Danish Waters Research Articles

Biosonar, dive, and foraging activity of satellite tracked harbor porpoises (Phocoena phocoena)

AbstractThis study presents bioacoustic recordings in combination with movements and diving behavior of three free‐ranging harbor porpoises (a female and two males) in Danish waters. Each porpoise was equipped with an acoustic data logger (A‐tag), a time‐depth‐recorder, a VHF radio transmitter, and a satellite transmitter. The units were programmed to release after 24 or 72 h. Possible foraging occurred mostly near the surface or at the bottom of a dive. The porpoises showed individual diversity in biosonar activity (<100 to >50,000 clicks per hour) and in dive frequency (6–179 dives per hour). We confirm that wild harbor porpoises use more intense clicks than captive animals. A positive tendency between number of dives and clicks per hour was found for a subadult male, which stayed near shore. It showed a distinct day‐night cycle with low echolocation rates during the day, but five times higher rates and higher dive activity at night. A female traveling in open waters showed no diel rhythm, but its sonar activity was three times higher compared to the males'. Considerable individual differences in dive and echolocation activity could have been influenced by biological and physical factors, but also show behavioral adaptability necessary for survival in a complex coastal environment.

Phenotypic and genetic divergence among harbour porpoise populations associated with habitat regions in the North Sea and adjacent seas

Determining the mechanisms that generate population structure is essential to the understanding of speciation and the evolution of biodiversity. Here, we investigate a geographical range that transects two habitat gradients, the North Sea to North Atlantic transition, and the temperate to subpolar regions. We studied the harbour porpoise (Phocoena phocoena), a small odontocete inhabiting both subpolar and temperate waters. To assess differentiation among putative populations, we measured morphological variation at cranial traits (N = 462 individuals) and variation at eight microsatellite loci for 338 of the same individuals from Norwegian, British and Danish waters. Significant morphological differentiation reflected the size of the buccal cavity. Porpoises forage in relatively shallow waters preying mainly on benthic species in British and Danish waters, and on mesopelagic and pelagic fish off the coast of Norway. We suggest that the observed differentiation may be explained by resource specialization and either adaptation or developmental responses to different local habitats.

Behavioural responses of harbour seals to human‐induced disturbances

ABSTRACT In Denmark, harbour seals, Phoca vitulina, were first protected in 1977, and since then a number of seal reserves have been established in Danish waters. The effectiveness of these reserves to prevent human‐induced disturbances to the seal population have, however, not been evaluated. To evaluate this, experimental disturbances were conducted in one of the most important seal reserves in Denmark (Anholt seal reserve). Specifically, the behavioural responses (alert distance, flight initiation distance, flee distances and flight duration) of harbour seals to approaching pedestrians and boats were studied. The project was conducted during three periods related to the breeding cycle of harbour seals. In all periods, harbour seals were alerted by boats at significantly greater distances compared with pedestrians (560–850 m and 200–425 m, respectively). Similar differences in the flight initiation distances were observed, 510–830 m for boats and 165–260 m for pedestrians. In most cases seals were alerted and began to flee when the approaching boat was outside the reserve, whereas seals did not respond to approaching pedestrians until after they had entered the reserve. Harbour seals exhibited weaker and shorter‐lasting responses during the breeding season. They were more reluctant to flee and returned to the haul‐out site immediately after being disturbed, in some cases even during the disturbance. This seasonal tolerance is most likely attributed to a trade‐off between fleeing and nursing during the breeding season, and hence not an indication of habituation. Based on the results of this study it is suggested that the reserve boundaries on land be placed at least 425 m from the haul‐out area and the boundary at sea should extend to at least 850 m from the haul‐out area in order to secure adequate year‐round protection from disturbances. Copyright © 2011 John Wiley & Sons, Ltd.

