Statistical analysis of mixed-motive shell borings in Ordovician, Silurian, and Devonian brachiopods from northern and eastern Canada

Hundreds of shell borings of different origin and displaying variable patterns were found in strophomenide, pentameride, and orthide brachiopods of five Paleozoic localities in northern and eastern Canada. The borings were analyzed using simple statistics as well as cluster and nonmetric multidimensional scaling analyses. At the Ordovician Anticosti Island locality, all borings are parasitic or post-mortem in origin, while at the Wenlock-Ludlow Baillie-Hamilton Island, almost all borings are predatory. At the remaining three localities, borings represent a mix of predatory, parasitic, and post-mortem domichnial borings in all three brachiopod taxa, the proportions of which were controlled largely by brachiopod shell morphology and paleoecology. For the strophomenides, predatory borings can be segregated from parasitic and post-mortem domichnial using simple and multivariate statistical analyses. Sowerbyella-type strophomenides have a higher proportion of predatory borings at the Lochkovian localities than at the Ordovician localities, while the reverse is true for the Strophomena-type strophomenides. In pentamerides and orthides, very few predatory borings are identified; most borings were emplaced by parasitic or post-mortem domichnial borers. In pentamerides, this is due to the internal structure of the shells, which elevated the muscles of the organism above the shell floor, rendering them inaccessible to boring predators. In orthides, more deliberate defense mechanisms such as toxins or external ornamentation may have deterred predation. This study indicates that throughout the Paleozoic interactions between borers and brachiopods were complex but can be elucidated using a large sample size and statistical analyses.