Alien Biospheres: Part 10 - Allopatry

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Every single episode of this has me thinking 2 things:

1: "What's gonna become the dominant species on the planet?"

&, 2. "When's he gonna throw a rock at it?"

👍︎︎ 18 👤︎︎ u/Mildly_OCD 📅︎︎ Mar 22 2021 🗫︎ replies

Polypodia gang

👍︎︎ 5 👤︎︎ u/Spiritual_Juice3500 📅︎︎ Mar 22 2021 🗫︎ replies

Mass extinction episode when?

👍︎︎ 4 👤︎︎ u/206yearstime 📅︎︎ Mar 22 2021 🗫︎ replies

I’m just a junkie waiting for his next fix

👍︎︎ 1 👤︎︎ u/SansCulture 📅︎︎ Mar 28 2021 🗫︎ replies
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This alien planet is brought to you by  Campfire Technology. If you’re a fan of the   Alien Biospheres series and looking to make  your own alien planet, or any other kind of   worldbuilding or writing project, Campfire Blaze  can be a big help in keeping your work organized.   Blaze comes with pages to help you chart out the  maps and specific locations within your world,   while the species page lets you delineate your  world’s native clades of flora and fauna, as well   as individual facts, statistics, and miscellaneous  details for every individual species. There are   also tools for fleshing out the human aspects of  your world, like cultures, politics, languages,   religions, and magic, along with pages for  narrative elements like characters, relationships,   timelines, and story arcs. Blaze’s free version  already includes most of these features, but   you can also individually purchase new modules to  let you catalogue your world in even more detail.   To sign up for Blaze or to learn more,  check out the link in the description.   Over the last few episodes, we’ve explored  each of the major biomes of our alien   planet and seen how their respective  ranges of temperature, precipitation,   and resource availability have shaped the  evolution of their resident species. But in   the enormous spans of time over which evolution  occurs, conditions rarely remain stable for long.   As the eastern and western tectonic plates  gradually drift away from each other,   the supercontinent that has existed since life  first moved onto dry land will finally break   apart into two separate landmasses. This will have  several immediate effects on the climate. What was   once the arid continental interior will now form  the new coastlines, and the proximity to the sea   and the resultant increase in precipitation will  turn the plains and deserts into tracts of forest.   Meanwhile, the mountain ranges along the exterior  coasts will continue to rise, which will lower   atmospheric carbon dioxide levels through silicate  weathering, and the snow and ice that accumulates   on their slopes will increase the planet’s albedo,  leading to a gradual decrease in average global   temperatures. And if these mountains, particularly  those in the northern plateau, see the formation   of ice caps and glaciers, then for the first time  in the history of terrestrial life, the planet   will have entered an ice age. Up until now, the  planet has been going through a greenhouse period,   in which there were no polar ice caps or  continental ice sheets, but as the climate cools,   a cycle of glacial periods may begin, wherein  for intervals spanning roughly 100,000 years,   the global temperature will drop by several  degrees, sea levels will fall by as much as   100 meters, and ice sheets will advance from  the poles and intrude into the temperate zones.   Due to subtle changes in the planet’s orbit,  these glacial periods will be punctuated by   relatively warm interglacial periods, each of  which may last several tens of thousands of years.   These drastic shifts in climate will have  numerous ramifications for many ecosystems   across the planet, but the breakup of the  continents has another important consequence:   with the exception of particularly mobile species,  the respective inhabitants of the two continents   are now cut off from each other, and thus  evolutionary developments that occur on one   continent won’t affect the other, meaning they’ll  have entirely separate evolutionary trajectories.   Such a situation is called allopatry, and occurs  whenever a population becomes separated by a   physical or geographical barrier, whether  it be a mountain range, a body of water,   or anything else that might prevent the two  halves of the population from interbreeding.   Since gene flow can no longer occur, a frequent  consequence of allopatry is that the separated   subpopulations evolve into different species,  which on a global scale will result in the   clades and ecosystems on the eastern and western  continents gradually diverging from each other. In   light of this, for the first time in this series,  we’re going to need to keep track of biogeography,   or the distribution and range of each of our  clades, particularly whether they exist on   the eastern or western continent, which will be  essential for determining how ecosystems form.   To begin with, the overall landscape of the  eastern continent will remain much the same as   it was before the continents split apart, since  it will remain largely within the tropics as it   drifts eastward, and even during an ice age the  equator will still experience high temperatures.   While stretches of tropical forest will occur  along the coast, the inland areas will stay   comparatively sparse, especially within  the rain shadow of the eastern mountains.   During the glacial periods, the forests  may diminish with the decrease in rainfall,   mainly surviving as patches of fragmented gallery  forest separated by stretches of open plains.   Environments like these allow plains-dwelling  animals to live alongside forest-dwelling ones,   a perfect habitat for large browsers like the  procerapods, as they’ll be able to continuously   feed on the tree tops while still having enough  space to move freely between areas of vegetation.   