Spatial trends of perfluorochemicals in harbor seals (Phoca vitulina) from Danish waters

Spatial trends of concentrations of perfluorinated chemicals (PFCs) were investigated in harbour seal liver tissue from seven locations in Denmark, ranging from the Wadden Sea in the southern North Sea to the Western Baltic. All samples were collected during the phocine distemper epizootic in 2002 which provided access to a large number of comparable samples over a short time period. PFOS was dominating (mean: 92% of ∑PFC) among the PFCs in the samples, followed by considerably lower concentrations of PFHxS (1.8%), PFDA (1.7%), PFNA (1.6%) PFUnA (1.5%), PFOA (0.9%) and PFOSA (0.5%). The concentrations of all the investigated compounds showed significant differences among the seven locations. PFOS showed the highest concentrations in the Wadden Sea, where high burdens have also been recorded in German seals. Most compounds showed a trend towards higher concentrations at one or both extremes of the geographic range. Two different patterns of relative PFC concentrations were detected; one in the inner Danish waters where PFOSA and PFUnA were more prevalent and another in the Wadden Sea and Limfjord where PFOA, PFHxS and PFNA were found in greater proportions. These patterns probably represent Baltic and North Sea contamination sources.

Geographic variation of skeletal ontogeny and skull shape in the harbour porpoise (Phocoena phocoena)

All extant porpoise species show paedomorphic skeletal traits. These traits are hypothesized to be connected with rapid growth to reproductive size and could thus show geographic variation according to ecological circumstances. To investigate this, skeletal ontogeny was compared among harbour porpoises from California, West Greenland, the inner Danish waters, and the Sea of Azov. Porpoises from California grew to larger sizes than Danish porpoises, which were again larger than Greenlandic porpoises, whereas Azov porpoises were smallest. Size differences were largely attributable to differences in timing of offset of growth. Expression of paedomorphosis followed the same pattern among populations and sexes as adult sizes; Californian porpoises were less paedomorphic than the other populations across all assessed traits, whereas porpoises from Azov were the most paedomorphic. We propose that the larger size and less profound paedomorphism seen in Californian porpoises are attributable to fluctuation of prey availability, owing to variation in upwelling on which productivity in Californian waters depends. Skull shapes after correction for allometry were significantly different among all populations, Sea of Azov porpoises being most divergent. There was no overlap of skull shapes between the Atlantic, the Pacific, and Azov, supporting the current division of harbor porpoises ( Phocoena phocoena (L., 1758)) into three subspecies ( Phocoena phocoena phocoena (L., 1758), Phocoena phocoena relicta Abel, 1905, and Phocoena phocoena vomerina Gill, 1865).

Mapping porosity anomalies in deep Jurassic sandstones – an example from the Svane-1A area, Danish Central Graben

Hydrocarbon-bearing Upper Jurassic sandstone reservoirs at depths of more than 5000 m may form a future exploration target in the Danish Central Graben (Fig. 1). The Upper Jurassic sandstone play in the Danish sector has historically been less successful than in the neighbouring Norwegian and British sectors of the North Sea. This is mainly due to poor reservoir quality of the sandstones. However, the discovery in 2001 of an oil accumulation at a depth of more than 5000 m in the Svane-1 well has triggered renewed interest in the Upper Jurassic High Temperature – High Pressure (HTHP) sandstone play in Danish waters. The Jurassic plays comprise sandstone reservoirs deposited in a variety of environments, ranging from fluvial to deep marine.

Improvement of hydrodynamic forecasting of Danish waters: impact of low-frequency North Atlantic barotropic variations

The Danish Maritime Safety Administration (DaMSA) provides forecast of elevations, currents, and other parameters to the maritime society. Accurate and reliable predictions are important to help navigate Danish waters in a safe manner, and the forecasts are routinely used by the Vessel Traffic Services in the Great Belt and the Sound. The DaMSA model setup includes three nested models, with coarse resolution in the North Atlantic and increasing to 600 m in the Belt Sea and South West Baltic. Observations of some special events in late 2009 drew attention to a possible relation between Atlantic-scale surge events and small-scale currents in the Danish Straits. During the special event with large-scale surge, the observed southward moving current in the Danish Straits was 0.5–2.0 m/s for several days, while the operational model showed a much smaller response. As a consequence, the entire DaMSA model complex was revised during 2010. Multi-annual reruns have showed that with the updated model, the explained variance of the current increases from 67% to 88%.

