Pulsed pressure: Fluctuating impacts of multifactorial environmental change on a temperate macroalgal community

Global change impacts marine organisms and communities mainly through ocean warming, acidification, deoxygenation, and changes in nutrient inputs and water circulation. To assess the ecological impacts of global change, the effects of multiple interacting environmental drivers, including their fluctuations, should be tested at different levels of biological organization. In an outdoor mesocosm study, we investigated the differential effects of three simulated upwelling events coupled with ocean warming (1–5°C above ambient) on a temperate benthic community in the Western Baltic Sea. Ocean warming, especially in summer when temperatures are close to or above the physiological optimum of many species, is likely to impose thermal stress with species-specific impacts. As the properties of deep water vary seasonally, so will the effects of upwelling. Upwelling of cooler deep water in midsummer may alleviate thermal stress, although this mitigation may be modulated by upwelling-associated shifts in other water-quality parameters such as salinity, nutrients, or late-summer hypoxia. This investigation showed that in the Western Baltic Ocean warming was rather beneficial in early and late summer but detrimental when ambient temperatures were highest in midsummer. The effects of upwelling in the absence of ocean warming were generally weakly beneficial, while this effect tended to vanish with intensifying imposed ocean warming. Hypoxia associated with the late summer upwelling impacted some of the grazer species but did not impact the macroalgae. We conclude that in coastal temperate benthic communities, ocean warming is the predominant stressor that may partially and seasonally be buffered by upwelling.