Worldwide patterns of mitochondrial DNA differentiation in the harbor seal (Phoca vitulina)

The harbor seal (Phoca vitulina) has one of the broadest geographic distributions of any pinniped, stretching from the east Baltic, west across the Atlantic and Pacific Oceans to southern Japan. Although individuals may travel several hundred kilometers on annual feeding migrations, harbor seals are generally believed to be philopatric, returning to the same areas each year to breed. Consequently, seals from different areas are likely to be genetically differentiated, with levels of genetic divergence increasing with distance. Differentiation may also be caused by long-standing topographic barriers such as the polar sea ice. We analyzed samples of 227 harbor seals from 24 localities and defined 34 genotypes based on 435 bp of control region sequence. Phylogenetic analysis and analysis of molecular variance showed that populations in the Atlantic and Pacific Oceans and east and west coast populations of these oceans are significantly differentiated. Within these four regions, populations that are geographically farthest apart generally are the most differentiated and often do not share genotypes or differ in genotype frequency. The average corrected sequence divergence between populations in the Atlantic and Pacific Oceans is 3.28% +/- 0.38% and those among populations within each of these oceans are 0.75% +/- 0.69% and 1.19% +/- 0.65%, respectively. Our results suggest that harbor seals are regionally philopatric, on the scale of several hundred kilometers. However, genetic discontinuities may exist, even between neighboring populations such as those on the Scottish and east English coasts or the east and west Baltic. The mitochondrial data are consistent with an ancient isolation of populations in both oceans, due to the development of polar sea ice. In the Atlantic and Pacific, populations appear to have been colonized from west to east with the European populations showing the most recent common ancestry. We suggest the recent ancestry of European seal populations may reflect recolonization from Ice Age refugia after the last glaciation.      

Congenital hemicerebral anomaly in a stranded Pacific harbor seal (Phoca vitulina richardsi)

A stranded 5-month-old female Pacific harbor seal (Phoca vitulina richardsi) was presented displaying tachypnea and diminished lung sounds. No neurological abnormalities were noted. The animal was treated for verminous pneumonia, but died 2 wk later. Gross necropsy examination revealed a severe obstructive verminous pneumonia associated with large numbers of Otostrongylus circumlitus. In addition, the majority of the right cerebral hemisphere was absent, with hypoplasia of the left cerebellar hemisphere, absence of the right pyramid, and malformation of the right occipital bone. Histopathologic findings included multifocal thrombosis and inflammation of pulmonary arteries, verminous pneumonia, and mild vacuolation of the subependymal white matter in the third ventricle representing swelling of myelin sheaths and edema. This is the first report of a hemicerebral anomaly in a marine mammal.     

Improved estimates of digestion correction factors and passage rates for harbor seal (Phoca vitulina) prey in the northeast Atlantic

Diet composition in pinnipeds is widely estimated using hard prey remains recovered from feces. To estimate the size and number of prey represented in fecal samples accurately, digestion correction factors (DCFs) must be applied to measurements and counts of fish otoliths and cephalopod beaks. In this study, 101 whole prey feeding trials were conducted with six harbor seals (Phoca vitulina) and 18 prey species. We derived species‐ and grade‐specific estimates of digestion coefficients (DCs) and species‐specific recovery rates (RRs) to account for partial and complete digestion, respectively. Greater than 98% of otoliths were passed within three days of consumption. RRs were smallest for Atlantic salmon smolts (RR = 0.306, SE = 0.031) and increasingly larger for sandeels (RR = 0.494, SE = 0.017), flatfish (RR = 0.789, SE = 0.033), and large gadoids (RR = 0.944, SE = 0.034). Species‐specific otolith width DCs were smallest for Trisopterus species (DC = 1.14, SE = 0.015) and increasingly larger for flatfish (DC = 1.27, SE = 0.045), large gadoids (DC = 1.32, SE = 0.067) and sandeels (DC = 1.57, SE = 0.035). RRs were similar to those from gray seals (Halichoerus grypus), but harbor seal species‐ and grade‐specific DCs were generally smaller. Differences in partial and complete digestion rates among prey species and between seal species highlight the importance of applying DCFs when reconstructing diet.     

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.     

An outbreak of a herpesvirus infection in harbor seals (Phoca vitulina)

During an outbreak of a herpesvirus infection in juvenile harbor seals, 11 out of 23 seals died. The duration of the disease in these 11 animals varied from 1-6 days. Nasal discharge, inflammation of the oral mucosa, vomiting, diarrhea and fever up to 40 degrees C were observed in the first days of the disease. In later stages coughing, anorexia and lethargy occurred. Severe necrosis of the liver and interstitial pneumonia were the most striking histopathological findings.     

