Habitat and developmental constraints drove 330 million years of horseshoe crab evolution

Abstract

<p>Records of evolutionary stasis over time are central to uncovering large-scale evolutionary modes, whether by long-term gradual change or via enduring stability punctuated by rapid shifts. The key to this discussion is to identify and examine groups with long fossil records that, ideally, extend to the present day. One group often regarded as the quintessential example of stasis is Xiphosurida, the horseshoe crabs. However, when, how and, particularly, why stasis arose in xiphosurids remain fundamental, but complex, questions. Here, we explore the protracted history of fossil and living xiphosurids and demonstrate two levels of evolutionary stability: developmental stasis since at least the Pennsylvanian and shape stasis since the Late Jurassic. Furthermore, shape and diversity are punctuated by two high-disparity episodes during the Carboniferous and Triassic – transitions that coincide with forays into habitation of marginal environments. In an exception to these general patterns, body size increased gradually over this period and, thus, cannot be described under the same, often-touted, static models of evolution. Therefore, we demonstrate that evolutionary stasis can be modular and fixed within the same group at different periods and in different biological traits, while other traits experience altogether different evolutionary modes. This mosaic in the tempo and mode of evolution is not unique to Xiphosurida but likely reflects variable mechanisms acting on biological traits, for example transitions in life modes, niche occupation and major evolutionary radiations.</p>