Tradeoffs of managing cod as a sustainable resource in fluctuating environments.

Sustainable human exploitation of living marine resources stems from a delicate balance between yield stability and population persistence to achieve socioeconomic and conservation goals. But our imperfect knowledge of how oceanic oscillations regulate temporal variation in an exploited species can obscure the risk of missing management targets. We illustrate how applying a management policy to suppress fluctuations in fishery yield in variable environments (prey density and regional climate) can present unintended outcomes in harvested predators and the sustainability of harvesting. Using Atlantic cod (Gadus morhua, an apex predatory fish) in the Barents Sea as a case study we simulate age-structured population and harvest dynamics through time-varying, density-dependent and density-independent processes with a stochastic, process-based model informed by 27-year monitoring data. In this model, capelin (Mallotus villosus, a pelagic forage fish), a primary prey of cod, fluctuations modulate the strength of density-dependent regulation primarily through cannibalistic pressure on juvenile cod survival; sea temperature fluctuations modulate thermal regulation of cod feeding, growth, maturation, and reproduction. We first explore how capelin and temperature fluctuations filtered through cod intrinsic dynamics modify catch stability and then evaluate how management to suppress short-term variability in catch targets alters overharvest risk. Analyses revealed that suppressing year-to-year catch variability impedes management responses to adjust fishing pressure, which becomes progressively out of sync with variations in cod abundance. This asynchrony becomes amplified in fluctuating environments, magnifying the amplitudes of both fishing pressure and cod abundance and then intensifying the density-dependent regulation of juvenile survival through cannibalism. Although these transient dynamics theoretically give higher average catches, emergent, quasicyclic behaviors of the population would increase long-term yield variability and elevate overharvest risk. Management strategies that overlook the interplay of extrinsic (fishing and environment) and intrinsic (life history and demography) fluctuations thus can inadvertently destabilize fish stocks, thereby jeopardizing the sustainability of harvesting. These policy implications underscore the value of ecosystem approaches to designing management measures to sustainably harvest ecologically connected resources while achieving socioeconomic security.

Synergies between climate and management for Atlantic cod fisheries at high latitudes.

The widespread depletion of commercially exploited marine living resources is often seen as a general failure of management and results in criticism of contemporary management procedures. When populations show dramatic and positive changes in population size, this invariably leads to questions about whether favorable climatic conditions or good management (or both) were responsible. The Barents Sea cod (Gadus morhua) stock has recently increased markedly and the spawning stock biomass is now at an unprecedented high. We identify the crucial social and environmental factors that made this unique growth possible. The relationship between vital rates of Barents Sea cod stock productivity (recruitment, growth, and mortality) and environment is investigated, followed by simulations of population size under different management scenarios. We show that the recent sustained reduction in fishing mortality, facilitated by the implementation of a "harvest control rule," was essential to the increase in population size. Simulations show that a drastic reduction in fishing mortality has resulted in a doubling of the total population biomass compared with that expected under the former management regime. However, management alone was not solely responsible. We document that prevailing climate, operating through several mechanistic links, positively reinforced management actions. Heightened temperature resulted in an increase in the extent of the suitable feeding area for Barents Sea cod, likely offering a release from density-dependent effects (for example, food competition and cannibalism) through prolonged overlap with prey and improved adult stock productivity. Management and climate may thus interact to give a positive outlook for exploited high-latitude marine resources.

Under-ice observations by trawls and multi-frequency acoustics in the Central Arctic Ocean reveals abundance and composition of pelagic fauna.

The rapid ongoing changes in the Central Arctic Ocean call for baseline information on the pelagic fauna. However, sampling for motile organisms which easily escape vertically towed nets is challenging. Here, we report the species composition and catch weight of pelagic fishes and larger zooplankton from 12 trawl hauls conducted in ice covered waters in the Central Arctic Ocean beyond the continental slopes in late summer. Combined trawl catches with acoustics data revealed low amounts of fish and zooplankton from the advective influenced slope region in the Nansen Basin in the south to the ice-covered deep Amundsen Basin in the north. Both arctic and subarctic-boreal species, including the ones considered as Atlantic expatriate species were found all the way to 87.5o N. We found three fish species (Boreogadus saida, Benthosema glaciale and Reinhardtius hippoglossoides), but the catch was limited to only seven individuals. Euphausiids, amphipods and gelatinous zooplankton dominated the catch weight in the Nansen Basin in the mesopelagic communities. Euphausiids were almost absent in the Amundsen Basin with copepods, amphipods, chaetognaths and gelatinous zooplankton dominating. We postulate asymmetric conditions in the pelagic ecosystems of the western and eastern Eurasian Basin caused by ice and ocean circulation regimes.

Polar cod in jeopardy under the retreating Arctic sea ice.

The Arctic amplification of global warming is causing the Arctic-Atlantic ice edge to retreat at unprecedented rates. Here we show how variability and change in sea ice cover in the Barents Sea, the largest shelf sea of the Arctic, affect the population dynamics of a keystone species of the ice-associated food web, the polar cod (Boreogadus saida). The data-driven biophysical model of polar cod early life stages assembled here predicts a strong mechanistic link between survival and variation in ice cover and temperature, suggesting imminent recruitment collapse should the observed ice-reduction and heating continue. Backtracking of drifting eggs and larvae from observations also demonstrates a northward retreat of one of two clearly defined spawning assemblages, possibly in response to warming. With annual to decadal ice-predictions under development the mechanistic physical-biological links presented here represent a powerful tool for making long-term predictions for the propagation of polar cod stocks.

Food web dynamics affect Northeast Arctic cod recruitment.

Proper management of ecosystems requires an understanding of both the species interactions as well as the effect of climate variation. However, a common problem is that the available time-series are of different lengths. Here, we present a general approach for studying the dynamic structure of such interactions. Specifically, we analyse the recruitment of the world's largest cod stock, the Northeast Arctic cod. Studies based on data starting in the 1970-1980s indicate that this stock is affected by temperature through a variety of pathways. However, the value of such studies is somewhat limited by the fact that they are based on a quite specific ecological and climatic situation. Recently, this stock has consisted of fairly young fish and the spawning stock has consisted of relatively few age groups. In this study, we develop a model for the effect of capelin (the cod's main prey) and herring on cod recruitment since 1973. Based on this model, we analyse data on cod, herring and temperature going back to 1921 and find that food-web effects explain a significant part of the cod recruitment variation back to around 1950.

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability.

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.

Density-dependent migratory waves in the marine pelagic ecosystem.

The migration of large aggregations of animals that sweep through the landscape is a phenomenon with large consequences in many ecosystems. It has been suggested that such migrations are mediated by resource depletion. Under this hypothesis it has been shown that simple foraging rules may generate density-dependent migratory waves (DDMW) in which the speed and amplitude increase with animal abundance. We tested these predictions on a 32-year data set of the spatial distribution of the two youngest age groups of a small pelagic schooling fish, the capelin (Mallotus villosus), by the end of their annual feeding migration in the Barents Sea. Our data suggest that the two age groups divided the Barents Sea by forming migratory waves that moved in opposite directions. The aggregation and spatial displacement of these waves increased with increasing age-specific abundance. However, possibly through social interactions, migratory pattern was modified by the abundance of the other age group.