Cooperation During the Designation and Management of the Dogger Bank

The Dogger Bank is a potential transboundary marine protected area (MPA) located in Dutch, German, Danish and UK waters. International and EU legal instruments contain provisions requiring cooperation during the designation and pro- tection/management of the Dogger Bank as an MPA. Many of the cooperation requirements are not specified; therefore, Member States are left with a certain discretion as to the extent and form of the implementation of the required cooperation. Since this cooperation began taking place, the required standards seem to be being met. However, the exact meaning of the requirement for cooperation included in the Marine Strategy Framework Directive (MSFD) is questionable; it remains to be seen whether the current cooperation complies with this Directive. In order to optimise the current cooperation practice, an increasing role for the European Commission (EC) and the Habitats Committee within the process of the designation and management of the Dogger Bank as an MPA could be strived for. Furthermore, when looking at the relevant legal instru- ments an amendment of Oslo-Paris (OSPAR) Recommendation 2003/3 provides an option to further establish a specific legal basis for cooperation.

Acoustic surveys confirm the high-density areas of harbour porpoises found by satellite tracking

AbstractSveegaard, S., Teilmann, J., Berggren, P., Mouritsen, K. N., Gillespie, D., and Tougaard, J. 2011. Acoustic surveys confirm the high-density areas of harbour porpoises found by satellite tracking. – ICES Journal of Marine Science, 68: 929–936. The population status of the harbour porpoise (Phocoena phocoena) has long been of concern in European waters. Consequently, the European Commission (EC) Habitats Directive obligates all EC member states to designate marine protected areas (MPAs) for harbour porpoises before 2012. These will be designated areas having the greatest density of porpoises. However, little is known about comparability between the monitoring methods used to examine porpoise distribution and density, and conflicting results may arise, especially when considering their varying sample size and temporal and spatial scales. Here, vessel-based acoustic surveys are seen as an independent method of testing the temporal and spatial permanence of previously identified areas of high density of harbour porpoises found by satellite-tracking them in inner Danish waters. Based on six acoustic surveys, a strong spatial accord was found between the number of acoustic detections of harbour porpoises and their density distribution obtained from 10 years of satellite tracking. The results confirm the presence and permanence of areas of high density of porpoises and validate the two methods for identifying and monitoring future MPAs for the species.

Temperature affects the timing of spawning and migration of North Sea mackerel

Climate change accentuates the need for knowing how temperature impacts the life history and productivity of economically and ecologically important species of fish. We examine the influence of temperature on the timing of the spawning and migrations of North Sea Mackerel using data from larvae CPR surveys, egg surveys and commercial landings from Danish coastal fisheries in the North Sea, Skagerrak, Kattegat and inner Danish waters. The three independent sources of data all show that there is a significant relationship between the timing of spawning and sea surface temperature. Large mackerel are shown to arrive at the feeding areas before and leave later than small mackerel and the sequential appearance of mackerel in each of the feeding areas studied supports the anecdotal evidence for an eastward post-spawning migration. Occasional commercial catches taken in winter in the Sound N, Kattegat and Skagerrak together with catches in the first quarter IBTS survey furthermore indicate some overwintering here. Significant relationships between temperature and North Sea mackerel spawning and migration have not been documented before. The results have implications for mackerel resource management and monitoring. An increase in temperature is likely to affect the timing and magnitude of the growth, recruitment and migration of North Sea mackerel with subsequent impacts on its sustainable exploitation.

