Behavioral stereotypy and some ecological consequences of entrance-shaft placement of the domichnium Sanctum laurentiensis in Ordovician trepostomate Bryozoa

Abstract The ichnofossil Sanctum laurentiensis from the Late Ordovician (Cincinnatian) of Laurentia is a domichnium tunneled into live colonies of ramose and frondose trepostome Bryozoa. A preferred entrance location was chosen by the trace-making endoskeletozoan. The organism chose to tunnel in positions near growing tips of branches where one of two conditions prevailed. At stereotypical sites, here considered “determinate”, domicile entrances were made where a trepostome branch was in the growth process of bifurcation; sites considered “indeterminate” were chosen where one blade or branch was impinging on another, or on some other nearby substrate, thus interfering with normal colony growth in a random, or non-ontogenetic, manner. This stereotypy is a rare example of demonstrable entrance-siting behavior by an unknown invertebrate organism. A skeleto-structural condition was present at both determinate and indeterminate locations on ramose trepostome colonies. Stereotypical entrance-site choices included a location where bryozoan freewall membrane was compromised or thinned and exozone was incompletely developed. Sanctum producers preferred incomplete exozone in order to access the endozone for excavation to form a domichnium. Middle and Late Ordovician diversification of ramose bryozoan forms stimulated escalation of bioeroder and symbiont taxa in living zoaria thus encouraging further trepostome evolution.