The harbour seal,Phoca vitulina,in the Oosterschelde: decline and possibilities for recovery

Within a timespan of a few decadesm the harbour seal almost completely disappeared from the estuaries in the south-west of the Netherlands. In 1960 a population of around 350 animals still lived in the Oosterschelde and Westerschelde area. About a quarter of this population lived in the Oosterschelde. At present less than 17 animals can be regularly observed in the whole area. Human influences are responsible for the rapid decline of the population. Initially a high hunting pressure and later environmental pollution are the main causes. Loss of habitat and disturbance at the resting places are additional important factors. The Oosterschelde still is a suitable habitat for seals. A short term natural development of a viable population in the area is not to be expected. Only with human help through active management, i.e. reintroduction of rehabilitated seals (preferably originating from that area) and strict conservation of the extant Oosterschelde seal population, accompanied by environmental sanitation of the neighbouring waters, can the current southern Dutch harbour seal population increase.     

Historical population size of the harbour seal,Phoca vitulina,in the Delta area,SW Netherlands

Seals in the Dutch Delta area have been subject to hunting pressure for centuries, promoted by a bounty system which generated a sort of hunting statistics. Hunting mortality is used to estimate historical population size. Based on ranges for most likely net recruitment rates, corresponding population trajectories are back calculated from an assessed population size of 350 seals in 1960. It is concluded that the size of the harbour seal population in the Delta area in 1900 will have been close to 11 500 animals. Significant loss of habitat has occurred due to closing off parts of the larger estuaries and the enlargement of the entrance to the harbour of Rotterdam in the 1960s and early 1970s. It is estimated that about 4000 harbour seals could inhabit the remaining Delta area under tidal influence. This outcome, based on retrospective population analysis, will be an important reference in defining management objectives for the recovery of the harbour seal population in the Delta area, which amounted to 18 seals in 1992.     

Quantitative aspects of harbour seal (Phoca vitulina) play

The play behaviour of harbour seals was analysed on videotape, which allowed the determination of rates of play per age class relative to the population of seals of the same age, and sex of the player in 190 episodes. The majority of the results of our earlier study were confirmed in that adult play was unusually common, and most play was solitary rather than social. However, the reason for the apparent predominance of juvenile play in our 1986 study was most likely because of a large number of juvenile seals present in the herd at that time. Similarly, mothers, pups and weaners proved to play at high rates when population size was taken into account. Adults' playing increased during the weaning/mating period, whereas juveniles and subadults played progressively less over time. Mothers and pups played more near weaning, after which time the youngsters played at a steady high rate, though never socially. Adult social play was only evident during the mating period. The play of males and females differed qualitatively and quantitatively.     

The lipid composition of serum lipoproteins in the harbour seal, Phoca vitulina

The density profile of serum lipoproteins and their lipid composition was studied in 12 adult, female harbour seals. The animals were sampled after an approximate 20 hr fast. The density profile of lipoproteins showed that the harbour seals displayed a distinct VLDL (density less than 1.006 g/ml) and HDL band (density about 1.125 g/ml), but no clear LDL band. There was a rather diffuse population of lipoproteins in the density range of 1.019-1.100 g/ml. Mean serum total cholesterol concentration was 5.7 mmol/l; about 60% of this cholesterol was located in the HDL fraction (density greater than 1.063 g/ml). The fasted seals were found to carry 4% of serum total lipids in chylomicrons. These lipoproteins consisted of 51% of triaclyglycerols (on the basis of total chylomicron lipids). The LDL (defined as heparin-manganese precipitable lipoproteins in VLDL and chylomicron-deficient serum) contained 49% of cholesterol and 43% of phospholipids (on the basis of total LDL lipids). The HDL (defined as heparin-manganese soluble lipoproteins in VLDL and chylomicron-deficient serum) contained 36% of cholesterol and 58% of phospholipids (on the basis of total HDL lipids).     

