Description: To understand the thermal plasticity of a coastal foundation species across its latitudinal distribution, we assess physiological responses to high temperature stress in the kelp Laminaria digitata in combination with population genetic characteristics and relate heat resilience to genetic features and phylogeography. We hypothesize that populations from Arctic and cold-temperate locations are less heat resilient than populations from warm distributional edges. Using meristems of natural L. digitata populations from six locations ranging between Kongsfjorden, Spitsbergen (79°N), and Quiberon, France (47°N), we performed a common-garden heat stress experi- ment applying 15°C to 23°C over eight days. We assessed growth, photosynthetic quantum yield, carbon and nitrogen storage, and xanthophyll pigment contents as response traits. Population connectivity and genetic diversity were analyzed with microsatellite markers. Results from the heat stress experiment suggest that the upper temperature limit of L. digitata is nearly identical across its distribution range, but subtle differences in growth and stress responses were revealed for three popu- lations from the species’ ecological range margins. Two populations at the species’ warm distribution limit showed higher temperature tolerance compared to other populations in growth at 19°C and recovery from 21°C (Quiberon, France), and pho- tosynthetic quantum yield and xanthophyll pigment responses at 23°C (Helgoland, Germany). In L. digitata from the northernmost population (Spitsbergen, Norway), quantum yield indicated the highest heat sensitivity. Microsatellite genotyping revealed all sampled populations to be genetically distinct, with a strong hierarchical structure between southern and northern clades. Genetic diversity was lowest in the isolated population of the North Sea island of Helgoland and highest in Roscoff in the English Channel. All together, these results support the hypothesis of moderate local differentiation across L. digitata's European distribution, whereas effects are likely too weak to ameliorate the species’ capacity to withstand ocean warming and marine heatwaves at the southern range edge.

Description: The sugar kelp Saccharina latissima experiences a wide range of environmental conditions along its geographical and vertical distribution range. Temperature and salinity are two critical drivers influencing growth, photosynthesis and biochemical composition. Moreover, interactive effects might modify the results described for single effects. In shallow water coastal systems, exposure to rising temperatures and low salinity are expected as consequence of global warming, increased precipitation and coastal run-off. To understand the acclimation mechanisms of S. latissima to changes in temperature and salinity and their interactions, we performed a mechanistic laboratory experiment in which juvenile sporophytes from Brittany, France were exposed to a combination of three temperatures (0, 8 and 15°C) and two salinity levels (20 and 30 psu (practical salinity units)). After a temperature acclimation of 7 days, sporophytes were exposed to low salinity (20 psu) for a period of 11 days. Growth, and maximal quantum yield of photosystem II (Fv/Fm), pigments, mannitol content and C:N ratio were measured over time. We report for the first time in S. latissima a fivefold increase in the osmolyte mannitol in response to low temperature (0°C) compared to 8 and 15°C that may have ecological and economic implications. Low temperatures significantly affected all parameters, mostly in a negative way. Chlorophyll a, the accessory pigment pool, growth and Fv/Fm were significantly lower at 0°C, while the de-epoxidation state of the xanthophyll cycle was increased at both 0 and 8°C compared to 15°C. Mannitol content and growth decreased with decreased salinity; in contrast, pigment content and Fv/Fm were to a large extent irresponsive to salinity. In comparison to S. latissima originating from an Arctic population, despite some reported differences, this study reveals a remarkably similar impact of temperature and salinity variation, reflecting the large degree of adaptability in this species.

Description: The Arctic climate is changing rapidly. The warming and resultant longer open water periods suggest a potential for expansion of marine vegetation along the vast Arctic coastline. We compiled and reviewed the scattered time series on Arctic marine vegetation and explored trends for macroalgae and eelgrass (Zostera marina). We identified a total of 38 sites, distributed between Arctic coastal regions in Alaska, Canada, Greenland, Iceland, Norway/Svalbard, and Russia, having time series extending into the 21st Century. The majority of these exhibited increase in abundance, productivity or species richness, and/or expansion of geographical distribution limits, several time series showed no significant trend. Only four time series displayed a negative trend, largely due to urchin grazing or increased turbidity. Overall, the observations support with medium confidence (i.e., 5–8 in 10 chance of being correct, adopting the IPCC confidence scale) the prediction that macrophytes are expanding in the Arctic. Species distribution modeling was challenged by limited observations and lack of information on substrate, but suggested a current (2000– 2017) potential pan-Arctic macroalgal distribution area of 820.000 km2 (145.000 km2 intertidal, 675.000 km2 subtidal), representing an increase of about 30% for subtidaland 6% for intertidal macroalgae since 1940–1950, and associated polar migration rates averaging 18–23 km decade−1 . Adjusting the potential macroalgal distribution area by the fraction of shores represented by cliffs halves the estimate (412,634 km2 ). Warming and reduced sea ice cover along the Arctic coastlines are expected to stimulate further expansion of marine vegetation from boreal latitudes. The changes likely affect the functioning of coastal Arctic ecosystems because of the vegetation’s roles as habitat, and for carbon and nutrient cycling and storage. We encourage apan-Arctic science- and management agenda to incorporate marine vegetation into a coherent understanding of Arctic changes by quantifying distribution and status beyond the scattered studies now available to develop sustainable management strategies for these important ecosystems.

