artiopoda – Nix Illustration

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.

Strange Symmetries #06: Trilobite Tridents

The genus Walliserops was one of the weirdest-looking trilobites, covered in numerous pointy spines and sporting a large three-pronged “trident” on the front of its face.

They also had some degree of asymmetry in their bodies. Their tridents often didn’t fork evenly, and their long forehead spines curved off to one side – possibly so they could lift their heads up without stabbing themselves in the back.

Walliserops hammii lived in what is now Morocco during the early-to-mid Devonian, about 403-392 million years ago. Around 5cm long (~2″) It was one of the “short trident” species of Walliserops, and its chunky forehead spine curved particularly strongly to the right.

The function of these trilobites’ elaborate tridents is still poorly understood. But an unusual individual of the long-tridented species Walliserops trifurcatushas been found with a lopsided four-pronged trident, and since it was able to grow to full maturity the shape of the structure probably wasn’t absolutely vital for survival, suggesting it wasn’t used for feeding or sensory purposes.

The tridents may instead have been used for combat with each other similar to the horns of some modern beetles. However, these sorts of features are usually only seen in males, and there’s currently no definite evidence for any significant sexual dimorphism in trilobites.

(Although perhaps like ceratopsid dinosaurs their ornaments were just present in both males and females, being also useful for species recognition, visual display, and defense against predators.)

Cambrian Explosion #54: Trilobita – Transform and Roll Up

Most trilobites were able to roll themselves up into a protective ball – a behavior known as enrollment or volvation – exposing just their heavily armored backs to attackers. They’re often found fossilized curled up like this, and rare preservation of soft tissues shows that they had a complex system of muscles to help them quickly achieve this pose while simultaneously tucking their antennae and all their limbs safely inside their enrolled shells.

Some species also developed sharp defensive spines and spikes that jutted out when they enrolled, making themselves even more daunting to potential predators in one of the earliest known examples of an evolutionary “arms race”.

Cambrian Explosion #53: Trilobita – A Prolific Paleozoic Posse

The biggest stars of the Cambrian euarthropods, and most of the Paleozoic Era, were of course the trilobites. Known from literally tens of thousands of species spanning over 270 million years, they’re some of the most recognizable and popular fossils.

Trilobites’ exact evolutionary origins and transitional forms are unknown, but they’re thought to have originated in Siberia in the very early Cambrian and their leg anatomy indicates they were a part of the artiopodan lineage. They made a sudden and dramatic entrance to the fossil record about 521 million years ago, appearing fully-formed and rapidly diversifying and spreading all around the world within just a couple of million years.

Their hard calcified exoskeletons made them much more likely to fossilize than soft-bodied animals, with a distinctive three-part body plan consisting of a head shield, three-lobed thorax segments, and a tail shield. Each individual regularly molted their carapace throughout their life, meaning that most trilobite remains are actually empty discarded shells rather than actual carcasses.

Along with being heavily armored arthropod tanks, most species were also able to roll themselves up to defend against predators, and some developed additional elaborate spines and spikes.

…And some were just weird.

Cambrian Explosion #52: Artiopoda – Who Needs A Thorax Anyway?

The nektaspids were one of the most unique-looking groups of artiopodans, with soft-shelled unmineralized bodies, no eyes, and large head and tail shields with very few actual body segments in between – varying from 6 all the way down to none at all.

First appearing in the fossil record around 518 million years ago, only a few different species are known but they appear to have been abundant animals distributed in outer shelf waters worldwide during the Cambrian.

Their classification has traditionally been uncertain but specimens with well-preserved limbs show very trilobite-like leg anatomy, helping to place them in the artiopodans as potentially some of the closest “trilobitomorph” relatives to the actual trilobites.

Cambrian Explosion #51: Artiopoda – Surprising Lookalikes

The aglaspidid artiopodans were a major lineage of early Paleozoic euarthropods – one of the most diverse after their cousins the trilobites, although far far behind them in terms of actual species numbers.

But despite their diversity and worldwide range actual fossils of them are incredibly rare, and for a long time they were considered to be a “problematic” wastebasket group of uncertain affinities, mainly interpreted as being related to the chelicerates. More recently evidence from preserved limb anatomy has instead placed them within the artiopodans in a grouping known as vicissicaudatans, closely related to forms like Sidneyia and the later cheloniellids.

Unusually for euarthropods they had a phosphatic exoskeleton, and they experienced their main burst of diversification in the late parts of the Cambrian period, after most of the actual evolutionary explosion had already settled.

They mainly inhabited shallow near-shore environments, and may actually have been some of the very first animals to venture onto land. Some examples of the trace fossil Protichnites might represent aglaspidids scuttling over the Cambrian shorelines to mate and lay their eggs in a similar manner to horseshoe crabs.

Cambrian Explosion #50: Artiopoda – More Than Just Trilobites

The dominant group of Cambrian euarthropods were the artiopodans, a hugely diverse and long-lasting lineage that included the familiar trilobites along with all their close relatives.

They were some of the first euarthropods to appear in the fossil record, with fully formed trilobites seeming to “suddenly” appear about 521 million years ago and quickly spread worldwide. With the ancestral euarthropods estimated to have arisen between 550 and 540 million years ago, and the ancestral artiopodans not long after that, this means there must have been a lot of very rapid evolution and diversification in the space of just 20-30 million years.

Artiopodans were generally seafloor-crawling animals with flattened bodies and wide flaring segments in a trilobite-like shape. Different species could range from about 1mm (0.04″) to around 70cm long (2’4″) – with the largest Cambrian forms reaching as much as 55cm (1’10”), rivalling some of the bigger radiodonts in size.

Unsolved Paleo Mysteries Month #06 – Tricky Trilobites

Trilobites are common and recognizable fossils, found around the world from the Early Cambrian to the Late Permian (521-250 mya), and ranging in size from 1mm to 72cm (0.03″ – 2′4″). They were some of the first organisms on Earth with complex eyes, and some groups also developed ornamentation like spines, horns, and tridents. The image above depicts a particularly elaborate genus known as Dicranurus.

Occasionally fossils have been found showing fine details of trilobite anatomy like antennae, legs, gills, and digestive organs, and we’ve even recently discovered their eggs.

And yet we don’t really know where they came from. Much like the pterosaurs we started the month off with, trilobites appear suddenly in the fossil record with no intermediate or ancestral forms to definitively link them to other groups. We know they were definitely arthropods, but which arthropods they were most closely related to is still uncertain.

They might be related to the chelicerates (arachnids, horseshoe crabs, and eurypterids), or they might be part of the mandibulates (crustaceans, insects, and myriapods). But the exact relationships of these major arthropod groups are still in dispute, too, and phylogenetic results can vary wildly depending on whether trilobites are included in the analysis or not.

It’s probably going to be some time before any sort of consensus is reached.