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AbstractMarine heatwaves are extreme events that can have profound and lasting impacts on marine species. Field observations have shown seaweeds to be highly susceptible to marine heatwaves, but the physiological drivers of this susceptibility are poorly understood. Furthermore, the effects of marine heatwaves in conjunction with ocean warming and acidification are yet to be investigated. To address this knowledge gap, we conducted a laboratory culture experiment in which we tested the growth and physiological responses of Phyllospora comosa juveniles from the southern extent of its range (43–31°S) to marine heatwaves, ocean warming and acidification. We used a ‘collapsed factorial design’ in which marine heatwaves were superimposed on current (today's pH and temperature) and future (pH and temperature projected by 2100) ocean conditions. Responses were tested both during the heatwaves, and after a 7‐day recovery period. Heatwaves reduced net photosynthetic rates in both current and future conditions, while respiration rates were elevated under heatwaves in the current conditions only. Following the recovery period, there was little evidence of heatwaves having lasting negative effects on growth, photosynthesis or respiration. Exposure to heatwaves, future ocean conditions or both caused an increase in the degree of saturation of fatty acids. This adjustment may have counteracted negative effects of elevated temperatures by decreasing membrane fluidity, which increases at higher temperatures. Furthermore, P. comosa appeared to down‐regulate the energetically expensive carbon dioxide concentrating mechanism in the future conditions with a reduction in δ13C values detected in these treatments. Any saved energy arising from this down‐regulation was not invested in growth and was likely invested in the adjustment of fatty acid composition. This adjustment is a mechanism by which P. comosa and other seaweeds may tolerate the negative effects of ocean warming and marine heatwaves through benefits arising from ocean acidification.

AbstractDissolved organic carbon (DOC) released by macroalgae is an intrinsic component of the coastal ocean carbon cycle, yet knowledge of how future ocean warming may influence this is limited. Temperature is one of the primary abiotic regulators of macroalgal physiology, but there is minimal understanding of how it influences the magnitude and mechanisms of DOC release. To investigate this, we examined the effect of a range of temperatures on DOC release rates and physiological traits of Ecklonia radiata, the most abundant and widespread kelp in Australia that represents a potentially significant contribution to coastal ocean carbon cycling. Juvenile sporophytes were incubated at eight temperatures (4–28°C) for 14 days, after which time, DOC concentrations and physiological traits (growth, photosynthesis, respiration, Fv/Fm, photosynthetic pigment content, and carbon, and nitrogen content) were analyzed using thermal performance curves (TPCs) or regression analyses. Thermal optima were 15.63°C for growth and 25.84°C for photosynthesis, highlighting vulnerability to future ocean warming. Dissolved organic carbon concentrations increased when the temperature was above ~22°C, being greatest at the highest temperature tested (28°C), which was likely driven by photosynthetic overflow and thermal stress. Mean Fv/Fm, total chlorophyll, and total fucoxanthin content were lowest at 28°C. The C:N ratio of blades increased linearly with temperature from 23.9 ± 1.30 at 4°C to 33.0 ± 1.22 at 28°C. We demonstrate increased DOC release by E. radiata under elevated seawater temperatures and discuss potential implications for coastal carbon cycling under future ocean warming given the complex and uncertain fate of macroalgal DOC in the marine environment.

Kelp forests are threatened by ocean warming, yet effects of co-occurring drivers such as CO 2 are rarely considered when predicting their performance in the future. In Australia, the kelp Ecklonia radiata forms extensive forests across seawater temperatures of approximately 7–26°C. Cool-edge populations are typically considered more thermally tolerant than their warm-edge counterparts but this ignores the possibility of local adaptation. Moreover, it is unknown whether elevated CO 2 can mitigate negative effects of warming. To identify whether elevated CO 2 could improve thermal performance of a cool-edge population of E. radiata , we constructed thermal performance curves for growth and photosynthesis, under both current and elevated CO 2 (approx. 400 and 1000 µatm). We then modelled annual performance under warming scenarios to highlight thermal susceptibility. Elevated CO 2 had minimal effect on growth but increased photosynthesis around the thermal optimum. Thermal optima were approximately 16°C for growth and approximately 18°C for photosynthesis, and modelled performance indicated cool-edge populations may be vulnerable in the future. Our findings demonstrate that elevated CO 2 is unlikely to offset negative effects of ocean warming on the kelp E. radiata and highlight the potential susceptibility of cool-edge populations to ocean warming.

