Relating environmental pressures to littoral biological water quality indicators in Western Balkan lakes: Can we fill the largest gaps?

Along six transects in each of six lakes across the Western Balkans, we collected data for three groups of littoral biological water quality indicators: epilithic diatoms, macrophytes, and benthic invertebrates. We assessed the relationships between them and three environmental pressures: nutrient load (eutrophication), hydro-morphological alteration of the shoreline, and water level variation, separating the effect of individual lakes and continuous explanatory variables. Lake water total phosphorus concentration (TP) showed substantial variation but was not related to any of the tested biological indicators, nor to any of the tested pressures. We suggest that this may be due to feedback processes such as P removal in the lake littoral zone. Instead, we found that a gradient in surrounding land-use towards increasing urbanization, and a land-use-based estimate of P run-off, served as a better descriptor of eutrophication. Overall, eutrophication and water level fluctuation were most important for explaining variation in the assessed indicators, whereas shoreline hydro-morphological alteration was less important. Diatom indicators were most responsive to all three pressures, whereas macrophyte biomass and species number responded only to water level fluctuation. The Trophic Diatom Index for Lakes (TDIL) was negatively related to urbanization and wave exposure. This indicates that it is a suitable indicator for pressures related to urbanization, although a confounding effect of wave exposure is possible. Invertebrate abundance responded strongly to eutrophication, but the indicator based on taxonomic composition (Average Score Per Taxon) did not. Our results suggest that our metrics can be applied in Western Balkan lakes, despite the high number of endemic species present in some of these lakes. We argue that local water management should focus on abating the causes of eutrophication and water level fluctuation, whilst preserving sufficient lengths of undeveloped shoreline to ensure good water quality in the long run.

Environmental effects of offshore produced water discharges: A review focused on the Norwegian continental shelf.

Produced water (PW), a large byproduct of offshore oil and gas extraction, is reinjected to formations or discharged to the sea after treatment. The discharges contain dispersed crude oil, polycyclic aromatic hydrocarbons (PAHs), alkylphenols (APs), metals, and many other constituents of environmental relevance. Risk-based regulation, greener offshore chemicals and improved cleaning systems have reduced environmental risks of PW discharges, but PW is still the largest operational source of oil pollution to the sea from the offshore petroleum industry. Monitoring surveys find detectable exposures in caged mussel and fish several km downstream from PW outfalls, but biomarkers indicate only mild acute effects in these sentinels. On the other hand, increased concentrations of DNA adducts are found repeatedly in benthic fish populations, especially in haddock. It is uncertain whether increased adducts could be a long-term effect of sediment contamination due to ongoing PW discharges, or earlier discharges of oil-containing drilling waste. Another concern is uncertainty regarding the possible effect of PW discharges in the sub-Arctic Southern Barents Sea. So far, research suggests that sub-arctic species are largely comparable to temperate species in their sensitivity to PW exposure. Larval deformities and cardiac toxicity in fish early life stages are among the biomarkers and adverse outcome pathways that currently receive much attention in PW effect research. Herein, we summarize the accumulated ecotoxicological knowledge of offshore PW discharges and highlight some key remaining knowledge needs.

Littoral eutrophication indicators are more closely related to nearshore land use than to water nutrient concentrations: A critical evaluation of stressor-response relationships.

Biological assessment metrics and water chemistry measurements are used to quantify the link between stressors and their effects on lake ecosystems, for the Water Framework Directive. However, correlations between metrics and water chemistry are often poor. This is seen as major weaknesses of Water Framework Directive-related monitoring and assessment. We analyzed macrophytes, benthic algae, benthic macroinvertebrates, water chemistry and sediment total phosphorus content in the littoral of six lakes in the Western Balkans and used CORINE land use data to estimate nutrient enrichment via runoff from the adjacent land. Lakes with a higher estimated phosphorus runoff from the adjacent land did not have higher littoral water nutrient concentrations, but littoral diatom assemblages indicated more eutrophic conditions. These lakes also had higher abundances of littoral benthic primary producers, which in turn were associated with low concentrations of dissolved nutrients, but only in autumn, not in spring. This is consistent with primary producers taking up nutrients during the summer growth season. In lakes with high abundances of benthic primary producers, it is likely that the littoral vegetation plays a large role in the transfer of nutrients from the water to the benthos. This process impairs correlations between biological metrics and water nutrient concentrations. Our results suggest that CORINE land cover may be more useful to characterize littoral nutrient enrichment than lake water chemistry. Increased benthic primary producer biomasses and "eutrophic" diatom indices may indicate littoral nutrient enrichment even if water nutrient concentrations are low.

