Migrations of young spotted seals (Phoca largha) from Peter the Great Bay, Sea of Japan/East Sea, and the pattern of their use of seasonal habitats.

We studied the migrations of young spotted seals during their annual cycle. In May 2017, we attached satellite tags (SPOT-293A) to three individuals (two underyearlings and one yearling) captured at their breeding ground in Peter the Great Bay, western Sea of Japan/East Sea. The operational time of the installed tags ranged from 207 to 333 days; a total of 27195 locations were uploaded. All three seals migrated east and further north along the coast of the mainland. The average daily migration speed of the seals ranged between 70 and 135 km/day. The yearling moved faster than the underyearlings. During early August, they arrived at their summer habitats, which were located in the northern part of the Tatar Strait (Sea of Japan/East Sea) for the underyearling seals and in Aniva Bay (Sea of Okhotsk) for the yearling seal. While moving from the place of tagging to the summer feeding grounds, the seals covered a distance of 2300 to 3100 km. From August to October, each seal permanently stayed within the same isolated area. The reverse migration of all three seals began in November. When the seals traveled south, they used the same routes by which they had moved north in the spring, but they moved at a faster speed. By December, two seals returned to their natal islands, where both stayed until their transmitters stopped sending signals (in March 2018).

A mark-recapture approach for estimating population size of the endangered ringed seal (Phoca hispida saimensis).

Reliable population estimates are fundamental to the conservation of endangered species. We evaluate here the use of photo-identification (photo-ID) and mark-recapture techniques for estimating the population size of the endangered Saimaa ringed seal (Phoca hispida saimensis). Photo-ID data based on the unique pelage patterns of individuals were collected by means of camera traps and boat-based surveys during the molting season in two of the species' main breeding areas, over a period of five years in the Pihlajavesi basin and eight years in the Haukivesi basin. An open model approach provided minimum population estimates for these two basins. The results indicated high survival rates and site fidelity among the adult seals. More accurate estimates can be obtained in the future by increasing the surveying effort both spatially and temporally. The method presented here proved effective for evaluating population size objectively, whereas the results of the current snow lair censuses are dependent on varying winter conditions, for instance. We therefore suggest that a photo-ID-based non-invasive mark-recapture method should be used for estimating Saimaa ringed seal abundances in order to ensure reliable, transparent population monitoring under changing climatic conditions.

Echoes from the past: Regional variations in recovery within a harbour seal population.

Terrestrial and marine wildlife populations have been severely reduced by hunting, fishing and habitat destruction, especially in the last centuries. Although management regulations have led to the recovery of some populations, the underlying processes are not always well understood. This study uses a 40-year time series of counts of harbour seals (Phoca vitulina) in the Wadden Sea to study these processes, and demonstrates the influence of historical regional differences in management regimes on the recovery of this population. While the Wadden Sea is considered one ecologically coupled zone, with a distinct harbour seal population, the area is divided into four geo-political regions i.e. the Netherlands, Lower Saxony including Hamburg, Schleswig-Holstein and Denmark. Gradually, seal hunting was banned between 1962 and 1977 in the different regions. Counts of moulting harbour seals and pup counts, obtained during aerial surveys between 1974 and 2014, show a population growth from approximately 4500 to 39,000 individuals. Population growth models were developed to assess if population growth differed between regions, taking into account two Phocine Distemper Virus (PDV) epizootics, in 1988 and 2002 which seriously affected the population. After a slow start prior to the first epizootic, the overall population grew exponentially at rates close to assumed maximum rates of increase in a harbour seal population. Recently, growth slowed down, potentially indicative of approaching carrying capacity. Regional differences in growth rates were demonstrated, with the highest recovery in Netherlands after the first PDV epizootic (i.e. 17.9%), suggesting that growth was fuelled by migration from the other regions, where growth remained at or below the intrinsic growth rate (13%). The seals' distribution changed, and although the proportion of seals counted in the German regions declined, they remained by far the most important pupping region, with approximately 70% of all pups being born there. It is hypothesised that differences in hunting regime, preceding the protection in the 1960's and 1970's, created unbalance in the distribution of breeding females throughout the Wadden Sea, which prevailed for decades. Breeding site fidelity promoted the growth in pup numbers at less affected breeding sites, while recolonisation of new breeding areas would be suppressed by the philopatry displayed by the animals born there. This study shows that for long-lived species, variable management regimes in this case hunting regulations, across a species' range can drive population dynamics for several generations.

