Physical habitat attribute mediates biotic resistance to non-indigenous species invasion

A soft-shelled non-indigenous clam, Nuttallia obscurata, has invaded coastal soft-sediment habitats of the northeastern Pacific. In a survey of 35 sites within the San Juan Islands, Washington, USA, Nuttallia was found almost exclusively in sandy substrates, higher in the intertidal than most native clams (>1 m above mean lower low water). The distinctive distribution of Nuttallia suggested that tidal height and sediment composition may be important physical factors that control its refuge availability, regulating its exposure to predation and ultimately the success of its invasion. I tethered Nuttallia for 24 h in the high intertidal where it is typically found and in the low intertidal at an elevation where it was never found. Clams restrained to the surface suffered high mortality from crab predation at both tidal heights, whereas control clams with unrestricted burrowing movement exhibited high mortality rates only in the low intertidal. In a second experiment, I transplanted sediment within and between the two intertidal heights to measure effects of tidal height and sediment type on survival and burial depth of Nuttallia. At both tidal heights all clams placed on mud-cobble substrate, naturally common in the low intertidal, suffered high mortality rates (>60% in 24 h). Nuttallia on loosely packed sand substrate, naturally found in the upper intertidal, survived much better, however, because they buried deeper than in the tightly packed mud. Caged control clams at both tidal heights suffered no mortality. Apparently native predators are mitigating community level impacts of an invader by excluding Nuttallia completely from some beaches with improper sediment characteristics or relegating it in others to a zone not often inhabited by native species, thereby reducing potential competitive interactions. These findings show that a physical habitat characteristic can mediate biotic resistance to an invader and thus control invasion success and community-level impacts. Generally, such physical-biological interactions may explain some of the reported site-to-site variability in invasion success, as well as the patchy distribution of many soft-sediment infaunal species.