Carrion Trilobites

Elsewhere in the world, fossil traces of trilobites living in freshwater, let alone venturing onto land, are scarce at best. On Ryl Madol, land trilobites are curiously quite common. In some respects, their evolution here was likely easy, in other ways hard. Like their cheliceratan uncles they were able to evolve simple book-lungs by invagination of their gill-arms into their body. However, what must have been a great challenge is their lack of true mouth-parts, compared to the wide range of mandibles and cheliceres of their cousins. The closest thing they have is a hypostome, which acts more like a backwards-pointed spade or shovel rather than a jaw. This is not a big problem when living on the ocean floor, where microscopic detritus constantly rains from above, but land-life does not grant such luxuries. As the myriapods prove, even feeding off leaf-litter requires mandibles. Facing jawed competition from not only centipedes and arachnids but also the great barage of insects, it is perhaps for this restraint why elsewhere on Earth trilobites were never able to make the big leap onto land, dooming them to an oceanic existence where they would eventually meet their end once marine environments became near-uninhabitable in the Permian. On Ryl Madol the trilobites were able to carve out a few terrestrial niches for themselves, sometimes literally. The reasons for this are unknown but it has been speculated that it may be potentially due to the island’s distinctive lack of derived Neoptera insects, which may have given archaic arthropods more room to expand and adapt.

While some of the rylmadolian trilobites have adapted to feed on soft-bodied worms and nematodes, the majority of terrestrial trilobites are still detritus-feeders, living off the soft excrements and droppings of various larger animals through simple sucking- and shoveling-motions. These basically fill out the same ecological niches as dung beetles and scarabs would. While this may seem like a pitiful existence from a human perspective, it is actually quite a respectable job, as many a great pathogen and toxin meets their end in the robust glabella-stomachs of these critters, keeping the ecosystem healthy and safe from serious disease outbreaks. Still, at least one group has taken up a more savoury diet. The carrion trilobites are a special clean-up-crew that appears anywhere where a large vertebrate carcass lies, usually deep in the thermal chasms. The little critters possess a strongly reinforced hypostome shaped like a wood-carver. They use this to scrape off meat into easily digestible chunks and they are especially good at removing the last bits of flesh from bones. A large swarm of these can turn an ungainly corpse into a beautifully stripped and clean skeleton in under two hours. Rumours of them allegedly doing the same to still-living stalkers are almost assuredly a myth.

Orthrocaris

Ryl Madol is not just a nightmare for its brave explorers but also for the average taxonomists, whether they study the animals live on the island or preserved in formaldehyde from the safety of their comfy university offices. Evolution has taken its very own unique path here, often with entire groups which are elsewhere only known from fossils. The end result are thus forms and combinations which defy all of our anatomical borders.

One of the most enigmatic of these is little Orthrocaris antebabylonensis, or Insanity Fish, curiously often found swimming within the pools and swamps which form at the center of the ancient, ruined plazas and their flooded sewers. There can be no other way than to simply describe it as a chimaera. At the front, a soft, bivalved, gill-bearing carapace, very much like in a phyllocarid crustacean, gives rise to two segmented arms and a head with two compound-eyes. So far, so arthropod. But then the mouth is ringed by four or more unsegmented little tentacles, very much like in a cephalopod. The hidden mouth itself is a single claw-shaped mandible similar to that of a velvet worm. Behind the carapace then emerges an elongated, eel-like body, with a dorsal and ventral fin running down its length. Though this body is soft, on the inside it contains a stabilizing organ which unmistakably resembles the chorda dorsalis of our own phylum. What can be said of such a creature which wildly mixes traits or arthropods, cephalopods and chordates? It is the invertebrate(?) equivalent of a platypus.