Hematodinium sp. (Alveolata, Syndinea) detected in marine decapod crustaceans from waters of Denmark and Greenland

Five decapod crustacean species were examined for presence of the parasitic dinoflagellate Hematodinium spp. (Alveolata, Syndinea) by morphological methods (colour and pleopod methods) as well as by PCR and nested PCR with Hematodinium-specific primers. Nephrops norvegicus, Pagurus bernhardus and Liocarcinus depurator were sampled by trawling in Danish waters and Chionoecetes opilio and Hyas araneus were sampled by trapping off the west coast of Greenland. The existence of Hematodinium has not previoiusly been documented in Danish waters, but it was detected in all 3 decapod species examined in the present study. Hematodinium sp. was also detected for the first time in H. araneus and the existence of Hematodinium sp. in Greenlandic C. opilio was documented by PCR. Analyses of 26 Hematodinium sp. ITS1 sequences, including sequences from all 5 host species sampled, revealed more than 95% sequence similarity between 24 of the sequences. Two Hematodinium sp. ITS1 sequences from C. opilio were only 81% similar to the 24 other ITS1 sequences. The nested PCR approach resulted in the highest reported percentages of positive samples for Hematodinium sp.in the hosts investigated (between 45 and 87.5%). However, no decapods were found to be infected with Hematodinium sp. based on morphological methods. Consequently, Hematodinium sp. may be more common than previously believed, and, assuming that the DNA found originated from viable and infectious parasite cells, infections may not always be fatal. We suggest that the hosts investigated may have been subject to latent infections that could develop into a fatal disease only if the hosts were physiologically stressed due to other factors.

Modelling spatial patterns in harbour porpoise satellite telemetry data using maximum entropy

The distribution of harbour porpoises in EU waters is poorly understood, and modelled predictions of their distributions could inform the strategic spatial planning of future exploitation of the marine environment to avoid potential conflicts. We analysed satellite telemetry data from 39 harbour porpoises Phocoena phocoena in inner Danish waters using a modelling tool rooted in maximum entropy: Maxent. Maxent does not require absence data and has been shown to be effective for data characterised by small sample size, sampling bias and locational errors. For each season we used an iterative bootstrapping procedure to randomly select among the most precise records from each of the 39 tagged individuals, and ran Maxent on pooled records based on explanatory environmental variables hypothesised to serve as good proxies for harbour porpoise prey abundance. Among our environmental variables, distance to coast and bottom salinity had the most explanatory power, and their response shapes were relatively consistent across most seasons. The predictive power of the models (assessed by ROC‐AUC) ranged from 0.70 to 0.86 within seasons. The southern Kattegat, the Belt Seas, most western part of the Baltic Sea and the Sound were predicted to have relatively high probabilities of occurrence across seasons. In contrast, the central part of Kattegat and the Baltic Sea south and east of Limhamn and Darss Ridge consistently showed low probabilities of occurrence. Areas with the lowest probabilities of occurrence were generally characterised by high predictive uncertainty. Our methods have implications for the analyses of satellite tagged animals in terrestrial and marine environments. By coupling a bootstrapping procedure with Maxent we circumvented some of the statistical challenges presented by satellite telemetry data to generate spatial predictions within the inner Danish waters.

Late Pleistocene and Holocene whale remains (Cetacea) from Denmark and adjacent countries: Species, distribution, chronology, and trace element concentrations

We describe and review the subfossil whale bones (mammalian order Cetacea) material from the southern Scandinavian area, that is, Skagerrak, Kattegat, the inner Danish waters and the southwestern Baltic Sea. Fifteen species were identified from the subfossil records of which all, except for the bowhead whale (Balaena mysticetus), have also been encountered in the modern times. Fifty-one specimens were radiocarbon dated covering 12 of the subfossil species. The dates fell in three distinct clusters with a few specimens before the last glacial maximum (LGM), a large group between LGM and the Pleistocene/Holocene boundary (ca. 17.0–11.7 cal. kyr BP), and another large group from ca. 8.0 cal. kyr BP onward. Seventeen of the radiocarbon dated specimens have been subjected to trace element analysis by Instrumental Neutron Activation Analysis. Cross plots of the concentrations of Fe and Zn, and Fe and Co show that it is possible to distinguish crayfish eaters from fish/squid eaters. This can be used as a novel and independent method for the determination to species of whale remains of otherwise uncertain speciation.

