invertebrate – Nix Illustration

Falcatacaris

The enigmatic thylacocephalans were a group of bizarre little arthropods, found in marine deposits all over the world from the late Ordovician (~435 million years ago) to the late Cretaceous (~85 million years ago). They had shield-like bivalved carapaces, large compound eyes, three pairs of spiny grasping limbs, and multiple pairs of small paddle-like swimming limbs, but details of their internal anatomy are poorly known and their evolutionary relationships to other arthropods are still very uncertain.

Traditionally they’ve been classified as crustaceans, possibly as close relatives of remipedes or malacostracans – but they’ve also recently been proposed as instead being part of a much more ancient branch of arthropods, potentially related to stem–mandibulates like Acheronauta.

Falcatacaris bastelbergeri was a thylacocephalan living during the late Jurassic, about 150 million years ago, in what is now Germany. Around 2.5cm long (~1″), its carapace had tiny interlocking square “teeth” resembling a zipper along the hinge line between the two valves, a ridge along each side, and a long pointed knife-shaped spine at the front.

Like other thylacocephalans it was probably a swimming predator, likely nocturnal or hunting in murky conditions based on its enlarged eyes, and would have captured smaller aquatic prey using its raptorial limbs.

Tachypleus syriacus

Tachypleus syriacus was a horseshoe crab from the late Cretaceous (~100-95 million years ago) of what is now Lebanon.

Closely related to modern tri-spine horseshoe crabs, it displayed a similar level of sexual dimorphism. Females grew to at least 25cm long (~10″), with rounded front edges to their carapaces and shorter rear spines, while males were around 30% smaller with a scalloped shape to the front of their carapaces.

One recently described female specimen also preserves distinctive nodules around the rim of its carapace, which may represent some sort of sensory structure.

This particular specimen is also unique for preserving a coprolite in the process of being expelled from the horseshoe crab’s body – that’s right, it died while pooping.

Hexameroceras

Hexameroceras panderi was a nautiloid cephalopod that lived during the late Silurian, about 425-423 million years ago, in what is now Czechia.

Around 5cm long (2″), it had a downwards-curving egg-shaped shell that preserved the original color pattern on one fossil specimen, showing closely-packed crisscrossing vertical and horizontal bands.

Like several of its close oncocerid and discosorid relatives, its shell also developed a highly constricted opening as it reached maturity. This eventually formed into a narrow visor-like shape with several lobes that probably correlated to the life positions of the eyes and arms, with a “spout” at the bottom for the siphon.

Diagram showing how the lobed "visor" formed in Octameroceras — Development of the “visor” in the related *Octameroceras sinuosum*
From fig 6 in Stridsberg (1981)

The function of this structure is still unclear. It may have been a defensive measure against predators – but it would have also severely limited the range of motion of the arms and the size of food that could be eaten through the mouth, suggesting that Hexameroceras may have specialized in very small prey, perhaps even filter-feeding.

Another possibility is that these visored nautiloids might represent brooding females, walling themselves into their shells to protect their eggs and dying after releasing the hatchlings through the tiny remaining gap.

Panacanthocaris

Panacanthocaris ketmenica* here was a member of an extinct group of crustaceans known as kazacharthrans – close relatives of modern tadpole shrimp known mainly from Central Asia during the mid-to-late Triassic (but with possible German relatives from both the late Triassic and further back in the late Paleozoic).

Fossils of Panacanthocaris have been found in Kazakhstan and northwest China, dating to about 235-221 million years ago. It was fairly big compared to most of its modern cousins, reaching at least 10cm in length (~4″), and had distinctive spines around the edges of its carapace and its telson.

It’s not clear if it had eyes – there’s a single opening near the front of its carapace that may have housed some, and so I’ve depicted it here with just one naupliar eye similar to the “third eye” of tadpole shrimp.

It probably had a fairly similar lifestyle to its modern relatives, living in shallow freshwater and temporary pools and opportunistically feeding on everything from algae to smaller aquatic animals.