Monitoring harbour seal (Phoca vitulina) in the Danish Wadden Sea

Monitoring of harbour seal (Phoca vitulina) has been carried out in the Danis part of the Wadden Sea during the past 8 years by the Fishery and Maritime Museum. In addition to this, field observations over three breeding seasons have been undertaken. The census shows a steady increase of approx. 14% per yeart. The distribution of the seals in the area can be split up into two different types of haulout localities, and a varied pattern of haulouts is observed during the year. The mass die-off in 1988 is discussed within the framework of the knowledge on the population, and it is recommended that more effort be put into monitoring and population studies of the harbour seal in the entire International Wadden Sea. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, 1–4 November 1988)     

Photoperiod control of birth timing in the harbour seal (Phoca vitulina)

The harbour seal ( Phoca vitulina ) has delayed implantation, precise annual birth timing, and significant latitudinal variation in birth timing. The birth timing patterns of four distinct groups of seals, including colonies of P. v. vitulina and colonies and captive individuals of P. v. richardsi , were examined using population-based photoperiod analysis to assess the role of photoperiod in setting annual birth timing. This analysis simultaneously determined the time, relative to birth, at which photoperiod response was likely to occur and the critical photoperiod.
Despite marked differences in birth timing patterns, a high level of agreement was found among groups for the timing of photoperiod response. The two subspecies, however, demonstrated significantly divergent critical photoperiods. Eastern Atlantic harbour seals were exposed to a common critical photoperiod of 11.7 h/day on the 268th pre-partum day. Wild Pacific harbour seals were exposed to 14.3 h/day on the 283rd pre-partum day. These times corresponded to the estimated occurrence of blastocyst implantation.
Using the above information, three small captive populations were subjected to artificially prolonged photoperiods during the period of embryonic diapause to test whether subsequent birth timing could be delayed. Technical difficulties invalidated results at two sites. At the third and largest colony, the mean pupping date of six individuals was significantly delayed by 10.7days.
The precision and latitudinal variation of annual birth timing in the harbour seal are due to a response to photoperiod which occurs immediately prior to blastocyst implantation. The critical photoperiod, however, is divergent among subspecies and, thus, has probably evolved allowing seasonal adaptation. Similar environmental signalling has been described for California sea lions and northern fur seals and represents the likely timing mechanism for most pinniped species.     

Chemical characterization of the oligosaccharides in milk of high Arctic harbour seal (Phoca vitulina vitulina)

Carbohydrates were extracted from high Arctic harbour seal milk, Phoca vitulina vitulina (family Phocidae). Free neutral oligosaccharides were separated by gel filtration and preparative thin layer chromatography, while free sialyl oligosaccharides were separated by gel filtration and then purified by ion exchange chromatography, gel filtration and high performance liquid chromatography. Oligosaccharide structures were determined by 1H-NMR spectroscopy. The structures of the neutral oligosaccharides were as follows: lactose, 2'-fucosyllactose, lacto-N-neotetraose, lacto-N-neohexaose, monofucosyl lacto-N-neohexaose and difucosyl lacto-N-neohexaose. Thus, all of the neutral saccharides contained lactose or lacto-N-neotetraose or lacto-N-neohexaose as core units and/or non-reducing alpha(1-2) linked fucose. These oligosaccharides have also been found in hooded seal milk. The structures of the silalyl oligosaccharides were: monosialyl lacto-N-neohexaose, monosialyl monofucosyl lacto-N-neohexaose, monosialyl difucosyl lacto-N-neohexaose and disialyl lacto-N-neohexaose. These oligosaccharides contained lacto-N-neohexaose as core units, and one or two alpha(2-6) linked Neu5Ac, and/or non-reducing alpha(1-2) linked Fuc. The Neu5Ac residues were found to be linked to GlcNAc or penultimate Gal residues. The acidic oligosaccharides are the first to have been characterized in the milk of any species of seal.     

Earlier pupping in harbour seals,Phoca vitulina

The annual reproductive cycle of most seal species is characterized by a tight synchrony of births. Typically, timing of birth shows little inter-annual variation. Here, however we show that harbour seals Phoca vitulina from the Wadden Sea (southeast North Sea) have shortened their yearly cycle, moving parturition to earlier dates since the early 1970s. Between 1974 and 2009, the birth date of harbour seals shifted on average by −0.71 d yr⁻¹, three and a half weeks (25 days) earlier, in the Dutch part of the Wadden Sea. Pup counts available for other parts of the Wadden Sea were analysed, showing a similar shift. To elucidate potential mechanism(s) for this shift in pupping phenology, possible changes in population demography, changes in maternal life-history traits and variations in environmental conditions were examined. It was deduced that the most likely mechanism was a shortening of embryonic diapause. We hypothesize that this could have been facilitated by an improved forage base, e.g. increase of small fishes, attributable to overfishing of large predator fishes and size-selective fisheries.     

