We use scanning electron microscopy imaging to examine the shell microstructure of fossil and living species in five families of caenogastropods (Strombidae, Volutidae, Olividae, Pseudolividae, and Ancillariidae) to determine whether parallel or convergent evolution is responsible for the development of a unique caenogastropod trait, the extreme parietal callus (EPC). The EPC is defined as a substantial thickening of both the spire callus and the callus on the ventral shell surface such that it covers 50% or more of the surface. Caenogastropods as a whole construct the EPC convergently, using a variety of low-density, poorly organized microstructures that are otherwise uncommon in caenogastropod non-callus shell construction. Within clades, however, we see evidence for parallelism in decreased regulation in both the shell and callus microstructure. Low-density and poorly ordered microstructure—such as used for the EPC—uses less organic scaffolding and is less energetically expensive than normal shell microstructure. This suggests the EPC functions to rapidly and inexpensively increase shell thickness and overall body size. Tests of functional ecology suggest that the EPC might function both to defend against crushing predation through increased body size and dissipation of forces while aiding in shell orientation of highly mobile gastropods. These interpretations hinge on the current phylogenetic placement of caenogastropod families, emphasizing the essential contribution of phylogeny when interpreting homoplasy.

The occupation of new environments by evolutionary lineages is frequently associated with morphological changes. This covariation of ecotype and phenotype is expected due to the process of natural selection, whereby environmental pressures lead to the proliferation of morphological variants that are a better fit for the prevailing abiotic conditions. One primary mechanism by which phenotypic variants are known to arise is through changes in the timing or duration of organismal development resulting in alterations to adult morphology, a process known as heterochrony. While numerous studies have demonstrated heterochronic trends in association with environmental gradients, few have done so within a phylogenetic context. Understanding species interrelationships is necessary to determine whether morphological change is due to heterochronic processes; however, research is hampered by the lack of a quantitative metric with which to assess the degree of heterochronic traits expressed within and among species. Here I present a new metric for quantifying heterochronic change, expressed as a heterochronic weighting, and apply it to xiphosuran chelicerates within a phylogenetic context to reveal concerted independent heterochronic trends. These trends correlate with shifts in environmental occupation from marine to nonmarine habitats, resulting in a macroevolutionary ratchet. Critically, the distribution of heterochronic weightings among species shows evidence of being influenced by both historical, phylogenetic processes and external ecological pressures. Heterochronic weighting proves to be an effective method to quantify heterochronic trends within a phylogenetic framework and is readily applicable to any group of organisms that have well-defined morphological characteristics, ontogenetic information, and resolved internal relationships.