When it comes to browsing herbivores, the larger  the animal is, the better its ability to reach   for food, and the more food the animal can  obtain, the larger it can afford to grow,   and in especially lush areas, like the periphery  of the equatorial forests, this feedback loop   may result in these already massive procerapods  growing even bigger. To help them reach upward,   their legs and pedipalps may elongate, and their  foreguts may expand to contain and process the   enormous volumes of nutrient-poor foliage  they feed on throughout the day. To support   such a huge cephalothorax, they may develop  stronger muscles and hardened tendons in their   back, which will attach to enlarged projections  along the dorsal surface of their limb girdles,   functioning much like the neural spines of many  large animals on earth. Being big also acts as a   deterrent to predators, and as the largest animals  on the eastern continent, these procerapods will   be virtually immune to attack once fully grown.  We’ll call these gigantic beasts stylopods.   Scurrying among the feet of these giants will  be a menagerie of smaller herbivores like the   leptopods. Most leptopods are adapted to live in  open habitats, but in this patchwork of plains and   fragmented forest, some clades may evolve to  occasionally wander among the stands of trees   in search of food. These clades may benefit from  more generalist diets than most other leptopods,   feeding not only on brachyphytes but also mixed  caulophyte species, necrophytes, and low tree   branches. The trees will also provide a degree  of shelter from large steppe-dwelling predators,   though they can still rely on their speed  to avoid attack when out in the open,   but many leptopods and other herbivores may  settle on another common form of defense.   We’ve brought up the concept of competition many  times in this series, but whenever we’ve mentioned   it before, we’ve been referring to competition  between different species, or interspecific   competition. However, intraspecific competition,  or competition between members of the same species   is an equally important factor in evolution. The  fewer resources are present in a species’ habitat,   the more intense the intraspecific  competition will be to gather those resources.   In environments where intraspecific competition is  especially high, a species may evolve territorial   behaviours, aggressively repelling other  individuals from their home range to keep as much   of the local resources to themselves as possible.  On the other hand, though, the presence of other   individuals does come with some benefits: the more  animals are around, the easier it will be for them   to collectively keep an eye out for danger,  and the less vulnerable each of them will be   to attack, since predators will be forced to pick  out and go after only one target. Because of this,   many species exhibit collective behaviours whereby  individuals instinctively congregate in groups,   which, depending on the clade and group structure,  can be described as herding, flocking, shoaling,   or various other terms. Along the periphery of the  coastal gallery forests of the eastern continent,   food will be abundant enough to keep  intraspecific competition to a minimum,   and so the resident leptopods can afford to  evolve collective behaviours for greater safety,   forming herds comprising anywhere from several  dozen to over a hundred members, giving these   leptopods, which we’ll call camptopods, a further  degree of protection from the many predators they   share their habitat with. The largest predators  on the eastern continent will be the onychodonts,   most of which will be steppe-dwelling  hypercarnivores preying on a variety of big game.   However, along the edges of the coastal forests,  many of the local titanopods like the stylopods   may be too large and formidable to risk  attacking, so many onychodont species may   specialize for hunting smaller, faster herbivores  like the camptopods and other leptopods.   To catch such nimble prey, they may adapt to  improve their already impressive weaponry.   The onychdonts’ greatest asset lies in their  centaurism, which lets them run faster and more   efficiently than other predators and frees  up their forelimbs to use for manipulation.   As they play an increasingly prominent  role in catching prey, these limbs may   evolve into specialized raptorial appendages,  developing pointed serrations along their   undersides to close over the prey like a vice  and minimizing the chance of it struggling free.   As these limbs take over the role of securing  prey, the pedipalps may reduce into a pair of   retractable fangs that can be jabbed into the  preys’ hide to kill it quickly and efficiently.   These eastern onychodonts,  which we’ll call cryptodonts,   will be adept pursuit hunters in open areas,  but in this landscape, the trees will be   plentiful enough to provide cover for ambush  predators like the deinognathans. The rivers   and watering holes may be ideal habitats for the  semi-aquatic trachygnathans and dolichognathans,   and the fully terrestrial ancestral forms  may thrive along the forest periphery.   Going after plains-dwelling herbivores may bring  them into competition with the onychodonts,   and thus the two clades may need to undergo  a degree of niche partitioning if they’re to   coexist in the same habitats. The deinognathans’  signature mandibles are so large and robust that   they can’t be used to chew or process food, this  job instead being relegated to the pedipalps,   which break up the food and pass it into the  throat, the exact opposite condition to that   of the onychodonts. Whereas the onychodonts  use their pedipalps to inflict precise,   puncturing bites to kill their prey quickly,  the deinognathans rely on their tremendously   powerful bite force to crush their prey, which may  serve as a preadaptation for new modes of feeding.   Durophagy is a feeding behaviour characterized by  specializations for cracking open bones, shells,   or other hard structures, for which having a  high bite strength is a fundamental prerequisite.   As such, while the onychodonts may only be able to  feed on the softer, fleshier parts of their prey,   some species of deinognathans may adapt feed on  the bones and other carrion they leave behind.   To most other carnivores, these parts of the  carcass would be inedible, and thus these   deinognathans will face very little competition  as they evolve into durophagous scavengers.   As briefly mentioned in part 6, scavenging  doesn’t yield as much food as active predation,   so these bone-eating deinognathans will likely be  smaller than their macropredatory cousins and will   still actively hunt prey when carrion is scarce.  