The Primal Scroungers: Squaboons and their Relatives

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285 million years post-establishment

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above: the squabgoblin, one of the most successful species of primal scrounger.

The clever and fearsome squabgoblin of the longdark swamp is not alone on its branch of the scrounger family tree. It has many relatives which are closer to it than to any other scrounger lineage, with the bogbeast sitting at the base of its branch, with a divergence date around 272 million years PE.  It also has several much more closely related genera, from which it split between 4 and 8 million years ago, which share varying degrees of commonality with it. These animals range in size, diet, appearance, and behavior dramatically, but all fall under one clade which is known as the primal scroungers. This subgroup is further divided into three subsets: the squabgoblin clade, which includes just two species; the closely-related squaboon clade, which is very diverse and includes more than 20 species (plus one outlier); and the bogbeast clade, the most basal of all, which includes around six species. A representative sampling of each group is presented below.

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The squabgoblin's closest living relative is the far larger and much less gregarious squabsquatch of northern Serinaustra, a dweller of thick wetland that is highly reclusive and difficult to observe. The squabsquatch closely resembles its cousin but is much bigger, reaching 6.5 feet high and a weight of 240 pounds. It is also omnivorous, unlike the squabgoblin, and most of its diet is vegetation, with hunting of land prey very infrequent but scavenging and fishing somewhat less so. It is rare and occurs at a very low population density, and where it does exist, it favors thickets of very thick vegetation growing over terrain that is difficult to traverse and extremely soggy; long toes with lobes help splay its weight over mud and muck. Its choice of the most remote and inhospitable swamps to make its home helps to keep it hidden from prying eyes; though it is not a small animal, it is a shy one, and the squabsquatch is very wary of unfamiliar things and places; once a territory is established, it forms well-worn trails to necessary feeding grounds and rarely strays from them unless forced, bedding down in the same places each night to roost and generally following a routine. It rarely leaves cover, avoiding clearings and bright sunlight, and if it must migrate long distances will do so under the cover of darkness. Its size helps protect it from some enemies, but its most feared one is its own kin: the super-social squabgoblin, which is capable of complex tool use that the squabsquatch seems unable or unwilling to utilize, will drive it away as a rival, or even kill it as prey when they meet. Its current shyness and widely dispersed population are likely recent adaptations related to the evolutionary direction its relative has taken in the last couple million years, becoming a highly successful predator that has already displaced many other carnivores in the Serinaustran ecosystem. 

The squabsquatch lives in comparatively small troops of ten to twenty-five, with the most fundamental associations being harems of females and their young along with a single male, which averages 30% heavier than the female but is otherwise similar in appearance. Though it may seem the male is in charge, the actual politics of squabsquatch sociality are more complex, and have more in common with the squabgoblin than it first seems. The male in these troops protects his group from predators and from rival males, but his social ties to the females are not overly strong, and by outnumbering him, they ultimately maintain control the group as a committee, with his role being to aid in protecting them and their young but not making the majority of decisions. If he behaves recklessly or does not seem to be taking his role seriously, the females have the power to expel him and choose another, fitter male. It is, therefore, in the male's best interest to try and keep his group away from rival males at all times, if only so that the females don't spot a better option to replace him, and so this species does not generally gather in groups larger than a single family. Males living with harems aggressively fight one another if they meet, but it rarely comes to this, as they scent mark their territories and proclaim ownership with bellowing calls that serve to discourage rivals from making a move on their group. Most males don't live with females, however - in the nature of polygamous species, a majority will always be left as bachelors, and these will form small, looser associations together for company and safety in the absence of mates. They are only cooperative when goals are shared, however, and will fight to gain breeding rights and claim a harem, demonstrating that their alliances are purely pragmatic. 

Only female squabsquatches have enduring social bonds throughout life, with daughters rarely leaving their troops but males dispersing at young adulthood, between three and five years old. In many harem-forming species, females must leave to avoid mating with their own father, but as female squabsquatches have a lot of power in their troops, they tend to replace their singular male every few years, so that by the time the young are mature they can mate with a new, unrelated partner.  

