Mercury,cadmium, zinc,copper and selenium in harbour porpoise (Phocoena phocoena) from West Greenland

Muscle, liver, kidney and skin samples taken from 78 harbour porpoises (Phocoena phocoena) were analysed for mercury, cadmium, zinc, copper and selenium. The highest concentrations of mercury were found in the liver (geometric mean 4.17 g/g wet weight), whilst the highest concentrations of cadmium were in the kidney (g.m. 13.2 g/g ww). The levels of cadmium were more than ten times higher than in harbour porpoises from the North Sea and the British NW coast, whilst the mercury levels were about the same. The importance of the cadmium content in the prey is discussed, but this attempt did not revealed the differences. Very high levels of zinc (g.m. 359 g/g ww) and selenium (g.m. 28.6 g/g ww) were found in skin samples, respectively seven and ten times more than in liver. A significant correlation was found between age and the level of mercury and cadmium in all organs. The concentration of mercury and selenium in liver and skin samples and of cadmium and zinc in kidney samples were highly correlated.     

Gas exchange and heart rate in the harbour porpoise, Phocoena phocoena

The respiratory physiology, heart rates and metabolic rates of two captive juvenile male harbour porpoises (both 28 kg) were measured using a rapid-response respiratory gas analysis system in the laboratory. Breath-hold durations in the laboratory (12 ± 0.3 s, mean ± SEM) were shorter than field observations, although a few breath-holds of over 40 s were recorded. The mean percentage time spent submerged was 89 ± 0.4%. Relative to similarly-sized terrestrial mammals, the respiratory frequency was low (4.9 ± 0.19 breaths · min⁻¹) but with high tidal volumes (1.1 ± 0.01 l), enabling a comparatively high minute rate of gas exchange. Oxygen consumption under these experimental conditions (247 ± 13.8 ml O₂ · min⁻¹) was 1.9-fold higher than predicted by standard scaling relations. These data together with an estimate of the total oxygen stores predicted an aerobic dive limit of 5.4 min. The peak end-tidal O₂ values were related to the length of the previous breath-hold, demonstrating the increased oxygen uptake from the lung for the longer dives. Blood oxygen capacity was 23.5 ± 1.0 ml · 100 ml⁻¹, and the oxygen affinity was high, enabling rapid oxygen loading during ventilation. Accepted: 11 August 1999     

Variation in harbour porpoise activity in response to seismic survey noise

Animals exposed to anthropogenic disturbance make trade-offs between perceived risk and the cost of leaving disturbed areas. Impact assessments tend to focus on overt behavioural responses leading to displacement, but trade-offs may also impact individual energy budgets through reduced foraging performance. Previous studies found no evidence for broad-scale displacement of harbour porpoises exposed to impulse noise from a 10 day two-dimensional seismic survey. Here, we used an array of passive acoustic loggers coupled with calibrated noise measurements to test whether the seismic survey influenced the activity patterns of porpoises remaining in the area. We showed that the probability of recording a buzz declined by 15% in the ensonified area and was positively related to distance from the source vessel. We also estimated received levels at the hydrophones and characterized the noise response curve. Our results demonstrate how environmental impact assessments can be developed to assess more subtle effects of noise disturbance on activity patterns and foraging efficiency.     

Population structure of bycaught harbour porpoise (Phocoena phocoena) in Norway

ABSTRACT

The preference for coastal habitats makes the harbour porpoise, Phocoena phocoena, vulnerable to fisheries conflicts and hence prone to die due to entangling in fishing nets. An opportunistic sampling of such casualties (134 individuals) in Norwegian waters was used to assess the genetic population structure of the species by SNP-genotyping at 78 loci. The results of genetic clustering obtained for these individuals failed to identify more than one genetic group. Likewise, the individually-based F did not meet an Isolation-by-Distance pattern, thus supporting the conclusion that harbour porpoise in Norway probably belongs to a single genetic group or population.     

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.     

