Dag Ø. Hjermann

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.

Landscape ecology of the wart-biter Decticus verrucivorus in a patchy landscape

1. Populations of wart-biter Decticus verrucivorus living on habitat islands in an intensively exploited agricultural landscape in south-eastern Norway were studied. Auditory recordings of singing males indicated that 27 (39%) of 70 habitat islands contained populations in the first year of the study, while 2 years (one wart-biter generation) later the number was 16 (23%). Dispersing male wart-biters were also observed, and the dispersal distances measured. 2. Predictors of island occupancy were identified by applying logistic regression models to the pattern of presence/absence, probing the predictive power of both landscape variables (island size and isolation) as well as various microhabitat quality variables (vegetation, aspect and slope). The isolation of habitat islands relative to surrounding wart-biter populations was measured by two types of connectivity indices: one based on the empirical probability distribution of dispersal distances, the other on the negative-exponential dispersal function. 3. Habitat area and the connectivity indices were by far the best predictors of occupancy. The probability of occupation increased with increasing area and decreasing isolation. Of the unoccupied habitat islands, the ones with observations of single males were less isolated than the ones without. From the pattern of occupancy, the average dispersal distance was estimated to be 40m, while the observed average dispersal distance of males was 37 m. 4. In addition to the effects of area and connectivity, the probability of occupation was positively influenced by increasing slope to the south, increasing amounts of the plant Achillea millefolium, and decreasing fraction of vegetation lower than 10 cm. The probability of local population extinction increased with declining habitat area and increasing fraction of low vegetation. 5. The situation of the wart-biter in this highly fragmented study area seems to largely fit a metapopulation model: extinctions and absence of wart-biters were adequately predicted by habitat island size and isolation, and observed dispersal distances closely matched estimated dispersal distances from the model.

Seasonal plankton dynamics along a cross-shelf gradient.

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.

Direct and indirect climate forcing in a multi-species marine system

Interactions within and between species complicate quantification of climate effects, by causing indirect, often delayed, effects of climate fluctuations and compensation of mortality. Here we identify direct and indirect climate effects by analysing unique Russian time-series data from the Norwegian Sea–Barents Sea ecosystem on the first life stages of cod, capelin, herring and haddock, their predators, competitors and zooplanktonic prey. By analysing growth and survival from one life stage to the next (eggs–larvae–juveniles–recruits), we find evidence for both bottom-up, direct and top-down effects of climate. Ambient zooplankton biomass predicts survival of all species, whereas ambient temperature mainly affects survival through effects on growth. In warm years, all species experienced improved growth and feeding conditions. Cohorts born following a warm year will, however, experience increased predation and competition because of increased densities of subadult cod and herring, leading to delayed climate effects. While climate thus affects early growth and survival through several mechanisms, only some of the identified mechanisms were found to be significant predictors of population growth. In particular, our findings exemplify that climate impacts are barely propagated to later life stages when density dependence is strong.

Interaction between seasonal density-dependence structures and length of the seasons explain the geographical structure of the dynamics of voles in Hokkaido : an example of seasonal forcing

The grey-sided vole (Clethrionomys rufocanus) is distributed over the entire island of Hokkaido, Japan, across which it exhibits multindash;annual density cycles in only parts of the island (the northndash;eastern part); in the remaining part of the island, only seasonal density changes occur. Using annual sampling of 189 greyndash;sided vole populations, we deduced the geographical structure in their secondndash;order density dependence. Building upon our earlier suggestion, we deduce the seasonal densityndash;dependent structure for these populations. Strong direct and delayed density dependence is found to occur during winter, whereas no density dependence is seen during the summer period. The direct density dependence during winter may be seen as a result of food being limited during that season: the delayed density dependence during the winter is consistent with volendash;specialized predators (e.g. the least weasel) responding to vole densities so as to have a negative effect on the net growth rate of voles in the following year. We conclude that the observed geographical structure of the population dynamics may be properly seen as a result of the length of the summer in interaction with the differential seasonal densityndash;dependent structure. Altogether, this indicates that the geographical pattern in multindash;annual density dynamics in the greyndash;sided vole may be a result of seasonal forcing.

Trophic interactions affecting a key ecosystem component: a multistage analysis of the recruitment of the Barents Sea capelin (Mallotus villosus)

The Barents Sea stock of capelin (Mallotus villosus) has suffered three major collapses (>90% reduction) since 1985 due to recruitment failures. As capelin is a key species in the area, these population collapses have had major ecosystem consequences. By analysing data on spawner biomass and three recruitment stages (larvae, 0-group, and 1-year-olds), we suggest that much of the recruitment failures are caused by predation from herring (Clupea harengus) and 0-group and adult Northeast Arctic cod (Gadus morhua). Recruitment is furthermore positively correlated with sea temperatures in winter and spring. Harvesting of maturing capelin on their way to the spawning grounds reduced the abundance of larvae significantly, but this reduction to a large extent is compensated for later in life, as mortality is strongly density-dependent between the larval stage and age 1. Altogether, our study indicates a very high importance of trophic interactions, consistent with similar findings in other high-latitude marine ec...

