species presence here. Both are stagmice, with the larger of the two being rather closely related to the giant cantercrabs. The Sifting Stagmouse (Glareofortipes timidus) grows to about eight centimeters long, and it shares the cantercrab's limb arrangement, but it eats only small, soft food. It uses its rake-like foreclaws to filter-feed, dragging them through the loose pebbles that cover much of the snapscrub ground. Its low-slung shell and three powerful pairs of weight-bearing legs allow it to quickly bury itself in this same substrate if a predator approaches. When the coast is clear, this stagmouse is often brave enough to search for another food source. The piles of debris that form beneath trees and scrubjade plants are much richer in edible detritus, but they are almost always patrolled by sucker-sheriffs, scrubsnatchers, and other Scansoriarthriforms not detailed in this article. To fend them off, the sifters travel in groups of siblings from the same clutch, with one or two individuals picking off the defenders while the rest of the herd feeds. The foraging herdmates share food readily, presenting it to the sentries on outstretched front legs. 

The much smaller False Herald (Microfortipes imitator) benefits from the good reputation of some of its cousins. Having evolved a ranger-like body shape, it mimics the chemical and visual signals that rangers use to alert other Scansoriarthriforms to the approach of Pestilarthrids. In exchange, it gets a reward of nectar and protein-rich fungi, which it eats quickly before fleeing the scene. By the time the defending isopods realize danger isn't coming, the culprit is long gone, off to secure its next meal elsewhere. In response, many arboreal communities have recently begun to form close partnerships with just one true ranger species, several of which have evolved bright white markings to help their mutualists recognize them. The reign of the false heralds is thus on the decline in the trees, but skystalks are still vulnerable, as they can't afford to form specialized partnerships. With hosts that are smaller, softer, and usually out in the open, the woodlice that defend species like scrubjade and floodflame must react to any indication of threat, even if that means accepting a high risk of false alarms.

Other Isopods

The ability to glide (or otherwise greatly slow one's descent rate) has evolved several times already on Apterra, in part due to its relative lack of flying clades compared to Earth. Two large solitary woodlice, both originally from much wetter environments, are now common on the snapscrub. Pseudelytrians (tribe Pseudelytriini) evolved in Loxodia, and most species are still restricted to the northern half of that continent, but the arboreal Bristly Pseudelytrians (Plumophysis) have used their gliding skills to spread much further. The Xeric Bristly Pseudelytrian (P. abeliensis) is at the furthest frontier of this ongoing expansion; it only reached the snapscrub about 30,000 years ago, and its descendants are bound to keep moving; in a few more tens of millennia, they will populate the western half of the continent. Its heavy exoskeleton hinders it somewhat when gliding, so adults don't go for maximum distance. They simply seek the closest hiding spot, often one pre-chosen directly below their perch. Juveniles, however, must disperse to find a territory of their own. With lighter, less sclerotized bodies, they can achieve a glide ratio of 20:1 or greater, using their bristly legs and tails to increase their effective wing area. 

Squashroaches are a far more ubiquitous group, found worldwide and numbering over 300 species (including about 120 of dodecipedes). Squashgliders (Aeropronarthrus) are an Abelian genus from the tropical rainforests along the Medithalassic coast. They've been able to expand into drier habitats thanks to a soft pouch that enshrouds the gills. In most squashglider species, this pouch only exists in females carrying eggs, keeping them safe from bumps and scrapes upon landing. In the Scrub Squashglider (A. xericus), all individuals are born with this pouch, which not only keeps the gills wet but prevents them from being torn apart by landing on coarse sand. This species' gliding style begins with a vertical drop to gain speed, during which all the limbs are tucked under the body to reduce drag. At about a meter off the ground, it levels out, shooting forward about seven times the distance it dropped from. Though this is a rather poor glide ratio, it allows the soft-bodied squashroach to avoid gliding through open air, where it is most vulnerable to predation.

