Low hunting costs in an expensive marine mammal predator.

Many large terrestrial mammalian predators use energy-intensive, high-risk, high-gain strategies to pursue large, high-quality prey. However, similar-sized marine mammal predators with even higher field metabolic rates (FMRs) consistently target prey three to six orders of magnitude smaller than themselves. Here, we address the question of how these active and expensive marine mammal predators can gain sufficient energy from consistently targeting small prey during breath-hold dives. Using harbor porpoises as model organisms, we show that hunting small aquatic prey is energetically cheap (<20% increase in FMR) for these marine predators, but it requires them to spend a large proportion (>60%) of time foraging. We conclude that this grazing foraging strategy on small prey is viable for marine mammal predators despite their high FMR because they can hunt near continuously at low marginal expense. Consequently, cessation of foraging due to human disturbance comes at a high cost, as porpoises must maintain their high thermoregulation costs with a reduced energy intake.

Rerouting of a major shipping lane through important harbour porpoise habitat caused no detectable change in annual occurrence or foraging patterns.

Shipping is one of the largest industries globally, with well-known negative impacts on the marine environment. Despite the known negative short-term (minutes to hours) impact of shipping on individual animal behavioural responses, very little is understood about the long-term (months to years) impact on marine species presence and area use. This study took advantage of a planned rerouting of a major shipping lane leading into the Baltic Sea, to investigate the impact on the presence and foraging behaviour of a marine species known to be sensitive to underwater noise, the harbour porpoise (Phocoena phocoena). Passive acoustic monitoring data were collected from 15 stations over two years. Against predictions, no clear change occurred in monthly presence or foraging behaviour of the porpoises, despite the observed changes in noise and vessel traffic. However, long-term heightened noise levels may still impact communication, echolocation, or stress levels of individuals, and needs further investigation.

Environmental impact of the explosion of the Nord Stream pipelines.

Armed conflicts have, in addition to severe impacts on human lives and infrastructure, also impacts on the environment, which needs to be assessed and documented. On September the 26th 2022, unknown perpetrators deliberately ruptured the two gas pipelines Nord Stream 1 and 2 with four coordinated explosions near a major chemical munition dump site near the Danish island of Bornholm in the Baltic Sea. While the massive release of natural gas into atmosphere raised serious concerns concerning the contribution to climate change-this paper assesses the overlooked direct impact of the explosions on the marine ecosystem. Seals and porpoises within a radius of four km would be at high risk of being killed by the shockwave, while temporary impact on hearing would be expected up to 50 km away. As the Baltic Proper population of harbour porpoises (Phocoena phocoena) is critically endangered, the loss or serious injury of even a single individual is considered a significant impact on the population. The rupture moreover resulted in the resuspension of 250000 metric tons of heavily contaminated sediment from deep-sea sedimentary basin for over a week, resulting in unacceptable toxicological risks towards fish and other biota in 11 km3 water in the area for more than a month.

Ship noise causes tagged harbour porpoises to change direction or dive deeper.

Shipping is the most pervasive source of marine noise pollution globally, yet its impact on sensitive fauna remains unclear. We tracked 10 harbour porpoises for 5-10 days to determine exposure and behavioural reactions to modelled broadband noise (10 Hz-20 kHz, VHF-weighted) from individual ships monitored by AIS. Porpoises spent a third of their time experiencing ship noise above ambient, to which they regularly reacted by moving away during daytime and diving deeper during night. However, even ships >2 km away (noise levels of 93 ± 14 dB re 1 µPa2) caused animals to react 5-9 % of the time (∼18.6 ships/day). Ships can thus influence the behaviour and habitat use of cetaceans over long distances, with worrying implications for fitness in coastal areas where anthropogenic noise from dense ship traffic repeatedly disrupt their natural behaviour.

Are European Blue Economy ambitions in conflict with European environmental visions?