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The next closest branch of scroungers to the squabgoblin are the squaboons. They comprise a large genus of scroungers closely related to squabgoblins and squabsquatches, but which are not as close to either as those species are to one another, forming a sister lineage to both. They are generally a little bigger than the squabgoblin but much smaller than the squabsquatch, with significant sexual dimorphism in size and sometimes in appearance - males are always larger, and may or may not have different coloration and ornamentation. This genus is omnivorous, though different species can have mostly plant- or animal-based diets. Like their close relatives, most squaboons are matrilineal and female-dominant socially, but this is not true for all of them.

The superb squaboon is one of the more dramatically varied species between male and female, with the male being twice the female's weight and completely different in plumage color, length, and distribution. Despite this, the female of this form is still in charge, forming the leadership of the largest social aggregations of all scroungers - rarely up to a thousand individuals, which exist in mostly stable groups (most, however, are more modest and contain between 250 - 400 animals.) The vast food resources required for so many animals make forming a territory unrealistic, and these scroungers travel constantly as a nomadic crowd, foraging as they go and rarely visiting the same site more than once in a year. This species favors uplands and avoids crossing through water too deep to wade in, traveling through the open woodlands and meadow clearings of Serinaustra's highest elevations, the clearview mountains. This region, really more a series of rolling hills than truly mountainous, is still the driest place on the continent as water tends to wash away to the wetlands at lower levels despite rain falling most days. They avoid the longdark swamp not only for its unfavorable terrain but also because they have poor night vision, and so are mainly diurnal. This does not entirely mean they avoid meeting their predatory relative that hunts in the night, for it has a wide tolerance of habitat and its range extends outside the polar regions, but unlike the squabsquatch it has little to fear from its rival thanks to its even larger numbers; it travels in such huge crowds and is belligerent enough besides that not even the squabgoblin, in its clans of a hundred or more, will face it head-on. If the squaboon were more skittish and were to flee from its rival when they met, this might give the squabgoblin an advantage, but the superb squaboon is aggressive and cohesive, working as one force to defend themselves from predators. Though their tool use is more primitive, they do utilize items in their environment as weapons, and their upland habitat uniquely provides them with an abundance of rocks with which they will very willingly bludgeon their enemies if the situation demands it. For the most part, the squabgoblin finds it is not a battle worth starting, particularly as the two species have diets with little overlap and so are not significant competitors. 

Female superb squaboons, weighing about 20 pounds, are the main fighting force of the troop, even though they are smaller than males, which also have significantly more impressive hook-like projections lining their facial tentacles that can deliver painful wounds. Like in the squabgoblin, female social bonds are stronger and more cohesive, allowing coalitions of many females to work together to fend off threats and collectively protect their offspring. They also maintain dominance this way over single males which operate less cooperatively. The different behavior between males and females is even more extreme in this species, in which the male is not usually even a permanent fixture of the clan, but rather an outside presence that only joins females when they are receptive to mate. His large size and more dangerous weapons exist more to fight other males than to defend the troop from danger, and males have their own separate social hierarchy while visiting the female group, with the largest and strongest males siring most of the young. The strongest, most mature male superb squaboons are brightly colored with long, flowing plumage on neck and tail and featherless red and blue faces which shine in the light with an iridescent purple sheen, their physical appearance advertising their fitness to females who do not typically spend long enough with them to observe their strength firsthand.