The other squashroaches that live here are in no way restricted to this habitat; both species have interconnected populations across Abeli. The Pocketpod Squashslider (Levipronarthrus cavernensis), as its name suggests, is affiliated with the pocketpod tree, and there is nowhere in the pocketpod's range where L. cavernensis cannot be found. Its role is to prevent decay and putrefaction in the tree's cavern; its young live and feed at the bottom where water collects, consuming rotten detritus that might otherwise cause infection in the tree or its other inhabitants. Layers of decaying twigs, leaves, and small animal carcasses build up over time, and these hinder the pocketpod's ability to absorb the water that pools in the same place. The tree's resident Scansoriarthriforms are rather sensitive to infectious diseases, but squashsliders are far more resilient. The genus first lived in matkelp rafts where rot and disease were never far away, and when they encountered early pocketpods growing on the shoreline, they took up residence there too. Their presence allowed pocketpods to grow much larger cavities without risking pathogen buildup. The two soon spread into environments with high disease risk, such as rainforests, and those where water scarcity necessitated large size, such as the snapscrub.

The Giant Pebblepede (Saxibates gigas) is solitary and long-lived, with some individuals reaching well into their third decade. It lives in poorly-vegetated microhabitats within the snapscrub, such as dry creekbeds, gravelly rubble piles, outcrops of bedrock, and occasionally the small sand dunes that form during the dry months. The sclerotized warty lumps on its back are mostly for camouflage, and with each molt they lighten or darken to match the surrounding substrate. This makes dispersal easier, as individuals can settle in an area and quickly begin to blend in, even if their ancestors lived with a completely different background for generations. Pebblepedes are ambush predators, and they may remain motionless for weeks or months between meals. Their most common prey are large flightless arthropods, but unwary juvenile birds and rats can also be taken when crossing barren ground.

Janitor bugs (subfamily Sturcopellinae) were the first of Apterra's animals to make a meal of others' waste, and though they are now joined by the likes of megafaunal coprovores, they still provide most of the planet's terrestrial dung disposal. Most janitors these days are cryptic and shy; coprovores are specialized for following great herds of animals, so the Sturcopellines rarely get a chance to take advantage of that food source anymore. Instead, they focus on smaller or non-herding animals, and they've become smaller themselves to get by on less. Only the Armored Janitors (Thyreopurgor), who live in arid regions where coprovore populations are low, have retained a large size. The snapscrub, of course, is not a full-on desert, so coprovores pass through certain spots many times per year, but they must tread carefully, as the Razor-Shelled Janitor Bug (T. debilifex) has evolved to fight back. Though not the largest of its genus, this species is the most heavily armored, with a series of tooth-like spines along its back. These start out dull to prevent damage during molting, but the thinner spines at the base of its tail can be scraped against them until they become as sharp as knives. 

During lean times, individuals establish their territories where herbaceous plants dominate, as these attract the highest volume of herbivores. Between closely-packed scrubjade plants, beneath thickets of tumblesnaps, and in the shade of bloatseed bushes, razor-shelled janitor bug burrows are a common sight. Each leads to a long corridor that ascends and descends repeatedly, ensuring that air pockets will always remain at the high points if floodwaters fill the tunnel. This sinusoidal pattern continues for up to four meters, and at the bottom of each trough is buried a small ball of dung, enough to feed about a dozen mancae until they're ready to leave and establish their own territory. Summer rainstorms trigger synchronized dispersal of juveniles from many burrows. This achieves predator satiation, and it has a secondary benefit: within the next couple days, the rain will promote plant growth, which will bring herds of grazers and browsers, which will in turn bring coprovores. They will arrive to a land dotted with half-grown spiny isopods, forcing them to avoid the small hazards. The nomadic herbivores and coprovores are thus restricted to tree-covered patches of the snapscrub where janitor bugs are rare. Local species, which have evolved to pay close attention to where they step during this dispersal event, can still move freely about the landscape.

