Geir Ottersen

An Overview of the North Atlantic Oscillation

The North Atlantic Oscillation (NAO) is one of the most prominent and recurrent patterns of atmospheric circulation variability. It dictates climate variability from the eastern seaboard of the United States to Siberia and from the Arctic to the subtropical Atlantic, especially during boreal winter, so variations in the NAO are important to society and for the environment. Understanding the processes that govern this variability is, therefore, of high priority, especially in the context of global climate change. This review, aimed at a scientifically diverse audience, provides general background material for the other chapters in the monograph, and it synthesizes some of their central points. It begins with a description of the spatial structure of climate and climate variability, including how the NAO relates to other prominent patterns of atmospheric circulation variability. There is no unique way to define the spatial structure of the NAO, or thus its temporal evolution, but several common approaches are illustrated. The relationship between the NAO and variations in surface temperature, storms and precipitation, and thus the economy, as well as the ocean and ecosystem responses to NAO variability, are described. Although the NAO is a mode of variability internal to the atmosphere, indices of it exhibit decadal variability and trends. That not all of its variability can be attributed to intraseasonal stochastic atmospheric processes points to a role for external forcings and, perhaps, a small but useful amount of predictability. The surface, stratospheric and anthropogenic processes that may influence the phase and amplitude of the NAO are reviewed.

Ecological effects of the North Atlantic Oscillation

Climatic oscillations as reflected in atmospheric modes such as the North Atlantic Oscillation (NAO) may be seen as a proxy for regulating forces in aquatic and terrestrial ecosystems. Our review highlights the variety of climate processes related to the NAO and the diversity in the type of ecological responses that different biological groups can display. Available evidence suggests that the NAO influences ecological dynamics in both marine and terrestrial systems, and its effects may be seen in variation at the individual, population and community levels. The ecological responses to the NAO encompass changes in timing of reproduction, population dynamics, abundance, spatial distribution and interspecific relationships such as competition and predator-prey relationships. This indicates that local responses to large-scale changes may be more subtle than previously suggested. We propose that the NAO effects may be classified as three types: direct, indirect and integrated. Such a classification will help the design and interpretation of analyses attempting to relate ecological changes to the NAO and, possibly, to climate in general.

The Response of Terrestrial Ecosystems to Climate Variability Associated with the North Atlantic Oscillation

Climatic factors influence a variety of ecological processes determining patterns of species density and distribution in a wide range of terrestrial ecosystems. We review the effects of the NAO on processes and patterns of terrestrial ecosystems, including both plants and animals. In plants, the NAO index correlates with date of first flowering, tree ring growth and with quality of agricultural crops (wheat and wine grapes). Also, breeding dates are earlier after high NAO index winters for amphibians and birds in Europe. Population dynamical consequences of the NAO have also been reported for birds, and the differential impact of the NAO on two similar species may prevent competitive exclusion. Different effects of the NAO on large herbivore populations have been reported for different regions, depending on limiting factors and the correlation with local weather parameters. The NAO synchronizes population dynamics of lynx and some other carnivore populations in the eastern U.S. Most effects are on an ecological time scale; the evolutionary consequences of long term trends in the NAO are poorly documented. Important for predator and prey dynamics is (1) the disruption of phenology (the match-mismatch hypothesis), (2) that there may be delayed effects (cohort-effects), and (3) that effects of the NAO may interact with other factors such as density. We discuss the challenges related to nonlinearity, of using different climate indices, and how we can progress using these pattern-oriented NAO studies at coarse scales to conduct better process-oriented small-scale experiments.

Ice-age survival of Atlantic cod: agreement between palaeoecology models and genetics

Scant scientific attention has been given to the abundance and distribution of marine biota in the face of the lower sea level, and steeper latitudinal gradient in climate, during the ice-age conditions that have dominated the past million years. Here we examine the glacial persistence of Atlantic cod (Gadus morhua) populations using two ecological-niche-models (ENM) and the first broad synthesis of multi-locus gene sequence data for this species. One ENM uses a maximum entropy approach (Maxent); the other is a new ENM for Atlantic cod, using ecophysiological parameters based on observed reproductive events rather than adult distribution. Both the ENMs were tested for present-day conditions, then used to hindcast ranges at the last glacial maximum (LGM) ca 21 kyr ago, employing climate model data. Although the LGM range of Atlantic cod was much smaller, and fragmented, both the ENMs agreed that populations should have been able to persist in suitable habitat on both sides of the Atlantic. The genetic results showed a degree of trans-Atlantic divergence consistent with genealogically continuous populations on both sides of the North Atlantic since long before the LGM, confirming the ENM results. In contrast, both the ENMs and the genetic data suggest that the Greenland G. morhua population post-dates the LGM.

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 worlds 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 cods 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.

DENSITY DEPENDENCE AND DENSITY INDEPENDENCE DURING THE EARLY LIFE STAGES OF FOUR MARINE FISH STOCKS

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.

Spawning stock and recruitment in North Sea cod shaped by food and climate

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.

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.

Productivity in the Barents Sea - Response to Recent Climate Variability

The temporal and spatial dynamics of primary and secondary biomass/production in the Barents Sea since the late 1990s are examined using remote sensing data, observations and a coupled physical-biological model. Field observations of mesozooplankton biomass, and chlorophyll a data from transects (different seasons) and large-scale surveys (autumn) were used for validation of the remote sensing products and modeling results. The validation showed that satellite data are well suited to study temporal and spatial dynamics of chlorophyll a in the Barents Sea and that the model is an essential tool for secondary production estimates. Temperature, open water area, chlorophyll a, and zooplankton biomass show large interannual variations in the Barents Sea. The climatic variability is strongest in the northern and eastern parts. The moderate increase in net primary production evident in this study is likely an ecosystem response to changes in climate during the same period. Increased open water area and duration of open water season, which are related to elevated temperatures, appear to be the key drivers of the changes in annual net primary production that has occurred in the northern and eastern areas of this ecosystem. The temporal and spatial variability in zooplankton biomass appears to be controlled largely by predation pressure. In the southeastern Barents Sea, statistically significant linkages were observed between chlorophyll a and zooplankton biomass, as well as between net primary production and fish biomass, indicating bottom-up trophic interactions in this region.

Horizontal distribution and overlap of planktivorous fish stocks in the Norwegian Sea during summers 1995-2006

Abstract The Norwegian Sea harbours several large pelagic fish stocks, which use the area for feeding during the summer. The period 1995–2006 had some of the highest biomass of pelagic fish feeding in the Norwegian Sea on record. Here we address the horizontal distribution and overlap between herring, blue whiting and mackerel in this period during the summers using a combination of acoustic, trawl and LIDAR data. A newly developed temperature atlas for the Norwegian Sea is used to present the horizontal fish distributions in relation to temperature. The centre of gravity of the herring distribution changed markedly several times during the investigated period. Blue whiting feeding habitat expanded in a northwestern direction until 2003, corresponding with an increase in abundance. Strong year classes of mackerel in 2001 and 2002 and increasing temperatures throughout the period resulted in an increased amount of mackerel in the Norwegian Sea. Mackerel was generally found in waters warmer than 8°C, while herring and blue whiting were mainly found in water masses between 2 and 8°C. The horizontal overlap between herring and mackerel was low, while blue whiting had a large horizontal overlap with both herring and mackerel. The changes in horizontal distribution and overlap between the species are explained by increasing stock sizes, increasing water temperature and spatially changing zooplankton densities in the Norwegian Sea.