Durophages often have blunt, rounded teeth,   an optimal shape for breaking  through hard materials like bone,   along with extra chambers within their digestive  tract and especially powerful stomach acid.   Among these deinognathans, these adaptations  may also let them crunch through the shells   of placostracans and other diplostomes, or  even to subsist on tough woody vegetation.   On account of having such powerful, crushing  jaws, let’s call these animals malleognathans.   On the far side of the continent lies the coastal  rainforest that we covered in the last episode,   which fortunately, will be largely  unaffected by the onset of the ice age,   as it will still receive plenty of rainfall  thanks to the warm ocean currents along the coast.   It is, however, blocked off from the  rest of the continent by the mountains,   meaning many of the local rainforest species will  exist nowhere else in the world. But a select   few clades will still be able to pass over the  mountains, most notably the flying clades like the   opisthopterans and pleuropterans, and since these  clades have co-evolved with and help disperse   the chromatophytes, they’ll carry these plants  with them as they expand their range westward,   effectively bringing their own food source into  their new habitats. The earliest pleuropterans   will be small, generalist frugivores, but the  forms that spread over the mountains and colonise   the rest of the continent may grow to larger sizes  and adapt for long-distance flight. The same rules   of wing design that applied to the opisthopterans  will also apply to the pleuropterans; the bigger   they get, the larger their wings will need to be  in proportion to their body to keep them airborne.   Many of these forms may retain a similar  diet to the ancestral pleuropterans,   feeding mainly on fruit and other mixed plant  matter as well as the occasional malacoform.   This radiation, which we’ll call platypterans,  will comprise the majority of pleuropteran   diversity outside the eastern rainforest. But  some species may spread into habitats where   the chromatophytes, their principle source of  food, may not be able to grow in any abundance,   forcing them to adapt to new food sources.  We mentioned last time how the pleuropterans   and opisthopterans would avoid competition with  each other by specializing for different diets.   However, while the opisthopterans only  have their elastic spines and simple   struts extending into the wing membrane, the  pleuropterans have a fully-developed skeleton,   which lends them enough internal support to  grow much larger than the opisthopterans can.   This means that once the pleuropterans reach a  certain size, they won’t face any competition if   they exploit carnivorous niches, which may result  in some of the more generalist clades evolving   predatory behaviours. Their wings may become more  pointed to give them bursts of speed when hunting,   and their teeth and pedipalps may sharpen  to snatch up and butcher their kills.   These forms, which we’ll call theropterans, will  feed on prey too big for any of the opisthopterans   to catch, such as large diplostomes, small  osteopods, or even other pleuropterans.   Many of the pleuropterans may also manage to  fly across the sea and find habitats on the   western continent as well, where they’ll find  a very different landscape to that of the east.   The dry ice age climate will turn the majority  of the continent into deserts and arid steppe,   with the newly created coastline being the  only area to support any sizable forests.   In the inland regions, the retreating woodlands  will give way to plains of tylophytes,   pagophytes, and any other brachyphytes that  can manage to withstand the dry conditions,   with the drought-resistant nodophytes forming  isolated pockets of forest around lasting   sources of freshwater. Such a huge change in  climate and vegetation and the oscillations in   temperature between glacial cycles will radically  restructure many ecosystems across the continent.   The most vulnerable animals will be ectotherms  like the ancestral platydonts, dromaeopods,   and deinognathans, as they’re unable to generate  their own heat to combat the effects of the cold,   and so many species may die off outside the  tropics. Even megafauna like the titanopods   may be forced into decline, as even though  they’re able to maintain a more stable internal   temperature than other ectotherms thanks to their  gigantothermy, this still may not be enough to   let them contend with the constant oppressive  cold and the lack of food in the inland deserts.   Only the pachypods may have any notable presence  in the temperate zone, as they’re better equipped   to deal with the cold than other titanopods due  to their thick layers of fat and stocky build,   but even they may struggle to survive beyond a  certain latitude. On the other hand, endotherms,   like the eurycheirids, leptopods, and onychodonts,  will have a huge advantage in the new climate,   and begin taking over the vacant niches of  the ectotherms in temperate areas. The bulky,   cold-adapted hadrodonts may remain the dominant  superpredators in the northern regions,   primarily hunting large, slow prey like the  pachypods, but other varieties of onychodonts,   especially those that are more gracile and  lightly-built, may have the advantage in   warmer areas. The cryptodonts would likely  thrive in these regions, but since they only   exist on the eastern continent, a separate clade  may evolve to exploit similar niches in the west,   though they may do so in a completely different  way. While the cryptodonts, along with most other   onychodonts, have vertically-aligned pedipalps  to bite down through their prey’s hide,   in this western clade, the pedipalps may instead  evolve to oppose each other and close laterally,   acting like a pincer to securely grab prey  while simultaneously inflicting lethal wounds.   