There is another smaller morph of male, however, which does not develop any of the characteristics of the typical dominant one but which closely resembles the female. As has occurred in a number of other polygamous birds with very different male and female morphs, these males may find reproductive success by behaving in a more feminine way, avoiding fights with other males entirely and spending all of their time within the female group. Here, they form more lasting ties with the females, even helping to protect their young. This form of male makes up approximately 15% of the male population and differs from the typical form mainly in having much less testosterone, which effectively prevents them from becoming fully mature, though they are still able to mate. Unlike others, these don't leave their natal clans at maturity, and are not driven off by the female leadership. The dominance hierarchy of these birds, as in squabgoblins, is passed down from mother to daughter with the further detail of males typically exiled near adulthood and then losing all rank. This means that the cryptic male morph, as it is not kicked out, can in rare cases be the only male to ever inherit a position of leadership within a tribe, it it happens to be born to a matriarchal female who passes away without an older, female successor. Though they will couple with them, female squaboons don't ever seem to regard these males as being male because they don't change coloration. They don't seem to view them as merely immature, either, because they rebuff adolescent males which attempt to mate within the clan fiercely. It turns out that one of the ways these scroungers maintain social bonds is through same-sex ritualized pairing; females which mate with cryptic males likely perceive them as another female in every respect, and so would not expect them to be able to father any offspring with them, as another female would not be able to. They therefore don't reject them in favor of stronger males. In this way, the cryptic male has perhaps perfected its role mimicking a female to the fullest extent of any similar species; while less than 20% of typical males may reproduce, 100% of cryptic males will do so. Why this morph has not totally outcompeted the other is related to the lower fertility of the cryptic; its lower hormone levels mean it mates less, and produces fewer fertile eggs than other males, and so remains in balance at a consistent but low level within the population.

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The stricken squaboon is in the same genus as the superb squaboon, and yet is remarkably unlike it. This 24 pound bird is only slightly sexually dimorphic, with males about 10% heavier and with slightly more prominent red wattles above the nostrils. It is not highly sociable in comparison, either, and occurs only in small bands of twenty or thirty - and they are led by the male, rather than by allied females. This scrounger lacks the confidence its cousin can flaunt thanks to its huge ranks, and lives its life in deep forest underbrush, where it does its best to remain unseen by its enemies. The stricken squaboon has many, however, and like the squabsquatch, one of the worst is the squabgoblin. While superb squaboons succeed in avoiding predation thanks to their numbers and cohesive nature, the stricken squaboon represents an alternative behavior pattern in which it tries to avoid its fearsome relative by being as inconspicuous as possible. Its plumage is subdued compared to other squaboons and mostly consists of brown and black blotches, and it is almost silent, having foregone vocalizations because of the risk of being overhead by unwanted ears. It differs from the equally timid squabsquatch though in that it is neither rare, nor avoidant of the longdark swamp where its predator is most abundant

The stricken squaboon has evolved to live in a way that we would perceive as grim: by reproducing quickly enough to survive as a population in spite of frequent predation by squabgoblins, which culls their numbers back harshly whenever they become too large to remain unnoticed, preferentially killing the adults but missing many of the smaller juveniles that can more easily escape them in the undergrowth. Chicks are born in litters of four to six, up to twice each year, and can run from birth. This species is unfortunately a favored prey item that is easily killed because it has lost its cultural tool use that once let it defend itself; the high mortality rate of adults means individuals rarely live long enough to develop new skills and teach them to others, requiring youngsters to  be fully able to survive on instinct alone in the not unlikely event they lose their parents before adulthood. This loss over million of years has gone hand in hand with an increasingly plant-based diet, and the stricken squaboon is now more than 90% vegetarian, and eats mostly grass and leaves, favoring fungi in the winter. Plants don't require tools or a big brain to subdue, and as all these factors come together, the stricken squaboon has become significantly less intelligent than its relatives. High predation also makes longterm social bonds typical of other scroungers difficult to forge, and without them, the larger males come to dominate groups by force.

