Ocean acidification (OA) is predicted to play a major role in shaping species biogeography and marine biodiversity over the next century. We tested the effect of medium-term exposure to OA (pH 8.00, 7.30 and 6.70 for 30 d) on acid -base balance in the decapod crab Necora puber -a species that is known to possess good extracellular buffering ability during short-term exposure to hypercapnic conditions. To determine if crabs undergo physiological trade-offs in order to buffer their haemolymph, we characterised a number of fundamental physiological functions, i.e. metabolic rate, tolerance to heat, carapace and chelae [Ca 2+ ] and [Mg 2+ ], haemolymph [Ca 2+ ] and [Mg 2+ ], and immune response in terms of lipid peroxidation. Necora puber was able to buffer changes to extracellular pH over 30 d exposure to hypercapnic water, with no evidence of net shell dissolution, thus demonstrating that HCO 3 -is actively taken up from the surrounding water. In addition, tolerance to heat, carapace mineralization, and aspects of immune response were not affected by hypercapnic conditions. In contrast, whole-animal O 2 uptake significantly decreased with hypercapnia, while significant increases in haemolymph [Ca 2+ ] and [Mg 2+ ] and chelae [Mg 2+ ] were observed with hypercapnia. Our results confirm that most physiological functions in N. puber are resistant to low pH/ hypercapnia over a longer period than previously investigated, although such resistance comes at the expenses of metabolic rates, haemolymph chemistry and chelae mineralization.KEY WORDS: Ocean acidification · Physiology · Metabolic depression · Necora puber · Thermal tolerance · Carbon capture and storage Resale or republication not permitted without written consent of the publisher

An organism's physiological processes form the link between its life-history traits and the prevailing environmental conditions, especially in species with complex life cycles. Understanding how these processes respond to changing environmental conditions, thereby affecting organismal development, is critical if we are to predict the biological implications of current and future global climate change. However, much of our knowledge is derived from adults or single developmental stages. Consequently, we investigated the metabolic rate, organic content, carapace mineralization, growth, and survival across each larval stage of the European lobster Homarus gammarus, reared under current and predicted future ocean warming and acidification scenarios. Larvae exhibited stage-specific changes in the temperature sensitivity of their metabolic rate. Elevated Pco2 increased C∶N ratios and interacted with elevated temperature to affect carapace mineralization. These changes were linked to concomitant changes in survivorship and growth, from which it was concluded that bottlenecks were evident during H. gammarus larval development in stages I and IV, the transition phases between the embryonic and pelagic larval stages and between the larval and megalopa stages, respectively. We therefore suggest that natural changes in optimum temperature during ontogeny will be key to larvae survival in a future warmer ocean. The interactions of these natural changes with elevated temperature and Pco2 significantly alter physiological condition and body size of the last larval stage before the transition from a planktonic to a benthic life style. Thus, living and growing in warm, hypercapnic waters could compromise larval lobster growth, development, and recruitment.

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Ocean acidification is predicted to have detrimental effects on many marine organisms and ecological processes. Despite growing evidence for direct impacts on specific species, few studies have simultaneously considered the effects of ocean acidification on individuals (e.g. consequences for energy budgets and resource partitioning) and population level demographic processes. Here we show that ocean acidification increases energetic demands on gastropods resulting in altered energy allocation, i.e. reduced shell size but increased body mass. When scaled up to the population level, long-term exposure to ocean acidification altered population demography, with evidence of a reduction in the proportion of females in the population and genetic signatures of increased variance in reproductive success among individuals. Such increased variance enhances levels of short-term genetic drift which is predicted to inhibit adaptation. Our study indicates that even against a background of high gene flow, ocean acidification is driving individual- and population-level changes that will impact eco-evolutionary trajectories.

Most studies assessing the impacts of ocean acidification (OA) on benthic marine invertebrates have used stable mean pH/pCO2 levels to highlight variation in the physiological sensitivities in a range of taxa. However, many marine environments experience natural fluctuations in carbonate chemistry, and to date little attempt has been made to understand the effect of naturally fluctuating seawater pCO2 (pCO2sw) on the physiological capacity of organisms to maintain acid–base homeostasis. Here, for the first time, we exposed two species of sea urchin with different acid–base tolerances, Paracentrotus lividus and Arbacia lixula, to naturally fluctuating pCO2sw conditions at shallow water CO2 seep systems (Vulcano, Italy) and assessed their acid–base responses. Both sea urchin species experienced fluctuations in extracellular coelomic fluid pH, pCO2, and [HCO3−] (pHe, pCO2e, and [HCO3−]e, respectively) in line with fluctuations in pCO2sw. The less tolerant species, P. lividus, had the greatest capacity for [HCO3−]e buffering in response to acute pCO2sw fluctuations, but it also experienced greater extracellular hypercapnia and acidification and was thus unable to fully compensate for acid–base disturbances. Conversely, the more tolerant A. lixula relied on non-bicarbonate protein buffering and greater respiratory control. In the light of these findings, we discuss the possible energetic consequences of increased reliance on bicarbonate buffering activity in P. lividus compared with A. lixula and how these differing physiological responses to acute fluctuations in pCO2sw may be as important as chronic responses to mean changes in pCO2sw when considering how CO2 emissions will affect survival and success of marine organisms within naturally assembled systems.