Trends of harbour porpoise (Phocoena phocoena) density in the southern North Sea

In the southern North Sea, harbour porpoise occurrence increased in recent years after a phase of low abundances during earlier decades. Only very few studies on porpoise presence in the southern German North Sea exist so far. As anthropogenic activities will strongly increase in this part of the North Sea during the next years it is most important to assess population level effects. This study focuses on the analysis of temporal and spatial trends in porpoise density in this area of recent change. Dedicated aerial line-transect distance sampling surveys were conducted in the southern German North Sea between May 2002 and June 2013 to assess porpoise density and distribution. Statistical inferences on porpoise population trends were made using a Markov Chain Monte Carlo (MCMC) technique. Two approaches were chosen to test for a trend in porpoise density and an additional model focused on the change in density of calves. During 55,820 km of survey effort 4377 porpoises including 140 calves were recorded. A significant effect of increasing spatial aggregation from the lower density areas in the south-eastern German Bight to hot spot areas in the western parts was detected. For the western part of the study area a significant increase in porpoise density between 2002 and 2013 was detected. Seasons were significantly different with highest porpoise density in spring and successively decreasing densities in summer and autumn. From 2008 onwards high densities were also observed in summer. Calf density increased during the study period and was significantly higher in the west. On the basis of this extensive and unique data set on porpoise occurrence in the southern German North Sea the findings clearly show that especially the south-western German North Sea serves as habitat of increasing importance for porpoises throughout the last decade. Definite reasons still remain unresolved. Changes in prey abundance or less favourable conditions in other areas could be important factors, which may also have caused a southward shift from high density areas in northern waters. On this baseline, further integrative approaches might lead to a sound understanding of the effect of anthropogenic activities on the future development of porpoise populations.     

Histological and ultrastructural aspects of the nasal complex in the harbour porpoise,Phocoena phocoena

During the evolution of odontocetes, the nasal complex was modified into a complicated system of passages and diverticulae. It is generally accepted that these are essential structures for nasal sound production. However, the mechanism of sound generation and the functional significance of the epicranial nasal complex are not fully understood. We have studied the epicranial structures of harbor porpoises (Phocoena phocoena) using light and electron microscopy with special consideration of the nasal diverticulae, the phonic lips and dorsal bursae, the proposed center of nasal sound generation. The lining of the epicranial respiratory tract with associated diverticulae is consistently composed of a stratified squamous epithelium with incomplete keratinization and irregular pigmentation. It consists of a stratum basale and a stratum spinosum that transforms apically into a stratum externum. The epithelium of the phonic lips comprises 70–80 layers of extremely flattened cells, i.e., four times more layers than in the remaining epicranial air spaces. This alignment and the increased number of desmosomes surrounding each cell indicate a conspicuous rigid quality of the epithelium. The area surrounding the phonic lips and adjacent fat bodies exhibits a high density of mechanoreceptors, possibly perceiving pressure differentials and vibrations. Mechanoreceptors with few layers and with perineural capsules directly subepithelial of the phonic lips can be distinguished from larger, multi‐layered mechanoreceptors without perineural capsules in the periphery of the dorsal bursae. A blade‐like elastin body at the caudal wall of the epicranial respiratory tract may act as antagonist of the musculature that moves the blowhole ligament. Bursal cartilages exist in the developmental stages from fetus through juvenile and could not be verified in adults. These histological results support the hypothesis of nasal sound generation for the harbor porpoise and display specific adaptations of the echolocating system in this species. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Synthetic communication signals influence wild harbour porpoise (Phocoena phocoena) behaviour