Persistence thresholds for phocine distemper virus infection in harbour seal Phoca vitulina metapopulations

1. This paper explores the concept of the critical community size for persistence of infection in wildlife populations. We use as a case study the 1988 epidemic of phocine distemper virus in the North Sea population of harbour seals, Phoca vitulina .
2. We summarize the available data on this epidemic and use it to parameterize a stochastic compartmental model for an infection spreading through a spatial array of patches coupled by nearest-neighbour mixing, with replacement of susceptibles occurring as a discrete annual event.
3. A combination of analytical and simulation techniques is used to show that the high levels of transmission between different seal subpopulations, combined with the small annual birth cohort, act to make persistence of infection impossible in this harbour seal population at realistic population levels. The well known mechanisms by which metapopulation structures may act to promote persistence can be seen to have an effect only at weaker levels of spatial coupling, and higher levels of host recruitment, than those empirically observed.     

Assessment of errors associated with harbour seal (Phoca vitulina) faecal sampling

Six harbour seals, ages 4–8 years, were held as pairs in a 10 times 20 times 2 m tank filled with sea water, and on 60 occasions were fed a meal of a specific species of fish or cephalopod of known size. The tank was drained periodically, and harbour seal faeces were collected on a 0.5 mm sieve. Number and size of otoliths and beaks found in faeces were determined. Fifty-eight percent of 670 fish and 37% of 36 cephalopods fed to harbour seals were represented by their otoliths or beaks in faeces. Estimated number of prey consumed was determined from the greatest number of left or right otoliths or upper or lower beaks collected in faeces. Estimated length ofprey was determined from measurements of otoliths and beaks recovered in the tank and relationships of otolith and beak measurements to prey length. Estimated number of fish eaten was not significantly different among pairs of harbour seals, but was different among species of fishes. Only 24–35% of fish species with small otoliths were represented in faeces, whereas more robust otoliths from other species were less apt to be completely dissolved. Estimated length of fishes was significantly less than lengths of fishes fed to harbour seals in 39 (76.5%) of 51 trials. Cephalopod beaks were not affected by passage through the harbour seal digestive tract. Amount of otolith dissolution was not related to species of fish; estimated fish length was underestimated by an average 27.5%. Although some (7.4%) of the otoliths were collected within 100 h after the fish were ingested, more than 90% were recovered within 24 h after the fish was eaten. Correction factors were developed which will allow researchers to estimate more reliably number and size of fish and cephalopod prey eaten by harbour seals.     

Clines revisited: The timing of pupping in the harbour seal (Phoca vitulina)

The regional variation in the pupping season of the harbour seal ( Phoca vitulina ) was reviewed using the birth periods reported for 65 colony sites distributed over a range from 30.4 to 78.5 North latitude. The birth timing of P. v. vitulina was not related to latitude, but birthing in P. v. concolor along eastern North America exhibited a latitudinal cline. The timing of birth in P. v. richardsi varied in three distinct patterns: (1) a significant unidirectional latitudinal cline extending between Baja California and the west coast of Washington; (2) a cluster comprised of Puget Sound, Washington and Vancouver Island, British Columbia colonies in which birthing occurred an average of 65 days later than on the Washington coast; and (3) a cluster from northern British Columbia and Alaska which did not demonstrate a latitudinal cline. Insufficient data were available for the analysis of P. v. mellonae or P. v. stejnegeri .
We found great regional variation in the timing of birth among all colonies, with mean birth dates occurring as early as 15 March and as late as 3 September. Little variation existed north of 50. To the south of 50, however, most of the variation could be attributed to correlation with latitude or to affiliation with the Puget Sound, Washington-Vancouver Island, British Columbia geographic area.
Clinal variation in pupping could result from: (1) geographic variations in a selective factor with perhaps gene exchange between contiguous populations playing a role in smoothing the variation; or (2) for populations between Mexico and the west coast of Washington, regional variation in a non-selective environmental variable, such as photoperiod.     