Possible adverse impact of contaminants on Atlantic cod population dynamics in coastal ecosystems.

While many in-laboratory ecotoxicological studies have shown the adverse impact of pollutants to the fitness of an individual, direct evidence from the field on the population dynamics of wildlife animals has been lacking. Here, we provide empirical support for a negative effect of pollution on Atlantic cod (Gadus morhua) population dynamics in coastal waters of Norway by combining unique time series of juvenile cod abundance, body size, environmental concentration of toxic contaminants and a spatially structured population dynamics model. The study shows that mercury concentration might have decreased the reproductive potential of cod in the region despite the general decline in the environmental concentration of mercury, cadmium and hexachlorobenzene since the implementation of national environmental laws. However, some cod populations appeared to be more resistant to mercury pollution than others, and the strength and shape of mercury effect on cod reproductive potential was fjord-specific. Additionally, cod growth rate changed at scales smaller than fjords with a gradient related to the exposure to the open ocean and offshore cod. These spatial differences in life-history traits emphasize the importance of local adaptation in shaping the dynamics of local wildlife populations. Finally, this study highlights the possibility to mitigate pollution effects on natural populations by reducing the overall pollution level, but also reveals that pollution reduction alone is not enough to rebuild local cod populations. Cod population recovery probably requires complementary efforts on fishing regulation and habitat restoration.

Levels and trends of tributyltin (TBT) and imposex in dogwhelk (Nucella lapillus) along the Norwegian coastline from 1991 to 2017.

The banning of organotin biocides, such as tributyltin (TBT), from use in marine antifouling paints is now leading to reproductive health recovery in marine gastropod populations all over the world. TBT induces so-called imposex (superimposition of male sexual characters onto females) in certain marine gastropods, such as the common dogwhelk Nucella lapillus. In this publication, the results of the Norwegian TBT and imposex monitoring in N. lapillus from the period 1991-2017 are presented. Significantly higher levels of TBT and imposex were measured in coastal areas close to shipping lanes along most of the coast prior to 2008 than afterwards. Levels started declining after restrictions were imposed on the use of TBT in all antifouling paint applications, with a total ban in 2008. In 2017, no sign of imposex was found in N. lapillus in any of the monitoring stations along the Norwegian coastline. Based on monitoring data shown herein, the importance of long-term biomonitoring and international chemical regulations, as well as the TBT and imposex story in general, are discussed.

Additive Models Reveal Sources of Metals and Organic Pollutants in Norwegian Marine Sediments.

We characterized spatial patterns of surface sediment concentrations of seven polychlorinated biphenyls (PCBs), seven polycyclic aromatic hydrocarbons (PAHs), three chlorinated pesticides, and five metals in Norwegian waters and Skagerrak. In total, we analyzed 5036 concentrations of 22 chemical substances that were measured between 1986 and 2014 at 333 sampling sites by means of generalized additive models (GAMs). We found that GAMs with organic carbon content of the sediment and latitude and longitude as co-variates explained as ca. 75% of the variability of the contaminant sediment concentrations. For metals, a predominantly hotspot-driven spatial pattern was found, i.e., we identified historical pollution hotspots (e.g., Sørfjord in western Norway) for mercury, zinc, cadmium, and lead. Highest concentrations of PAHs and PCBs were found close to densely populated and industrialized regions, i.e., in the North Sea and in the Kattegat and Skagerrak. The spatial pattern of the PCBs suggests the secondary and diffuse atmospheric nature of their sources. Atmospheric inputs are the main sources of pollution for most organic chemicals considered, but north of the Arctic circle, we found that concentrations of PAHs increased from south to north most likely related to a combination of coal-eroding bedrock and the biological pump. The knowledge acquired in the present research is essential for developing effective remediation strategies that are consistent with international conventions on pollution control.