Description: In recent years, kelp populations worldwide have faced decline and extirpation at their equatorward limits, while models predict a poleward shift of kelp ecosystems during climate change. To gain an understanding of local thermal adaptation and response plasticity in a forest-forming kelp species, we assessed populations of Laminaria digitata along its entire European distribution range for their capacity to withstand high temperature stress, and analysed population structure and diversity with microsatellite markers (n=12). We sampled wild meristematic L. digitata material (n=30) at six locations ranging from Kongsfjorden, Spitsbergen, to the southernmost distribution limit in Quiberon, France. In a heatwave experiment, we subjected samples from all locations to the same, sublethal temperature treatments (15–23°C for eight days including acclimation) and assessed growth, storage compounds, photosynthetic efficiency and pigment contents as response traits. Recovery was assessed following seven days at 15°C. Microsatellite genotyping revealed all sampled populations to be genetically distinct entities, underlying strong regional structuring between southern and northern clades. Genetic diversity was highest at the southern distribution limit in Quiberon and lowest in the geographically isolated population on the island of Helgoland in the North Sea. The physiological response of L. digitata to temperature was similar over the entire distribution range and did not reflect the mean temperature gradient along the latitudinal gradient. However, material from Spitsbergen and Helgoland presented subtle differentiations in their temperature responses, which reflect long-term local temperature histories at these sites. Finally, a heatwave reaching 23°C for five days led to a cessation of growth, from which none of the sampled populations recovered. Our results suggest that the heat stress response of L. digitata is generally stable across its distribution range, despite strong genetic structuring of the populations. Slight local differentiation occurred in populations from the most distinct thermal environments, but 23°C posed a growth limit for all populations. This implies that local adaptation in trailing edge populations of L. digitata might not alleviate detrimental effects of global warming.

Description: Kelps are important foundation species in coastal ecosystems currently experiencing pronounced shifts in their distribution patterns caused by ocean warming. While some populations found at species’ warm distribution edges have been recently observed to decline, expansions of some species have been recorded at their cold distribution edges. Reduced population resilience can contribute to kelp habitat loss, hence, understanding intraspecific variations in physiological responses across a species’ latitudinal distribution is crucial for its conservation. To investigate potential local responses of the broadly distributed kelp Saccharina latissima to marine heatwaves in summer, we collected sporophytes from five locations in Europe (Spitsbergen, Bodø, Bergen, Helgoland, Locmariaquer), including populations exposed to the coldest and warmest local temperature regimes. Meristematic tissue from sporophytes was subjected to increasing temperatures of 1C2, 1C4 and 1C6�C above the respective mean summer temperatures (control, 1�0�C) characteristic for each site. Survival and corresponding physiological and biochemical traits were analyzed. Vitality (optimum quantum yield, Fv/Fm) and growth were monitored over time and biochemical responses were measured at the end of the experiment. Growth was highest in northern and lowest in southern populations. Overall, northern populations from Spitsbergen, Bodø and Bergen were largely unaffected by increasing summer temperatures up to 1C6�C. Conversely, sporophytes from Helgoland and Locmariaquer were markedly stressed at 1C6�C: occurrence of tissue necrosis, reduced Fv/Fm, and a significantly elevated de-epoxidation state of the xanthophyll cycle (DPS). The variations in phlorotannins, mannitol and tissue C and N contents were independent of temperature treatments and latitudinal distribution pattern. Pronounced site-specific variability in response to increasing temperatures implies that exceeding a threshold above the mean summer temperature exclusively affect rear-edge (southernmost) populations.