A non-native bryozoan, Schizoporella errata, forms extensive patches of free-living balls and reef-like structures (bryoliths) on the mudflats in south San Francisco Bay, California. The ball-like bryoliths range from 2 to 20 cm in diameter, and the reef-like structures can be nearly 1 m across. While S. errata is known to form bryoliths in other locations, free-living aggregations like these have not been reported. Colony morphology appears to be a plastic trait as analysis of relationships among forms using cytochrome oxidase subunit I (COI) nucleotide sequence data revealed no genetic separation. We recorded > 50 species of algae and invertebrates living on and in the bryoliths and determined the invasion status for 34 of the 50 species. Of the 34, 25 (74%) were non-natives and included fouling species that require hard substrate. The bryoliths may thus facilitate colonization by invaders on the mudflats and serve as stepping stones between the limited hard substrate habitats in the Bay.
KEY WORDS: Bryoliths · Bryozoans · Schizoporella · Habitat modification · Invasive species · FacilitationResale or republication not permitted without written consent of the publisher Mar Ecol Prog Ser 412: 129-139, 2010 in the Hawaiian Islands, grows attached to the reef but also forms unattached 'tumbleweeds' on sand flats that may aid dispersal through viable fragments (C. J. Zabin pers. obs.).Non-native marine species that create 3-dimensional structures in soft-bottom habitats can also have significant ecological impacts, facilitating the presence of obligate hard-substrate organisms, creating living space for other organisms, and in some cases altering the quality of the nearby benthos (e.g. Posey 1988, Ricciardi et al. 1997, Crooks & Khim 1999, Bruno et al. 2005, Rodriguez 2006. In a literature review, Bruno et al. (2005) found that non-native species facilitated the presence of native species at least as often as that of other non-native species. However, it might be reasoned that when a non-native species creates a novel substrate type, other non-natives may be more likely than natives to either (1) be pre-adapted to use the new substrate or (2) display the environmental plasticity to do so. For example, Heiman et al. (2008) found that reefs created by the Australian tubeworm Ficopomatus enigmaticus in a predominantly soft-bottom estuary in Central California supported abundant communities of non-native amphipods and polychaetes. Native species associated with native oysters also inhabited the worm reefs, but non-native species were much more abundant in the tubeworm reefs than on oysters, suggesting differential facilitation of non-native species by this new foundation species (Heiman et al. 2008).In 2006, we became aware of another organism typically thought of as hard-substrate dependent, living as an unattached form on the mudflats in San Francisco Bay, California. Spheroidal colonies of a bryozoan, ranging from golf ball to football size (~2 to 20 cm), were found in large numbers in the shallow subt...