Horseshoe crabs are famous examples of “living fossils“, having changed their external appearance very little over hundreds of millions of years. But some fossil species were much more varied in shape than their morphologically conservative modern relatives, such as Austrolimulus fletcheri here.

Living in freshwater environments in what is now New South Wales, Australia, during the Middle Triassic (~247-242 million years ago), Austrolimulus had incredibly long spines on each side of its head, reaching a span of around 18cm (7″) – wider than its total body length!

The function of these spines is unclear, but they may have acted like a hydrofoil in fast-moving currents, or they may have served a defensive purpose by making Austrolimulus‘ carapace too wide and unwieldy for some predators to deal with.


The synziphosurines were ancient marine chelicerate arthropods that were traditionally thought to be early representatives of the horseshoe crab lineage. But more recent studies have shown them to occupy a slightly more basal position on the chelicerate evolutionary tree, instead being related to the common ancestor of sea scorpions, horseshoe crabs, and arachnids.

(Also horseshoe crabs might actually be part of the arachnids, closely related to ricinuleids!)

Venustulus waukeshaensis was one of the earliest known synziphosurines, living in what is now Wisconsin, USA during the early Silurian, about 436 million years ago. It grew to around 8cm long (~3.2″) and had six pairs of appendages on the underside of its body, with the first pair modified into chelicerae and the rest being walking legs.

It also appears to have been blind, lacking any evidence of eyes despite its fossils being fairly well-preserved – suggesting it lived in conditions where vision wasn’t much use, such as dark murky water or burrowing around in seafloor sediment.


Nicknamed “Santa Claws”, Sanctacaris uncata was a marine arthropod from the Middle Cambrian (~505 mya) Burgess Shale deposits of Canada. Its exact evolutionary relationships are unclear, but it’s thought to have been very closely related to or part of an early branch of the chelicerates – the lineage that includes modern arachnids and horseshoe crabs.

Measuring up to about 9cm long (3.5″), it had forward-facing eyes and five pairs of grasping appendages on the underside of its head, adaptations that suggest it was an active predator convergently similar to anomalocaridids. It probably swam around grabbing onto whatever small prey items it could catch, trapping them in its “limb basket” while it ate them.


Vaderlimulus tricki, a horseshoe crab from the Early Triassic of Idaho, USA (~251-247 mya). Named for its resemblance to the shape of Darth Vader’s helmet, it’s the earliest known Mesozoic horseshoe crab from North America and was closely related to another oddly-shaped form from Australia.

It was much smaller than its modern relatives, only about 10cm long (4″), and probably lived in a brackish estuary environment where seawater and freshwater met.