Yet while the stricken has in some ways become the cow for the squabgoblin's slaughter, it still demonstrates some intelligent problem-solving, and has had to innovate new ways of surviving on a diet unlike most scroungers which is both less nutritious than meat and often mildly poisonous. Stricken squaboons must eat large quantities of clay to absorb plant toxins from their diet, which would otherwise make them sick, for they have evolved to be plant-eaters quickly over only a short evolutionary time and lack the acquired immunity long-term herbivores will often acquire. An even more extraordinary survival strategy may sometimes be employed by the female of the species - nest parasitism. By sneaking an egg into the nest of another more dominant scrounger species - most often the superb squaboon - she can ensure a safe upbringing for her chick where its chances of reaching adulthood are vastly improved and predation is much less likely. This is a double-edged sword, though, for though the chicks raised by surrogate mothers will have higher survival rates, they will imprint on their foster parent and only interbreed with them. Male hybrids will not typically mate at all, being unable to compete with larger purebred males, but females will do so, producing second generation hybrids and so on. This still allows some genes of the female who dumps her egg in another nest to be passed on to other generations, even if they are ultimately absorbed into the gene pool of another species. The genes for improved plant digestion may have come to the superb squaboon from the stricken squaboon in this way, and so ultimately helped it to survive and become so successful as an omnivore. This sort of low-level hybridization over the long term can skew genetic relationships, keeping the two parent species genetically similar enough to continue to produce fertile offspring even after millions of years of divergence. This sort of intermixing mirrors the wumpos of Serinarcta, another group of tentacle birds which muddy the boundaries of taxonomy though frequent crossing with their cousins; its existence in both is likely a result of selective pressures toward strong sociability in close-related animals that continue to interact in their environments.

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The boomer squaboon is the largest member of this genus. What this very dimorphic scrounger lacks in bright colors or elaborate plumage is made up for by the specialized left and right facial tentacles of the male, which are wide, flattened, and able to fold into a tubular shape with a widened, funnel-like opening, and are used to amplify its low-pitched croaking call like a megaphone. Unlike any other species, boomer males form leks which are perused by strictly female social bands of up to 50 animals. Males have no place in the clan structure at all, living entirely solitary lives outside of mating season, and serve neither to protect the females nor have anything to do with their own young once they hatch. The size difference is the most drastic of all the squaboons - robust males can grow to three and a half feet high and weigh 50 pounds, while lighter-built females are a foot shorter and half as heavy. 

Boomer squaboons are a species adapted to open wetlands, especially grassy marshes with little tree cover to provide hiding places to predators. With webbed feet, they can swim and often do so to cross expanses of water between vegetated islands. They are nearly perfect 50/50 omnivores which eat a variety of fish, invertebrates, and the seeds, tubers and crisp new stems of grasses. Complex tool use occurs in this scrounger, which is very intelligent, perhaps moreso than any other of its genus and even rivaling the squabgoblin in ingenuity. Their most specialized skill is using their dexterous tentacles to weave reed baskets, which have many uses. They can be used to transport food, but also are used to hold young chicks which can't walk very far or swim well for several weeks after hatching. They are also are used as fish traps, with the squaboons even baiting them with scraps of food; females teach this learned trade to their young, but it is more refined in the female, as the adult male's larger, specialized tentacles make it a more clumsy artisan. Males don't interact with female groups when not at the lek site and are not tolerated by them, but bonds between females in the clan are long-lasting and vital to cooperative nesting and chick-rearing behavior that ensures young are protected and provided for. Females form pair bonds between each other and share incubation duties, while once chicks are a few days old they join a communal creche and gain the protection of the entire group. Males leave the clan at age two, but females don't disperse unless the band becomes very large and then fissures into two smaller ones. 

Boomer squaboons hybridize, too. Occasional crosses occur with the superb squaboon, its nearest relative within the genus, where their ranges narrowly overlap, and despite very different behaviors and appearances of the sexes in each species, they remain genetically close. This has likely gone on at low levels since the two species mostly speciated and formed separate breeding pools, with the occasional influx of genes from the ancestors of the boomer influencing the size and enlarged tentacles of the male superb squaboon - or perhaps vice versa. While superb squaboons may also interbreed with stricken squaboons - and such crossing may have influenced the longer plumage of the superb male - hybridization between the stricken and the boomer is not viable. Though both of these species share common genetic markers with the superb species through long-term hybridization events, with their totally separated ranges and longer, more total separation, neither is compatible enough with each other to bear hybrid young any longer.