Comparison of caged and native blue mussels (Mytilus edulis spp.) for environmental monitoring of PAH, PCB and trace metals.

Contaminant bioaccumulation was studied in blue mussels (Mytilus edulis spp.) using the harbor waters of Kristiansand (Norway) as a case study. A suite of chemical contaminants (trace metals, PAHs and PCBs) was analyzed in caged and native mussels as well as in passive samplers (Diffusive Gradients in Thin films (DGT)-devices and silicone rubbers) placed alongside the mussels for estimation of contaminant concentrations in water and uptake rates and bioaccumulation factors (BAFs) in mussels during a six-months deployment period. Estimated logBAFs were in the ranges 2.3-5.5, 3.8-5.2 and 3.2-4.4 for metals, PCBs and PAHs, respectively. Contaminant levels in caged mussels increased rapidly to stable levels for trace metals, whereas for hydrophobic organic contaminants the increase was steady but slow and for many compounds did not reach the levels observed in native mussels. Some key issues related to mussel caging design, such as mussel deployment time and confounding influence from seasonal fluctuations, are discussed herein.

Mercury concentration trend as a possible result of changes in cod population demography.

Mercury (Hg) in Atlantic cod (Gadus morhua) is one of many parameters that are monitored through OSPAR's Joint Assessment and Monitoring Programme. Time series for cod in the Inner Oslofjord (Norway) go back to 1984. Until 2014, annual median Hg-concentrations in cod from the Inner Oslofjord showed both significant upward long-term (whole time series) and short-term (recent 10 years) trends (when 2015 was included, the short-term trend was not significant). However, the median length of the cod sampled also showed upward trends. This may have been caused by low cod recruitment in the area since the start of the 2000s, as indicated by beach seine surveys. To investigate how length would impact the trend analysis, the Hg-concentrations in the cod were normalised to 50 cm. No significant short-term trend in Hg-concentrations could be detected for length-normalised concentrations. The results indicated that most of the upward trend in Hg-concentrations could be attributed to the sampling of larger fish. The reasons for the apparent change in the cod population demography are not conclusive, however, sampling bias must also be considered.

Rapid expansion of the invasive oyster Crassostrea gigas at its northern distribution limit in Europe: Naturally dispersed or introduced?

The Pacific oyster, Crassostrea gigas, was introduced to Europe for aquaculture purposes, and has had a rapid and unforeseen northward expansion in northern Europe. The recent dramatic increase in number of C. gigas populations along the species' northern distribution limit has questioned the efficiency of Skagerrak as a dispersal barrier for transport and survival of larvae. We investigated the genetic connectivity and possible spreading patterns between Pacific oyster populations on the southern Norwegian coast (4 localities) and Swedish and Danish populations by means of DNA microsatellite analysis of adult oysters, and by simulating larvae drift. In the simulations we used a 3D oceanographic model to explore the influence of recent climate change (1990-2010) on development, survival, and successful spreading of Danish and Swedish Pacific oyster larvae to Norwegian coastal waters. The simulations indicated adequate temperature conditions for development, survival, and settlement of larvae across the Skagerrak in warm years since 2000. However, microsatellite genotyping revealed genetic differences between the Norwegian populations, and between the Norwegian populations and the Swedish and Danish populations, the latter two populations being more similar. This patchwork pattern of genetic dissimilarity among the Norwegian populations points towards multiple local introduction routes rather than the commonly assumed unidirectional entry of larvae drifted from Denmark and Sweden. Alternative origins of introduction and implications for management, such as forecasting and possible mitigation actions, are discussed.

Cascading effects of mass mortality events in Arctic marine communities.

