Increasing water temperatures are predicted around the globe, with high amplitudes of warming in Subarctic and Arctic regions where Atlantic cod (Gadus morhua) populations currently flourish. We reconstructed oxygen isotope and temperature chronologies from Icelandic cod otoliths, one of the largest cod stocks in the world, to determine if cod moved or migrated over the last 100 years to avoid increasing water temperatures. For δ18Ootolith analysis, individual annual growth increments from immature and mature life history stages were micromilled from adult otoliths, which were collected in southern Iceland. Linear mixed-effect models confirmed that stable oxygen isotope time series of immature and mature cod differ significantly between both life stages (p < 0.001). Overall, cod otolith δ18O was significantly correlated with water temperature (sea surface temperature: p < 0.001, temperature at 200 m depth: p < 0.01), indicating that Atlantic cod were exposed to fluctuating water temperatures during the past 100 years and did not move as a response to increasing ocean temperatures. All of the alternate physical factors that were considered for the isotope-based variation in the temperature exposure of Icelandic cod were rejected.
The isotopic composition of inorganic carbon in otoliths (δ13Coto) can be a useful tracer of metabolic rates and a method to study ecophysiology in wild fish. We evaluated environmental and physiological sources of δ13Coto variation in Icelandic and Northeast Arctic (NEA) cod (Gadus morhua) over the years 1914–2013. Individual annual growth increments of otoliths formed at age 3 and 8 were micromilled and measured by isotope-ratio mass spectrometry. Simultaneously, all annual increment widths of the otoliths were measured providing a proxy of fish somatic growth. We hypothesized that changes in the physiological state of the organism, reflected by the isotopic composition of otoliths, can affect the growth rate. Using univariate and multivariate mixed-effects models we estimated conditional correlations between carbon isotopic composition and growth of fish at different levels (within individuals, between individuals, and between years), controlling for intrinsic and extrinsic effects on both otolith measurements. δ13Coto was correlated with growth within individuals and between years, which was attributed to the intrinsic effects (fish age or total length). There was no significant correlation between δ13Coto and growth between individuals, which suggests that caution is needed when interpreting δ13Coto signals. We found a significant decrease in δ13Coto through the century which was explained by the oceanic Suess effect-admixture of isotopically light carbon from fossil fuel. We calculated the proportion of the respired carbon in otolith carbonate (Cresp) using carbon isotopic composition in diet and dissolved inorganic carbon of the seawater. This approach allowed us to correct the values for each stock in relation to these two environmental baselines. Cresp was on average 0.275 and 0.295 in Icelandic and NEA stock, respectively. Our results provide an insight into the physiological basis for differences in growth characteristics between these two cod stocks, and how that may vary over time.
Global warming is increasing ocean temperatures, forcing marine organisms to respond to a suite of changing environmental conditions. The stable oxygen isotopic composition of otoliths is often used as an index of temperature exposure, but the accuracy of the resulting temperature reconstructions in wild, free-swimming Atlantic cod (Gadus morhua) has never been groundtruthed. Based on temperatures from data storage tags (DST) and corresponding salinity values, the stable oxygen isotope (δ18O) value was predicted for each month of tagging and compared with δ18Ootolith values measured in situ with secondary ion mass spectrometry (SIMS). Paired-sample Wilcoxon tests were applied to compare measured and predicted δ18O values. The difference between measured and predicted mean and maximum δ18Ootolith values was not significant, suggesting a good correspondence between SIMS-measured and DST-predicted δ18Ootolith values. However, SIMS-measured and predicted minimum δ18Ootolith values were significantly different (all samples: p < 0.01, coastal and frontal cod: p < 0.05), resulting in overestimation of maximum temperatures. Our results confirm that otoliths are well-suited as proxies for mean ambient temperature reconstructions. A possible matrix effect and the absence of a reliable aragonite standard for SIMS measurements appeared to cause a small divergence between measured and predicted δ18Ootolith values, which affected the estimation accuracy of absolute temperature. However, relative temperature changes were accurately estimated by SIMS-analyzed δ18Ootolith values.
Large‐scale, climate‐induced synchrony in the productivity of fish populations is becoming more pronounced in the world's oceans. As synchrony increases, a population's “portfolio” of responses can be diminished, in turn reducing its resilience to strong perturbation. Here we argue that the costs and benefits of trait synchronization, such as the expression of growth rate, are context dependent. Contrary to prevailing views, synchrony among individuals could actually be beneficial for populations if growth synchrony increases during favorable conditions, and then declines under poor conditions when a broader portfolio of responses could be useful. Importantly, growth synchrony among individuals within populations has seldom been measured, despite well‐documented evidence of synchrony across populations. Here, we used century‐scale time series of annual otolith growth to test for changes in growth synchronization among individuals within multiple populations of a marine keystone species (Atlantic cod, Gadus morhua). On the basis of 74,662 annual growth increments recorded in 13,749 otoliths, we detected a rising conformity in long‐term growth rates within five northeast Atlantic cod populations in response to both favorable growth conditions and a large‐scale, multidecadal mode of climate variability similar to the East Atlantic Pattern. The within‐population synchrony was distinct from the across‐population synchrony commonly reported for large‐scale environmental drivers. Climate‐linked, among‐individual growth synchrony was also identified in other Northeast Atlantic pelagic, deep‐sea and bivalve species. We hypothesize that growth synchrony in good years and growth asynchrony in poorer years reflects adaptive trait optimization and bet hedging, respectively, that could confer an unexpected, but pervasive and stabilizing, impact on marine population productivity in response to large‐scale environmental change.
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