Pockmarkers (Excisarthrus) are smaller than average even for Late Arthrocene janitor bugs, never exceeding a centimeter in length. The local species is a generalist; it takes advantage of all sorts of fallen foods, from dung to carrion to the tattered remains of bloatseed berries after they explode. Shy and weakly armored, the Sand-Sliding Pockmarker (E. xericus) must contend with a variety of larger and scarier scavengers, so it makes away with its food as quickly as it can. It uses its kitchen-knife-like forelegs to smoothly cut out a chunk up to half its own body size, then scurries away to find a safe place to stash it. It is most active during the driest months, when carrion is abundant and small dunes are scattered across the snapscrub. These provide much-needed refuge for a pockmarker fleeing conflict; it dives in and "swims" with its paddle-like pleon. The legs and head tuck beneath the body, keeping its food safe from damage, and the cephalothoracic hump becomes the front of the body. Its curved shape allows it to serve as a scoop, pushing away obstructions. At the bottom of the dune, where sand turns to hardened earth or gravel, is where the pockmarker stashes its collection of morsels. Males are semi-nomadic and usually store only a few scraps at a time, but females are territorial, as they must build up an extensive larder to feed their young. A well-fed manca will molt at least three times before exhausting its food source and making its journey to the surface. It then takes up temporary residence beneath a parasol grove and feeds on the droppings of small vertebrates until it reaches adulthood. 

The Desert Geoglyph (Geoglyph aridicus) is the largest terrestrial crustacean on Apterra, with elderly females growing to over 40cm long and three kilograms in mass. It belongs to the castlebug family, and its diet is that of a generalist herbivore. Its gigantism evolved as a water retention mechanism; it can go several days without drinking or eating succulent plants for hydration. This species has no gill-protective coverings like so many other isopods in its range. Instead, it has a series of small glands on its underside that drip water onto its gills whenever they start to dry out. When resting, it can further conserve water by digging a shallow pit and hunkering down. It lines this temporary nest with thick mucus excreted from the same glandular system, allowing it to maintain a chamber of high humidity beneath its body. Young mancae stay with their mother for several months, taking advantage of this protective environment until they are large enough to build their own.

Though its top speed is only about a kilometer per hour, G. aridicus spends about fourteen hours a day on the move, foraging as it goes. This allows it to cross huge distances without drawing attention to itself, though it leaves a conspicuous trail in its wake. Its great size and low-slung body cause it to plow through sediment as it walks, resulting in long U-shaped paths of disturbed sand, gravel, or dirt. These lines, which often take on complex geometric shapes that reflect the geoglyph's foraging patterns, are the basis of its common name. In true desert environments, geoglyph trails crisscross the surface at a high density, as there are few other herbivores competing for food. Here, though far from rare, these trails are usually separated by many meters, and adults rarely stumble across one another. This species also tends to be smaller in the snapscrub, rarely exceeding 30cm in length, as water is less scarce and competition is fiercer. 

Melodiopterygines

Songbugs (subfamily Melodiopteryginae) have exploded in diversity worldwide since the Middle Arthrocene, with eight species commonly found on the snapscrub. Their defining features are derived parental care, complex stridulatory communication, and multi-jointed jaws that allow for fine manipulation at the expense of the strength possessed by other Pugilopsids. Some species even have hypertrophied maxillary palps branching off about halfway down their maxillae, giving them a pair of weak pincers on their face. Their venom is usually weak, and most species no longer use it to kill prey; it functions primarily to soften food so it can be slurped up through the proboscis.

1) Snapscrub True Shufflefly (Collembolaphaga eucollembolaphaga xerica): Of the three shufflefly species in this habitat, only this one retains its ancestral diet of soil-dwelling springtails. Adults and larvae alike feed on the small hexapods, which are easy to find but evade capture well with their furcula. The shufflefly's size is an advantage, allowing it to create no noticeable vibrations as it approaches its target. When it strikes, it immediately lifts its prey off the ground to prevent it from escaping. It then uses its sharp proboscis to stab the base of the furcula, injecting a tiny amount of venom to incapacitate the jumping organ. With its main defense rendered useless, the springtail can be safely carried off, alive, to the shufflefly's nest. The clutch of 10-20 larvae usually requires two or three full-grown springtails per instar, so the parents need to catch about a dozen over the course of their young's 72-hour childhood. Adults feed less frequently, as they have little need for protein-rich animal prey, but they may visit scrubjade flowers for carbohydrates.