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The final squaboon we meet today is the bloodbreasted squaboon, a mid-sized (around 25-30 lb) species which is found in north-central Serinaustra, in an intermediate eco-region roughly between the longdark swamp and the saltswamp. The closest relative to the stricken squaboon, this brightly marked and confident species shares none of that species' timid nature and acceptance of its place low in the food chain. It is a bold, remarkably smart animal which dwells in only one specialized sort of grassland, the saltwater savannah, in which it has few predators to worry about. This region is perhaps Serinaustra's closest analogue to the northern soglands, being a vast, damp plain - but differs in being pockmarked with shallow saltwater ponds and rivers, for it is periodically inundated in water saltier than the ocean by the overflowing banks of the great blue salt lake directly to the west. This prevents the growth of trees, save for a few specialists that don't form closed forests, but favors the survival of hardy salt-adapted puffgrasses which form an expansive monoculture over the sandy soil. This is an environment that is rich in food despite seeming inhospitable - as long as one knows where to look. The vegetation here is not good to eat, being disgustingly high in sodium, and so even though it is closest related to the vegetarian stricken squaboon, the bloodbreasted is mostly carnivorous, and such a diet not only helps to fuel the demands of a bigger brain, but is also harder to find, encouraging higher problem solving ability. It eats mainly crustaceans - crabs are diverse and widespread here, but spend much of their lives burrowed below the (salty) water table, and the bloodbreasted squaboon spends much of its time unearthing them. It does so not only its own claws but also with tools that are, arguably, the most complex in nature of any scroungers: shovels, that it may carve out of animal bones or driftwood with sharp flakes of rock that are themselves tools, produced by smashing smooth stones from the shore of the lake against one another.  This two-staged manufacturing technique, in which a tool is used to manufacture an additional tool, is indicative of a high degree of ingenuity and foresight. In addition to crustaceans, these animals will eat the large, fat-rich seeds and "fruit" of the salt grass, which grow inside small air-filled pods adapted to break free and float away during flood periods; the hollow berries have an appealing crunch when consumed and contain less sodium than the rest of the plant's tissues. 

Bloodbreasted squaboons live in mixed-sex clans in which males live year-round, but have a separate hierarchy to females, and in which females outnumber males substantially. They are nearly the same size, males being about 5% heavier, and have similar plumage with subtle blotches of reddish-gold and black. Males and females alike have featherless faces, but mature males alone have a prominent featherless throat which is revealed to be stunningly bright red when they part the long plumage of their necks to reveal it, filled with air and swollen into a heart-like shape; the intensity of this color is determined by how many crustaceans the male has consumed, for it is produced by carotenoid pigments in the skin. It is display of fitness and strength, as well as foraging success, and is most intense in the most dominant males. Two types of clans exist, breeding and bachelor, the former comprising several related males as a coalition and as many as twenty females, the latter being comprised of only younger, older, or less fit males that cannot compete to join a harem. Males fight over breeding rights, and bachelor males will regularly instigate fights with breeding males and try to take their place. Females, for the most part, will not intervene - they will want to mate with the strongest males to have the strongest chicks. 

If they happen to dislike an individual male, however, even if he is strong, they will gang up to drive it and its group away and defend their current males. Because females have strongly-defined bonds and help one another raise their young collectively, infanticide does not exist in these or most squaboons, even if new males take over a clan. The females would sooner kill the male than allow it to harm their young, and he best never try. 

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More basal than the squaboons, but still related to them, is a monotypic species that forms its own genus with no relatives closer to it than to any other species.Though physically nondescript in many respects, this outlying species of primal scrounger is immediately memorable for being both very small and for its unusual habit of climbing trees with its feet, often swinging between perches upside down. Unlike the extinct boras - the fully-arboreal softbill birds of the mid-Ultimocene - it does not use its tentacles to grasp branches, and they are short and unspecialized in this species. Only weighing two or three pounds, twigswingers are one of the smallest scroungers, but use this to their advantage in living in a habitat that is not otherwise easily accessible to them. Because it is so small, this scrounger can jump very far for its size and climb up tree trunks using its legs and no arms, without having to evolve specific anatomical changes to allow it. Its toes are long and have hooked claws to grasp bark, and strong legs effectively let it run right up vertical surfaces with ease. Once in the canopy it usually moves by rapidly brachiating, head-down and toes up, like an inverted ape. It stops often to twitch its head right-side up and get a right side up view of its surroundings, but it can see perfectly well upside down and adjust its jumping to compensate for reversed gravity relative to its point of view compared to jumping in the same way on the ground. It feeds on high energy food sources - sap, nectar and insects that will sustain its rapid and almost constant explorations.