This means that unlike their eastern relatives,  these forms won’t need to use their forelimbs   in restraining prey, so these limbs may shrink  to allow the animal to support larger, stronger   pedipalps. Let’s call these forms ensidonts. But  the animals that are best-suited for the ice age   will be the thylacopods and thecopods, both of  which have specialized integument to complement   their endothermy, and with such efficient  thermoregulation, they’ll thrive as the   climate cools and other clades are driven from the  temperate zone. But as they proliferate and spread   across the continent, they’ll need to face new  challenges: while the tundra where the thylacopods   first evolved was too cold and desolate  for any predators other than the thecopods,   as they move further south they may come into  contact with much larger predators like the   hadrodonts and ensidonts, which may incentivize  them to invest in some defensive adaptations.   Since the majority of the western continent  consists of open landscape, they can’t rely on the   cover of vegetation like their camptopod cousins,  so their speed and stamina will be their primary   means of evading attack, and in the arms race  against the swift onychodonts, they may adapt to   become even faster. The eight-legged gait of most  osteopods provides a stable base, but if an animal   is going to invest in speed, any superfluous legs  may get in the way and present a waste of energy.   Long ago, centaurism evolved in the ancestors of  the onychodonts, granting them increased speed and   coordination, and now, perhaps these thylacopods  may convergently undergo the same development for   a similar reason. Even among tetrapods on earth,  centaurism has independently evolved many times,   so it’s likely to be widespread on this  alien planet as well. With only six legs,   they’ll be able to run more efficiently, and  their newly freed-up forelimbs can now be used   for manipulation and gathering food along  with the pedipalps. With this development,   along with their unrivalled ability to tolerate  the cold, this clade of thylacopods, which we’ll   give the name allobrachids, may become some of  the most successful megafauna within the temperate   latitudes. Where there’s a relative abundance of  food, like the nodophyte forests along the north   eastern coast, some species may exploit the  niches left vacant by the titanopods, growing   considerably larger and using their forelimbs and  lengthened cephalothorax to reach for foliage.   Their size will not only increase their feeding  envelope and discourage predators, but will also   increase the amount of body heat they generate,  making them even more resistant to the cold. These   animals, which we’ll call teleobrachids,  will be the largest of all leptopods,   browsing on nodophytes and other trees within the  northern forests. Other allobrachids may retain   more generalist grazing behaviours, taking over  the niches of some of the basal leptopods. Like   their camptopod relatives in the east, species  living in areas with enough food to support   a large number of animals may evolve herding  behaviours as a further defense against predators,   but the structure of and interactions within these  herds may be influenced by the thylacopods’ unique   physiology. Back in part 5, we introduced the  dichotomy between R-selection, maximizing the   quantity of offspring with minimal parental  investment, and K-selection, producing smaller   numbers of offspring but investing more heavily in  them to increase their chances of survival. Where   a species falls on the spectrum between R and K  selection is a key aspect of its life history,   or the pattern of survival and reproduction  throughout its lifecycle, which will be influenced   by a variety of factors. Typically, clades  will transition from R-selection to K-selection   as they evolve larger body sizes and as their  populations reach carrying capacity, so by this   point in history, most species of megafaunal  osteopods will likely tend towards k-selection.   The thylacopods will be more k-selected than most,  since the female protects her young from the cold   while they’re within the ootheca by cradling them  with her gonopods, which gives them a greater   chance of surviving through their development,  at the expense of encumbering the mother. Note,   however, that only the female bears this burden,  which ties into another aspect of life history.   One of the only universally applicable differences  between males and females across all clades is   that, by definition, female gametes are larger and  require more energy to produce than male gametes,   and therefore, males usually have  a higher fecundity than females.   This is especially true in species with internal  fertilization, since once a female is fertilized,   she’ll be unable to mate again while  she carries the developing young,   further limiting the potential number of  offspring she can have during her lifetime.   This results in what’s known as Bateman’s  principle, the concept that reproduction   is a greater investment for females than for  males, and therefore, while males are usually   incentivized to mate with as many females as  possible to maximize their reproductive output,   females need to make sure that every attempt  at reproduction will be worth the investment.   This means that females will often only choose to  mate with males that display an adequate degree   of fitness, and who will therefore pass on their  fitness to the offspring, meaning the males will   need to compete with each other for the right to  mate, another form of intraspecific competition.   However, in the osteopods, there’s the  confounding factor of sequential hermaphoriditism.   The earliest ancestors of the osteopods were  capable of changing sex during their lifecycle,   which many modern clades may retain, but exactly  how their hermaphroditism operates will vary based   on their life history strategy. Sequential  hermaphrodites can be either protandrous,   capable of changing from male to female, or  protogynous, changing from female to male.   However, this change in sex will normally only  occur if it presents a potential increase the   individual’s reproductive success. For example,  in a population with a biased sex ratio,   switching to the less common sex would present  more opportunities for reproduction, and therefore   a high degree of competition between males for  access to females will favour protandry. Keeping   all of this in mind, among the thylacopods, the  burden of carrying the ootheca while the young   develop increases the cost of reproduction  for the female, which will incentivize her   to be more selective when it comes to mate choice,  which will increase competition among males, and   ultimately lead to protandry being the dominant  form of hermaphroditism among these species.   