2) Bloatborer (Collembolaphaga frugiphila xerica): This shufflefly has an aberrant parasitic lifestyle. Five million years ago, its ancestors were specialized to feed on springtails that congregated around rotting fruitgrass berries. As time went on, larger frugivorous arthropods evolved and began to carry off these fruits in massive numbers, so this shufflefly lineage adapted to arrive earlier to freshly-fallen berries. As they showed up sooner and sooner, the flies found fewer and fewer springtails, so they began to feed on the fruits themselves, even laying their eggs within the flesh instead of building nests. Other frugivorous insects, especially those of the carryberry genus, still posed a risk, as they were happy to steal a shufflefly-inhabited berry for their own young. With so many of their larvae being eaten by other songbug species, this berry-boring shufflefly nearly went extinct, but one group survived with a different strategy. Its eggs did not hatch within the berry at all; instead, they remained dormant in a tough capsule until consumed by a larger insect larva. Within the foregut, the eggs would hatch, and one or more shufflefly larvae would begin eating their host from the inside out. These stowaways gave rise to the modern Boring Shufflefly (C. frugiphila), which has already branched into new subspecies in many different biomes. Each uses a particular fruitgrass and a particular insect host, and many are branching out to target adult frugivores as well. The snapscrub subspecies sticks to larvae, infecting the young of buryberries, podstashers, and rarely certain dustflies. Adults no longer eat at all, living only long enough to mate and deposit eggs into as many as a hundred berries.

3) Parasol Mancamasq (Collembolaphaga infiltratrix umbeliphila): This shufflefly's diet has changed to a more moderate degree; it now eats young isopods instead of springtails. In larval form, it can target nearly any social woodlouse, as it can adjust its scent to mimic many different hosts. A single egg is laid on a gravid Scansoriarthriform, Pestilarthrid, or any other species that cares for its young after they hatch. The young larva is an unassuming grey-white oval that bears only a passing visual resemblance to a woodlouse manca, but it quickly detects their scent and begins to produce the same volatile molecules from a patch on its face. The mother isopod is thus fooled and begins to protect the brood-parasite as one of its own offspring. Meanwhile, the mancamasq quietly kills one of its "siblings" after another. It eventually grows big enough to slip away and pupate, and females will almost always lay their eggs on the same species they grew up with. Males, however, have no preference, so they serve to maintain gene flow between females that specialize on different isopods.

4) Buryberry (Fructicoliculex fodiens): This flightless species of carryberry lives only in dry environments with low predation pressure and lots of exposed ground. It plucks only mature, unblemished fruits in hopes of avoiding parasites. Each berry is rolled across the surface to an inconspicuous spot, then buried at least 10 centimeters deep. A single egg is laid inside, and if all goes well, the larva will get all the energy it needs to reach maturity in the safety of this burrow. It pupates and emerges as a subimago, and if it is not infected with any bloatborers, it immediately molts again into the imago stage. If it is infected, the parasites emerge from its anus at this stage and flee for the surface. The subimaginal buryberry, internally scarred and having lost much of its energy to the shuffleflies, must then search for food at ground level to finish its growth. Retaining larviform mandibulate mouthparts, it is a generalist, eating anything it can find as soon as it leaves its natal burrow. Though such individuals face a high risk of predation, their final molt gives them a chance to heal their wounds and emerge as fully-functional adults, which go on to eat adult bloatborers and other insects that parasitize bloatseed berries.

5) Scrub Podstasher (Fructicoliculex maximus xericus): The largest of the carryberries, this one is a strong flier and yet another species with a distribution matching that of the pocketpod tree. It is not picky with the bloatseed berries it takes, as it can transport dozens in a single day to its nest inside a tree chamber. Its young thus face far greater parasite and disease loads, and up to 75% are lost to bloatborers alone. The survivors can recover in a similar fashion to the buryberry; this healing ability has evolved many times in the Amphibiinsects as more endoparasites have evolved over the epochs. Every part of the body can be refreshed in this final molt, except for the wings, which are retained from subimago to imago. Adults spend most of their time around the pocketpod, protecting the nest from airborne predators. 

6) Swiping Songbug (Pseudofalcoculex vulgaris): This species in the swooping songbug genus is not one of those that outcompeted large falconflies. It instead feeds by gleaning arthropods of 1-4 cm in length from exposed surfaces like leaves, logs, and rocks. Such a prey base is unusually broad for a songbug, allowing it to achieve success all across the Abelian and Loxodian continents. 