Mass mortality events caused by pulse anthropogenic or environmental perturbations (e.g., extreme weather, toxic spills or epizootics) severely reduce the abundance of a population in a short time. The frequency and impact of these events are likely to increase across the globe. Studies on how such events may affect ecological communities of interacting species are scarce. By combining a multispecies Gompertz model with a Bayesian state-space framework, we quantify community-level effects of a mass mortality event in a single species. We present a case study on a community of fish and zooplankton in the Barents Sea to illustrate how a mass mortality event of different intensities affecting the lower trophic level (krill) may propagate to higher trophic levels (capelin and cod). This approach is especially valuable for assessing community-level effects of potential anthropogenic-driven mass mortality events, owing to the ability to account for uncertainty in the assessed impact due to uncertainty about the ecological dynamics. We hence quantify how the assessed impact of a mass mortality event depends on the degree of precaution considered. We suggest that this approach can be useful for assessing the possible detrimental outcomes of toxic spills, for example oil spills, in relatively simple communities such as often found in the Arctic, a region under increasing influence of human activities due to increased land and sea use.

Toxicokinetics of pyrene in the freshwater alga Chara rudis.

Chara has been suggested a good model to study uptake of xenobiotics into cytoplasm due to their large internode cells surrounded by a layer of cortex cells. We studied the uptake and elimination of pyrene (nominal concentration of 5 µg L(-1)) in the freshwater alga Chara rudis during 22 days in two treatments mimicking epilimnetic (warm and light) and hypolimnetic (cold and dark) conditions. The growth of Chara during the exposure was higher in epilimnetic conditions (40%) compared to both hypolimnetic pyrene exposed Chara and controls (epilimnetic and hypolimnetic, no pyrene). In the water, a more rapid dissipation of pyrene was observed in epilimnetic conditions, possibly as a result of the increased algal growth. In the cortex, pyrene, 1-OH-pyrene (minor metabolite) and an unknown hydrophobic major metabolite was measured. Pyrene amounts decreased over time, while amounts of the unknown metabolite increased. In internode cytoplasm, pyrene and 1-OH-pyrene showed initially increasing followed by decreasing trends, while the unknown metabolite was not detected. The total mass balance showed that we were able to account for the applied pyrene until 4 days of exposure. However, after this time, there was a significant decrease in amounts accounted for by fluorescence, suggesting that the metabolism of pyrene involves degradation of the ring structure. The degradation was larger in epilimnetic than hypolimnetic conditions.

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.

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.

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

Population growth, and hence the population's 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.

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.

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.

Timing and abundance as key mechanisms affecting trophic interactions in variable environments.

Climatic changes are disrupting otherwise tight trophic interactions between predator and prey. Most of the earlier studies have primarily focused on the temporal dimension of the relationship in the framework of the match-mismatch hypothesis. This hypothesis predicts that predator's recruitment will be high if the peak of the prey availability temporally matches the most energy-demanding period of the predators breeding phenology. However, the match-mismatch hypothesis ignores the level of food abundance while this can compensate small mismatches. Using a novel time-series model explicitly quantifying both the timing and the abundance component for trophic relationships, we here show that timing and abundance of food affect recruitment differently in a marine (cod/zooplankton), a marine-terrestrial (puffin/herring) and a terrestrial (sheep/vegetation) ecosystem. The quantification of the combined effect of abundance and timing of prey on predator dynamics enables us to come closer to the mechanisms by which environment variability may affect ecological systems.

Competition among fishermen and fish causes the collapse of Barents Sea capelin.

The vast majority of the world's fisheries are typically managed within a single-species perspective, ignoring the dynamic feedback mechanisms generated by the ecological web of which they are a part. Here we show that the dynamics of the Barents Sea capelin (Mallotus villosus), the world's largest stock of this species, is strongly influenced by both within-system ecological feedback mechanisms and the impact of harvesting. Both overexploitation and predation by herring (Clupea harengus) can cause the population to collapse, whereas predation by cod (Gadus morhua) is demonstrated a delay in the stock's recovery after a collapse. Such collapses, which have occurred twice in 20 years, affect the entire Barents Sea ecosystem, a region that for ages has provided food for all of Europe.