The difference in reproductive priorities between  the sexes may also lead to differences in their   physiology as well. Sexual selection occurs when  traits evolve based on their role in competing   for and acquiring mates, often arising from a  preference for the trait by the opposite sex.   Sexual selection frequently operates differently  between the two sexes, sometimes resulting in   sexual dimorphism, wherein males and females of a  species exhibit different characteristics beyond   their reproductive system. The thylacopods already  exhibit sexual dimorphism in the shape of their   gonopods, as only the female has the distinct,  pouch-like gonopods to help her carry the ootheca,   while the males have more typical gonopods  to help them fertilize the females,   but sexual selection may end up driving further  dimorphism. One very common dimorphic trait is   body size, which is also highly correlated with  mate competition; typically, the more intense   the competition between males, the larger the  males will be in comparison to the females,   which may therefore also be the case in these  allobrachids. If competition is particularly   fierce, males may also evolve horns, antlers,  or other structures with which to spar with   each other over access to females, which can also  double as a defence against predators. In these   allobrachids, the equivalent of these structures  may evolve from their recently freed-up forelimbs.   While the females and the juveniles may simply use  these limbs for manipulation and gathering food,   in the mature males these limbs may strengthen  and grow hardened club-like ends to turn them   into powerful punching arms, with which the  males may batter each other into submission.   To withstand the blows from these limbs, they may  also develop reinforced bones within their skull   and further armor around their cephalothorax.  Due to this distinctive armament, we’ll give   them the name ceratobrachids. However, even if  a male ceratobrachid wins a contest for a mate,   he may still sustain injuries during the  battle, which may reduce his chances of   winning further contests, or, in extreme cases,  may even result in him dying shortly thereafter,   so these fights may end up limiting the males’  long-term reproductive success. Because of this,   another clade of allobrachids may settle for a  less violent way of competing for mates. Another   common sexually dimorphic trait is the presence of  frills, crests, or vibrant coloration, which males   use to advertise their fitness without needing to  engage in physical confrontations. One clade of   allobrachids may opt for this strategy, with males  evolving a large, colorful crests on their arms   and cephalothorax to use in courtship displays.  Structures like these usually have no function   outside of attracting mates, and therefore  come about purely through sexual selection,   providing no direct benefit, or sometimes, even  posing an active disadvantage to the individual.   In a process called Fisherian runaway, a  sexually-selected trait may become so exaggerated   that it reduces the individual’s overall fitness,  such as how the enormous tails of peacocks and   widowbirds demand a lot of energy to grow and  maintain and hamper their ability to escape   predators. At first, this seems to contradict the  fundamental premise of natural selection; we’d   expect males that are hindered by overly large or  costly ornamentation to be less likely to survive   and reproduce, and to be outcompeted by males  with more reasonable, less encumbering displays,   so how is it that such exuberant and costly traits  can persist? The answer lies in the fact that,   if a male has survived to maturity despite  having such a huge and obvious handicap,   that, in and of itself, serves as proof of his  fitness, and so females’ may maintain a preference   for the trait even if the males that possess it  have lower survival rates than those without it,   creating a feedback loop that selects for  increasingly exaggerated ornamentation.   Among these allobrachids, fisherian runaway  may select for larger and larger crests, which,   despite the fact that they’ll get in the way  while feeding and make the male more visible and   vulnerable to predators, will be maintained purely  because of the females’ preference for them.   In these animals, which we’ll call  corythobrachids, a male’s reproductive   success will be primarily determined by the size  of his crests, while less successful males may   become female as a way of easing the intensity  of competition, shedding their crests as they   do so. Along with these clades of thylacopods,  the thecopods will also come to thrive in the   temperate latitudes, though they’ll still live  in the shadow of much larger onychodonts like   the hadrodonts and ensidonts. Most species may  avoid competition with these clades by remaining   comparatively small and inhabiting areas with  too little prey to support larger predators.   Where prey is especially scarce, like the cold  deserts and barren scrubland that forms the   majority of the continental interior, the smaller,  more generalist species will have the upper hand.   To help them handle a wider variety of food, their  mandibles, which in other onychodonts are simply   designed for cutting meat into pieces small enough  to be swallowed, may acquire additional grinding   surfaces to help them chew vegetation as well,  while the role of shearing off chunks of flesh   may be increasingly played by the pedipalps, which  may therefore acquire additional cusps and cutting   edges, an innovation broadly equivalent  to heterodonty in earth’s vertebrates.   But beyond these physical adaptations, the  limited availability of resources in their   habitat may also have implications for their life  history. Once again, Bateman’s principle posits   that the difference in investment between the  two sexes during reproduction will incentivize   females to be selective in mate choice and  males to mate with as many females as possible.   However, the lower a species’ population density,  the fewer opportunities there’ll be for mating,   so at some point, instead of wasting time and  energy searching for multiple females, a male may   ultimately achieve greater reproductive success  by investing more heavily in a single mate.   