(* Sometimes also called P. ketmenia. May also be the same thing as Iliella spinosa, but until that paper is officially published the current name still stands.)

Triopus

Although Triopus draboviensis here might look like an isopod or a trilobite, this small arthropod was actually part of a rather rare group called cheloniellids.

Known from the early Ordovician to the early Devonian (~480-408 million years ago), only about 7 different species of cheloniellid have been described so far. Their evolutionary relationships were uncertain and controversial for a long time, but currently they’re thought to be distant cousins of trilobites within the Artiopoda.

Living in what is now Czechia during the late Ordovician, about 460-450 million years ago, Triopus is only known from two partial fossils. It was around 4cm long (~1.6″), and like other cheloniellids it had a body made up of wide radiating exoskeleton segments that fully covered its legs, and probably also a pair of whip-like appendages at the rear.

Its body was more domed than those of its relatives, who were generally very flattened, suggesting it was specialized for a slightly different lifestyle or habitat. Without any preserved appendages it’s not clear what its ecological role was, but since other cheloniellids had horseshoe-crab-like feeding structures it may have been a similar sort of generalist, preying on small invertebrates and scavenging carrion on the seafloor.

Glossoceras

Although the only nautiloids living today have characteristic tightly coiled shells, earlier in their evolutionary history these cephalopods were much more diverse.

And Glossoceras gracile here is an example of one of the more unusual groups of nautloids: the ascocerids.

Living during the Late Silurian, about 422 million years ago, in wheat is now Gotland, Sweden, Glossoceras was only around 5cm long as an adult (~2″). Like other ascocerids it started out its life looking like a fairly standard early nautiloid, with a long straight shell that curved slightly upwards, but as it approached maturity things got weird – the front part of the shell grew out into a much more bulbous flask-like shape, and the old juvenile section broke off entirely.

The gas-filled buoyancy chambers of its adult shell were positioned directly above its body chamber rather than behind like in other nautiloids, giving it very good stability in the water. The shell walls were also very thin and lightweight, which would have made it a much more maneuverable swimmer.

Spectember/Spectober 2023 #11: A Large Spider

An anonymous submission requested a “spider the size of a coconut crab”:

A shaded sketch of a speculative giant herbivorous descendant of jumping spiders. It's a big stocky spider covered in fuzz, with thick legs ending in cloven-hoof-like claws. It has the characteristic large main pair of eyes of jumping spiders, with the other three pairs more spread out around the front and sides of its wide head, and it also has two pairs of "horns". Its abdomen is wide and round, somewhat flattened on the top, with several spiky structures around the edge.

Ceratohispidus aspectus is a distant descendant of jumping spiders living on an Aotearoa-like landmass, isolated with no mammalian predators.

This particular lineage is notable for both their extreme gigantism (with their larger size and weight causing them to lose the ability to jump) and for having taken up herbivory in a similar manner to one modern species. Most of these big plant-eating spiders are around the size of wētāpunga, and occupy a similar ecological niche, but Ceratohispidus is the largest of them by far – rivalling the modern coconut crab with a body length of up to 40cm (~1’4″) and a legspan of almost 1m (~3’3″).

After reaching sexual maturity at 5-10 years old, adults grow very slowly, molting only once every year or two and taking several decades to actually get anywhere close to their maximum size.

Ceratohispidus’ thick legs end in hoof-like claws, and it selectively browses on vegetation by snipping off pieces with its pincer-like palps. A gizzard-like structure in its digestive system helps to grind up fibrous plant material with small gastroliths, and its wide abdomen houses both large book lungs and a tracheal system with air sacs that can contract and expand to provide a small amount of active ventilation.

While the “horns” and spikes ornamenting its body may provide some defense from the few avian and reptilian predators in its habitat, they’re mainly used as part of highly elaborate visual displays between individuals.

