CHELICERATA: EURYPTERIDS, SPIDERS AND HORSESHOE CRABS

Sanctacaris uncata

This lovely is Sanctacaris uncata — a wonderful example of Chelicerata.

We first see them emerge in our ancient oceans in the Middle Cambrian, some 508 million years ago, as the arthropod Sanctacaris uncata (Briggs & Collins, 1988) known from the Glossopleura Zone, Stephen Formation of Mount Stephen in the Burgess Shale, British Columbia, Canada. 

Sanctacaris is proof positive that chelicerates, although rare, were present in the Middle Cambrian sea. Even at this early stage of evolution, Sanctacaris had the number and type of head appendages found in modified form in the eurypterids and xiphosurids, the major Palaeozoic groups that succeeded it. Even more interesting is that Sanctacaris had all the characteristics of later chelicerates except chelicerae — placing this early arthropod in a primitive sister group of all other chelicerates.

An extinct marine creature half a billion years old may sound otherworldly, but you know some of their more well-known marine brethren — sea spiders, the sexy eurypterids, chasmataspidids and horseshoe crabs — and some of their terrestrial cousins — spiders, scorpions, harvestmen, mites and ticks. 

They are grouped together because, like all arthropods, they have a segmented body and segmented limbs and a thick chitinous cuticle called an exoskeleton. Add those characteristics to a body system with two body segments — a cephalothorax and an abdomen. 

Like all arthropods, chelicerates' bodies and appendages are covered with a tough cuticle made mainly of chitin and chemically hardened proteins. 

Since this cannot stretch, the animals must moult to grow. In other words, they grow new but still soft cuticles, then cast off the old one and wait for the new one to harden. 

Until the new cuticle hardens the animals are defenceless and almost immobilized.  This also helps to explain why you find so many cephalons or moulted head shields — or whatever else our good arthropod friends shed and regrow — in the field and far fewer body fossils of the whole animal.

Some chelicerate are predatory animals that patrol the warm waters near thermal vents. They can be found feeding upon other predators and fish. Although the group were originally solely predatory, they have diversified to use all sorts of feeding strategies: predation, parasitism, herbivory, scavenging and dining on bits of decaying organic matter. 

Although harvestmen can digest solid food it is more akin to a mashed pulp by the time they do. The guts of most modern chelicerates are too narrow to digest solid food, instead, they generally liquidize their chosen meal by grinding it with their chelicerae and pedipalps then flooding it with digestive enzymes. 

To conserve water, air-breathing chelicerates excrete waste as solids that are removed from their blood by Malpighian tubules, structures that also evolved independently in insects — another case of convergent evolution.

The evolutionary origins of chelicerates from the early arthropods have been debated for decades. And although there is considerable agreement about the relationships between most chelicerate sub-groups, the inclusion of the Pycnogonida in this taxon has recently been questioned and the exact position of scorpions is still controversial, though they were long considered the most primitive or basal of the arachnids. 

We still have much to explore to sort out their evolutionary origins and placement within the various lineages but we will get there with time.

Image One: Reconstruction of Sanctacaris uncata, a Cambrian Habeliidan arthropod (stem-Chelicerata: Habeliida). by Junnn11 @ni075; Image Two: Chelicerata by Fossil Huntress

Aria C, Caron JB (December 2017). "Mandibulate convergence in an armoured Cambrian stem chelicerate". BMC Evolutionary Biology. 17 (1): 261. doi:10.1186/s12862-017-1088-7. PMC 5738823. PMID 29262772.

Legg DA (December 2014). "Sanctacaris uncata: the oldest chelicerate (Arthropoda)". Die Naturwissenschaften. 101 (12): 1065–73. doi:10.1007/s00114-014-1245-4. PMID 25296691.

Briggs DE, Collins D (August 1988). "A Middle Cambrian chelicerate from Mount Stephen, British Columbia" (PDF). Palaeontology. 31 (3): 779–798. Archived from the original (PDF) on July 16, 2011. Retrieved April 4, 2010.

Briggs DE, Erwin DH, Collier FJ (1995). Fossils of the Burgess Shale. Washington: Smithsonian Institution Press. ISBN 1-56098-659-X. OCLC 231793738.