7) Stooping Songbug (Pseudofalcoculex canipugnax): This species belongs to a group of aerial hunters that target prey their own size or larger. On the snapscrub, this means large dustflies and a few other songbugs are on the menu. Members of this clade (subgenus canipugnax) are also known as Dogfighters, and they have contributed more than any other lineage to the decline of most Falcoculicids. There are over fifty species of dogfighter songbugs rapidly spreading across most continents, with even the islands of Choeropica and Post-Abeli sporting recently-arrived endemics. 

8) Longhorn Spiny Songbug (Spinoculex microviridiphagus): A descendant of the five-spined songbug, this armored species has evolved to rapidly process the soft greenery that appears for just a few days after it rains. Its highest population density is here on the snapscrub, where rain is common enough to support thousands per hectare, yet there are relatively few larger grazers competing for fresh-grown grass. Of course, the adult spiny songbug does not have chewing mouthparts, so it relies on its young for assistance. Larvae of this species have small chewing mandibles that serve to pulverize soft vegetation. Each time the parents return with a fresh meal of grass, the larvae regurgitate a portion of pre-chewed pulp. The adult songbugs spit venom on this slurry, reducing it further to a soupy green liquid than can flow easily through the proboscis. When not actively rearing young, the adults can sustain themselves on plant nectar alone; the additional nutrition from grass is only necessary for egg production and the increased energy demands involved in courtship and rearing a clutch. 

Other Insects

Aside from songbugs, several other clades in the order Flagelloculiciformes live on the snapscrub - taken as a whole, the whiplashers and harpoonsjaws outnumber all other non-aquatic mosquitoes combined. Some, like the dipterachnids, have yet to falter in their rate of diversification, while others have seen their niches become restricted over the past few million years, mainly due to competition with derived Pugilopsoids. The falconflies (Falcoculicidae) are one such case; for a time, they were the planet's only acrobatic aerial predators, but now that role has been filled by the swooping songbug genus. Though a handful of large falconfly species are still holding out, their days are numbered; the family's future now rests on its smallest members. These belong to the genus Nanofalco, which first branched off as far back as ten million years ago. Because large falconflies target a wide range of prey sizes, these smaller ones, who were restricted to small targets only, found limited success. Songbugs, on the other hand, tend to specialize heavily for a narrow range of prey, leaving a wider niche space open for small, inconspicuous falconflies like the Motehawk (Nanofalco vectiraptor). Ever-abundant small dustflies make up most of its diet, but it retains the generalist behavior of all falconflies; it also eats a wide variety of Flagelloculiciform flies smaller than its own half-centimeter-long self.

Dipterachnids, which belong to whiplasher side of the Flagelloculiciform tree, are present here in large numbers across three species. The largest and least populous is the Dancing Clawshield (Unguioscillopteryx xericus), which both scavenges and actively hunts small animals in semiarid and arid regions across Abeli. The backward-hooked claws around the margins of its asymmetrical wings don't do much to deter predators; their main purpose is to entangle with those of another individual during territorial fights or mating displays. Latched together, two or more clawshields spin and jump from side to side, each aiming to flip its opponent onto its back. The loser is often cannibalized, especially if the fight is between two males competing for a female. These messy battles can attract the attention of larger predators, forcing the combatants to disentangle themselves and flee. Luckily, like many Oscillopterines, the dancing clawshield can use its rigid wings to jump distances of around half a meter. 

Flapaways like the Pod Pocketprickle (Sonoretropteryx cavernensis) can also leave the ground under wing-based power; when threatened, they take off backwards and can remain aloft for as long as five meters. Some flapaways can go further, but pocketprickles belong to a genus that has sacrificed some of its flight ability for a different form of defense. The halteres of this species no longer serve a sensory purpose; they are now noise-makers that scrape and drum against the sides of the body, generating a loud rattle to scare away predators. Other prey arthropods, especially those that share its pocketpod home, have become accustomed to this sound, and they may even congregate around pocketprickles to take advantage of the relative safety they provide. The flapaway must eventually leave the tree to hunt, as it will be killed if it tries to find prey within the pocketpod. It often patrols just a meter or two away, and whenever it catches a small insect or isopod, it immediately takes off and lands just outside the entrance to its home, leaving itself less vulnerable to kleptoparasites and its own predators.

