Invited reviewCategories of architectural designs in trace fossils: A measure of ichnodisparity

Ichnodisparity has been recently introduced as a concept to assess the variability of morphologic plans in biogenic structures, revealing major innovations in body plan, locomotory system and/or behavioral program. Whereas ichnodiversity is measured in terms of the number of ichnotaxa (i.e. ichnogenera or ichnospecies), ichnodisparity is evaluated based on the identification of categories of architectural design. Seventy-nine categories of architectural designs (58 for bioturbation structures and 21 for bioerosion structures), encompassing 523 ichnogenera (417 for bioturbation structures and 106 for bioerosion structures), are defined. They are restricted to invertebrate ichnotaxa, whereas vertebrate trace fossils were not included. Although the scheme is designed to be comprehensive, the proposed categories are necessarily works in progress because of the state of flux in ichnotaxonomy and the need to adjust the definitions of categories according to the scope and scale of the analysis. Although it may be said that the establishment of categories of architectural design is to a certain degree a subjective enterprise, this is not different from ichnotaxonomy because classifying trace fossils from a taxonomic perspective implies observing the morphology of the trace and interpreting it in terms of behavior. The concept of ichnodisparity is free of some of the vagaries involved in ichnotaxonomy. The fact that ichnodiversity and ichnodisparity exhibit different trajectories during the Phanerozoic underscores the importance of adding the latter to the ichnologic toolkit.