That nobody has been quite able to shoehorn this little monster into any taxonomical drawer is not surprising. Some claim the arthropodal characteristics outweigh the others, arguing that this may be some archaic member of that phylum that simply evolved the other traits through convergent evolution. Others argue that a chorda dorsalis classifies Orthrocaris automatically within the Chordata. The truth may literally lie somewhere in-between. Some have noted a resemblance to the Cambrian organism Nectocaris, which used to be similarly reconstructed as a chordate with the head of a bug. However, these reconstructions have long been deemed erroneous, as better-preserved fossils of the organism instead suggest that Nectocaris looked more like a flattened, two-tentacled squid. The resemblance to old reconstructions may thus only be a coincidence. Others have suggested a possible affinity of Orthrocaris to the Vetulicolia, who did greatly resemble an odd mix of arthropod and chordate. The cladistic placement of Vetulicolia is itself highly debated and no known vetulicolian fossil is known to have possessed limbs or tendrils as Orthrocaris has, so this affinity also remains entirely speculative. Said limbs most resemble the scourges of Cambrian legless arthropods like Leanchoilia. If recent genomic studies turn out to be true and Deuterostomia is indeed not a true group, then Orthrocaris may be the descendant of some unknown basal group of the alternative chordate-protostome clade Centroneuralia. In other words, it could, in a sense, literally be the ancestor of all groups discussed. Perhaps it could even be the result of some hybridisation accident, back in some dark Cambrian days when the genomes of all the phyla were still similar enough for such taxonomic bastards to come into existence.

Regardless of its classification, contact with Orthrocaris should be avoided at all costs unless one is properly protected. Although the organism is only as large as an index finger, the bite of some individuals is lethal. Not because it is venomous. For some unknown reason, the saliva of the organism carries a prion inside it. Prions are misfolded proteins, which can cause other proteins to misfold in the same way, allowing them to pathogenically reproduce almost like viruses. There is no known treatment method for prion diseases, so 100% of all infected end up either dead or severely disabled for life.

Like mad-cow-disease, the Orthrocaris-prion is neurodegenerative, meaning it attacks the human brain. It can lead to insanity, followed eventually by brain-death. Some infected have been reported to have escaped their containment and in a mad rave wandered out into the island, only to return days later, with almost no scratch on their body. The animals on the island (who seem to be immune to the prion) show no interest in attacking infected humans. Most shockingly, those patients who escaped without a protective mask on suffered no ill effects from the island’s toxic spores. It seems almost like a sadistic twist that only this disease grants humans the ability to survive on Ryl Madol, but at the price of a short life of insanity. The Headtakers, never too fond of outsiders, have by now discovered the disease’s cause and effect and weaponized it. Using the snake-like glottis in their lower jaw, Headtakers are capable of using blowpipes. When they are feeling especially sadistic, they will carry a leathern waterbag full with little, living Orthrocaris with them, remove the “fish’s” tiny arms and shoot them through the pipes at the necks of unsuspecting humans.

The most bizarre aspect of the prion is just how it cripples the brain. In most cases it mainly affects the areas responsible for language processing, making the infected eventually incapable of talking in coherent words or sentences. Instead, they all start spouting completely unintelligible gibberish. However, all those infected speak the same gibberish and seem to be able to understand each other to some degree, while suddenly incapable of understanding their healthy colleagues. In the early stages of the disease, before they lose the capability of speech completely, they can be heard in their quarantine cells seemingly conversing with each other in undecipherable but still distinctive words and phrases like “Oadriax!”, “Dilzmo!”, “Cordziz”, “Agtoltorn parach asimp!”, “Pappe hasatan, pappe hasatan aleppe” and “Rahel mai ameche zabi almi”. I imagine this must surely be of interest for neurolinguists, but they have not returned my calls.

Sealacanth

As the sun sets and the waters calm down, a strange creature hauls itself onto a beach. Its limbs end in fins, yet they already have joints which foreshadow elbows and knees. Its two dorsal fins now hang to the sides, useless outside the sea. It breathes heavily, almost strained, but it is doing better than the other water-dwellers that have been swept here, dead ammonites, clams and fish, which have been crushed by the gravity and heat. It succeeds where others fail. Beyond the beach lies a lush rainforest, filled with big invertebrates. To the victor will go the spoils.

Only the presence of scaly, bipedal creatures on the beach and the distant cries of seabirds betray that this is not an ancient scene from the middle of the Devonian period. This is the very modern life of a particular fish on the island of Ryl Madol. The sealacanth (Erpetolepis littoralis) looks like an ancient holdover but is in many ways also something quite new, perhaps among the most advanced of the walking fish that live on Earth.