Possibly the most extreme case of this  phenomenon occurs in ceratioid angler fish,   which live at such low population densities  that when a male encounters a female,   he may never get another chance at reproduction,  and so he permanently attaches himself to her,   providing her with a steady input  of sperm to fertilize her eggs.   While the thecopods are unlikely to be driven  to such an extreme, the scarcity of resources   in their habitat will inevitably result in  low population densities, which may select for   increased reproductive investment in the males and  ultimately culminate in the evolution of monogamy,   staying with only a single mate at a time.  Monogamy is quite rare as a reproductive strategy,   but may evolve when females are widely scattered  or if the offspring are likely to die without   investment from both parents. When a male  fertilizes a female, he may remain with her   while she’s pregnant, assisting in hunting and  helping to care for the young once they’re born.   He may only leave once the young are  old enough to survive on their own,   at which point he may begin looking for another  female, a behaviour called serial monogamy.   Another consequence of resource scarcity will  be increased intraspecific competition, so to   protect their access to local food sources, these  thecopods might exhibit territorial behaviours.   However, in the search for a mate, the  male will be forced to wander far and wide,   and thus it may not be feasible for him to  defend a territory of his own, instead sharing   a territory with the females he mates with. But  the female may not accept his presence lightly;   having a male within her territory will mean  she’ll have to share her resources with him,   which she may only tolerate if she judges  him as a decent mate, violently attacking   him and chasing him off if he doesn’t meet her  standards. Because of this heightened interest   in territorial defence, the females may be  larger and more aggressive than the males,   and since the process of acquiring a mate is  a lot more hazardous for the male, find his   own territory and becoming female may present  a potential increase in reproductive success,   and therefore protandry will once again  be the dominant form of hermaphroditism   among these animals. With this life  history strategy, these thecopods,   which we’ll call amphidonts, will be some of the  most highly k-selected animals yet to evolve,   letting them maintain stable populations  as the ice age encroaches. But while   the cold-adapted thecopods and thylacopods  will have an advantage in the new climate,   so will the clades that once inhabited the  central desert, whose ability to survive on very   little food and water will let them considerably  expand their range as the continent dries out.   The amblypods may begin outcompeting other small  herbivores in arid areas within the tropics,   while the lystrocheirids, being adaptable enough  to eat just about anything, may be able to spread   into the temperate plains and scrublands. As  they arrive in these more fertile areas, some   lystrocheirid species may increase the proportion  of plant matter in their diet, with their broad,   multipurpose teeth enlarging to help gnaw  through brachyphyte bulbs and tree roots.   As they rely less and less on scavenging, their  pedipalps, previously adapted for reaching inside   carcasses to probe for fresh meat, may lose their  role in feeding and assume an entirely sensory   function, essentially becoming a pair of antennae.  Both their fossoriality and their nocturnality   will be major assets, having originally evolved as  a way of escaping the desert heat, but now serving   to help them avoid the higher levels of predation  they’ll encounter as they colonize new ecosystems.   But one of the most exceptional aspects  of their biology is in their reproduction:   While the offspring of most other osteopods are  precocial, more or less fully-developed at birth,   the lystrocheirids are highly altricial,  undergoing a significant portion of their   development after they’re born, with their young  being blind, boneless, and largely immobile.   For these young to have any chance of survival,  they’ll need at least some help from the parents,   which in the basal desert-dwelling species  came in the form of birthing the offspring   within a carcass to serve as a first meal, but  in these new species, this may evolve into a   more dedicated form of parental care, such as  keeping the young safe within a deep burrow   and providing them with food as they develop.  Once again, however, Bateman’s principle means   that a male will see greater reproductive  success from mating with multiple females,   and spending time and energy caring for the  young will limit his ability to look for mates,   so unlike the amphidonts, whose females are widely  scattered enough to justify the male assisting   in a parental care, the role of nurturing the  young may fall solely on the female lystrocheirid.   However, while multiple matings will maximize the  male’s fitness, searching for mates will expose   them to many of the steppe-dwelling predators,  and so an individual’s reproductive potential   may be cut short by an early death. If a species  has an especially high mortality rate, then rather   than relying on a decently long lifespan to  provide multiple opportunities for mating,   an animal may benefit from investing in only a  single clutch of offspring early in life while   it still can. Therefore, while most osteopods  are iteroparous, producing multiple clutches of   offspring over the course of their lives, these  lystrocheirids may evolve to become semelparous,   investing all their energy in exactly one  reproductive event and dying soon afterward.   Semelparity is most commonly seen in R-selected  species, and is correlated with small sizes, short   lifespans, low survival rates, and large clutch  sizes. Semelparity is quite rare in clades with   internal fertilization, especially tetrapods, but  it does happen occasionally. Likewise, iteroparity   is likely to be the norm for most osteopods, but  at least some of these lystrocheirids, especially   smaller species with high growth rates, may be  some of the few osteopods to evolve semelparity.   