The Social Squirewing (Microthyreopteryx geoglyphagus), despite measuring less than a centimeter long, is the only regular predator of adult geoglyph castlebugs. It accomplishes this through relentless mobbing; up to a hundred individuals share a territory, and though they don't work cooperatively, they form a disorganized swarm when a geoglyph approaches. Each possesses a finely-serrated proboscis that can saw through layers of exoskeleton, though it usually takes several individuals to cut all the way through to the soft tissue below. They can now begin injecting venom, which may take up to an hour to take effect. Throughout the entire attack, the geoglyph will fight back. Its first reaction is to buck and leap off the ground, hoping to intimidate the dipterachnids. If they don't back down, its only effective defense is to wriggle its body in a serpentine motion, catching one squirewing at a time in the the spaces where its armor plates overlap. The trapped insect is then sliced apart by the woodlouse's many spines and cutting edges. Social squirewings consequently have a per-meal death rate of nearly one in three, even during attacks that ultimately prove successful. Those nimble enough to avoid being torn to pieces are rewarded handsomely, as a fully-grown geoglyph provides enough meat for thousands of larvae. The surviving adults pair off, eat their fill, then lay eggs in the carcass. The larvae live in the rotting meat for several weeks, then emerge as a new swarm of adults. Many are allowed to join their parents' swarm, replacing the lost fighters, but many more are driven away to search for a new territory. Some may join other groups they pass along the way, but most are taken by insectivores during this nomadic phase. In many cases, the swarm dies out before it can settle down, but if they find an unoccupied territory, the young dipterachnids can found a fighting force that lasts many generations, continuously replacing its losses with other newly-eclosed flies.

The remaining snapscrub Flagelloculiciforms belong to a harpoonjaw subclade called the Chewing Flies (Gnathoculicoidea), a superorder on a similar scale to the Pugilopsoids. There are three Gnathoculicoid families, all descended from the Early Arthrocene locust reaper. Foliopters and gleanerflies have both been discussed previously, and the snapscrub-dwelling members of these groups are little changed from their Middle Arthrocene ancestors. They are important folivores in this environment, especially when migratory vertebrates aren't around. The Podgleaner (Parviscabognathus cavernophagus) is a more dedicated herbivore than the earliest gleanerflies, feeding on the exterior or pocketpods, and the Palmsnap Foliopter (Xerifoliopteryx australis) eats the leaves of any palmsnap that still has them. Sister to the foliopter-gleanerfly clade is the Sky Plague (Aeropestilus vulgaris), which belongs to the monotypic family Aeropestilidae. Sky plague functions similarly to Plague isopods: it congregates in groups of thousands or millions, and it eats mostly herbaceous vegetation less than a meter above ground level. But while Pestilarthrids are mainly grazers of woodlouse-grasses, Aeropestilids eat lower-quality plant matter like basket-grasses and wellstalks. They've been a factor in the success of caudiciform plants like well-whips and rhizomatous ones like loop-grasses, as such adaptations allow plants to survive complete defoliation. 

Sky plague can cross distances of many hundreds of kilometers each generation, allowing it to maintain a single cohesive gene pool worldwide, but it isn't found in all environments. In forests, its gleanerfly and foliopter cousins are the dominant insect folivores, as these possess both chemical resistance and behavioral adaptations against the defenses of palm-grasses. In productive grasslands, Pestilarthrids are far more common because skystalks compose most of the plant community. Only in floral assemblages with relatively few woodlouse-grasses can sky plague reach high densities. This includes arid or semiarid places like the snapscrub, high mountainsides, subarctic steppes, and certain low-productivity subtropical savannas. Gregarity allows this species to thrive in these wildly different environments; for example, in the colder areas of its range, swarms can generate body heat before takeoff by piling on top of each other and vibrating their maxillae.