Temperature effects on copepod egg hatching: does acclimatization matter?

This report investigates female sizes, egg sizes and egg hatching rates in relation to temperature for the near-shore calanoid copepod Acartia tonsa cultured at 6, 9, 14 and 24°C for several generations to achieve acclimatization. Inverse size relationships of eggs and females were revealed with increasing temperature. Eggs produced at 6°C were 85 ± 4 μm in diameter, but decreased to 80 ± 3 μm at 24°C. Female cephalothorax length was 840 ± 52 and 692 ± 39 μm at 9 and 24°C, respectively Parallel hatching experiments were performed between non-acclimatized and acclimatized cultures across a range of temperatures reflecting natural conditions in Danish waters. A greater fraction of eggs enter quiescence as temperature declines. Eggs were able to hatch at temperatures as low as 1.5°C. Final egg hatching success increased with temperature. Acclimatization of the copepods resulted in a lower maximum hatching rate, but a higher final hatching success at 9 and 6°C compared with egg hatching from non-acclimatized copepods. Eggs develop significantly better at low temperatures when compared with females in culture. This adaptation suggests that the observed annual population fluctuations in, for example, Danish waters can be explained as follows: the population of copepods in the pelagic virtually die out during the autumn―winter months; the population the following year emerges from resting eggs from the sediment and is produced by the pelagic copepods the previous year(s).

Nematode infections of maricultured and wild fishes in Danish waters: A comparative study

Rainbow trout ( Oncorhynchus mykiss) from a Danish mariculture system (a total of 166 fish) and from a freshwater system (a total of 40 fish before transfer to the marine system) were investigated for the presence of nematode larvae through a 1 year cycle. Correspondingly, wild fishes (a total of 160 fish) (cod, whiting, saithe, haddock, hake, flounder, turbot, common sole, lemon sole, witch, plaice, herring, mackerel, garfish, angler fish, and European eel) obtained from the same geographic region were examined. Morphological and molecular techniques were applied for diagnosis. No nematode larvae were recovered from the aquacultured fish. In contrast, anisakid larvae ( Anisakis simplex) occurred frequently in wild fishes (body cavity and musculature) although at varying intensity. Other anisakid species ( Pseudoterranova decipiens and Contracaecum sp.) were also found in the wild fishes but at lower frequency. Examination of the stomach content from the cultured trout confirmed that aquacultured fish preferentially feed on pelleted dry feed although algae, invertebrates and small fish occurred in a few cases. The background for the absence of nematode larvae in maricultured rainbow trout (low infection risk due to feeding using heat-treated feed without any infective parasite larvae) and the implications for food safety issues (lower likelihood of infected fish from mariculture systems) are discussed.

Seasonal succession and composition of the zooplankton community along an eutrophication and salinity gradient exemplified by Danish waters

The seasonal distribution of the major components of the zooplankton community, protozooplankton, copepods and cladocerans, along a eutrophication gradient were examined in order to establish if eutrophication through increases in phytoplankton biomass and productivity has an impact on biomass and composition of the zooplankton community. Data on salinity, temperature, inorganic nutrients, chlorophyll a, phytoplankton biomass and production, protozooplankton and mesozoo-plankton biomass were collected during routine sampling at seven stations representing a eutrophication gradient from open to estuarine waters. Mesozooplankton biomass varied seasonally in all areas with a unimodal pattern characterized by a peak between mid-spring and mid-summer. Total mesozooplankton biomass as well as copepod and cladoceran biomass, decreased with DIN concentration. Salinity and the type of ecoystem (well-mixed shallow estuaries versus stratified open waters) were the major factors controlling the biomass proportion of copepod species. Annual production of the total copepod community as well as the total grazing impact of copepods on primary production was higher in open waters than in estuarine waters. In estuarine type ecosystems, phytoplankton production is underexploited by copepod grazing whereas in the open type ecosystems, the phytoplankton production alone could not satisfy the carbon demand of copepods stressing the potential importance of protozoans in the copepod diet.