Aerial low-frequency hearing in captive and free-ranging harbour seals (Phoca vitulina) measured using auditory brainstem responses.

The hearing sensitivity of 18 free-ranging and 10 captive harbour seals (Phoca vitulina) to aerial sounds was measured in the presence of typical environmental noise through auditory brainstem response measurements. A focus was put on the comparative hearing sensitivity at low frequencies. Low- and mid-frequency thresholds appeared to be elevated in both captive and free-ranging seals, but this is likely due to masking effects and limitations of the methodology used. The data also showed individual variability in hearing sensitivity with probable age-related hearing loss found in two old harbour seals. These results suggest that the acoustic sensitivity of free-ranging animals was not negatively affected by the soundscape they experienced in the wild.

[The reactivity of liver monoamine oxidase in the ringed seal Phoca hispida ladogensis].

The goal of the work consisted in study of substrate and inhibitor specificity of liver monoamine oxidase (MAO) of the freshwater Ladoga subspecies of the ringed seal Phoca hispida ladogensis. The studied enzyme has been established to have large substrate specificity by deaminating, apart from eight classic substrates of MAO of terrestrial mammals, also histamine, the diamine oxidase substrate. It is found out that the deamination rates of benzylamine, beta-phenylethylamine, and N-methylhistamine almost one order higher than rates of deamination of serotonin and noradrenaline. MAO of the seal liver does not deamnate putrescine and cadaverine and is not sensitive to 10(-2) M semicarbazide. We calculated bimolecular constants of interaction rates of inhibitors chlorgiline, deprenyl, berberine, sanguinarine, chelidonine, and four derivatives of acridine with the enzyme at deamination of nine substrates. By the method of the substrate-inhibitor analysis, we showed the enzyme heterogeneity, i. e., the existence in the seal liver of at least two different MAO.

The effect of diet and feeding level on survival and weight gain of hand-raised harbor seal pups (Phoca vitulina).

Hundreds of stranded harbor seals pups (Phoca vitulina) are brought to wildlife rescue centers every year. Typical hand-rearing diets include artificial milk-replacers and diets based on macerated fish fed via gavage, but weight gains are often low and mortality rates can be high. This study compared survival and weight gain of orphaned seal pups fed either artificial milk-replacer or fish-formula. Pups admitted to the facility in summer 2007 (n=145) and 2008 (n=98) were randomly assigned to one of two diets and fed by gavage until weaning. In 2007, pups fed milk-replacer gained more (43 ± 12 g/d) than those fed fish-formula (loss of 13 ± 6 g/d; P<0.002). In 2008, when intake was increased from 8 to 11% of body weight daily, weight gain improved for both diets but remained higher in pups fed milk-replacer (123 ± 12 g/d, vs. loss of 6 ± 8 g/day; P<0.001). Pup survival to weaning was significantly higher in 2008 than 2007 (P<0.001) and was higher for pups on milk-replacer compared with those on fish-formula (P<0.05). Survival was also correlated with body weight at admittance (P<0.001). Although neither diet achieved the weight gains recorded in mother-raised pups (400-800 g/d), the artificial milk-replacer was clearly more successful, and pups fared better in the second year of the study when intake was higher.

Geography and stage of development affect persistent organic pollutants in stranded and wild-caught harbor seal pups from central California.

Persistent organic pollutants have been associated with disease susceptibility and decreased immunity in marine mammals. Concentrations of polychlorinated biphenyls (PCBs), dichlorodiphenyltrichloroethane and its metabolites (DDTs), polybrominated diphenyl ethers (PBDEs), chlordanes (CHLDs), and hexachlorocyclohexane isomers (HCHs) were evaluated in terms of stage of development and likely exposure routes (in utero, suckling, fasting) in the blubber of 202 stranded and wild-caught, primarily young of the year (n=177), harbor seals (Phoca vitulina) in the central California coast. This is the first report of HCH concentrations in the blubber of California seals. Lipid normalized concentrations ranged from 200 to 330,000 ng/g for sum PCBs, 320-1,500,000 ng/g for sum DDTs, 23-63,000 ng/g for sum PBDEs, 29-29,000 ng/g for sum CHLDs, and 2-780 ng/g for sum HCHs. The highest concentrations were observed in harbor seal pups that suckled in the wild and then lost mass during the post-weaning fast. Among the pups sampled in the wild and those released from rehabilitation, there were no differences in mass, blubber depth, or percent lipid although contaminant concentrations were significantly higher in the pups which nursed in the wild. When geographic differences were evaluated in a subset of newborn animals collected near their birth locations, the ratio of sum DDTs to sum PCBs was significantly greater in samples from an area with agricultural inputs (Monterey), than one with industrial inputs (San Francisco Bay). A principal components analysis distinguished between seals from San Francisco Bay and Monterey Bay based on specific PCB and PBDE congeners and DDT metabolites. These data illustrate the important influence of life stage, nutritional status, and location on blubber contaminant levels, and thus the need to consider these factors when interpreting single sample measurements in marine mammals.