We report the outcomes of a comprehensive study of the potential consequences of the implementation of the EU Maritime Spatial Planning Directive (MSPD) in Danish waters. The analyses are anchored in a framework developed in support of data-driven Ecosystem-Based Maritime Spatial Planning. The data for the models include not only human stressors but also information on the distribution of ecosystem components ranging from planktonic communities over benthic communities to fish, seabirds and marine mammals. We have established a baseline, based on state-of-the-art data sets, with respect to combined effects upon ecosystem components. Future scenarios for the developments in human stressors were estimated for 2030 and 2050 based on information on existing policies, strategies and plans and were compared to the baseline. In addition, we developed a scenario for implementation of the Marine Strategy Framework Directive (MSFD), i.e. working towards meeting the objectives of Good Environmental Status. Our results indicate that (1) combined human stressors will possibly increase in 2030 and 2050 compared to the baseline, (2) increased combined human stressors are likely to lead to a worsening of the environmental and ecological status sensu the Marine Strategy Framework Directive and the Water Framework Directive (WFD), and (3) the MSPD implementation process appears to conflict with the MSFD and WFD objectives. Accordingly, we are sceptical of claims of an untapped potential for Blue Growth in Danish marine waters.

Screening for Influenza and Morbillivirus in Seals and Porpoises in the Baltic and North Sea.

Historically, the seals and harbour porpoises of the Baltic Sea and North Sea have been subjected to hunting, chemical pollutants and repeated mass mortalities, leading to significant population fluctuations. Despite the conservation implications and the zoonotic potential associated with viral disease outbreaks in wildlife, limited information is available on the circulation of viral pathogens in Baltic Sea seals and harbour porpoises. Here, we investigated the presence of the influenza A virus (IAV), the phocine distemper virus (PDV) and the cetacean morbillivirus (CeMV) in tracheal swabs and lung tissue samples from 99 harbour seals, 126 grey seals, 73 ringed seals and 78 harbour porpoises collected in the Baltic Sea and North Sea between 2002-2019. Despite screening 376 marine mammals collected over nearly two decades, we only detected one case of PDV and two cases of IAV linked to the documented viral outbreaks in seals in 2002 and 2014, respectively. Although we find no evidence of PDV and IAV during intermediate years, reports of isolated cases of PDV in North Sea harbour seals and IAV (H5N8) in Baltic and North Sea grey seals suggest introductions of those pathogens within the sampling period. Thus, to aid future monitoring efforts we highlight the need for a standardized and continuous sample collection of swabs, tissue and blood samples across Baltic Sea countries.

Forecasting shifts in habitat suitability of three marine predators suggests a rapid decline in inter-specific overlap under future climate change.

Understanding how environmental and climate change can alter habitat overlap of marine predators has great value for the management and conservation of marine ecosystems. Here, we estimated spatiotemporal changes in habitat suitability and inter-specific overlap among three marine predators: Baltic gray seals (Halichoerus grypus), harbor seals (Phoca vitulina), and harbor porpoises (Phocoena phocoena) under contemporary and future conditions. Location data (>200 tagged individuals) were collected in the southwestern region of the Baltic Sea; one of the fastest-warming semi-enclosed seas in the world. We used the maximum entropy (MaxEnt) algorithm to estimate changes in total area size and overlap of species-specific habitat suitability between 1997-2020 and 2091-2100. Predictor variables included environmental and climate-sensitive oceanographic conditions in the area. Sea-level rise, sea surface temperature, and salinity data were taken from representative concentration pathways [RCPs] scenarios 6.0 and 8.5 to forecast potential climate change effects. Model output suggested that habitat suitability of Baltic gray seals will decline over space and time, driven by changes in sea surface salinity and a loss of currently available haulout sites following sea-level rise in the future. A similar, although weaker, effect was observed for harbor seals, while suitability of habitat for harbor porpoises was predicted to increase slightly over space and time. Inter-specific overlap in highly suitable habitats was also predicted to increase slightly under RCP scenario 6.0 when compared to contemporary conditions, but to disappear under RCP scenario 8.5. Our study suggests that marine predators in the southwestern Baltic Sea may respond differently to future climatic conditions, leading to divergent shifts in habitat suitability that are likely to decrease inter-specific overlap over time and space. We conclude that climate change can lead to a marked redistribution of area use by marine predators in the region, which may influence local food-web dynamics and ecosystem functioning.

Estimating the abundance of the critically endangered Baltic Proper harbour porpoise (Phocoena phocoena) population using passive acoustic monitoring.

