Overgrowth competition between clades: implications for interpretation of the fossil record and overgrowth indices

Overgrowth interactions (2693 in total) were observed among three major groups (arguably clades) of bryozoans--cheilostomatids (57 species), ctenostomatids (3 species), and cyclostomatids (14 species). The bryozoans studied here occur in shallow water at high-temperate polar latitudes where they encrust hard substrata such as rock piles. The main study site was the intertidal and infralittoral zones of Kodiak Island, Alaska, but observations were also made in similar zones of South Georgia Island and the Falkland Islands in the South Atlantic Ocean. Cheilostomatids dominated the number of species, individuals, and interactions at all depths. Intraclade interactions formed 73.7% of the encounters for cheilostomatids, 1.6% for ctenostomatids, and 5.7% for cyclostomatids. The competitive ranking of the three clades was broadly ctenostomatids > cyclostomatids > cheilostomatids. Significantly, these results contradict all previous quantitative studies of bryozoan overgrowth, in which cheilostomatids are reported to overgrow cyclostomatids at a higher rate. From these studies and the literature, we calculated win indices to vary from 0 to 0.42 for living cyclostomatids, from 0.08 to 0.9 for living cheilostomatids, and from 0.25 to 0.75 for living ctenostomatids. The win indices of cyclostomatid and cheilostomatid clades show significantly more variation in living assemblages than in fossil assemblages. This disparity may be due to differential preservation (polar and subpolar assemblages last less than 4 years). The diversity was very high in terms of both species richness and interaction types (outcomes between competitor pairs). Comparison with the literature suggests the possibility that nearshore diversity of bryozoans may be bimodal (have two peaks) between high arctic and antarctic latitudes. Indices of success in overgrowth competition have been constructed in various ways. For cheilostomatids, the method of calculation had little influence on the ranking of representatives. In contrast, the apparent success of ctenostomatids and cyclostomatids varied hugely with how the index was calculated. This inconsistency is due to the use of very different strategies in overgrowth competition; among the two latter groups, many interactions involve tied outcomes.