Additional notes on the food of the Common porpoise (Phocoena phocoena)

Investigation of the food (through stomach contents) of Common porpoises caught in Scottish waters, discussed in an earlier paper, has been continued at the Marine Laboratory. The results confirm the earlier conclusions and stress the prominence of fish in the food. They also indicate the comparatively small size of the fish eaten and the predominantly pelagic or semi-pelagic nature of the majority of species in the diet.     

Structural fiber reinforcement of keel blubber in harbor porpoise (Phocoena phocoena)

This study investigated the functional morphology of the blubber that forms the caudal keels of the harbor porpoise (Phocoena phocoena). Blubber is a pliant biocomposite formed by adipocytes and structural fibers composed of collagen and elastic fibers. Caudal keels are dorsally and ventrally placed triangular wedges of blubber that define the hydrodynamic profile of the porpoise tailstock. Mechanical tests on carcasses demonstrate that when keels are bent, they strain nonuniformly along their lengths, with highest strains just caudal to the dorsal fin and lowest at the insertion of the flukes. Therefore, caudal keels undergo nonuniform longitudinal deformation while maintaining a stable, triangular cross-sectional shape. Polarizing and transmitted light microscopy techniques were used to investigate blubber's 3D fiber architecture along the length of the dorsal keel. The triangular cross-sectional shape of the keel appears to be maintained by structural fibers oriented to act as tensile stays. The construction of the blubber composite is regionally specific :structural fiber densities and diameters are higher in the relatively stiff caudal region of the keel than in the more deformable cranial keel region. The orientations of structural fibers also change along the length of the keel. Cranially, no fibers are oriented along the long axis, whereas a novel population of longitudinally oriented fibers reinforces the keel at the insertion of the flukes. Thus, differences in the distribution and orientation of structural fibers contribute to the regionally specific mechanical properties of the dorsal keel.     

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.     

Isolation and characterization of Brucella from the lungworms of a harbor porpoise (Phocoena phocoena)

Adult female nematodes identified as Pseudalius inflexus were collected from the lungs of a juvenile male harbor porpoise (Phocoena phocoena) found dead on a beach in Cornwall, UK. Classic and molecular typing methods, immunologic and electron microscopy immunolabeling techniques, provided evidence of Brucella sp. infection within the uterine tissue of nematodes of this marine mammal. This finding presents further evidence to suggest parasites should be considered as a potential means of transfer of bacterial infection in marine mammals and highlights the zoonotic implications for humans exposed to marine mammals through occupation or leisure.     

Evoked potentials of the auditory cortex of the porpoise,Phocoena phocoena

Summary Evoked potential (EP) recordings in the auditory cortex of the porpoise,Phocoena phocoena, were used to obtain data characterizing the auditory perception of this dolphin. The frequency threshold curves showed that the lowest EP thresholds were within 120–130 kHz. An additional sensitivity peak was observed between 20 and 30 kHz. The minimal EP threshold to noise burst was 3·10^–4–10^/s-3 Pa. The threshold for response to modulations in sound intensity was below 0.5 dB and about 0.1% for frequency modulations. Special attention was paid to the dependence of the auditory cortex EP on the temporal parameters of the acoustic stimuli: sound burst duration, rise time, and repetition rate. The data indicate that the porpoise auditory cortex is adapted to detect ultrasonic, brief, fast rising, and closely spaced sounds like echolocating clicks.Abbreviation EP evoked potential     

Mitochondrial DNA sequence variation and phylogeographic patterns in harbour porpoises (Phocoena phocoena) from the North Atlantic