Twigswingers are much less social than squaboons, their social structure reduced to its basest elements: usually, a male and a female, plus their young of one or two years, though polygamy still occurs with multiple males which may live with a single female in a reverse harem. Such accessory males are usually brothers of the dominant breeding male, and may sire offspring when he is not watching, but primarily they assist in childcare for their relative and may inherit his position if he dies. Twigswingers are unusual among scroungers for carrying their newborn chicks with them for prolonged periods - or rather, letting the young cling to them with their claws; this frees them from attending a vulnerable nest while the chicks are too uncoordinated to keep up with their fast-paced parents as they vault from one tree to another, many stories above the ground. This is almost always the role of the male, and is coveted enough that males may fight over who gets to carry the baby, as doing so may earn them respect in the female's eyes. Twigswingers sleep communally in long-lasting grassy nests built in small tree holes, and alternate around 6 hours of wakefulness and 6 hours of sleep on a circadian rhythm  independent of the time of day. This is common for polar animals for whom the day can be either all dark or all light depending on the season - the twigswinger is able to forage successfully in both light and dark conditions, with good night vision and acute daylight vision, though poor perception of color. This makes finding fruit more difficult, so that this scrounger has evolved a much more refined sensitivity to the ethylene gas given off by ripe fruit, allowing it to forage by scent without being able to visually distinguish red berries from green, and to do so in both summer and winter. 

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Lastly, we reach the earliest-diverging primal scrounger lineage, relatives of the bogbeast known as brawlers. They are a small genus of very big animals, which appear to be intermediate in morphology between the bogbeast (which like the twigswinger is the only species in its genus) and the squaboon-squabgoblin clade. They sit closer to the former, having split from the ancestors of the bogbeast around 8 million years ago, but more physically resemble squaboons in many respects except for size - they are the second largest scrounger genus of their era, reaching a weight of 2,700 pounds and a length of fifteen feet. Brawlers are more social than bogbeasts but less than squaboons, and  living in small groups of three to six, each group has only a single larger male which is always distinguishable by its featherless face and, in the largest species like the belligerent brawler, by its enlarged cheek flanges that hang around its face looking kind of like deflated beach balls - a sort of thick waddle. This serves both makes them appear bigger and to protect the face during fights with other males in which each attempts to stab the other with spiky horns on top of their heads, which have a common origin with the smaller nodules along the face of the bogbeast. Brawlers can move their flanges upward through muscle attachments, and use them to cover their eyes when facing a strike from an opponent. 

These scroungers are mostly natives of northern Serinaustra, in low-lying forest and swamp habitats. They are carnivorously-inclined omnivores, though about half the diet is leafy vegetation, mostly aquatic plants as they are softer and more easily consumed. They hunt rarely for themselves, but often steal carcasses from other predators as there are few that are larger or stronger than themselves, especially as a group. As their name suggests brawlers have bad temperaments, and in addition to the aggressive fighting between males, they will face up against almost any other animal that confronts them, even the bogbeast, and usually come out dominant in altercations due to their cooperative behavior. Far from timid, these powerful scroungers don't shy from conflict and have very little fear, often foraging in open areas as well as in thicker forest with little worry about predators. Groups with young are the most dangerous, as females will fight to their own death to protect their small, fuzzy chicks, and will often not stop attacking a perceived enemy until it is dead and dismembered to the point of being unrecognizable... at which time the clan will usually eat it. 

Phylogeny of the Primal Scroungers