Despite the name and appearance, Erpetolepis is not a close relative of the coelacanths which also swim around in the Indian Ocean. The presence of internal nostrils and labyrinthodont teeth identify it not as an actinistian but as a tetrapodomorph, though closer to basal forms such as Eusthenopteron than to more famous fossils like Tiktaalik or the true tetrapods. Its many adaptations towards littoral life seem to be parallel developments. Unlike archaic lobe-finned fish, the sealacanth has clearly defined limb-joints, though not yet true fingers, instead just a flexible fan of lepidotrichia like other fish. It is a very subtle difference, but the shoulder-girdle is also not fully connected anymore to the skull, allowing it to move its head independently of the body, an important development which in true tetrapods led to the evolution of a true neck. It also sports fully developed lungs in addition to gills.

Actual seals are largely absent from the coastlines of Ryl Madol, on account of the presence of the toxic spores. While there is no shortage of aquatic and semiaquatic amphibians and reptiles on these beaches, very few of them have seemingly managed to utilized the resulting opportunity quite as successfully as these clumsy-looking fish, earning them their name. Like pinnipeds, sealacanths hunt in the shallow and open waters for food but when they want to sleep and rest, they come onto land and sometimes even sleep in groups. Though gregarious, they are not exactly social, as the lack of ears limits their ability to communicate. Towards human explorers they are largely apathetic, though some degree of curiosity is reported on occasion.

The prey of sealacanths consists of slow-swimming or crawling invertebrates and small vertebrates, largely fish, lizard-sized marine reptiles and washed-up carcasses. Sometimes sealacanths can be observed digging up the rocky nests of Spheniscodapedon, the penguin-like reptile seen here in the background, in order to feed on their eggs. This does not always go well, as these “rhynchoguins” tend to guard their nests and have pretty mean hooked beaks, more than capable of ripping chunks of flesh out of even a large fish such as this. Though it is larger and stronger, if a flock of these reptiles gangs up on a sealacanth, they can very easily rip it to shreds. The further the fish crawls into the island, the more dangerous the threats become and so too the farther out into the sea it swims. Perhaps being stuck, ecologically and evolutionarily, on the water’s edge is good enough, at least for now.

Castorosaurus

Ryl Madol is able to support a large amount of amphibious and aquatic life thanks to its many pools, ponds and streams which pay tribute to large lakes and rivers. For many of the tetrapods and arthropods, these are vital for their reproduction, as their eggs and larvae lack shells and need to be laid in water. However, many of these breeding pools are not necessarily provided by the island topography. They are made by the wildlife itself, though, ironically, not by amphibians.

Encountered in almost every waterway are waterlogged wooden structures, which early explorers at first mistook as as bridges or fish-traps created by the island’s intelligent natives. But no. These are dams built by an aquatic reptile called Castorosaurus, whose name literally translated to “beaver-lizard”. Castorosaurs are derived members of the Captorhinidae, a group of Palaeozoic reptiles that, on account of missing postorbital windows, were traditionally grouped with the “Anapsida” alongside the turtles, though recent work has recovered them as being eureptiles close to the Diapsida. Even more recent work has declared them stem-amniotes, though this remains controversial.

Just like beavers, castorosaurs are ecosystem-engineers who prolifically build dams along narrow waterways, creating many pools throughout the island in which other organisms can thrive and reproduce. Their dams usually have a hollow chamber that can only be accessed from underwater. Here the castorosaurs lay their eggs and raise their young. The dam structure prevents terrestrial or aerial predators from easily gaining access to the nest, though the parents still tend to be on the look-out for any of the creeping amphibians that may dive and slither below.

Unlike the terrestrial captorhinids of the Palaeozoic, castorosaurs feature many aquatic adaptations such as webbed feet and a dorso-ventrally flattened tail like a platypus. This tail-shape is of course quite curious, as reptiles usually swim through sideways undulation and so one would expect a laterally flattened tail, but the tail seems to be mainly used as a sort of hammer in dam-building. What castorosaurs have retained from their ancestors are the long, kinked snouts with specialised teeth at the front. Said teeth have become quite large and spiky. Many people easily make the mistake of assuming that castorosaurs used their large front-teeth just like beavers to gnaw at wood, but that is in fact incorrect. They are entirely unsuited for such a task and are instead used to capture fish.