The complete mitochondrial DNA sequence of the harbor seal,Phoca vitulina

Summary The nucleotide sequence of the mitochondrial DNA (mtDNA) of the harbor seal, Phoca vitulina, was determined. The total length of the molecule was 16,826 bp. The organization of the coding regions of the molecule conforms with that of other mammals, but the control region is unusually long. A considerable portion of the control region is made up of short repeats with the motif GTACAC particularly frequent. The two rRNA genes and the 13 peptide-coding genes of the harbor seal, fin whale, cow, human, mouse, and rat were compared and the relationships between the different species assessed. At ordinal level the 12S rRNA gene and 7 out of the 13 peptide-coding genes yielded a congruent topological tree of the mtDNA relationship between the seal, cow, whale, human, and the rodents. In this tree the whale and the cow join first, and this clade is most closely related to the seal.Offprint requests to: Ú. Árnason     

First report of the giant kidney worm (Dioctophyme renale) in a harbor seal (Phoca vitulina)

A male harbor seal (Phoca vitulina) was found moribund on the coast of New Jersey in January of 2003 and died a few hours later in the Marine Mammal Stranding Center. On necropsy, a single female Dioctophyme renale was recovered from the peritoneal cavity, and a tissue mass was found adjacent to the pelvic urethra and urinary bladder. Within this tissue mass were found D. renale ova. This is the first report of this nematode in the harbor seal and in a North American marine mammal.     

Behavioral effects of environmental enrichment on harbor seals (Phoca vitulina concolor) and gray seals (Halichoerus grypus)

Zoos and aquariums have been incorporating environmental enrichment into their animal care programs for the past 30 years to increase mental stimulation and promote natural behaviors. However, most attempts to document the effects of enrichment on animal behavior have focused on terrestrial mammals. Staff at the National Aquarium in Baltimore conducted an investigation of the behavioral effects of enrichment on the seven harbor seals and two gray seals housed in the aquarium's outdoor seal exhibit. We expected that enrichment would change the amount of time the animals spent engaged in specific behaviors. The behaviors recorded were: resting in water, resting hauled out, maintenance, breeding display, breeding behavior, aggression, pattern swimming, random swimming, exploration, and out of sight. Activity levels (random swimming and exploration) were expected to increase, while stereotypic behaviors (pattern swimming) were expected to decrease. The frequency and duration of behaviors were documented for 90 hr in both the control phase (without enrichment) and the experimental phase (with enrichment). Statistically significant differences (P<0.05) in the time spent in pattern swimming, random swimming, exploration, and out of sight were observed between the two phases. With enrichment, pattern swimming and out of sight decreased, while random swimming and exploration behavior increased. These findings demonstrate that enrichment can promote behaviors (random swimming and exploration) that are likely to be normal for phocids in the wild, and that may contribute to the behavioral complexity of these seals in captivity. Zoo Biol 21:375–387, 2002. © 2002 Wiley‐Liss, Inc.     

Hematology and serum chemistry comparisons between free-ranging and rehabilitated harbor seal (Phoca vitulina richardsi) pups

The objectives of this study were to compare the hematology and serum chemistry values between free-ranging and stranded harbor seal (Phoca vitulina richardsi) pups and to ascertain how blood values of stranded pups changed during the rehabilitation process. Coincident with these comparisons, reference values were obtained for free-ranging pups. Stranded harbor seal pups (n = 28) recovered from areas between Pebble Beach and Moss Landing, California (USA) were admitted to The Marine Mammal Center, Sausalito, from March to May 1995, 1996, and 1998. Blood samples were collected from harbor seal pups before and after rehabilitation. As a control group, wild harbor seal pups were captured at Pebble Beach and Elkhorn Slough (n = 42) during the 1995, 1996, and 1998 pupping seasons. Mean eosinophil and calcium values of wild pups were significantly greater than those of newly admitted pups, whereas mean bands, aspartate aminotransferase, alanine aminotransferase, total bilirubin, and chloride values were significantly lower (P < or = 0.05). Mean neutrophil, band, lymphocyte, eosinophil, basophil, calcium, phosphorus, blood urea nitrogen, potassium, total protein, and globulin values of rehabilitated pups increased significantly after 2-3 mo in captivity, whereas, mean red blood cell, hemoglobin, hematocrit, cholesterol, and total bilirubin values decreased significantly (P < or = 0.05).     