Oxygen deficiency impacts on burying habitats for lesser sandeel, Ammodytes tobianus, in the inner Danish waters

Starting in 1980s, the inner Danish waters have yearly been exposed to seasonal oxygen deficiency (hypoxia). Through spatial–temporal interpolation of monitoring data (1998–2005), we investigated oxygen deficiency impacts on suitable burying habitats for lesser sandeel ( Ammodytes tobianus ). Furthermore, the consequences of a predicted 4 °C temperature increase within this century were investigated. Maps of bottom oxygen deficiency (oxygen saturation ≤ Scrit of sandeel) were overlaid on maps of sediment composition. Throughout the study period (1998–2005), about 8% of the suitable sediments were affected by oxygen deficiency during an average year and 23% in the most severe year. Regional differences underlay the interannual variations. The extent of oxygen deficiency in enclosed regions varied from 20% to 33% of the suitable seabed being affected, whereas in open-water regions oxygen deficiency problems were limited during average years. However, large areas of the open-water seabed experienced oxygen deficiency during severe years. In such years, under a 4.0 °C temperature increase scenario, the extent of oxygen deficiency on open-water suitable patches was predicted to increase from 25% to about 40%.

Genetic diversity and phylogenetic analysis of the attachment glycoprotein of phocine distemper viruses of the 2002 and 1988 epizootics

To investigate the possible origin and spread of the dramatic re-emergent 2002 distemper epizootic observed among seals in Danish Waters, we have sequenced wild-type genes of the attachment (H) glycoproteins of viruses from both the 2002 and 1988 epizootics. Phylogenetic analysis of the H genes of phocine distemper virus (PDV) together with other morbilliviruses, suggests that the re-emergent 2002 PDV is more closely related to a putative recent ancestral PDV than the 1988 PDV isolates. Moreover, upsurges of distemper disease in land-living carnivores linked in time and locality to the 2002 seal epizootic in Danish Waters was investigated and determined to be caused by canine distemper virus, the closest relative of PDV, revealing no direct epidemiological link to the seal epizootics.

Reference conditions for phytoplankton at Danish Water Framework Directive intercalibration sites

Phytoplankton is one of the biological quality elements included in the EU Water Framework Directive (WFD). Classification of water quality according to the WFD is based on the deviation of the present conditions from reference conditions. Given the lack of data from pristine conditions, this study used approximately 100-year-old measurements of Secchi depths from Danish waters in combination with relationships between Secchi depth and chlorophyll a (as a proxy for phytoplankton biomass) obtained from recent monitoring to calculate ‘historical’ or reference chlorophyll a (Chl-a) concentrations. Historical Secchi depth data were available for 9 out of the 11 Danish WFD intercalibration sites. At eight of the sites, reference summer (May-September) Chl-a concentrations were in the range 0.7–1.2 µg l−1. At one site, west of Bornholm in the western Baltic Sea, historical Secchi depth measurements date back to only the late 1950s corresponding to a calculated Chl-a concentration of 1.3 µg l−1. This value cannot be considered representative of reference conditions.

Assimilating temperature and salinity profile observations using an anisotropic recursive filter in a coastal ocean model

In order to improve the ocean forecasting in the North Sea and Baltic Sea, an assimilation scheme based on a bottom-topography-dependent anisotropic recursive filter has been used in this study. This scheme can stretch or flatten the shape of a local representative contour surface of the background error covariance function into the form of an ellipse. Furthermore, the computing efficiency has been largely improved due to implicit computation of the background error covariance. A two-month experiment has been used for verifying the impact of assimilating ocean profile observations on ocean forecasting. The results indicate that the use of temperature and salinity profiles can largely improve the oceanic forecasting. The root mean square differences between the forecasts and observations for temperature and salinity have been reduced by 36% and 18% in the experiment period, respectively. Moreover, it is found that the anisotropic recursive filter approach is especially efficient in areas with complex coastlines and sharp fronts, e.g., inner Danish waters. The results also show that the propagation of observation information from an observation position to its neighboring grid points is closely related to currents.