Invasion of the southern Gulf of St. Lawrence by the clubbed tunicate ( Styela clava Herdman): Potential mechanisms for invasions of Prince Edward Island estuaries

Abstract

All but one of the nine non-native marine species that established populations in the southern Gulf of St. Lawrence (sGSL) in the past decade initially invaded the sGSL via coastal and estuarine waters of Prince Edward Island (PEI). Almost half of these species are tunicates, and all but one still occur only in PEI. Recent introductions include Styela clava Herdman in 1997, Botryllus schlosseri (Pallas) in 2001, Botrylloides violaceus Oka in 2002, and Ciona intestinalis (Linnaeus) in 2004. The goal of this paper was to investigate which characteristics of PEI estuaries may have resulted in their being more susceptible to tunicate invasions than estuaries elsewhere in the sGSL. At least one genus that recently established viable populations in PEI was previously introduced to the Gulf of St. Lawrence, apparently without establishing permanent populations. This implies that either propagule pressure has increased or environmental factors are more conducive to establishment now than they were previously. The fluctuating resource availability model predicts increased invasibility of environments that experience pulses of resources such as space or nutrients. Intense development of both agriculture and aquaculture in PEI, and high population density compared to other areas of the sGSL, are associated with high and fluctuating estuarine nutrient levels and a large surface area of artificial substrates (mussel socks) that is kept relatively free of competitors, and is replaced regularly. Changes in nutrient loading and the development of aquaculture have also occurred within the past two to three decades. The provision of artificial structure is likely a critical factor in the successful establishment of tunicates in PEI, because natural hard substrates are scarce. Facilitation by green crabs ( Carcinus maenas L.) may be a contributing factor in the spread of Styela. Only one estuary lacking green crabs has an established population of Styela, and at least two known inoculations of Styela into estuaries without green crabs have failed. A likely mechanism for facilitation is the consumption by green crab of the snail Astyris lunata, a known Styela predator.