Segmentation in early Xiphosura and the evolution of the thoracetron

Abstract

Xiphosuran chelicerates, also known as horseshoe crabs, are a long-lived clade characterized by a highly distinctive morphology and are a classic example of supposed evolutionary stasis. One key feature of horseshoe crabs is the fusion of the opisthosomal segments into a single sclerite referred to as a thoracetron. There has been historical uncertainty as to whether the thoracetron originated once or multiple times within the clade. Here we review criteria for determining whether segments are fused and apply them to a broad census of taxa for which their ontogeny is known or the adult status of specimens can be reasonably asserted to explore the evolution of the thoracetron within a developmental framework. Our findings indicate that the thoracetron evolved once in the common ancestor to Xiphosura. However, subsequent independent loss of the thoracetron segment boundaries is identified and shown to be the result of heterochronic processes acting on a shared developmental pathway. The multiple cases of effacement of the thoracetron within Limuloidea are cases of peramorphically driven parallelism, while the effacement of the thoracetron in the pedomorphic Belinurina is a case of convergence. Xiphosurids therefore represent an interesting case study for recognizing parallelism and convergence on the same structure within closely related lineages. We also demonstrate that somite VII has been incorporated into the prosoma multiple times within the chelicerate lineage, which has implications for interpreting the ground pattern of the group.