mandibulata – 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.

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.

Antarcticarcinus

Euthycarcinoids were a group of arthropods that lived between the mid-Cambrian and the mid-Triassic – but despite existing for over 250 million years their fossil record is incredibly sparse, and it’s only within the last decade that they’ve been recognized as being close relatives of modern centipedes and millipedes.

The earliest members of this group were marine, living in shallow tidal waters, but they quickly specialized into brackish and freshwater habitats and were even some of the very first animals to walk on land. Fossil trackways show they were amphibious, venturing out onto mudflats to feed on microbial mats, avoid aquatic predators, and possibly lay their eggs in a similar manner to modern horseshoe crabs.

Most euthycarcinoid species are known from tropical and subtropical climates, but Antarcticarcinus pagoda here hints that these arthropods were much more widespread and diverse than previously thought. Discovered in fossil deposits in the Central Transantarctic Mountains of Antarctica, it lived in freshwater lakes during the Early Permian (~299-293 million years ago), at a time when the region was in similar polar latitudes to today with a cold icy subarctic climate.

About 8.5cm long (3.3″), it would have had a similar three-part body plan to other euthycarcinoids – with a head, a limb-bearing thorax, and a limbless abdomen ending in a tail spine – but its most distinctive feature was a pair of large wing-shaped projections on the sides of its carapace. These may have helped to stabilize its body when resting on soft muddy surfaces, spreading out its weight, or they might even have functioned as a hydrofoil generating lift while swimming.

Cambrian Explosion #61: Crustacea – Little Wigglers

We’re finally at the end of this series, and to finish off let’s look at one of the few types of Cambrian true crustaceans that are known only from fully mature adults: the skaracarids.

These tiny soft-bodied meiofaunal animals are known from late Cambrian areas of “Orsten-type preservation” in Sweden and South China, with a possible additional fragmentary occurrence in Poland – suggesting that they had a global distribution.

Cambrian Explosion #60: Crustacea – Larvae Larvae Everywhere

One of the characteristic features of the crustacean lineage are their larval forms, passing through various tiny larval stages. They often look nothing like their eventual adult forms and historically weren’t even recognized as being the same species, with their complex lifecycles not being properly recognized until the late 1800s.

A lot of Cambrian crustaceans are only known from their larvae, preserved in exquisite microscopic detail in sites of “Orsten-type preservation”. Only disarticulated fragments of larger-bodied forms have been found in a few places, and it isn’t until much later in the Paleozoic that fossil crustaceans actually seem to become abundant in marine ecosystems.

It’s not clear why there’s such a bias in their early fossil record compared to most other arthropods, but possibly they were just very very rare animals early on. Adult forms may have mostly lived in places where they just didn’t fossilize, while their tiny larvae sometimes dispersed into different environments with a better chance of preservation.

Cambrian Explosion #59: Stem-Crustacea – Actual Ancient Aliens & Bivalved Buddies

The majority of known fossils of Cambrian crustaceans are in the form of minuscule microfossils with “Orsten-type preservation” – formed in oxygen-poor seafloor mud and exceptionally well-preserved in three-dimensional detail. They can only be discovered and studied after dissolving away the rock around them with acid and picking through the residue under a microscope, then they’re scanned with an electron microscope to see their fine details.

And it turns out some of these tiny early crustaceans looked really weird.

Cambrian Explosion #58: Hymenocarina

The pancrustaceans are a grouping of mandibulates that contains all of the crustaceans and hexapods (insects and their closest relatives) along with their various stem-relatives.

They’re critical components of most ecosystems on the planet, and are major parts of the nutrient cycle. In aquatic environments the crustaceans dominate, with modern copepods and krill being some of the most abundant living animals and making up enormous amounts of biomass providing vital food sources for larger animals. On the land springtails and ants are especially numerous, and the air is full of flying insects, the only invertebrates to ever develop powered flight. Some groups of insects have also co-evolved complex mutualistic partnerships with flowering plants and fungi.

Hexapods and insects don’t appear in the fossil record until the early Devonian, but they’re estimated to have first diverged from the crustaceans* in the early Silurian (~440 million years ago), around the same time that vascular plants were colonizing the land.

(* Hexapods are crustaceans in the same sort of way that birds are dinosaurs. They originated from within one of the major crustacean lineages with their closest living relatives possibly being the enigmatic remipedes.)

But crustaceans and their pancrustacean ancestors go back much further into the Cambrian, and we’ll be finishing off this month and this series with some of those early representatives.

Cambrian Explosion #57: Tuzoiida

What were tuzoiids?

We don’t know.*

Despite hundreds of specimens having been found, and around 20 different species being described, these arthropods are an ongoing puzzle.

They’re known from between about 518 and 505 million years ago, in deposits associated with tropical and subtropical regions all around the world. They had large spiny bivalved carapaces up to 18cm long (7″), shaped like an upside-down domed taco shell, with a distinctive reticulated net-like surface ornamentation – but the rest of their ecology and anatomy is very unclear.

Most fossils are just empty carapaces, which appear to have been made of unmineralized chitin. Rare examples of soft-part preservation show they had a pair of stalked eyes sticking out the front, and a pair of short simple antennae, but impressions of the rest of their bodies are fragmentary and indistinct enough to not be particularly helpful.

Cambrian Explosion #56: Euthycarcinoidea

The euthycarcinoids were a group of euarthropods known from the mid-Cambrian to the mid-Triassic (~500-254 million years ago), surviving through multiple mass extinctions including the devastating “Great Dying” at the end of the Permian that finished off the trilobites. But despite an evolutionary history spanning around 250 million years they have a very sparse fossil record, extremely rare and known from less than 20 species across their entire time range.

For a long time their affinities were uncertain, and they’ve been variously suggested to have been crustaceans, trilobites, or chelicerates, or even to have been a lineage of earlier stem-euarthropods. But since the early 2010s better understanding of their anatomy has placed them in the mandibulates, probably as the closest relatives of the myriapods and helping to close the gap between the aquatic ancestors of that group and their earliest known terrestrial forms.

Cambrian Explosion #55: Fuxianhuiida

In the final stretch of this month we finally come to the last of the major groupings of euarthropods: the mandibulates, which include the modern myriapods (centipedes and millipedes) and pancrustaceans (crustaceans and insects), along with several extinct groups.

Characterized by possessing mandible mouthparts, mandibulates are by far the largest lineage of arthropods and the most successful group of animals on Earth. Over a million living species are known (most of of which are insects, particularly beetles) and an estimated six-to-ten times more than that are probably still undiscovered.

Mandibulates probably diverged from their chelicerate cousins around the start of the Cambrian 540 million years ago. If the trilobites and their artiopodan relatives were early or stem-mandibulates then the earliest known fossils of the group are about 521 million years old, otherwise the first records come from a few million years later in the Chinese Chengjiang fossil deposits (~518 million years ago).