Macroalgae contribute to nested mosaics of pH variability in a subarctic fjord

The Arctic Ocean is considered the most vulnerable ecosystem to ocean acidification, and large-scale assessments of pH and the saturation state for aragonite (O_arag) have led to the notion that the Arctic Ocean is already close to a corrosive state. In high-latitude coastal waters the regulation of pH and O_arag is, however, far more complex than offshore because increased biological activity and input of glacial meltwater affect pH. Effects of ocean acidification on calcifiers and non-calcifying phototrophs occupying coastal habitats cannot be derived from extrapolation of current and forecasted offshore conditions, but they require an understanding of the regimes of pH and O_arag in their coastal habitats. To increase knowledge of the natural variability in pH in the Arctic coastal zone and specifically to test the influence of benthic vegetated habitats, we quantified pH variability in a Greenland fjord in a nested-scale approach. A sensor array logging pH, O₂, PAR, temperature and salinity was applied on spatial scales ranging from kilometre scale across the horizontal extension of the fjord; to 100 m scale vertically in the fjord, 10–100 m scale between subtidal habitats with and without kelp forests and between vegetated tidal pools and adjacent vegetated shores; and to centimetre to metre scale within kelp forests and millimetre scale across diffusive boundary layers of macrophyte tissue. In addition, we assessed the temporal variability in pH on diurnal and seasonal scales. Based on pH measurements combined with point samples of total alkalinity, dissolved inorganic carbon and relationships to salinity, we also estimated variability in O_arag. Results show variability in pH and O_arag of up to 0.2–0.3 units at several scales, i.e. along the horizontal and vertical extension of the fjord, between seasons and on a diel basis in benthic habitats and within 1 m³ of kelp forest. Vegetated intertidal pools exhibited extreme diel pH variability of > 1.5 units and macrophyte diffusive boundary layers a pH range of up to 0.8 units. Overall, pelagic and benthic metabolism was an important driver of pH and O_arag producing mosaics of variability from low levels in the dark to peak levels at high irradiance generally appearing favourable for calcification. We suggest that productive coastal environments may form niches of high pH in a future acidified Arctic Ocean

An objective framework to test the quality of candidate indicators of good environmental status

Large efforts are on-going within the EU to prepare the Marine Strategy Framework Directive's (MSFD) assessment of the environmental status of the European seas. This assessment will only be as good as the indicators chosen to monitor the 11 descriptors of good environmental status (GEnS). An objective and transparent framework to determine whether chosen indicators actually support the aims of this policy is, however, not yet in place. Such frameworks are needed to ensure that the limited resources available to this assessment optimize the likelihood of achieving GEnS within collaborating states. Here, we developed a hypothesis-based protocol to evaluate whether candidate indicators meet quality criteria explicit to the MSFD, which the assessment community aspires to. Eight quality criteria are distilled from existing initiatives, and a testing and scoring protocol for each of them is presented. We exemplify its application in three worked examples, covering indicators for three GEnS descriptors (1, 5, and 6), various habitat components (seaweeds, seagrasses, benthic macrofauna, and plankton), and assessment regions (Danish, Lithuanian, and UK waters). We argue that this framework provides a necessary, transparent and standardized structure to support the comparison of candidate indicators, and the decision-making process leading to indicator selection. Its application could help identify potential limitations in currently available candidate metrics and, in such cases, help focus the development of more adequate indicators. Use of such standardized approaches will facilitate the sharing of knowledge gained across the MSFD parties despite context-specificity across assessment regions, and support the evidence-based management of European seas

Long photoperiods sustain high pH in Arctic kelp forests

Concern on the impacts of ocean acidification on calcifiers, such as bivalves, sea urchins, and foraminifers, has led to efforts to understand the controls on pH in their habitats, which include kelp forests and seagrass meadows. The metabolism of these habitats can lead to diel fluctuation in pH with increases during the day and declines at night, suggesting no net effect on pH at time scales longer than daily. We examined the capacity of subarctic and Arctic kelps to up-regulate pH in situ and experimentally tested the role of photoperiod in determining the capacity of Arctic macrophytes to up-regulate pH. Field observations at photoperiods of 15 and 24 hours in Greenland combined with experimental manipulations of photoperiod show that photoperiods longer than 21 hours, characteristic of Arctic summers, are conducive to sustained up-regulation of pH by kelp photosynthesis. We report a gradual increase in pH of 0.15 units and a parallel decline in pCO2 of 100 parts per million over a 10-day period in an Arctic kelp forest over midsummer, with ample scope for continued pH increase during the months of continuous daylight. Experimental increase in CO2 concentration further stimulated the capacity of macrophytes to deplete CO2 and increase pH. We conclude that long photoperiods in Arctic summers support sustained up-regulation of pH in kelp forests, with potential benefits for calcifiers, and propose that this mechanism may increase with the projected expansion of Arctic vegetation in response to warming and loss of sea ice.The study was funded by the Danish Environmental Protection Agency within the Danish Cooperation for Environment in the Arctic. It is also a contribution to the Greenland Ecosystem Monitoring program (www.G-E-M.dk) and the Arctic Science Partnership (www.asp-net.org). M.S.-M. was supported by a Fundación “La Caixa” fellowship (Spain). We acknowledge support by the CSIC Open Access Publication Initiative through its Unit of Information Resources for Research (URICI).Peer reviewe

Distribution, structure and function of Nordic eelgrass (Zostera marina) ecosystems: implications for coastal management and conservation

This paper focuses on the marine foundation eelgrass species, Zostera marina, along a gradient from the northern Baltic Sea to the north-east Atlantic. This vast region supports a minimum of 1480 km2 eelgrass (maximum >2100 km2), which corresponds to more than four times the previously quantified area of eelgrass in Western Europe. Eelgrass meadows in the low salinity Baltic Sea support the highest diversity (4–6 spp.) of angiosperms overall, but eelgrass productivity is low (<2 g dw m-2 d-1) and meadows are isolated and genetically impoverished. Higher salinity areas support monospecific meadows, with higher productivity (3–10 g dw m-2 d-1) and greater genetic connectivity. The salinity gradient further imposes functional differences in biodiversity and food webs, in particular a decline in number, but increase in biomass of mesograzers in the Baltic. Significant declines in eelgrass depth limits and areal cover are documented, particularly in regions experiencing high human pressure. The failure of eelgrass to re-establish itself in affected areas, despite nutrient reductions and improved water quality, signals complex recovery trajectories and calls for much greater conservation effort to protect existing meadows. The knowledge base for Nordic eelgrass meadows is broad and sufficient to establish monitoring objectives across nine national borders. Nevertheless, ensuring awareness of their vulnerability remains challenging. Given the areal extent of Nordic eelgrass systems and the ecosystem services they provide, it is crucial to further develop incentives for protecting them