We used our novel and programmable Porpoise Alarm (PAL, patd.) to synthesize life-like, electronic harbour porpoise communication signals based on those described for captive animals. In the Little Belt, Denmark, we employed PAL (source level 158 ± 1 dB p–p re 1 μPa@1 m; centroid frequency 133 ± 8.5 kHz) to synthesize three aggressive click train types termed “A”, “F3” and “M1” to naive, free-living harbour porpoises. Via theodolite tracking (372 h of total visual effort spread over 10 expeditions) we found that, depending on signal type, porpoises either avoid or become attracted to PAL: Signal types “A” and “F3” are slight deterrents, porpoises increasing minimum range (+23 to 32 m, respectively), whereas “M1” attracts porpoises, reducing range (by ? 29 m). As determined via archival acoustic detectors (AADs), both signals “F3” and “M1” led the animals to significantly intensify their click rate (by +10% and 68%, respectively) while signal “A” led to a significant reduction ( ? 59%). We propose that equipping fishing gear with PAL emitting signal “F3” could potentially reduce porpoise by-catch by increasing (1) awareness through enhanced echolocation and (2) distance to the nets. Detection probability and radius of PAL/AAD tandems could be improved by emitting signal “M1” to focus porpoise echolocation signals on the AAD. The signal may also be useful in luring animals away from hazards, which may be helpful for conservation measures prior to the onset of harmful acoustic activities such as pile-driving, seismic exploration or ammunition clearance.     

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena)

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I–VI), and 1.4 ms (II–IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal’s size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.     

Granulosa cell tumour in a harbour porpoise (Phocoena phocoena) from German waters

Neoplastic diseases in harbour porpoises Phocoena phocoena have rarely been described, and there are no reported gonadal stromal tumours. A 12 yr old female harbour porpoise was stranded on the North Sea coast of Schleswig-Holstein, Germany. Necropsy findings included a severe granulomatous pneumonia, pregnancy and a left ovarian tumour. Respiratory insufficiency was the likely cause of death. There was a multinodular mass composed of cords with peripherally palisading cells within the left ovary. The histological and cytological appearance of the neoplasm was suggestive of a granulosa cell tumour; supportive immunohistochemical stains, including those for vimentin, cytokeration, carcinoembryonic antigen, c-kit, chromogranin and α-smooth muscle action, were negative.     

Keeping returns optimal: gain control exerted through sensitivity adjustments in the harbour porpoise auditory system

Animals that use echolocation (biosonar) listen to acoustic signals with a large range of intensities, because echo levels vary with the fourth power of the animal's distance to the target. In man-made sonar, engineers apply automatic gain control to stabilize the echo energy levels, thereby rendering them independent of distance to the target. Both toothed whales and bats vary the level of their echolocation clicks to compensate for the distance-related energy loss. By monitoring the auditory brainstem response (ABR) during a psychophysical task, we found that a harbour porpoise (Phocoena phocoena), in addition to adjusting the sound level of the outgoing signals up to 5.4 dB, also reduces its ABR threshold by 6 dB when the target distance doubles. This self-induced threshold shift increases the dynamic range of the biosonar system and compensates for half of the variation of energy that is caused by changes in the distance to the target. In combination with an increased source level as a function of target range, this helps the porpoise to maintain a stable echo-evoked ABR amplitude irrespective of target range, and is therefore probably an important tool enabling porpoises to efficiently analyse and classify received echoes.     

Stratification and age-related differences in blubber fatty acids of the male harbour porpoise (Phocoena phocoena)

Fatty acid composition of blubber was determined at four body sites of 19 male harbour porpoises. A total of 65 fatty acids were quantified in each sample. The array of fatty acids contained in harbour porpoise blubber was similar to those found in other marine mammals. While chemical composition of total blubber was uniform over the body, with the exception of the caudal peduncle, vertical stratification was evident between the deep (inner) and superficial (outer) blubber layers. Fatty acids with chain lengths shorter than 18 carbons were present in significantly greater amounts in the outer blubber layer, while the longer-chain unsaturated fatty acids were more prevalent in the inner layer. This distribution suggests that the inner blubber layer is more active metabolically than the outer layer in terms of lipid deposition and mobilization. The degree of stratification between the two layers appears to increase with age, indicating a predictable turnover in the blubber layer of male porpoises. Harbour porpoise blubber contained high levels (2–27%) of isovaleric acid in the outer blubber layer, and these levels were positively correlated with age.Abbreviations Caud caudal dorsal body site - GC gas chromatograph - FA fatty acid(s) - IUPAC International Union of Pure and Applied Chemistry - PUFA polyunsaturated fatty acid(s) - II dor II dorsal body site - III dor III Dorsal body site - II Ven II ventral body site     