A combination of hydrodynamical and statistical modelling reveals non-stationary climate effects on fish larvae distributions

Biological processes and physical oceanography are often integrated in numerical modelling of marine fish larvae, but rarely in statistical analyses of spatio-temporal observation data. Here, we examine the relative contribution of inter-annual variability in spawner distribution, advection by ocean currents, hydrography and climate in modifying observed distribution patterns of cod larvae in the Lofoten–Barents Sea. By integrating predictions from a particle-tracking model into a spatially explicit statistical analysis, the effects of advection and the timing and locations of spawning are accounted for. The analysis also includes other environmental factors: temperature, salinity, a convergence index and a climate threshold determined by the North Atlantic Oscillation (NAO). We found that the spatial pattern of larvae changed over the two climate periods, being more upstream in low NAO years. We also demonstrate that spawning distribution and ocean circulation are the main factors shaping this distribution, while temperature effects are different between climate periods, probably due to a different spatial overlap of the fish larvae and their prey, and the consequent effect on the spatial pattern of larval survival. Our new methodological approach combines numerical and statistical modelling to draw robust inferences from observed distributions and will be of general interest for studies of many marine fish species.

The Influence of Physical Factors on Kelp and Sea Urchin Distribution in Previously and Still Grazed Areas in the NE Atlantic

The spatial distribution of kelp (Laminaria hyperborea) and sea urchins (Strongylocentrotus droebachiensis) in the NE Atlantic are highly related to physical factors and to temporal changes in temperature. On a large scale, we identified borders for kelp recovery and sea urchin persistence along the north-south gradient. Sea urchin persistence was also related to the coast-ocean gradient. The southern border corresponds to summer temperatures exceeding about 10°C, a threshold value known to be critical for sea urchin recruitment and development. The outer border along the coast-ocean gradient is related to temperature, wave exposure and salinity. On a finer scale, kelp recovery occurs mainly at ridges in outer, wave exposed, saline and warm areas whereas sea urchins still dominate in inner, shallow and cold areas, particularly in areas with optimal current speed for sea urchin foraging. In contrast to other studies in Europe, we here show a positive influence of climate change to presence of a long-lived climax canopy-forming kelp. The extent of the coast-ocean gradient varies within the study area, and is especially wide in the southern part where the presence of islands and skerries increases the area of the shallow coastal zone. This creates a large area with intermediate physical conditions for the two species and a mosaic of kelp and sea urchin dominated patches. The statistical models (GAM and BRT) show high performance and indicate recovery of kelp in 45–60% of the study area. The study shows the value of combining a traditional (GAM) and a more complex (BRT) modeling approach to gain insight into complex spatial patterns of species or habitats. The results, methods and approaches are of general ecological relevance regardless of ecosystems and species, although they are particularly relevant for understanding and exploring the corresponding changes between algae and grazers in different coastal areas.

Environmental toxicology: population modeling of cod larvae shows high sensitivity to loss of zooplankton prey.

Two factors determine whether pollution is likely to affect a population indirectly through loss of prey: firstly, the sensitivity of the prey to the pollutants, and secondly, the sensitivity of the predator population to loss of prey at the given life stage. We here apply a statistical recruitment model for Northeast Arctic cod to evaluate the sensitivity of cod cohorts to loss of zooplankton prey, for example following an oil spill. The calculations show that cod cohorts are highly sensitive to possible zooplankton biomass reductions in the distribution area of the cod larvae, and point to a need for more knowledge about oil-effects on zooplankton. Our study illustrates how knowledge about population dynamics may guide which indirect effects to consider in environmental impact studies.

Northeast Arctic cod population persistence in the Lofoten-Barents Sea system under fishing.

Population growth, and hence the populations persistence, is affected by several factors such as climate, species interaction, and harvesting pressure. Proper resource management requires an understanding of these factors. We apply techniques based upon age-structured population matrices to analyze estimated stock sizes derived from annual bottom trawl sampling in the winter feeding area of northeast Arctic cod (Gadus morhua L.) from 1981 to 2003. We run generalized additive models to explain population growth rate by different explanatory variables. Cod population growth was found to be positively related to the abundance of capelin (Mallotus villosus Miller), negatively related to the number of cannibalistic cod with a two-year lag, and marginally positively related to the winter North Atlantic Oscillation index (NAO). This model remains true independently from the population status (i.e., fished or non-fished). Capelin abundance is the main variable that to some degree can be adjusted in order to maintain the population size at a given level of cod harvesting. Our results point to the importance of managing conjointly cod and capelin stocks.