Like many animals, the lystrocheirids may time  their reproduction to occur during the part of   the year when food is most abundant. During this  time, the males may become extremely aggressive   and compete furiously to find and  fertilize as many females as possible,   with the less successful males taking advantage  of protandry to bypass the competition.   The fertilized females may then dig a  burrow in which to incubate the offspring,   of whom they’ll be fiercely protective until  they’re developed enough to survive on their own.   Both the males and females will begin to die  off once the breeding season ends and winter   approaches. The death of the adults may also  help the young survive by limiting intraspecific   competition between adults and juveniles. If  older individuals are able to more effectively   gather resources than younger ones, which will  almost certainly be the case for altricial species   like the lystrocheirids, the juveniles may end up  being outcompeted before they can reach adulthood.   Having the older generation die after the  juveniles leave their mothers’ protection   will help to relieve this competition, but some  species, especially those that retain iteroparity,   may further avoid competition by undergoing  a form of age-dependent niche partitioning.   These lystrocheirids are generalist omnivores,  but the ability to digest vegetation requires a   large foregut, which the young won’t have  developed yet by the time they’re born.   Because of this, the younger animals may be more  inclined to feed on meat and other food sources   that are relatively easy to digest, growing  expanded foreguts and thickened teeth as they age,   gradually transitioning from carnivory to  omnivory. This unusual life history will give   these animals, which we’ll call trypanocheirids,  a bizarre mix of r-selected and k-selected traits,   since they simultaneously invest in large  clutch sizes and high growth rates while also   exhibiting parental care. As mentioned earlier,  K-selection tends to become increasingly favoured   as ecosystems become more heavily populated  and as clades reach larger body sizes,   but there are some niches where R-selection is  quite likely to be maintained: Parasitism is a   form of symbiosis in which one species benefits  at the expense of another, usually characterized   by the parasite living on or within a host from  which it derives part or all of its nourishment.   On earth, parasitism is seen in a huge number of  entirely unrelated clades, although curiously,   it’s virtually non-existent among vertebrates,  possibly because their physiology and life   history strategies are often the opposite of what  would be optimal for parasitism. Parasites tend   toward small body sizes, which let them attach  to hosts more easily and less conspicuously,   and are usually strongly r-selected, as the  biggest obstacle to their reproductive success   is usually the challenge of dispersing to new  hosts, which may only be overcome by producing   offspring in vast numbers. Contrary to this,  most clades of vertebrates are biased toward   relatively large sizes, which usually goes  hand in hand with k-selection, putting them   at a disadvantage in competing for parasitic  niches. Likewise, the osteopods share similar   specializations to vertebrates, so they too  may take up parasitism less readily than other,   smaller clades. In particular, the malacoformes  are some of the smallest animals on the planet,   and have largely maintained the r-selection of  the ancestral lophostomes, making them prime   candidates to evolve parasitism. The transition  to parasitism may begin with phoresis, or phoresy,   using another organism simply as a means of  transport, usually without any negative impact   on the host. This may be ubiquitous among the tiny  malacoforms, as hitching a ride on a larger animal   will let them travel much further than they could  possibly crawl on their own. But perhaps some of   these malacoforms adapt to use their host not just  for dispersal, but also for feeding, subsisting on   the organic debris, glandular secretions, or even  dead skin that builds up on the animal’s hide. If   these malacoforms are especially vigorous in their  feeding, they may even open wounds on the host’s   body, which may result in some species evolving  hematophagy, or blood-drinking behaviours. Blood   has a very high nutrient content, making it  an enticing food source for many animals.   While in facultative hematophages, blood  forms only part of the animal’s diet, obligate   hematophages are specialized for feeding on blood  exclusively. These specializations may involve   optimizing the mouthparts for blood-sucking and  the production of chemical agents to stop the   blood from clotting while the animal drinks it,  maintaining a constant flow of blood as they feed.   This may be accomplished in these malacoforms with  sharp, toothy tentacles to bite through the host’s   skin and simultaneously inject anticoagulants  into the wound. Hematophagy doesn’t automatically   qualify as parasitism, as the interaction between  a hematophage and its host is often quite brief,   while parasitism typically  entails a long-term association.   To be considered a full-fledged parasite, the  animal must be adapted to remain attached to   the host for a significant part of its lifecycle,  which these malacoforms may achieve by evolving   powerful, spine-covered tentacles to  securely anchor themselves to the host,   and a long proboscis to bore into  the skin and reach the bloodstream.   The specializations a parasite undergoes will be  contingent on the exact physiology of the host,   resulting in co-evolution between the two. As  such, many parasites are extremely host-specific,   only adapted for parasitizing a single or a  small number of closely related host species,   so this clade of parasitic malacoforms, which  we’ll call myzognathans, may contain many   hundreds or even thousands of different species,  each one adapted to its own unique host species.   But another clade of malacoforms may evolve  parasitism in an entirely different way.   