Interannual differences in postrelease movements of rehabilitated harbor seal pups (Phoca vitulina richardii) in the Salish Sea

Despite the large number of harbor seals (Phoca vitulina richardii) rehabilitated worldwide, few studies have been conducted on postrelease movement and behavior of rehabilitated harbor seal pups. We compared interannual differences in movements and survival of 24 rehabilitated seal pups released in the Salish Sea in 2010 (n = 10), 2012 (n = 5), 2013 (n = 5), and 2014 (n = 4). We also compared the postrelease movement of these seals to the movement of 10 wild seal pups tracked in the same ecosystem in 2010. Transmission duration, total cumulative distance, and average daily distance varied annually. Maximum linear distance traveled from the release site was similar for the rehabilitated seal groups. Compared to wild seals (n = 10), and consistent with prior studies, rehabilitated pups (n = 24) traveled significantly farther daily and cumulatively than wild weaned pups. Unlike in a prior study in this ecosystem, we found no significant difference between transmission duration in wild and rehabilitated pups.     

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.     

Postrelease movement of rehabilitated harbor seal (Phoca vitulina richardii) pups compared with cohort‐matched wild seal pups

Harbor seal (Phoca vitulina richardii) populations in the inland waters of Washington and British Columbia are at or near carrying capacity. Stranded pups often are collected and admitted to rehabilitation centers, and then released when they reach a weight of 22 kg and meet a variety of preestablished health and release conditions. While rehabilitation is common practice, it is unclear if rehabilitated seal pups behave like wild weaned pups. Using satellite transmitters, we compared movement patterns of 10 rehabilitated pups with 10 wild weaned pups. When released, rehabilitated seals were longer and heavier than wild pups, while wild pups had a larger mean axillary girth. No clinically different blood parameters were detected. On average, rehabilitated harbor seal pups traveled nearly twice as far cumulatively, almost three times as far daily, and dispersed over three times as far from the release site compared to wild weaned seals. Additionally, wild harbor seals transmitted nearly twice as long as did rehabilitated seals. These patterns suggest that learned behavior during the brief 3–4 wk nursing period likely enables wild harbor seal pups to move less daily and remain closer to their weaning site than rehabilitated pups.     

Metabolic indicators in the skeletal muscles of harbor seals (Phoca vitulina)

The goal of this study was to determine the distribution of citrate synthase (CS), beta-hydroxyacyl coenzyme A dehydrogenase (HOAD), and lactate dehydrogenase (LDH) activities and myoglobin (Mb) concentration in the locomotor muscles (epaxial muscles) and heart of harbor seals. The entire epaxial musculature, which produces most of the power for submerged swimming, was removed and weighed, and three transverse sections (cranial, middle, and caudal) were taken along the muscle bundle. Multiple samples were taken along points on a circular grid using a 6-mm biopsy. A single sample was taken from the left ventricle of the heart. Muscle groups of similar function were taken from three dogs as a control. Mean values were calculated for four roughly equal quadrants in each transverse section of the epaxial muscles. There were no significant differences among the quadrants within any of the transverse sections for the three enzymes or Mb. However, there were significant differences in the mean enzyme activities and Mb concentrations along the length of the muscle. The middle and caudal sections had significantly higher mean levels of CS, LDH, and Mb than the cranial section, which may be correlated with power production during swimming. The enzyme ratios CS/HOAD and LDH/CS exhibited no variation within transverse sections or along the length of the epaxial muscles. Relative to the dog, the epaxial muscles and heart of the harbor seal had higher HOAD levels and lower CS/HOAD, which, taken together, indicate an increased capacity for aerobic lipid metabolism during diving.     

A captive feeding study with the Pacific harbor seal (Phoca vitulina richardii): Implications for scat analysis

Seven prey species ( n _total > 2,700) were fed to seven captive male Pacific harbor seals ( Phoca vitulina richardii ) in 177 experimental meals to quantify biases associated with scat analysis and current consumption models. Hard parts from an individual meal were recovered in an average of 3.8 ± 1.8 scats (range 1–10; mean ± SD). Overall, 57.7 ± 33.2% of otoliths and 89.5 ± 15.5% of squid beaks were recovered. Recovery rates varied, and prey with smaller, fragile otoliths were recovered in lesser quantities than prey with larger, robust otoliths. Recovery rates of all prey except pink salmon were improved by a mean of 31.7% when all diagnostic structures were included in estimates. Estimated recovery of pink salmon was 9.5 times that fed seals based on the all-structure technique. Mean length reduction of recovered otoliths was 20.4 ± 10.1%. Correction factors calculated from average length reduction improved length estimates for all fish species. Grade-specific length correction factors (gLCFs) reduced variability in all of the estimates and significantly improved estimates of prey with highly eroded otoliths including Pacific hake and shortbelly rockfish. The Biomass Reconstruction (BR) model accurately predicted biomass consumption within 4% of known consumption, whereas estimates based on frequency of occurrence were inaccurate.     

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)