The harbour porpoise (Phocoena phocoena)experiences high rates of incidental mortalityin commercial fisheries, and in some areasthese rates are sufficiently high to justifyconcern over population sustainability. Giventhe high incidental mortality, the resolutionof population structure will be important toconservation and management, but in the NorthAtlantic the relationships among many of theputative populations remain unclear. Aprevious genetic study demonstrated substantialgenetic differences between eastern and westernNorth Atlantic populations, however thelocation of this break remained unresolved. Inthe present study, we addressed this issue byincluding new samples from Iceland. Toinvestigate population structure, variation inthe mitochondrial DNA of 370 porpoises wascompared among six locations corresponding toseveral of the putative populations (Gulf ofMaine, Gulf of St. Lawrence, Newfoundland, WestGreenland, Iceland, Norway). The first 342base pairs of the control region were sequencedand genetic variation investigated by analysisof molecular variance (F_ST and_ST) and ² withpermutation. Although some fine scalepopulation structure was detected, porpoisesfrom Iceland were found to be more similar tothe western populations (W. Greenland, Gulf ofSt. Lawrence, Newfoundland, Gulf of Maine) thanto Norway. Furthermore, porpoises from Norwaywere different from all other regions. Thesepatterns suggest the existence of adiscontinuity between Iceland and Norway,possibly the result of isolating events causedby repeated range contractions and expansionsthroughout Quaternary glaciation events withinthe North Atlantic. These results suggest thatharbour porpoise populations within the NorthAtlantic are distinguishable, but patterns mustbe interpreted in light of their historicalbiogeography.     

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.     

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     

Development and rudimentation of the peripheral olfactory system in the harbor porpoise Phocoena phocoena (Mammalia: Cetacea)

Serial sections of 13 embryos and fetuses of the harbor porpoise from 10 mm crown-rump length up to 167 mm total length were studied. Unlike the adult animals, ontogenetic stages of 18–27 mm crown-rump length still show a typical mammalian olfactory bulb. The olfactory bulb primordium is penetrated by olfactory nerve fibers, the latter passing through the cribriform plate. However, the olfactory bulb anlage is gradually reduced in later stages, its placodal component being largely uncoupled from the telencephalon. As a ganglionlike structure, the remains of the placodal component stay in contact with the nasal septum and mucosa via thin bundles of nerve fibers. The ganglion and plexus can be traced within the meninges until the adult stage of the porpoise. There is strong evidence that they represent the material of the terminalis system, which cannot be distinguished from the olfactory system in earlier stages. A vomeronasal organ could not be detected in the embryonal and fetal material investigated.     

Patterns of ossification in the manus of the harbor porpoise (Phocoena phocoena): hyperphalangy and delta-shaped bones

With the transition from terrestrial to aquatic habitats, cetacean forelimbs have undergone significant modifications in bone morphology and soft tissue distribution. Some, but not all, of these modifications are also demonstrated in other lineages of extant and extinct secondarily aquatic tetrapods. This study examines the ontogenetic pattern of ossification of the manus of the harbor porpoise (Phocoena phocoena), using plain film radiography. Two modifications examined are hyperphalangy (number of phalanges per digit increased beyond the typical mammalian number) and the morphology of delta-shaped bones. Hyperphalangy in Phocoena phocoena is apparent in digits 2 and 3. Phalangeal counts in all digits are variable (sometimes between the right and left flippers of the same individual) and are not necessarily correlated with age. Phalangeal ossification and epiphyseal fusion proceeds along the proximo-distal axis within each digit. In addition, digits 2 and 3 are at a more advanced stage of ossification than more abaxial digits. Delta-shaped bones appear to be a normal stage in the ossification of phalanges in all digits except the third, and may persist in the adult in certain digits. In humans, this morphology is a developmental anomaly usually associated with other malformations, such as polydactyly or syndactyly. Delta-shaped bones in the cetacean manus display a consistent orientation and the process by which they are formed may be similar to that in extinct marine reptiles.     

Conservation genetics of harbour porpoises, Phocoena phocoena, in eastern and central North Atlantic

We examined polymorphism at 12 microsatelliteloci in 807 harbour porpoises , Phocoenaphocoena, collected from throughout thecentral and eastern North Atlantic to theBaltic Sea. Multilocus tests for allelefrequency differences, assignment tests,population structure estimates (F_ST) andgenetic distance measures (D_LR andD_C) all indicate six geneticallydifferentiated populations/sub-populationsafter pooling sub-samples within regions.Harbour porpoises from West Greenland, theNorwegian Westcoast, Ireland, the British NorthSea, the Danish North Sea and the inland watersof Denmark (IDW) are all geneticallydistinguishable from each other. A sample ofharbour porpoises collected off the Dutch coast(mainly during winter) was geneticallyheterogeneous and likely comprised a mixture ofindividuals of diverse origin. A mixed stockanalysis indicated that most of the individualsin this sample (77%) were likely migrantsfrom the British and Danish North Sea.     

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.     

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.     

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.     

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.