Unexpected reproductive traits of Grateloupia turuturu revealed by its resistance to bleach-based biosecurity protocols

A non-indigenous alga, Grateloupia turuturu (Halymeniales, Rhodophyta), was discovered in the Damariscotta Estuary (Maine, USA) in 2017, over 200 km north of its last reported location. Because of the presence of coastal facilities (aquaculture, marine laboratories) among potential vectors, we evaluated a mandated biosecurity protocol, namely, seawater treated with bleach at 50 ppm (50 mg l(-1)) free chlorine for >= 2 h and measured using test strips, for lethality against G. turuturu and several native algae. We report unexpected resilience to bleach-treatment (Mastocarpus > Grateloupia > Palmaria). Holdfasts of G. turuturu and Mastocarpus stellatus survived 50 and 1000 ppm free chlorine, respectively. Cystocarps on cultured blades of G. turuturu were particularly resistant to bleach-treatment; they produced outgrowths on fragmenting blades that became fertile tetrasporophytes in culture, suggesting reproductive adaptations of cystocarps to stressful conditions that may have broader evolutionary significance. Juvenile, microscopic crusts of G. turuturu were also bleach-resistant if covered by diatoms, and developed upright axes. Free chlorine test strips were inaccurate in seawater, and their use could increase the risk of failed biosecurity as a vector for invasion. Because chlorination at economic and environmentally safe levels is insufficient, we recommend a combination of treatments for comprehensive biosecurity.