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Harbour porpoises (Phocoena phocoena) stranding in large numbers around the southern North Sea with fatal, sharp-edged mutilations have spurred controversy among scientists, the fishing industry and conservationists, whose views about the likely cause differ. The recent detection of grey seal (Halichoerus grypus) DNA in bite marks on three mutilated harbour porpoises, as well as direct observations of grey seal attacks on porpoises, have identified this seal species as a probable cause. Bite mark characteristics were assessed in a retrospective analysis of photographs of dead harbour porpoises that stranded between 2003 and 2013 (n = 1081) on the Dutch coastline. There were 271 animals that were sufficiently fresh to allow macroscopic assessment of grey seal-associated wounds with certainty. In 25% of these, bite and claw marks were identified that were consistent with the marks found on animals that had tested positive for grey seal DNA. Affected animals were mostly healthy juveniles that had a thick blubber layer and had recently fed. We conclude that the majority of the mutilated harbour porpoises were victims of grey seal attacks and that predation by this species is one of the main causes of death in harbour porpoises in The Netherlands. We provide a decision tree that will help in the identification of future cases of grey seal predation on porpoises.

Reproductive failure in mammals due to exposure to polychlorinated biphenyls (PCBs) can occur either through endocrine disrupting effects or via immunosuppression and increased disease risk. To investigate further, full necropsies and determination of summed 25 polychlorinated biphenyls congeners (∑PCBs lipid weight) in blubber were undertaken on 329 UK-stranded female harbour porpoises (1990-2012). In sexually mature females, 25/127 (19.7%) showed direct evidence of reproductive failure (foetal death, aborting, dystocia or stillbirth). A further 21/127 (16.5%) had infections of the reproductive tract or tumours of reproductive tract tissues that could contribute to reproductive failure. Resting mature females (non-lactating or non-pregnant) had significantly higher mean ∑PCBs (18.5 mg/kg) than both lactating (7.5 mg/kg) and pregnant females (6 mg/kg), though not significantly different to sexually immature females (14.0 mg/kg). Using multinomial logistic regression models ΣPCBs was found to be a significant predictor of mature female reproductive status, adjusting for the effects of confounding variables. Resting females were more likely to have a higher PCB burden. Health status (proxied by \"trauma\" or \"infectious disease\" causes of death) was also a significant predictor, with lactating females (i.e. who successfully reproduced) more likely to be in good health status compared to other individuals. Based on contaminant profiles (>11 mg/kg lipid), at least 29/60 (48%) of resting females had not offloaded their pollutant burden via gestation and primarily lactation. Where data were available, these non-offloading females were previously gravid, which suggests foetal or newborn mortality. Furthermore, a lower pregnancy rate of 50% was estimated for \"healthy\" females that died of traumatic causes of death, compared to other populations. Whether or not PCBs are part of an underlying mechanism, we used individual PCB burdens to show further evidence of reproductive failure in the North-east Atlantic harbour porpoise population, results that should inform conservation management.

The population structure of the highly mobile marine mammal, the harbor porpoise (Phocoena phocoena), in the Atlantic shelf waters follows a pattern of significant isolation-by-distance. The population structure of harbor porpoises from the Baltic Sea, which is connected with the North Sea through a series of basins separated by shallow underwater ridges, however, is more complex. Here, we investigated the population differentiation of harbor porpoises in European Seas with a special focus on the Baltic Sea and adjacent waters, using a population genomics approach. We used 2872 single nucleotide polymorphisms (SNPs), derived from double digest restriction-site associated DNA sequencing (ddRAD-seq), as well as 13 microsatellite loci and mitochondrial haplotypes for the same set of individuals. Spatial principal components analysis (sPCA), and Bayesian clustering on a subset of SNPs suggest three main groupings at the level of all studied regions: the Black Sea, the North Atlantic, and the Baltic Sea. Furthermore, we observed a distinct separation of the North Sea harbor porpoises from the Baltic Sea populations, and identified splits between porpoise populations within the Baltic Sea. We observed a notable distinction between the Belt Sea and the Inner Baltic Sea sub-regions. Improved delineation of harbor porpoise population assignments for the Baltic based on genomic evidence is important for conservation management of this endangered cetacean in threatened habitats, particularly in the Baltic Sea proper. In addition, we show that SNPs outperform microsatellite markers and demonstrate the utility of RAD-tags from a relatively small, opportunistically sampled cetacean sample set for population diversity and divergence analysis.