First indication of gas embolism in a harbour porpoise (Phocoena phocoena) from German waters

In the last decade, a number of cetacean strandings have gas embolic pathology analogous to decompression sickness in man and experimental animals. Acute gas and fat embolic lesions have also been found in mass-stranded beaked whales causally linked to high-intensity anthropogenic mid-frequency sonar activities. Sporadic chronic gas embolic lesions have also been described. This communication describes a first case of numerous gas-filled fibrous cavities in the markedly enlarged liver of a dead adult male harbour porpoise stranded at the North Sea coast of Schleswig-Holstein, Germany. Additional pathological findings consisted of chronic parasitic granulomatous cholangiohepatitis, hepatic vein thrombosis, parasitic infestations of the bronchial tree and pulmonary blood vessels associated with bronchopneumonia and severe parasitic burden in both ear sinuses. The hepatic cavernous lesions are similar to those described in chronic gas bubble disease in other cetaceans, most likely resulting from decompression-related tissue supersaturation with nitrogen.     

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.     

Timing of epiphyseal development in the flipper skeleton of the harbour porpoise (Phocoena phocoena) as an indicator of paedomorphosis

Epiphyseal development was investigated on X‐rays of flippers from 158 harbour porpoises from Danish waters. Development followed a proximodistal pattern similar to what is known in other cetacean species. Ossification of epiphyses was rare in the phalanges of the first and fifth digits and in the more distal phalanges of the second, third and fourth digits. Along with the morphology of the first metacarpal and the more distal phalanges this suggested paedomorphosis relative to delphinids. Male and female porpoises showed similar progression of epiphyseal development until approximately the sixth year. From then on, female porpoises showed more progressed development than males. This suggests a higher level of paedomorphosis in the male porpoise. The mechanism behind phocoenid paedomorphosis seems to be progenesis, probably as an adaptation towards a high reproductive rate relative to the delphinids.     

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.     

Mitochondrial Control Region and microsatellite analyses on harbour porpoise (Phocoena phocoena) unravel population differentiation in the Baltic Sea and adjacent waters

The population status of the harbour porpoise (Phocoena phocoena) in the Baltic area has been a continuous matter of debate. Here we present the by far most comprehensive genetic population structure assessment to date for this region, both with regard to geographic coverage and sample size: 497 porpoise samples from North Sea, Skagerrak, Kattegat, Belt Sea, and Inner Baltic Sea were sequenced at the mitochondrial Control Region and 305 of these specimens were typed at 15 polymorphic microsatellite loci. Samples were stratified according to sample type (stranding vs. by-caught), sex, and season (breeding vs. non-breeding season). Our data provide ample evidence for a population split between the Skagerrak and the Belt Sea, with a transition zone in the Kattegat area. Among other measures, this was particularly visible in significant frequency shifts of the most abundant mitochondrial haplotypes. A particular haplotype almost absent in the North Sea was the most abundant in Belt Sea and Inner Baltic Sea. Microsatellites yielded a similar pattern (i.e., turnover in occurrence of clusters identified by STRUCTURE). Moreover, a highly significant association between microsatellite assignment and unlinked mitochondrial haplotypes further indicates a split between North Sea and Baltic porpoises. For the Inner Baltic Sea, we consistently recovered a small, but significant separation from the Belt Sea population. Despite recent arguments that separation should exceed a predefined threshold before populations shall be managed separately, we argue in favour of precautionary acknowledging the Inner Baltic porpoises as a separate management unit, which should receive particular attention, as it is threatened by various factors, in particular local fishery measures.