Male mating success in an aquatically mating pinniped, the harbour seal (Phoca vitulina), assessed by microsatellite DNA markers

Similar to many other pinniped species, harbour seals ( Phoca vitulina ) mate exclusively at sea. Here we present the first attempt to measure male mating success in an aquatically mating pinniped. Male mating success was estimated by paternity analysis in two cohorts of pups born at Sable Island, Nova Scotia, Canada, using microsatellite DNA markers. The genotypes of 275 pups born in 1994 and 1995 were compared to those of 90 candidate males at six microsatellite loci using a likelihood approach to resolve paternity. Paternity could be assigned for two, 22, 40 and 85 pups at confidence levels of 95, 80, 65 and 50%, respectively. Most successful males were assigned the paternity of a single offspring, suggesting a low variance in male mating success relative to most pinniped species. The proportion of paternal half sibs within cohorts and between maternally related sibs estimated by maximum likelihood were not significantly different from zero. It is thus unlikely that most offspring were sired by a small number of highly successful unsampled males, and that female harbour seals do not usually exhibit fidelity to the same male in sequential breeding seasons. A low level of polygyny in Sable Island harbour seals is consistent with predictions based on their breeding ecology, as females are highly mobile and widely dispersed in the aquatic mating environment at Sable Island.     

Respiratory chemosensitivity during wake and sleep in harbour seal pups (Phoca vitulina richardsii)

In this study, we examined the cardiorespiratory patterns of harbour seal pups under normoxic/normocarbic (air), hypoxic/normocarbic (15%, 12%, and 9% O2 in air), and normoxic/hypercarbic (2%, 4%, and 6% CO2 in air) conditions while awake and sleeping on land. Animals were chronically instrumented to record electroencephalogram (EEG), electromyogram (EMG), and electrocardiogram (EKG) signals, which, along with respiration (whole-body plethysmography) and oxygen consumption (VO2), were recorded from animals breathing each gas mixture for 2-4 h on separate days. Our results show that for animals breathing air, VO2 was not significantly lower during slow-wave sleep (SWS; 7.71 +/- 0.39 mL O2 min(-1) kg(-1); all measurements are mean +/- SEM) than during wakefulness (WAKE; 8.80 +/- 0.25 mL O2 min(-1) kg(-1)) and was unaffected by changes in respiratory drive. Although there was no significant fall in VO2 associated with a decrease in arousal state, breathing frequency (f(R)) did decrease (from 18.80 +/- 1.50 breaths min(-1) in WAKE to 10.40 +/- 0.49 breaths min(-1) in SWS), while the incidence of long apneas (>20 s) increased (12.76 +/- 4.06 apneas h(-1) in WAKE and 31.95 +/- 2.37 apneas h(-1) in SWS). Breathing was rarely seen during rapid eye movement (REM) sleep. Tachypnea was present at all levels of increased respiratory drive; however, hypoxia induced a dramatic bradycardia regardless of arousal state, while hypercarbia produced a tachycardia in SWS only. The hypoxic and hypercarbic chemosensitivities of harbour seal pups were similar to those of terrestrial mammals; however, unlike terrestrial mammals, where hypoxic and hypercarbic sensitivities are often reduced during SWS, the sensitivity of harbour seal pups to hypoxia and hypercarbia remained unchanged during the decrease in arousal state from WAKE to SWS.     

Sex differences in the nursing relationship between mothers and pups in the Atlantic harbour seal, Phoca vitulina concolor

Sex differences in the dynamics of the relationship between harbour seal mothers and their pups were investigated. An index of pair proximity governance suggested that pups became increasingly responsible for maintaining the distance between the pair over the course of the nursing period. Male pups demonstrated a stronger tendency to control initiation distance than female pups. A Nursing Index utilizing suckling bout initiations and terminations revealed that, as weaning approached, pups became progressively more responsible for the continuance of nursing. Male pups exhibited more control than female pups, primarily through initiating a greater portion of the bouts. On-teat times averaged 4.08 min per bout, with no significant differences between male and female pups. Male pups, however. nursed a greater portion of haul-out time due to higher bout frequencies obtained through more bout initiations. Mothers did not preferentially reject attempts to nurse by pups of either sex. These results suggest that the tenet of parental investment theory suggesting male pups enjoy preferential investment may be misleading. In harbour seals. male pups seem actively to seek out resources, as opposed to any inherent strategy on the par1 of the mother.     