Knowing the abundance of a population is a crucial component to assess its conservation status and develop effective conservation plans. For most cetaceans, abundance estimation is difficult given their cryptic and mobile nature, especially when the population is small and has a transnational distribution. In the Baltic Sea, the number of harbour porpoises (Phocoena phocoena) has collapsed since the mid-20th century and the Baltic Proper harbour porpoise is listed as Critically Endangered by the IUCN and HELCOM; however, its abundance remains unknown. Here, one of the largest ever passive acoustic monitoring studies was carried out by eight Baltic Sea nations to estimate the abundance of the Baltic Proper harbour porpoise for the first time. By logging porpoise echolocation signals at 298 stations during May 2011-April 2013, calibrating the loggers' spatial detection performance at sea, and measuring the click rate of tagged individuals, we estimated an abundance of 71-1105 individuals (95% CI, point estimate 491) during May-October within the population's proposed management border. The small abundance estimate strongly supports that the Baltic Proper harbour porpoise is facing an extremely high risk of extinction, and highlights the need for immediate and efficient conservation actions through international cooperation. It also provides a starting point in monitoring the trend of the population abundance to evaluate the effectiveness of management measures and determine its interactions with the larger neighboring Belt Sea population. Further, we offer evidence that design-based passive acoustic monitoring can generate reliable estimates of the abundance of rare and cryptic animal populations across large spatial scales.

Integrating disparate datasets to model the functional response of a marine predator: A case study of harbour porpoises in the southern North Sea.

Quantifying consumption and prey choice for marine predator species is key to understanding their interaction with prey species, fisheries, and the ecosystem as a whole. However, parameterizing a functional response for large predators can be challenging because of the difficulty in obtaining the required data on predator diet and on the availability of multiple prey species.This study modeled a multi-species functional response (MSFR) to describe the relationship between consumption by harbour porpoises (Phocoena phocoena) and the availability of multiple prey species in the southern North Sea. Bayesian methodology was employed to estimate MSFR parameters and to incorporate uncertainties in diet and prey availability estimates. Prey consumption was estimated from stomach content data from stranded harbour porpoises. Prey availability to harbour porpoises was estimated based on the spatial overlap between prey distributions, estimated from fish survey data, and porpoise foraging range in the days prior to stranding predicted from telemetry data.Results indicated a preference for sandeels in the study area. Prey switching behavior (change in preference dependent on prey abundance) was confirmed by the favored type III functional response model. Variation in the size of the foraging range (estimated area where harbour porpoises could have foraged prior to stranding) did not alter the overall pattern of the results or conclusions.Integrating datasets on prey consumption from strandings, predator foraging distribution using telemetry, and prey availability from fish surveys into the modeling approach provides a methodological framework that may be appropriate for fitting MSFRs for other predators.

A risk assessment of the effects of mercury on Baltic Sea, Greater North Sea and North Atlantic wildlife, fish and bivalves.

A wide range of species, including marine mammals, seabirds, birds of prey, fish and bivalves, were investigated for potential population health risks resulting from contemporary (post 2000) mercury (Hg) exposure, using novel risk thresholds based on literature and de novo contamination data. The main geographic focus is on the Baltic Sea, while data from the same species in adjacent waters, such as the Greater North Sea and North Atlantic, were included for comparative purposes. For marine mammals, 23% of the groups, each composing individuals of a specific sex and maturity from the same species in a specific study region, showed Hg-concentrations within the High Risk Category (HRC) and Severe Risk Category (SRC). The corresponding percentages for seabirds, fish and bivalves were 2.7%, 25% and 8.0%, respectively, although fish and bivalves were not represented in the SRC. Juveniles from all species showed to be at no or low risk. In comparison to the same species in the adjacent waters, i.e. the Greater North Sea and the North Atlantic, the estimated risk for Baltic populations is not considerably higher. These findings suggest that over the past few decades the Baltic Sea has improved considerably with respect to presenting Hg exposure to its local species, while it does still carry a legacy of elevated Hg levels resulting from high neighbouring industrial and agricultural activity and slow water turnover regime.

Environmental drivers of harbour porpoise fine-scale movements.

Quantifying intraspecific variation in movement behaviour of marine predators and the underlying environmental drivers is important to inform conservation management of protected species. Here, we provide the first empirical data on fine-scale movements of free-ranging harbour porpoises (Phocoena phocoena) in their natural habitat. Data were obtained from six individuals, tagged in two areas of the Danish North Sea, that were equipped with Global Positioning System (GPS) and dive recorder units (V-tags). We used multi-model inference and model averaging to evaluate the relative importance of various static and dynamic environmental conditions on the movement characteristics: speed, turning angle, dive duration, dive depth, dive wiggliness (a proxy for prey chasing behaviour), and post-dive duration. Despite substantial individual differences in horizontal and vertical movement patterns, we found that all the tracked porpoises responded similar to variation in environmental conditions and displayed movements that indicate a higher likelihood of foraging behaviour in shallower and more saline waters. Our study contributes to the identification of important feeding areas for porpoises and can be used to improve existing movement-based simulation models that aim to assess the impact of anthropogenic disturbance on harbour porpoise populations.

