#mandibulata

Waukartus muscularis was a small marine arthropod that lived during the early Silurian, about 437 million years ago, in an equatorial inland sea covering what is now Wisconsin, USA.

It was a member of the myriapods, related to modern centipedes and millipedes – but it represents a very early offshoot of this lineage, with its ancestors branching off sometime before the amphibious euthycarcinoids.

Growing up to about 3cm long (~1.2"), Waukartus had a head with four pairs of small appendages and what may be a pair of small stalked eyes, eleven body segments each with one pair of legs, and a telson with a pair of blade-like projections.

It appears to have been fully aquatic, but its unbranched limbs closely resemble those of terrestrial myriapods, suggesting that these arthropods initially evolved their walking legs for use on the seafloor and only later exapted them for land.

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Tanzanian blue ringleg centipede (Scolopendra morsitans), juvenile (top) and adult (bottom)

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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.

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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.

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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.

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