Molecular methods for differentiating grey seal (Halichoerus grypus) and harbour seal (Phoca vitulina) scat

Pinniped dietary information is crucial for understanding marine ecosystems; however, in the North Atlantic, grey and harbour seals haulout concomitantly and their faeces are visually indistinguishable. Therefore, we developed molecular methods to differentiate the species' scat. PCR primers were created that amplify a portion of mitochondrial 16S ribosomal DNA in grey and harbour seals. The samples were amplified and the resulting products were digested using the restriction enzyme Tsp509I, producing diagnostic banding patterns. These tools provide a mechanism by which separate dietary analysis can be achieved for grey and harbour seals at mixed haulouts in New England.     

Diving heart rate development in postnatal harbour seals, Phoca vitulina

Harbour seals, Phoca vitulina, dive from birth, providing a means of mapping the development of the diving response, and so our objective was to investigate the postpartum development of diving bradycardia. The study was conducted May-July 2000 and 2001 in the St. Lawrence River Estuary (48 degrees 41'N, 68 degrees 01'W). Both depth and heart rate (HR) were remotely recorded during 86,931 dives (ages 2-42 d, n = 15) and only depth for an additional 20,300 dives (combined data covered newborn to 60 d, n = 20). The mean dive depth and mean dive durations were conservative during nursing (2.1 +/- 0.1 m and 0.57 +/- 0.01 min, range = 0-30.9 m and 0-5.9 min, respectively). The HR of neonatal pups during submersion was bimodal, but as days passed, the milder of the two diving HRs disappeared from their diving HR record. By 15 d of age, most of the dive time was spent at the lower diving bradycardia rate. Additionally, this study shows that pups are born with the ability to maintain the lower, more fully developed dive bradycardia during focused diving but do not do so during shorter routine dives.     

Integrating genetic data and population viability analyses for the identification of harbour seal (Phoca vitulina) populations and management units

Identification of populations and management units is an essential step in the study of natural systems. Still, there is limited consensus regarding how to define populations and management units, and whether genetic methods allow for inference at the relevant spatial and temporal scale. Here, we present a novel approach, integrating genetic, life history and demographic data to identify populations and management units in southern Scandinavian harbour seals. First, 15 microsatellite markers and model‐ and distance‐based genetic clustering methods were used to determine the population genetic structure in harbour seals. Second, we used harbour seal demographic and life history data to conduct population viability analyses (PVAs) in the vortex simulation model in order to determine whether the inferred genetic units could be classified as management units according to Lowe and Allendorf's (Molecular Ecology, 19, 2010, 3038) ‘population viability criterion’ for demographic independence. The genetic analyses revealed fine‐scale population structuring in southern Scandinavian harbour seals and pointed to the existence of several genetic units. The PVAs indicated that the census population size of each of these genetic units was sufficiently large for long‐term population viability, and hence that the units could be classified as demographically independent management units. Our study suggests that population genetic inference can offer the same degree of temporal and spatial resolution as ‘nongenetic’ methods and that the combined use of genetic data and PVAs constitutes a promising approach for delineating populations and management units.     

Seasonal and regional variations in the feeding habits of the harbour seal, Phoca vitulina, in the Skagerrak and the Kattegat

Studies of the feeding of harbour seals have been carried out at the Tjärnö Marine Biological Laboratory since 1977. The studies are based on fish otoliths found in faeces at seal haulouts. The present paper compares feeding habitats at two different localities. Three families of fish, gadoids, pleuronectids and clupeoids were predominant in the seals' diet at a rocky shore habitat. Pleuronectids made up 75% of the diet at a sandy shore habitat. Temporal variations in feeding habits are also examined. The results indicate that harbour seals are opportunists in their choice of prey species, but some locally abundant species do not appear in their diet.