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Kelp forest declines have been linked to warming ocean temperatures worldwide. Ocean warming rarely occurs in isolation, so multiple stressor studies are necessary to understand the physiological responses of kelp to climate change. The canopy-forming bull kelp, Nereocystis luetkeana, is going locally extinct in areas of the Salish Sea that are seasonally warm and nutrient poor, while the understory kelp, Saccharina latissima, persists at those sites. Further, nitrogen availability can alter physiological responses of kelps to temperature stress, including alleviating warming stress. We compared the physiological responses of kelp sporophytes to high temperature stress and nitrogen limitation between two populations of N. luetkeana with different environmental histories (warm and nutrient poor vs. cold and nutrient rich) and between two species, N. luetkeana and S. latissima. Using laboratory mesocosms, we tested the interactive effects of short term (8-9 day) exposure of kelp blades to different temperatures: low (9, 13°C), moderate (15, 16°C), and warm (21°C) at two different nitrogen concentrations: low (1-3 μM) vs. high (>10 μM). We examined a wide array of physiological responses: blade growth, photosynthesis, respiration, photosynthetic yield, nutrient uptake, and tissue C:N. Both kelp species responded negatively to elevated temperatures, but not to low nitrogen levels. Blades of both species showed signs of metabolic stress and reduced growth in the warmest temperature treatment (21°C), at both high and low nitrogen levels, suggesting that N. luetkeana and S. latissima are susceptible to thermal stress over short time periods. Populations of N. luetkeana from warm, nutrient poor and cool, nutrient rich areas were equally susceptible to the effects of ocean warming. Our results suggest that nutrient additions may actually reduce kelp performance at supra-optimal temperatures, and a thorough understanding of kelp responses to coastal temperature and nutrient dynamics is needed to guide conservation and restoration actions. 

Warming ocean temperatures have been linked to kelp forest declines worldwide, and elevated temperatures can act synergistically with other local stressors to exacerbate kelp loss. The bull kelp Nereocystis luetkeana is the primary canopy‐forming kelp species in the Salish Sea, where it is declining in areas with elevated summer water temperatures and low nutrient concentrations. To determine the interactive effects of these two stressors on microscopic stages of N. luetkeana, we cultured gametophytes and microscopic sporophytes from seven different Salish Sea populations across seven different temperatures (10–22°C) and two nitrogen concentrations. The thermal tolerance of microscopic gametophytes and sporophytes was similar across populations, and high temperatures were more stressful than low nitrogen levels. Additional nitrogen did not improve gametophyte or sporophyte survival at high temperatures. Gametophyte densities were highest between 10 and 16°C and declined sharply at 18°C, and temperatures of 20 and 22°C were lethal. The window for successful sporophyte production was narrower, peaking at 10–14°C. Across all populations, the warmest temperature at which sporophytes were produced was 16 or 18°C, but sporophyte densities were 78% lower at 16°C and 95% lower at 18°C compared to cooler temperatures. In the field, bottom temperatures revealed that the thermal limits of gametophyte growth (18°C) and sporophyte production (16–18°C) were reached during the summer at multiple sites. Prolonged exposure of bull kelp gametophytes to temperatures of 16°C and above could limit reproduction, and therefore recruitment, of adult kelp sporophytes. 

Abstract Susan Lynn Williams (1951–2018) was an exceptional marine ecologist whose research focused broadly on the ecology of benthic nearshore environments dominated by seagrasses, seaweeds, and coral reefs. She took an empirical approach founded in techniques of physiological ecology. Susan was committed to applying her research results to ocean management through outreach to decision-makers and resource managers. Susan’s career included research throughout the USA in tropical, temperate, and polar regions, but she specialized in tropical marine ecology. Susan’s scholarship, leadership, and friendship touched many people, leading to this multi-authored paper. Susan’s scholarship was multi-faceted, and she excelled in scientific discovery, integration of scientific results, application of science for conservation, and teaching, especially as a mentor to undergraduate and graduate students and postdoctoral scholars. Susan served in a variety of leadership positions throughout her career. She embodied all facets of leadership; leading by example, listening to others, committing to the “long haul,” maintaining trust, and creating a platform for all to shine. Susan was an important role model for women in science. Susan was also a loyal friend, maintaining friendships for many decades. Susan loved cooking and entertaining with friends. This paper provides an overview of the accomplishments of Susan in the broad categories of scholarship, leadership, and friendship.