Owing to its extraordinary lifespan and wide geographical distribution along the continental margins of the North Atlantic Ocean, the ocean quahog Arctica islandica may become an important indicator species in environmental change research. To test for applicability and ''calibrate'' the Arctica-indicator, metabolic properties of A. islandica specimens were compared across different climatic and oceanographic regions. Fully saline populations from Iceland to the North Sea as well as animals from polyhaline and low salinity, environments, the White Sea and the Baltic were included in the study. This calibration centrally includes recordings of growth-age relationships in different populations. Shells were used as age recorders by counting annual growth bands. As a result of this study, we propose a general respiration model that links individual metabolic rates of A. islandica from five populations: Norwegian coast, Kattegat, Kiel Bay (Baltic Sea), White Sea and German Bight (North Sea), to body mass, water temperature and site. Temperature exerts distinct site specific effects on respiration rate, which is indicated by Q 10 values ranging from 4.48 for German Bight to 1.15 for Kiel Bay animals. Individual age, occurrence of apneal respiratory gaps, parasite infestation and salinity do not affect respiration rate. Respiration of Arctica islandica is significantly below the average of 59 bivalve species when compared at the same temperature and animal mass. This respiration model principally enables the coupling of A. islandica life history and population dynamics to regional oceanographic temperature models.

We compared lifetime and population energy budgets of the extraordinary long-lived ocean quahog Arctica islandica from 6 different sites-the Norwegian coast, Kattegat, Kiel Bay, White Sea, German Bight, and off northeast Iceland-covering a temperature and salinity gradient of 4-10°C (annual mean) and 25-34, respectively. Based on von Bertalanffy growth models and size-mass relationships, we computed organic matter production of body (P SB ) and of shell (P SS ), whereas gonad production (P G ) was estimated from the seasonal cycle in mass. Respiration (R) was computed by a model driven by body mass, temperature, and site. A. islandica populations differed distinctly in maximum life span (40 y in Kiel Bay to 197 y in Iceland), but less in growth performance (f# ranged from 2.41 in the White Sea to 2.65 in Kattegat). Individual lifetime energy throughput, as approximated by assimilation, was highest in Iceland (43,730 kJ) and lowest in the White Sea (313 kJ). Net growth efficiency ranged between 0.251 and 0.348, whereas lifetime energy investment distinctly shifted from somatic to gonad production with increasing life span; P S /P G decreased from 0.362 (Kiel Bay, 40 y) to 0.031 (Iceland, 197 y). Population annual energy budgets were derived from individual budgets and estimates of population mortality rate (0.035/y in Iceland to 0.173/y in Kiel Bay). Relationships between budget ratios were similar on the population level, albeit with more emphasis on somatic production; P S / P G ranged from 0.196 (Iceland) to 2.728 (White Sea), and P/B ranged from 0.203-0.285/y. Life span is the principal determinant of the relationship between budget parameters, whereas temperature affects net growth efficiency only. In the White Sea population, both growth performance and net growth efficiency of A. islandica were lowest. We presume that low temperature combined with low salinity represent a particularly stressful environment for this species.

Environmental factors such as temperature and salinity regimes shape lifespan in marine ectotherms. We investigated whether the effect occurs through modification of metabolic reactive oxygen species (ROS)-producing processes and is thus in line with the rate of living-free radical theory of aging. We compared 6 biogeographically and climatically distinct populations of the extremely long-lived ocean quahog Arctica islandica for age-dependent differences in metabolic rates and antioxidant capacities (superoxide dismutase, catalase activity and total glutathione concentration). The temperature and salinity ranges covered by the sampling locations (Norwegian coast, White Sea, Iceland, Kattegat, Kiel Bay and German Bight) were 3.7 to 9.3°C and 20 to 35 ppt. Bivalve shells were used as age recorders by counting annual growth bands. Maximum determined age in different populations varied between 29 and 192 yr. Extreme longevity observed in some North Atlantic A. islandica populations seems to be based on their very low lifetime mass-specific respiration, in combination with stable maintenance of antioxidant protection throughout life in mature specimens. While the antioxidant capacity was similar among all populations, the shorter-lived populations exhibited the highest metabolic rates and showed no metabolic response (Q 10 ) when warmed to higher temperature. Low and fluctuating salinity in the Baltic may further exert stress, which enhances respiration rates and reduces longevity in the Baltic Sea population. The exceptionally long lifespan of A. islandica cannot be exclusively explained by a well-established antioxidant defense system, and the long lifespan of some populations may rather be a function of low ROS formation (low metabolic rate) and high damage repair/removal capacities.

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