Three-dimensional kinematics of euchelicerate limbs uncover functional specialization in eurypterid appendages

Sea scorpions (Euchelicerata: Eurypterida) explored extreme limits of the aquatic euchelicerate body plan, such that the group contains the largest known marine euarthropods. Inferences on eurypterid life modes, in particular walking and eating, are commonly made by comparing the group with horseshoe crabs (Euchelicerata: Xiphosura). However, no models have been presented to test these hypotheses. Here, we reconstruct prosomal appendages of two exceptionally well-preserved eurypterids, Eurypterus tetragonophthalmus and Pentecopterus decorahensis, and model the flexure and extension of these appendages kinematically in three dimensions (3D). We compare these models with 3D kinematic models of Limulus polyphemus prosomal appendages. This comparison highlights that the examined eurypterid prosomal appendages could not have moved prey items effectively to the gnathal edges and would therefore not have emulated the motion of an L. polyphemus walking leg. It seems that these eurypterid appendages were used primarily to walk or grab prey, and other appendages would have moved prey for mastication. Such 3D kinematic modelling highlights how eurypterid appendage morphologies placed substantial limits on their function, suggesting a high degree of specialization, especially when compared with horseshoe crabs. Such three-dimensional kinematic modelling of these extinct groups therefore presents an innovative approach to understanding the position of these animals within their respective palaeoecosystems.