Integrated physiological response by four species of Rhodophyta to submarine groundwater discharge reveals complex patterns among closely-related species

Algal physiological ecology on submarine groundwater discharge (SGD) influenced reefs is likely shaped by intermittent, tidally-driven estuarine conditions that occur with SGD fluxes of fresh-to-brackish groundwater from the subterranean estuary to reef ecosystems. SGD is a common inconspicuous feature worldwide on reefs of basaltic high islands and continental margins. Yet, SGD-driven dynamics of algal physiology are not well understood. To understand how invasive species have physiologically outcompeted native species on many SGD-influenced reefs, physiology in tissue water potential (TWP) regulation, photosynthesis, nitrogen storage, and cellular anatomy were measured across a gradient of SGD-influence, for four Rhodophyte species. Compared with non-SGD conditions, SGD was associated with higher TWP, larger medulla cells with thinner walls, and thinner cortical cell walls for two invasives, Gracilaria salicornia and Acanthophora spicifera, higher photosynthetic rates in G. salicornia, greater nitrogen concentration for A. spicifera and G. salicornia, and increased $\delta$ ¹⁵N ratios for A. spicifera, G. salicornia, and native Laurencia dendroidea. Distinct physiological strategies were measured for the two invasive species across the gradient of SGD-influence, and for L. dendroidea and Gracilaria perplexa offshore. This study illuminates species-specific physiological response, and how introduced opportunistic species may outcompete native species under conditions of SGD.