Accumulation features of trace elements in mass-stranded harbor seals (Phoca vitulina) in the North Sea coast in 2002: the body distribution and association with growth and nutrition status.

Body distribution and growth- and nutritional status-dependent accumulation of 21 trace elements were investigated in harbor seals (Phoca vitulina) stranded in the North Sea coast in 2002. Higher concentrations and burdens of Mn, Se, Mo, Ag, Sn, Hg, and Bi in the liver, Cd in the kidney, As in the blubber, and Co, Sr, and Ba in the bone were observed. Significant positive correlations of hepatic Se, Mo, Ag, Cd, Sn, Hg, Tl, and Bi with standard body length were found, while significant negative relationships were detected for Mn, As, Rb, Sr, and Sb in the liver. Concentrations of Co, Se, Sr, Sn, Hg, and Bi in the liver, V, Sr, Ag, Sn, and Hg in the kidney, V, Mn, Co, Rb, Sr, Sn, Ba, and Pb in the blubber increased with decreasing blubber thickness of harbor seals, indicating enrichment of these elements in the target tissue by emaciation.

A computational model of teeth and the developmental origins of morphological variation.

The relationship between the genotype and the phenotype, or the genotype-phenotype map, is generally approached with the tools of multivariate quantitative genetics and morphometrics. Whereas studies of development and mathematical models of development may offer new insights into the genotype-phenotype map, the challenge is to make them useful at the level of microevolution. Here we report a computational model of mammalian tooth development that combines parameters of genetic and cellular interactions to produce a three-dimensional tooth from a simple tooth primordia. We systematically tinkered with each of the model parameters to generate phenotypic variation and used geometric morphometric analyses to identify, or developmentally ordinate, parameters best explaining population-level variation of real teeth. To model the full range of developmentally possible morphologies, we used a population sample of ringed seals (Phoca hispida ladogensis). Seal dentitions show a high degree of variation, typically linked to the lack of exact occlusion. Our model suggests that despite the complexity of development and teeth, there may be a simple basis for dental variation. Changes in single parameters regulating signalling during cusp development may explain shape variation among individuals, whereas a parameter regulating epithelial growth may explain serial, tooth-to-tooth variation along the jaw. Our study provides a step towards integrating the genotype, development and the phenotype.

Postnatal development of muscle biochemistry in nursing harbor seal (Phoca vitulina) pups: limitations to diving behavior?

Adult marine mammal muscles rely upon a suite of adaptations for sustained aerobic metabolism in the absence of freely available oxygen (O(2)). Although the importance of these adaptations for supporting aerobic diving patterns of adults is well understood, little is known about postnatal muscle development in young marine mammals. However, the typical pattern of vertebrate muscle development, and reduced tissue O(2) stores and diving ability of young marine mammals suggest that the physiological properties of harbor seal (Phoca vitulina) pup muscle will differ from those of adults. We examined myoglobin (Mb) concentration, and the activities of citrate synthase (CS), beta-hydroxyacyl coA dehydrogenase (HOAD), and lactate dehydrogenase (LDH) in muscle biopsies from harbor seal pups throughout the nursing period, and compared these biochemical parameters to those of adults. Pups had reduced O(2) carrying capacity ([Mb] 28-41% lower than adults) and reduced metabolically scaled catabolic enzyme activities (LDH/RMR 20-58% and CS/RMR 29-89% lower than adults), indicating that harbor seal pup muscles are biochemically immature at birth and weaning. This suggests that pup muscles do not have the ability to support either the aerobic or anaerobic performance of adult seals. This immaturity may contribute to the lower diving capacity and behavior in younger pups. In addition, the trends in myoglobin concentration and enzyme activity seen in this study appear to be developmental and/or exercise-driven responses that together work to produce the hypoxic endurance phenotype seen in adults, rather than allometric effects due to body size.

