Non-indigenous ascidians in southern California harbors and marinas

Southern California's many large harbors form an important coastal ecosystem, yet they are also a␣major destination for thousands of pleasure craft and cargo vessels that have often traveled great distances. Many groups of marine organisms, including ascidians, have now been documented as undergoing range extensions as a consequence of rapid ship-transport between distant harbors phenomenon. This has resulted in a rapid increase in the rate of introductions of non-indigenous species worldwide, yet these effects of boat traffic remain largely unstudied in southern California. Ascidians are sessile marine filter-feeders, hermaphroditic, and often self-fertilizing; many species are tolerant of a wide range of environmental conditions, can reach sexual maturity in just a few weeks, and have a long breeding season. This paper documents the arrival of 14␣non-indigenous species in southern California harbors␣during this century, 13 of which have persisted:␣four prior to the 1960s (Cionaintestinalis, Styelaclava, S.␣plicata, Botryllusschlosseri), another by 1972 (S.␣canopus, formerly S. partita), and 8 since 1983 [C.␣savignyi, Ascidia zara, Ascidia sp., Polyandrocarpa zorritensis, Symplegma brakenhielmi (formerly S. oceania, and S. reptans, Microcosmus squamiger, and Molgula␣manhattensis)]. We estimate the relative abundance and seasonal fluctuations of both non-indigenous and native ascidians in all harbors in southern California from San Diego to Santa Barbara based upon the historical record, our 35 yr of field notes, and our recent surveys carried out during fall 1994, spring and fall 1995, fall 1996 and spring 1997. Possible points of origin of the exotics and predictions on further U.S. Pacific coast range-extensions are included. The concomitant decline in numbers and species of native ascidians in the harbors of southern California during this century is also reviewed.