AbstractOcean acidification (OA) is the reduction in seawater pH due to the absorption of human-released CO2 by the world’s oceans. The average surface oceanic pH is predicted to decline by 0.4 units by 2100. However, kelp metabolically modifies seawater pH via photosynthesis and respiration in some temperate coastal systems, resulting in daily pH fluctuations of up to ±0.45 units. It is unknown how these fluctuations in pH influence the growth and physiology of the kelp, or how this might change with OA. In laboratory experiments that mimicked the most extreme pH fluctuations measured within beds of the canopy-forming kelp Ecklonia radiata in Tasmania, the growth and photosynthetic rates of juvenile E. radiata were greater under fluctuating pH (8.4 in the day, 7.8 at night) than in static pH treatments (8.4, 8.1, 7.8). However, pH fluctuations had no effect on growth rates and a negative effect on photosynthesis when the mean pH of each treatment was reduced by 0.3 units. Currently, pH fluctuations have a positive effect on E. radiata but this effect could be reversed in the future under OA, which is likely to impact the future ecological dynamics and productivity of habitats dominated by E. radiata.

AbstractKelps are in global decline due to climate change, which includes ocean warming. To identify vulnerable species, we need to identify their tolerances to increasing temperatures and determine whether tolerances are altered by co‐occurring drivers such as inorganic nutrient levels. This is particularly important for those species with restricted distributions, which may already be experiencing thermal stress. To identify thermal tolerance of the range‐restricted kelp Lessonia corrugata, we conducted a laboratory experiment on juvenile sporophytes to measure performance (growth, photosynthesis) across its thermal range (4–22°C). We determined the upper thermal limit for growth and photosynthesis to be ~22–23°C, with a thermal optimum of ~16°C. To determine if elevated inorganic nitrogen availability could enhance thermal tolerance, we compared the performance of juveniles under low (4.5 μmol · d−1) and high (90 μmol · d−1) nitrate conditions at and above the thermal optimum (16–23.5°C). Nitrate enrichment did not enhance thermal performance at temperatures above the optimum but did lead to elevated growth rates at the thermal optimum. Our results indicate L. corrugata is likely to be extremely susceptible to moderate ocean warming and marine heatwaves. Peak sea surface temperatures during summer in eastern and northeastern Tasmania can reach up to 20–21°C, and climate projections suggest that L. corrugata's thermal limit will be regularly exceeded by 2050 as southeastern Australia is a global ocean‐warming hotspot. By identifying the upper thermal limit of L. corrugata, we have taken a critical step in predicting the future of the species in a warming climate.

AbstractIncreased plant biomass is observed in terrestrial systems due to rising levels of atmospheric CO2, but responses of marine macroalgae to CO2 enrichment are unclear. The 200% increase in CO2 by 2100 is predicted to enhance the productivity of fleshy macroalgae that acquire inorganic carbon solely as CO2 (non‐carbon dioxide‐concentrating mechanism [CCM] species—i.e., species without a carbon dioxide‐concentrating mechanism), whereas those that additionally uptake bicarbonate (CCM species) are predicted to respond neutrally or positively depending on their affinity for bicarbonate. Previous studies, however, show that fleshy macroalgae exhibit a broad variety of responses to CO2 enrichment and the underlying mechanisms are largely unknown. This physiological study compared the responses of a CCM species (Lomentaria australis) with a non‐CCM species (Craspedocarpus ramentaceus) to CO2 enrichment with regards to growth, net photosynthesis, and biochemistry. Contrary to expectations, there was no enrichment effect for the non‐CCM species, whereas the CCM species had a twofold greater growth rate, likely driven by a downregulation of the energetically costly CCM(s). This saved energy was invested into new growth rather than storage lipids and fatty acids. In addition, we conducted a comprehensive literature synthesis to examine the extent to which the growth and photosynthetic responses of fleshy macroalgae to elevated CO2 are related to their carbon acquisition strategies. Findings highlight that the responses of macroalgae to CO2 enrichment cannot be inferred solely from their carbon uptake strategy, and targeted physiological experiments on a wider range of species are needed to better predict responses of macroalgae to future oceanic change.