Spectember 2023 #03: Raptorial Isopod

Modzilla07 asked for a “eurypterid or anomalocarid-esque isopod”:

A shaded sketch of a speculative swimming predatory isopod. Its shaped similarly to an extinct sea scorpion with a streamlines segmented body that ends in a fin-like horizontal tail fan. It has short antennae, a large pair of spiny grabbing front appendages similar to an anomalocarid, large compound eyes on short thick stalks, and two pairs of its legs are modified into long paddle-like structures.

Agriopterus modzillaseptenorum is descended from scavenger-predator intertidal cirolanids. At about 10cm long (~4″) it’s a giant compared to most other isopods, but not nearly as big as some of the radiodonts and eurypterids it convergently resembles.

Adapted for a free-swimming lifestyle, its second pair of antennae have been modified into spiny raptorial appendages and its first two pairs of legs have become flat swimming paddles. It’s a voracious little predator, usually snatching small fast-moving prey from the water and raking up soft-bodied animals from the seafloor – but groups will sometimes opportunistically swarm on much larger dead, dying, or injured targets.

Tuzoia

The tuzoiids were an enigmatic group of Cambrian invertebrates known mostly just from their spiny bivalved carapaces. Although hundreds of fossils of these arthropods were discovered over the last century or so, only vague fragments of the rest of their bodies have been found even in sites usually known for preserving soft tissue impressions.

…Until late 2022, when several new specimens from the Canadian Burgess Shale deposits (~508 million years ago) were described showing tuzoiid anatomy in exceptional detail, finally giving us an idea of what they looked like and where they fit into the early arthropod evolutionary tree.

Tuzoiids like Tuzoia burgessensis here would have grown up to about 23cm long (~9″). They had large eyes on short stalks, a pair of simple antennae, a horizontal fluke-like tail fan, and twelve pairs of appendages along their body – with the front two pairs at the head end being significantly spinier, and most (or all) of these limbs also bearing paddle-like exopods.

The large carapace enclosed most of the body, and was ornamented with protective spines and a net-like surface pattern that probably increased the strength of the relatively thin chitinous structure.

Together all these anatomical features now indicate that tuzoiids were early mandibulates (part of the lineage including modern myriapods, crustaceans, and insects), and were probably very closely related to the hymenocarines.

Tuzoiids seem to have been active swimmers that probably cruised around just above the seafloor, with their stout legs suggesting they could also walk around if they flexed their valves open. The arrangement of their spiny front limbs wasn’t suited to grabbing at fast-swimming prey, but instead may have been used to capture slower seafloor animals or to scavenge from carcasses.

Ursactis

Soft-bodied annelid worms only very rarely fossilize, so the group’s origins during the Cambrian Period are still rather poorly understood. So far about thirteen different species have been found in sites of exceptional preservation, showing that even very early on in their evolution these worms had already diversified into a wide range of ecologies including bottom-feeders, carnivores, swimmers, tube-builders, and even symbiotes sharing living space with early acorn worms.

Ursactis comosa here adds a fourteenth species to the list. Found in a newly-discovered outcrop of the 508-million-year-old Burgess Shale fossil deposits in western Canada, it’s known from nearly 600 specimens clustered together in several large groups, making it the current best-known and most numerous of all Cambrian annelids.

Up to about 1.5cm long (~0.6″), it was a polychaete-like worm bearing bundles of long bristles. There was a pair of large sensory palps on its head, and its body was made up of an unusually small number of segments – just 10, with larger individuals just increasing the size of their segments instead of adding on additional ones like most modern annelids.

Unlike other Cambrian annelids it also shows some evidence of basic tagmatization, differentiating some of the rear segments of its body with much longer bristles.

The large numbers of Ursactis found preserved in one place suggests these worms were exhibiting some sort of swarming behavior. Since ages from juveniles to fully-grown adults are represented together, and their anatomy indicates they were crawling detritivores, they were probably all taking advantage of a particularly nutrient-rich patch of seafloor at the time they were abruptly buried in a mudslide.