The harbour porpoise is the only cetacean species native to German waters. Since human pressures are suggested to shorten their reproductive lifespan, basic knowledge on reproduction is strongly required. One parameter is the onset of sexual maturity in female harbour porpoises. Therefore, we investigated the first signs of sexual maturity for a period of almost two decades (1990-2016). Ovaries from 111 female harbour porpoises from the German North Sea and Baltic Sea were examined for the presence and morphological structure of follicles, corpora lutea and corpora albicantia. Based on the ovarian characteristics we performed the first model-based estimation of age at sexual maturity for harbour porpoises from German waters. Additionally, we produced a demographical age structure based on all female strandings and bycatches from German coasts. Our results showed that corpora lutea and corpora albicantia as signs of former ovulation could be found in individuals at an age of 4.95 (± 0.6) years. No significant differences between specimens from the North Sea and Baltic Sea were detected. However, the average age at death differed significantly with 5.70 (± 0.27) years for North Sea animals and 3.67 (± 0.30) years for those in the Baltic Sea. Growing evidence exists that the shortened lifespan of Baltic Sea harbour porpoises is linked to an anthropogenically influenced environment with rising bycatch mortalities due to local gillnet fisheries. Thus, our findings support the idea of local management plans based on a model-based detection of age at sexual maturity and considering the anthropogenic impacts on the population for effective protection of harbour porpoises and the North Sea and Baltic Sea.

Odontocetes have evolved a rich diversity of prey- and habitat-specific foraging strategies, which allows them to feed opportunistically on locally and temporally abundant prey. While habitat-specific foraging strategies have been documented for some odontocete species, this is less known for the harbour porpoise (Phocoena phocoena). We collected multiple years of acoustic data using echolocation click loggers to analyse porpoise occurrence and buzzing behaviour, indicating feeding, in the German Wadden Sea (North Sea). Seasonal, diel and tidal effects were studied using Generalised Estimating Equations (GEE-GAMs). Locally season, time of day and tidal time significantly influenced the probability of porpoise detections and detection of foraging sequences (buzzes). Hunting strategies, and therefore frequency of buzzes, were likely affected by prey distribution and large differences between POD locations indicated that porpoises used highly specific behaviour adapted to tide and time of day to efficiently feed on the available prey. Strong seasonal and spatial variation in diel and tidal effects underline the importance of long-term observations. Studies on porpoise behaviour are often based on short-term observations and might rather reflect a seasonal than a general pattern. The results of this study show clearly that significant changes in porpoise behaviour can be found in short and long-term observations. Here some features are based on short term determinants and others are stable over years and care should be taken about drawing general conclusions based on local patterns. Highly variable spatio-temporal patterns indicate a high flexibility of porpoises in a highly variable environment and address a challenge for complex conservation management plans.

Marine traffic is threatening cetaceans on a local and global scale. The Istanbul Strait is one of the busiest waterways, with up to 2,500 vessels present daily. This is the first study to assess the magnitude of short- and long-term behavioural changes of the endangered Black Sea harbour porpoises (Phocoena phocoena relicta) in the presence of marine vessels within the Istanbul Strait. Markov chains were used to investigate the effect of vessel presence on the transition probability between behavioural states (diving, surface-feeding and travelling), and to quantify the effect on the behavioural budget and bout length (duration of time spent in a given state) of porpoises. Further, the changes on swimming directions of porpoises in relation to vessel speed and distance was investigated using generalized linear models. In vessel presence, porpoises were less likely to remain in a given behavioural state and instead more likely to switch to another state. Because of this, the bout length of all three behavioural states decreased significantly in the presence of vessels. The vessel effect was sufficiently large to alter the behavioural budget, with surface-feeding decreasing significantly in the presence of vessels. However, when taking into account the proportion of time that porpoises were exposed to vessels (i.e. 50%), the measured effect size was not large enough to significantly alter the animals' cumulative (diurnal) behavioural budget. Additionally, vessel speed and distance had a significant effect on the probability of porpoises showing a response in their swimming directions. The southern and middle sections of the Istanbul Strait, which have the heaviest marine traffic pressure, had the lowest porpoise sightings throughout the year. Conversely, northern sections that were exposed to a lesser degree of marine traffic hold the highest porpoise sightings. The effect shown in this study in combination with increasing human impacts within the northern sections should be considered carefully and species-specific conservation actions, including establishment of protected areas, should be put in place to prevent the long-term consequences of marine traffic on the Black Sea harbour porpoise population.