The dustfly family has seen near-continuous diversification since Polliniculex first evolved in the Middle Muricene, with a bout of especially rapid radiation during the first half of the Arthrocene. All species feed on energy-rich liquid food, usually nectar, though some have branched out significantly. They have four lifting wings, though the hind pair is small and lacks veins, betraying its history as a set of sensory halteres. These hindwings cannot fold, as they are constructed from a single sheet of chitin, and any color patterns take the form of pigment deposits on their surface. The forewings, on the other hand, have more complex features, including structural colors and pterostigmata in most large species. In the smallest dustflies, they also support long filaments that greatly increase lifting area with little added weight.

One such tiny Polliniculicid is the Dustmote (Parvivector vulgaris), which despite its size (usually less than 3mm across) is nearly cosmopolitan. It can be found wherever jade sweetstalks grow, including specialists like scrubjade, as all members of the tribe Clarisacchareae have similar spikelet structures. This consists of a pair of small supporting glumes, a fertile bisexual floret, and a showier sterile floret that attracts the dustflies. With nectaries around the periphery of the lodicules, the fertile floret is an easily accessible food source, so its primary pollinator does not face pressure to specialize for a single jade species. 

The Bloatbuzzer (Cavitofructophilus volafructensis) measures up to eight millimeters long and feeds on a different sugar-rich food: bloatseed berries. As a member of the Lapping Dustfly tribe (Corvoculicini), the bloatbuzzer has a softer, more mobile proboscis than other Polliniculicids. In this genus, the mandibles separate from the other stylets at the very tip, forming two finger-like appendages that serve to grab and tear an opening into the skin of fruit. Dustflies ancestrally had a typical mosquito-like larva, but many groups have since evolved more specialized larval forms to avoid competition with gillywogs. Bloatbuzzers and other members of their genus deposit larvae inside the same fruit they eat as adults, increasing competition intraspecifically in exchange for reducing it interspecifically. However, they have a strategy to mitigate this competition. When laying eggs, this species usually goes for very young, sometimes even underripe, fruits, leaving alone the large fragrant ones. This helps prevent the larvae from being eaten by vertebrate frugivores or infected with bloatborer larvae, and adult bloatbuzzers can feed from older berries without taking food from their young. In fact, the larvae leave behind scars on the berries wherever their siphons poke above the surface for air, providing access points for the adults to drink from later. 

The Lace Dustflies (Vittavector) are generalist pollinators, though far more diverse than dustmotes, with over a hundred species worldwide. This is because the minuscule dustmotes can reproduce in nearly any body of water, needing only about a milliliter for less than 24 hours. Lace dustflies, on the other hand, grow slowly and attain a size of several grams, so they have to specialize for large, gillywog-free bodies of water that last a long time and provide lots of food. Luckily, their larvae don't require full immersion in water; they can keep their gills functional even when living in a mass of moist solid matter. This means that, instead of the typical phytotelmata and small puddles that many Apterran flies use, most lace dustflies have specialized to deposit their larvae in places like riverside sand banks and low-lying forest soils. Scrubjade debris piles serve this role on the snapscrub; they contain few threats and competitors, and they can retain water for several weeks after a rain. The Scrubjade Lace Dustfly (V. xericus) breeds exclusively in such piles, and its young essentially parasitize the scrubjade-Scansoriarthriform community, feeding on the detritus that they rely on for their own water storage. 

The Floodflame Dustdisc (Quadiscus diluviophilus) is the world's largest non-Halteropterin dustfly, with a wingspan of just over twenty centimeters. The dustdisc genus, though relatively young, has about 250 species across the Panapterran continents, each of which feeds on one or a few species of sweetstalk flowers. It also has the widest range of sizes among all dustfly genera, with its smallest species reaching less than 1% the mass of this one. Its most diverse feature, however, is its proboscis, which is hard and immobile, allowing it to evolve into a shape suited for one particular flower. In the case of floodflame, the flower has two large, brightly-colored glumes whose cup-like shape holds the equally bright fertile floret. The nectaries are sandwiched between the glumes and the palea, forcing pollinators to rest on the floret to access the sweet liquid. The dustdisc can then slip its curved, flattened proboscis into the narrow gap. In the process, the dustdisc's belly gets covered in pollen, which it then unknowingly deposits on the next flower it visits. 

