Unraveling the Gordian Knot: Eight testable hypotheses on the effects of nutrient enrichment on tidal wetland sustainability.

The position of tidal wetlands at the land-sea interface makes them especially vulnerable to the effects of nutrient discharges and sea level rise (SLR). Experimental studies of coastal wetland nutrient additions report conflicting results among and within habitats, highlighting the importance of site-specific factors, and how spatial and temporal scaling modulates responses. This suite of influences as SLR accelerates creates a "Gordian Knot" that may compromise coastal habitat integrity. We present eight testable hypotheses here to loosen this knot by identifying critical modulators about nutrient form, soil type and porosity, physiochemical gradients, and eco-evolutionary responses that may control the impacts of nutrient enrichment on coastal wetland sustainability: (1) the delivery and form of the nutrient shapes the ecosystem response; (2) soil type mediates the effects of nutrient enrichment on marshes; (3) belowground responses cannot be solely explained by phenotypic responses; (4) shifting zones of redox and salinity gradients modulate nutrient enrichment impacts; (5) eco-evolutionary processes can drive responses to nutrient availability; (6) nutrient enrichment leads to multiple changed ecosystem states; (7) biogeography trumps a plant's plastic responses to nutrient enrichment; and, (8) nutrient-enhanced wetlands are more susceptible to additional (and anticipated) anthropogenic changes. They provide a framework to investigate and integrate the urgently needed research to understand how excess nutrients threaten the sustainability of coastal wetlands, and wetlands in general. While there is no single 'right way' to test these hypotheses, including a combination of complex and simple, highly-replicated experiments is essential.