Odontocetes produce a range of different echolocation clicks but four groups in different families have converged on producing the same stereotyped narrow band high frequency (NBHF) click. In microchiropteran bats, sympatric species have evolved the use of different acoustic niches and subtly different echolocation signals to avoid competition among species. In this study, we examined whether similar adaptations are at play among sympatric porpoise species that use NBHF echolocation clicks. We used a six-element hydrophone array to record harbour and Dall's porpoises in British Columbia (BC), Canada, and harbour porpoises in Denmark. The click source properties of all porpoise groups were remarkably similar and had an average directivity index of 25 dB. Yet there was a small, but consistent and significant 4 kHz difference in centroid frequency between sympatric Dall's (137±3 kHz) and Canadian harbour porpoises (141±2 kHz). Danish harbour porpoise clicks (136±3 kHz) were more similar to Dall's porpoise than to their conspecifics in Canada. We suggest that the spectral differences in echolocation clicks between the sympatric porpoises are consistent with evolution of a prezygotic isolating barrier (i.e., character displacement) to avoid hybridization of sympatric species. In practical terms, these spectral differences have immediate application to passive acoustic monitoring.

Assessments of the impact of offshore energy developments are constrained because it is not known whether fine-scale behavioural responses to noise lead to broader-scale displacement of protected small cetaceans. We used passive acoustic monitoring and digital aerial surveys to study changes in the occurrence of harbour porpoises across a 2000 km2 study area during a commercial two-dimensional seismic survey in the North Sea. Acoustic and visual data provided evidence of group responses to airgun noise from the 470 cu inch array over ranges of 5–10 km, at received peak-to-peak sound pressure levels of 165–172 dB re 1 µPa and sound exposure levels (SELs) of 145–151 dB re 1 µPa2 s−1. However, animals were typically detected again at affected sites within a few hours, and the level of response declined through the 10 day survey. Overall, acoustic detections decreased significantly during the survey period in the impact area compared with a control area, but this effect was small in relation to natural variation. These results demonstrate that prolonged seismic survey noise did not lead to broader-scale displacement into suboptimal or higher-risk habitats, and suggest that impact assessments should focus on sublethal effects resulting from changes in foraging performance of animals within affected sites.

Offshore windfarms provide renewable energy, but activities during the construction phase can affect marine mammals. To understand how the construction of an offshore windfarm in the Maryland Wind Energy Area (WEA) off Maryland, USA, might impact harbour porpoises (Phocoena phocoena), it is essential to determine their poorly understood year-round distribution. Although habitat-based models can help predict the occurrence of species in areas with limited or no sampling, they require validation to determine the accuracy of the predictions. Incorporating more than 18 months of harbour porpoise detection data from passive acoustic monitoring, generalized auto-regressive moving average and generalized additive models were used to investigate harbour porpoise occurrence within and around the Maryland WEA in relation to temporal and environmental variables. Acoustic detection metrics were compared to habitat-based density estimates derived from aerial and boat-based sightings to validate the model predictions. Harbour porpoises occurred significantly more frequently during January to May, and foraged significantly more often in the evenings to early mornings at sites within and outside the Maryland WEA. Harbour porpoise occurrence peaked at sea surface temperatures of 5°C and chlorophyll a concentrations of 4.5 to 7.4 mg m-3. The acoustic detections were significantly correlated with the predicted densities, except at the most inshore site. This study provides insight into previously unknown fine-scale spatial and temporal patterns in distribution of harbour porpoises offshore of Maryland. The results can be used to help inform future monitoring and mitigate the impacts of windfarm construction and other human activities.

Effects of constructing the DanTysk offshore wind farm (German Bight, 80 turbines, 6 m diameter foundations) were studied by passive acoustic monitoring of pile-driving noise and harbour porpoise Phocoena phocoena echolocation. An acoustic deterrence device (seal scarer) was used to protect porpoises from hearing loss and bubble curtains were used to attenuate the pile-driving noise. Porpoise occurrence, quantified by echolocation signals, decreased when the seal scarer was engaged, during pile driving and up to 5 h after pile driving stopped. This effect extended out to 12 km, less than the 18–25 km reported from other pile drivings performed without bubble curtains. The bubble curtains thus effectively reduced the temporary habitat loss and risk of hearing loss. The 2 bubble curtains each attenuated the noise by between 7 and 10 dB, when used separately, and 12 dB when used together. Attenuation was most pronounced above 1 kHz, where the pile-driving noise at larger distances was comparable to or lower than ambient noise. This suggests that noise regulation should be based on frequency-weighted sound levels in addition to broadband levels, to assure that mitigation measures are effective in reducing impact on animals and not only in fulfilling legal requirements. The strong reaction to the seal scarer raises concern that it may surpass the reactions to the pile-driving noise itself, when operating with bubble curtains, and calls for a re-evaluation of the specifications of seal scarer sounds.