Epibiont communities: Recruitment and competition on North American Carboniferous brachiopods

Abstract

Shell-encrusting assemblages have uniquely high ecological fidelity among paleocommunities because adjacent epibionts coexisted in ecological time, and the abundances and spatial relationships of the various members are accurately preserved. This study presents a quantitative reconstruction of communities on brachiopod shells (Composita and alate spirifers) from the Carboniferous of North America. Thirty-one epibiont communities, comprised of a total of 1180 encrusted brachiopods, are analyzed in terms of epibiont abundance (e.g., percent areal cover), ecological diversity (e.g., Shannon-Wiener Index), and the role of larval recruitment and spatial competition in structuring the communities.Mississippian epibiont communities contain from seven to 14 encrusting taxa that occupy up to 25 percent of host shell surfaces. These communities are spatially dominated by trepostome bryozoans, but also include common lower Paleozoic epibionts, such as encrusting ctenostome bryozoans, hederellids, cornulitids, and edrioasteroids. Pennsylvanian epibiont communities contain between eight and 16 epizoan taxa that occupy up to 34 percent of shell surfaces. They are dominated by fistuliporoid or trepostome bryozoans and increasing proportions of erect bryozoans (e.g., fenestellids), encrusting foraminiferans, and articulate brachiopods.Carboniferous epibionts, like their modern counterparts, exist in loosely organized ecological communities that vary widely in abundance and composition in space and geological time. There is little evidence for tight controls on community structure. Much of the variation is attributed to vagaries in the magnitude and selective nature of larval recruitment. Evidence of competition between community members is rare, except where a high percentage of the shell surface is covered. In these cases, sheetlike bryozoans are usually most abundant. Solitary epibionts are surprisingly successful competitors in one-on-one encounters, but apparently lose areal cover to longer-lived colonial encrusters during ecological succession.