Fine-scale movement responses of free-ranging harbour porpoises to capture, tagging and short-term noise pulses from a single airgun.

Knowledge about the impact of anthropogenic disturbances on the behavioural responses of cetaceans is constrained by lack of data on fine-scale movements of individuals. We equipped five free-ranging harbour porpoises (Phocoena phocoena) with high-resolution location and dive loggers and exposed them to a single 10 inch3 underwater airgun producing high-intensity noise pulses (2-3 s intervals) for 1 min. All five porpoises responded to capture and tagging with longer, faster and more directed movements as well as with shorter, shallower, less wiggly dives immediately after release, with natural behaviour resumed in less than or equal to 24 h. When we exposed porpoises to airgun pulses at ranges of 420-690 m with noise level estimates of 135-147 dB re 1 µPa2s (sound exposure level), one individual displayed rapid and directed movements away from the exposure site and two individuals used shorter and shallower dives compared to natural behaviour immediately after exposure. Noise-induced movement typically lasted for less than or equal to 8 h with an additional 24 h recovery period until natural behaviour was resumed. The remaining individuals did not show any quantifiable responses to the noise exposure. Changes in natural behaviour following anthropogenic disturbances may reduce feeding opportunities, and evaluating potential population-level consequences should be a priority research area.

Assessing auditory evoked potentials of wild harbor porpoises (Phocoena phocoena).

Testing the hearing abilities of marine mammals under water is a challenging task. Sample sizes are usually low, thus limiting the ability to generalize findings of susceptibility towards noise influences. A method to measure harbor porpoise hearing thresholds in situ in outdoor conditions using auditory steady state responses of the brainstem was developed and tested. The method was used on 15 live-stranded animals from the North Sea during rehabilitation, shortly before release into the wild, and on 12 wild animals incidentally caught in pound nets in Denmark (inner Danish waters). Results indicated that although the variability between individuals is wide, the shape of the hearing curve is generally similar to previously published results from behavioral trials. Using 10-kHz frequency intervals between 10 and 160 kHz, best hearing was found between 120 and 130 kHz. Additional testing using one-third octave frequency intervals (from 16 to 160 kHz) allowed for a much faster hearing assessment, but eliminated the fine scale threshold characteristics. For further investigations, the method will be used to better understand the factors influencing sensitivity differences across individuals and to establish population-level parameters describing hearing abilities of harbor porpoises.

Comparing Distribution of Harbour Porpoises (Phocoena phocoena) Derived from Satellite Telemetry and Passive Acoustic Monitoring.

Cetacean monitoring is essential in determining the status of a population. Different monitoring methods should reflect the real trends in abundance and patterns in distribution, and results should therefore ideally be independent of the selected method. Here, we compare two independent methods of describing harbour porpoise (Phocoena phocoena) relative distribution pattern in the western Baltic Sea. Satellite locations from 13 tagged harbour porpoises were used to build a Maximum Entropy (MaxEnt) model of suitable habitats. The data set was subsampled to one location every second day, which were sufficient to make reliable models over the summer (Jun-Aug) and autumn (Sep-Nov) seasons. The modelled results were compared to harbour porpoise acoustic activity obtained from 36 static acoustic monitoring stations (C-PODs) covering the same area. The C-POD data was expressed as the percentage of porpoise positive days/hours (the number of days/hours per day with porpoise detections) by season. The MaxEnt model and C-POD data showed a significant linear relationship with a strong decline in porpoise occurrence from west to east. This study shows that two very different methods provide comparable information on relative distribution patterns of harbour porpoises even in a low density area.

Ultra-High Foraging Rates of Harbor Porpoises Make Them Vulnerable to Anthropogenic Disturbance.