While ectoparasites like the myzognathans  attach themselves to the outside of the host,   endoparasites live inside their host’s body,  where they’re protected from the environment   and are less likely to be removed. This requires  a way of entering the host, which very small,   especially microbial parasites may achieve  by infiltrating the respiratory tract,   burrowing through the skin, or even by using  another parasite or hematophage as a vector,   but one of the most common methods is  by allowing themselves to be eaten.   Among the malacoforms, many species may lay  their eggs along or under leaves, where they   may end up being unwittingly swallowed by foraging  herbivores. If one clade evolves to produce eggs   that are durable enough to survive the digestion  process, some of the larvae may hatch out while   still inside the herbivore, where they’ll be  able to feed on the contents of the animal’s gut.   Offering ample nutrients and devoid of predators,  the herbivore will serve as an ideal incubator,   prompting some malacoforms to spend  their entire lives within their host.   However, one drawback to this lifestyle is the  difficulty it presents when it comes to dispersal;   being unable to survive outside the host  means that the only way to transmit their   offspring to a new host will be to disperse  their eggs in the host’s excrement, in the   hope that they land somewhere where they have a  chance of being ingested by another herbivore.   To maximize the chances of this happening, most  endoparasites once again tend towards R-selection,   producing enormous quantities of eggs  of which the vast majority will die.   While their reproductive systems expand, many  endoparasites also show a severe reduction   or simplification in other body systems, as  passively feeding on the host’s gut doesn’t   require a great deal of bodily complexity, to the  point that many endoparasites, like tape worms,   thorny-headed worms, and horsehair worms, have  no functioning digestive or respiratory tracts,   simply absorbing oxygen and nutrients  through their outer surface.   Similarly, in these malacoforms, the eyestalks and  other sensory organs, being essentially useless   inside a host, may atrophy completely, along with  the limbs and the majority of the internal organs,   while the oviduct may grow so large that its  extends into the spine-covered proboscis,   with which they anchor themselves to  their wall of their host’s intestines.   This lifestyle is likely to have first evolved  very early on in the history of terrestrial life,   with the parasitic clades diversifying in  tandem with their hosts. Like in ectoparasites,   co-evolution with the host results in most  endoparasites being highly host-specific,   but with one major caveat: being utterly reliant  on a single host means that if the host dies,   then so does the parasite, which is an especially  big problem if the host species experiences a high   rate of predation. Because of this, unlike  ectoparasites, which are often monoxenous,   completing their entire lifecycle on a single  host species, many endoparasites are heteroxenous,   developing inside one or more intermediate  hosts before being transmitted to the final,   or definitive host, only then maturing into  their adult form and commencing reproduction.   On the barren scrublands of the western continent,  one clade of these malacoforms may coevolve with   a family of xerostracans, which will serve  as prey for some species of theropterans.   When a theropteran kills and feeds on one  these xerostracans, it may unwittingly ingest   any malacoforms infesting in, which, if they  survive, may adapt to use the theropteran as   their definitive host. This will have the added  benefit of massively increasing the range of   their dispersal, since the theropterans  can travel much further than the slow,   ponderous placostracans can. This heteroxeny means  that to complete their lifecycle, endoparasites   like these often rely on their intermediate hosts  being eaten, so to maximize the chances of this   happening, they may produce chemical agents  that alter the morphology or behaviour of the   host to make it more susceptible to predation,  such as how the nematode myrmeconema causes the   abdomen of the ants it infests to turn bright  red, making them more noticeable to predators,   while lancet liver flukes manipulate their  ant hosts into climbing blades of grass,   where they’re more likely to be inadvertently  swallowed by cows, which serve as the flukes’   definitive host. In these malacoforms, when  the larvae reach maturity, they may secrete   chemicals that cause their xerostracan hosts to  lose their fear of predators, and to forage for   food during the time of day when the theropterans  are most active. Only once the larvae pass into   the theropteran’s gut will they develop their  expanded oviducts and begin producing eggs.   This may be only one of hundreds of different  lifecycles that develop among this clade,   with every species adapting for its own sequence  of hosts within its local ecosystem. We’ll call   them echinostomes. While it would be impossible  to account for all the changes that occur as the   continents break apart, the clades we’ve discussed  here will constitute some of the major groups that   will evolve over the course of about 30 million  years as life in the east and west diverges.   But beyond the two continents, there are some  habitats on this planet that present opportunities   for far more extreme forms of allopatry to  occur, and whose biogeography demands a more   thorough examination. In the next episode, we’ll  explore how life adapts to colonize the many   islands across the sea, and explore the unique and  highly unusual ecosystems that arise as a result.   Thanks to all the artists over on discord who  contributed artwork for this episode, and who   save me an enormous amount of time and effort when  making these videos while simultaneously making   them look a hell of a lot better in the process.  Links to the main server and the alien biospheres   fan server in the description. And once again, a  massive thanks to all the patrons, whose continued   support makes videos like this possible. Thanks  for watching, and I’ll see you in the next video.
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Channel: Biblaridion
Views: 375,786
Rating: 4.9775624 out of 5
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Length: 48min 19sec (2899 seconds)
Published: Mon Mar 22 2021
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