Finally, the snapscrub is home to three terrestrial neotenic mosquitoes, all belonging to the airgill clade. The first is a large basal form called the Annular Airgill (Trichovermis grandis), which ranges across most non-forested areas of the Abelian continent. This species eats mostly well-rotten carrion that few other animals can stomach, but it can also kill small game opportunistically. Like most airgills, the annular airgill spends its time in the substrate - in this case, usually loose gravel - and is rarely visible from above. Large airgills like this one possess hairs along their body; these are stiff and lie atop a dermal muscle that controls their position. Instead of wriggling through soil, these hairs allow the airgill to move inconspicuously, transporting individual particles down the length of its body to clear space ahead of itself. 

The Sap-Tapping Gillopod (Bradycaudocrus abeliensis) is a slow-moving tree parasite from the gummygrub subfamily. It descends from the sap-lapping gillopod, which already had a slow-paced lifestyle. Relying on a diet of toxic tree sap, it allowed its meals to sit in its crop for as long as three weeks before digestion. This reduced the potency of the trees' chemical defenses, but it also forced this lineage to become more and more sessile over time. Its present-day descendant rarely moves from a single secluded spot on a tree trunk, usually just below a leaf or branch. Here it inserts a long proboscis (not homologous with that of adult non-neotenic mosquitoes) formed from its labrum. This proboscis is not hollow, so it is not used directly to drink sap. It instead takes the form of a fine blade, which carves a diagonal channel for sap to flow from the stem. The cut is so small and clean that it provokes barely any chemical response from the tree, delaying the arrival of its Scansoriarthriform defenses. The other mouthparts, small and inconspicuous, drink the sugary liquid that leaks slowly from the incision. This species' stomach can store its entire body mass in fluid, turning the thorax into a water-balloon-like pouch when full. In most cases, the gummygrub is noticed by arboreal isopods before it can eat its fill. Luckily, it retains the ability to make a quick escape by jumping with its spring-loaded gills. If this happens during the day, the gillopod must find a new host quickly, as it has little resistance to the scorching heat of the ground. At night, the ground is hospitable enough that adult gillopods willingly descend to meet and mate; the plant-derived poisons they store in their bodies keep them safe from predation.

Another gummygrub parasite lives on the snapscrub - and all warm regions of Panapterra - but feeds on animals instead of plants. A member of the hematophagous dermopod group, the Clotfly (Sanguiculex spp.) lives in the flesh of vertebrates. Over the last eight million years, the life cycle of these flies has changed dramatically. Their eggs are nearly ubiquitous across the landscape and can lay dormant for a year or more. Development is kick-started by the chemical signals of a living body, so an egg will only hatch once it comes to rest on a potential food source. Each clotfly species targets only a few species of larger animals, so most newly-hatched nymphs die early from an unsuitable host. The lucky ones burrow into the skin, eating through the layers as they go. This meal gives them the energy for their first molt, at which point they swap their chewing mouthparts for a blood-sucking proboscis. In typical hematophagous dermopod fashion, they then search for a blood vessel to feed from for a short period. Unlike other members of their group, the clotflies sequester hemoglobin in their own bodies. This turns them red and allows them to sever their connection to the outside world, as they can get all the oxygen they need from the inside of their host. At this stage, males cease growing. They enter a blood vessel and join the bloodflow, passing through the heart and ultimately getting stuck in the small blood vessels of the lungs. They turn into small cysts and remain dormant for months or years, waiting for the arrival of larger females. But the females have a lot of growing left to do; they too enter a vein, but they station themselves behind a valve to avoid being swept away. They feed passively within the bloodstream, even absorbing certain nutrients through the skin of their long prolegs. This stage lasts several months, and the female dermopod may grow to two centimeters or longer. Her body eventually fills with eggs, and she detaches from her perch to make her way to the lungs. She probes around with her prolegs until she finds a dormant male, who she awakens by chewing through his protective mucus casing. They mate, the male dies, and the female sets herself to the new task of getting her eggs out of the host's body. She squeezes into the smallest arteriole possible, then digs the rest of the way out. As the host hacks and coughs, she releases ten thousand eggs or more. The next generation thus escapes and awaits a new host, while the mother dies and is eventually ejected from the old host's body.