If they cannot gnaw at trees and branches, where do castorosaurs get the wood for their dams then? The answer is that they simply pick it up from the forest or lake floor and so do not need to saw up fresh wood. But why it is that simple for them is actually interesting. Vast areas of Ryl Madol are covered in thick carpets of wood-litter that degrade into smaller pieces over time as animals trample over them, but never truly rot and decay. Some pools and moors have as a consequence devolved into large, malignant peat bogs in which unwary animals may stumble and get trapped. It is highly unusual and not something seen in any other forests in the world. Only recently has the reason behind this been revealed by botanists and microbiologists: Ryl Madol entirely lacks any organisms, be they bacteria or fungi, that can digest lignin, the main polymer that makes up wood. This used to be true for all of Earth in the Carboniferous period, from which huge coal deposits still survive. While it explains the primordial nature of Ryl Madol’s forests, it just opens up many more questions. Why has lignin-digestion never evolved or arrived on Ryl Madol? Or did it exist here once but the respective microorganisms have since somehow gone extinct? The lack of notable coal deposits in the island’s geological formations may be evidence of the latter, pointing towards this being a fairly recent development. It seems almost like time on the island is moving backwards.

Walky Tully

Ryl Madol is already home to a variety of animals which by all rights should not exist, at least not anymore, but even among them the walky tully (Micrormus holidayi) stands out, as it has evaded proper classification to this day.

What really can be said about an animal whose jaw sits at the end of an extendable proboscis, has eyes on stalks like a snail, a body like a tadpole and three clumsy legs? It has a backbone, so at the very least it can be considered some kind of vertebrate, but that is where consensus ends. Unlike any other vertebrate, there is no direct bone-connection between the jaws and the cranium, the “neck” is just a floppy tube made of cartilage and muscle, attached to what resembles the hyomandibular bone of sharks. Despite living in water, it has lungs but no gills. Its “fingers” have no resemblance to those of tetrapods, instead having evolved from fin-rays, the internal anatomy of the forelimbs somewhat resembling the alternating bone-structure of lungfish-fins. The single hindlimb is located behind the cloaca, meaning that it is possibly homologous with the anal fin found in most fish. However, in the vertebrae, the pleurocentrum dominates, which is a trait more typical of stegocephalian tetrapods than of fish.

With its proboscis and stalked eyes, many researchers have obviously tried linking this animal to the notorious fossil Tullimonstrum gregarium of Illinois, which is where its common name comes from. However, the classification of Tullimonstrum is itself controversial, as it is not even clear if this organism was a vertebrate or an invertebrate of some kind. If Tullimonstrum was a vertebrate, it would have been one of the most basal cyclostome kinds, a relative of lampreys and hagfish. It notably has no fins whatsoever, at least none that were ever able to be identified from the fossils. If Micrormus is indeed a descendant or close relative of the tully monster, then it must have evolved limbs, lungs, a loss of gills and various other characteristics that are not present in its Carboniferous ancestor independently of other vertebrates.

This suggests that the resemblance is merely due to convergent evolution and that Micrormus is some kind of highly aberrant bony fish, possibly sarcopterygian in origin. Due to its amphibious characteristics, the most radical proposal has been that it may descend from some type of tetrapodomorph that, like the coelacanth, still possessed a muscular anal fin but for whatever reason had lost its pelvic fins, which in true tetrapods evolved into our hindlegs. This hypothesis is not at all popular, but other suggestions have not been less crazy, such as the idea that Micrormus is a vertebrate-mimicking cephalopod or a relative of the dancing worms of Turkana. Genetic studies that could shed more light on the matter are unfortunately lacking.

In contrast, the actual life habits of the walky tully are surprisingly unspectacular. It is a small animal, able to fit comfortably inside a human hand. Like most fish on the island it has developed an amphibious lifestyle and spends a lot of time crawling around or even sleeping on lake shores. When “walking”, the single hindlimb is not used merely as support for the forelimbs but also helps the animal push forward, earning it the alternative name “mud-tripod”. In the water it hunts smaller fish and tadpoles, such as the mantiraptor larva seen here. Some researchers have proposed that it actually is a specialized tadpole-predator, but there is no conclusive data that it prefers this prey over any other small aquatic critters. Among its own enemies are various stegocephalians, predatory fish and the stork-like gruisaurs.

Tullys reproduce through external fertilization and lay their spawn inside protected alcoves along riverbanks. The young hatch as miniature adults without first going through a larval stage, which is why gleaning its evolutionary history from embryological data has also proven difficult.