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.
…But this group is also one of the biggest wastebaskets in paleontology.
First established in the early 20th century, the ptychopariids seemed to have some fairly good defining characteristics based on their facial sutures, large thoraxes, and relatively small pygidia. But the group quickly became a dumping ground for a massive amount of Cambrian trilobites, eventually containing numerous different families, hundreds of genera, and many more individual species.
Actually figuring out their internal evolutionary relationships also turned out to be extremely difficult – so much so that some paleontologists working on them just gave up trying and arranged the genera names alphabetically instead!
Even cladistic studies from the 1970s onward struggled to make sense of these highly “problematic” trilobites, and any larger-scale analysis was a daunting task due to how huge and diverse the ptychopariid wastebasket had become over the years. Worse, some of the anatomical features the group had been based around were starting to look more like the result of a lot of convergent evolution across multiple lineages than any actual shared ancestry.
Efforts were still made at breaking up the mess, however, with better-understood sub-groups like the Proetida, Harpida, Asaphida, Trinucleida, and Olenida being gradually split off into their own separate orders over the course of the last few decades.
But even by the early 2010s what remained of the Ptychopariida was still paraphyletic at best, more of an “evolutionary grade” of early trilobites than a single lineage, with most of its constituent families also rather poorly defined. There was even a proposal to abandon the group entirely, stating that “it serves no scientific purpose” and that its orphaned contents should be considered “order uncertain” until their actual relationships can be untangled.
Today the “ptychopariids” are in dire need of a full revision – since they were the ancestors of many other major groups they’re still crucial for understanding early trilobite evolution. There may be a salvageable single lineage somewhere in the remains of this wastebasket, even if it’s restricted to just close relatives of the genus Ptychoparia, but until somebody tackles them properly they’re stuck in taxonomic limbo with their name only being used in a loose sense.
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”.
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.
Trilobites were one of the most successful groups of early animals, existing for over 300 million years – and during that time they developed a huge diversity of weird heads, with various arrangements of spines, horns, eyestalks, and even long snouts and tridents.
But perhaps one of the oddest was the genus Odontocephalus, known mainly from the early-to-mid Devonian and represented here by Odontocephalus aegeria.
Living about 390 million years ago in northeast North America, this trilobite grew up to around 9cm long (3.5″). And although it wasn’t overall very elaborately ornamented, the front margin of its head had a row of extensions that flared out to meet at their tips, forming something resembling the cowcatchers used on trains.
The actual function of this structure is unknown. It might have been purely used for visual display since trilobites had excellent vision – but Odontocephalus was also a fast-moving bottom-dweller, and its “cowcatcher” may have served the same sort of purpose as its modern equivalent, deflecting small obstacles in its path as it trundled along the seabed.
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.