Community assembly in a modular organism: the impact of environmental filtering on bryozoan colony form and polymorphism

Understanding community assembly is a key goal in community ecology. Environmental filtering influences community assembly by excluding ill-adapted species, resulting in communities with similar functional traits. An RLQ (a four-way ordination) analysis incorporating spatial data was run on a data set of 642 species of cheilostomes (Bryozoa) from 779 New Zealand sites, and results were compared to trends in other sessile, epibenthic taxa. This revealed environmental filtering of colony form: encrusting-cemented taxa were predominant in shallow environments with hard substrata (<200 m), while erect-rooted taxa characterized deeper environments with soft substrata (>200 m). Furthermore, erect taxa found in shallow environments with high current speeds were typically jointed. Polymorphism also followed environmental gradients. External ovicells (brood chambers) were more common in deeper, low-oxygen water than immersed and internal ovicells. This may reflect the oxygen needs of the embryo or increased predation intensity in shallow environments. Bryozoans with costae tended to be found in deeper water as well, while bryozoans with calcified frontal shields were found in shallow environments with a higher concentration of CaCO₃ . Avicularia did not appear to be related to environmental conditions, and changes in pivot bar structure with depth likely represent a phylogenetic signal. The importance of substratum type as a strict environmental filter suggests that anchoring structures, like rootlets, may be "key innovations" for other sessile, epibenthic taxa like sponges and ascidians.