Symbiosis, competition, and physical disturbance in the growth histories of Pliocene cheilostome bryoliths

Free-living (unattached) subspherical bryozoan masses (bryoliths) in Pliocene tidal channel deposits of the Imperial Formation of southeastern California show complex intra- and interspecific interactions during their accretionary growth. Ranging up to 10 cm in length, the bryoliths are composed almost exclusively of the anascan cheilostome Biflustra commensale. Approximately 50% of the bryoliths are nucleated on cerithiid or muricoid gastropod shells; secondary occupants (presumably pagurid crabs) determined the subspherical growth and associated epibionts of all of these specimens. Evidence for crab occupation includes the thick and relatively symmetrical bryozoan overgrowths that form short tubes extending from the aperture, thinning and pinchout of laminae on the undersides of bryoliths (wear facets), and the distinctive borings (Helicotaphrichnus) of symbiotic worms. In some instances, shells were infested by bryozoans and other encrusters before death of the gastropod, but these overgrowths are thin relative to hermit-associated bryozoan colonies. Episodic hermit abandonment, indicated by extensive erosion of the bryolith and/or its colonization by a more diverse epibiont assemblage including oysters and serpulids, was more frequent among bryoliths nucleated on the largest and most fouled gastropod shells; it was also more frequent among bryoliths in the relatively high-energy tidal channel thalweg than among those associated with oyster thickets on muddy channel margins. Bryoliths nucleated on other shell substrata are similarly thick, but have more irregular stratigraphies including more sedimentary inclusions, more borings, and fewer encrusting epibionts. Pebbles of crystalline basement rock are also encrusted by B. commensale, but only thinly. All of these bryoliths not inhabited by crabs are limited to the channel thalweg. As many as four distinct colonies of B. commensale could coexist on a single bryolith; lines of competitive standoff between colonies are marked by mineralized walls and topographic ridges on the bryolith exterior, and by teepee-like structures in cross-section. These standoffs were preferred sites of infestation by other epibionts and were remarkably stable in position on bryoliths with continuous hermit occupation. Bryoliths that suffered repeated abandonment by hermits, or that depended entirely upon chance reorientation, are characterized by highly unstable standoff positions, reflecting scramble competition under less predictable conditions. These circumstances were most common among large bryoliths and among those in channel thalwegs.