The question of how individuals acquire and allocate resources to maximize fitness is central in evolutionary ecology. Basic information on prey selection, search effort, and capture rates are critical for understanding a predator's role in its ecosystem and for predicting its response to natural and anthropogenic disturbance. Yet, for most marine species, foraging interactions cannot be observed directly. The high costs of thermoregulation in water require that small marine mammals have elevated energy intakes compared to similar-sized terrestrial mammals [1]. The combination of high food requirements and their position at the apex of most marine food webs may make small marine mammals particularly vulnerable to changes within the ecosystem [2-4], but the lack of detailed information about their foraging behavior often precludes an informed conservation effort. Here, we use high-resolution movement and prey echo recording tags on five wild harbor porpoises to examine foraging interactions in one of the most metabolically challenged cetacean species. We report that porpoises forage nearly continuously day and night, attempting to capture up to 550 small (3-10 cm) fish prey per hour with a remarkable prey capture success rate of >90%. Porpoises therefore target fish that are smaller than those of commercial interest, but must forage almost continually to meet their metabolic demands with such small prey, leaving little margin for compensation. Thus, for these "aquatic shrews," even a moderate level of anthropogenic disturbance in the busy shallow waters they share with humans may have severe fitness consequences at individual and population levels.

Impacts of Underwater Noise on Marine Vertebrates: Project Introduction and First Results.

The project conducts application-oriented research on impacts of underwater noise on marine vertebrates in the North and Baltic Seas. In distinct subprojects, the hearing sensitivity of harbor porpoises and gray seals as well as the acoustic tolerance limit of harbor porpoises to impulsive noise from pile driving and stress reactions caused by anthropogenic noise is investigated. Animals are equipped with DTAGs capable of recording the actual surrounding noise field of free-swimming harbor porpoises and seals. Acoustic noise mapping including porpoise detectors in the Natura 2000 sites of the North and Baltic Seas will help to fully understand current noise impacts.

Review of Low-Level Bioacoustic Behavior in Wild Cetaceans: Conservation Implications of Possible Sleeping Behavior.

Shallow, low-activity, low-biosonar parabolic-shaped dives were observed in biologging data from tagged harbor porpoises in Danish waters and identified as potential sleeping behavior. This behavioral state merits consideration in assessing the context for noise exposure and passive acoustic monitoring studies. Similar dives have also been reported for other cetacean species. The existence of low-level bioacoustic dives that may represent that sleeping has implications for the mitigation of not only noise exposure but also of bycatch as well as legal repercussions given the protected status of sleeping, as a part of resting, under many legislative regimes.

Underwater noise emissions from a drillship in the Arctic.

Wideband sound recordings were made of underwater noise emitted by an active drillship, Stena Forth, working in 484 m of water in Baffin Bay, western Greenland. The recordings were obtained at thirty and one-hundred meters depth. Noise was recorded during both drilling and maintenance work at ranges from 500 m to 38 km. The emitted noise levels were highest during maintenance work with estimated source levels up to 190 dB re 1 µPa (rms), while the source level during drilling was 184 dB re 1 µPa (rms). There were spectral peaks discernible from the background noise to ranges of at least 38 km from the drillship with the main energy below 3 kHz. M-weighted sound pressure levels were virtually identical to broadband levels for low-frequency cetaceans and about 5 dB lower for high-frequency cetaceans. Signals from the dynamic positioning system were clearly detectable at ranges up to two km from the drillship.

Investigating the potential use of environmental DNA (eDNA) for genetic monitoring of marine mammals.

The exploitation of non-invasive samples has been widely used in genetic monitoring of terrestrial species. In aquatic ecosystems, non-invasive samples such as feces, shed hair or skin, are less accessible. However, the use of environmental DNA (eDNA) has recently been shown to be an effective tool for genetic monitoring of species presence in freshwater ecosystems. Detecting species in the marine environment using eDNA potentially offers a greater challenge due to the greater dilution, amount of mixing and salinity compared with most freshwater ecosystems. To determine the potential use of eDNA for genetic monitoring we used specific primers that amplify short mitochondrial DNA sequences to detect the presence of a marine mammal, the harbor porpoise, Phocoena phocoena, in a controlled environment and in natural marine locations. The reliability of the genetic detections was investigated by comparing with detections of harbor porpoise echolocation clicks by static acoustic monitoring devices. While we were able to consistently genetically detect the target species under controlled conditions, the results from natural locations were less consistent and detection by eDNA was less successful than acoustic detections. However, at one site we detected long-finned pilot whale, Globicephala melas, a species rarely sighted in the Baltic. Therefore, with optimization aimed towards processing larger volumes of seawater this method has the potential to compliment current visual and acoustic methods of species detection of marine mammals.