Xiphosurid from the Tournaisian (Carboniferous) of Scotland confirms deep origin of Limuloidea

Horseshoe crabs are archetypal marine chelicerates with a fossil record extending from the Lower Ordovician to today. The major horseshoe crab groups are thought to have arisen in the middle to late Palaeozoic. Here we present the oldest known limuloid from the lower Carboniferous (Tournaisian stage, c. 350 million years ago) of Scotland: Albalimulus bottoni gen. et sp. nov. A comprehensive phylogenetic analysis supports the placement of A. bottoni as a representative of the extant family Limulidae and 100 million years older than any other limulid taxon. The use of geometric morphometric analyses corroborate the erection of the new taxon and illustrates the exploitation of morphospace by xiphosurids. This new taxon highlights the complex evolutionary history of xiphosurids and the importance of documenting these unique Palaeozoic individuals.