Phagocytosis in pup and adult harbour, grey and harp seals.

Knowledge on pinniped immunology is still in its infancy. For instance, age-related and developmental aspects of the immune system in pinnipeds need to be better described. The present study examined the phagocytic activity and efficiency of harbour, grey and harp seal leukocytes. In the first part of the study, peripheral blood was collected from captive female harbour seals of various ages. Data showed an age-related decrease in phagocytosis in female harbour seals from sub-adult to adulthood. In the second part of the study, changes in phagocytosis were quantified during lactation in wild newborn harbour, grey and harp seals and in their mothers (harp and grey seals). In newborns of the same age, leukocytes of harbour and harp seals phagocytosed less than those of grey seal pups. The phagocytic activity and efficiency increased significantly from early to mid-lactation in newborn harbour seals, and from early to late lactation in newborn grey seals, which could suggest that the transfer of phagocytosis-promoting factor(s) in colostrum is an important feature of temporary protection for pups. In contrast, no changes in phagocytic activity and efficiency were observed in lactating females of the two seal species, harp and grey, examined. At late lactation, phagocytic activity in both grey and harp seal pups and phagocytic efficiency in grey seal pups were significantly higher than in their mothers. These results could reflect either the capacity of phagocytes of the newborn harp and grey seals to respond to pathogens. Results from this study suggest that the phagocytosis of the seal species examined is not fully developed at birth as it generally increases in pups during lactation. Thereafter, the phagocytic activity of seals appears to decrease throughout adulthood.

Re-alimentation in harbor seal pups: effects on the somatotropic axis and growth rate.

The metabolic hormones, growth hormone (GH) and insulin-like growth factor (IGF)-I, together with IGF binding proteins (IGFBP), have been well studied in domestic species and are the primary components of the somatotropic axis. This hormone axis is responsive to nutrient intake, associated with growth rate, and accretion of protein and adipose. However, this relationship has not been evaluated in species that rely heavily on adipose stores for survival, such as pinnipeds. The primary objectives of this research were to investigate the response of the somatotropic axis to reduced nutrient intake and re-alimentation in rehabilitated harbor seal pups, and to assess if these hormones are related to nutritional status and growth rate in harbor seals. Stranded harbor seal pups (n=24) arrived at the rehabilitation facility very thin after fasting for several days (nutritional nadir). Throughout rehabilitation nutrient intake increased and pups gained mass and body condition. Concentrations of GH and IGFBP-2 decreased with re-alimentation, while IGF-I and IGFBP-3 concentrations increased. Overall, GH and IGFBP-2 were negatively associated and IGF-I and IGFBP-3 were positively associated with growth rate and increased body condition of harbor sea pups. Further, the magnitude of the growth response was related to the magnitude in response of the somatotropic axis to varied levels of intake. These data suggest that multiple components of the somatotropic axis may be used to assess the energy status of individuals and may also provide information on the level of feed intake that is predictive of growth rate.

Growth variation in skull morphology of Kuril harbor seals (Phoca vitulina stejnegeri) and spotted seals (Phoca largha) in Hokkaido, Japan.

We examined morphological growth variations in skull features between the Kuril harbor seal and the spotted seal in Hokkaido, Japan. Skulls from 80 Kuril harbor seals and 41 spotted seals were collected, and we measured 29 metric and 6 non-metric cranial characteristics. Three growth classes were defined according to the postnatal developmental stage: pups (0 year), subadults (1-4 years old) and adults (more than 5 years old). We detected sexual dimorphism in Kuril harbor seal pups, subadults, and adults. Although interspecies differences were detected in each growth class, Kuril harbor seals were larger and more massive than spotted seals; this feature was already detectable in pups. We did not detect certain cranial characteristics with which to identify the two species, but it was possible to identify any unknown specimens to their species, sex, and growth class using the cranial data generated in this study. Using 6 non-metric cranial characteristics, we identified significant interspecies differences with regard to the shape of the temporozygomatic suture and the extent of the nasal-incisive suture; the shape of the temporozygomatic suture and the shape of the nares were indicators of growth class in Kuril harbor seals. Although non-metric cranial characteristics have a lower discriminating power than metric characteristics, they are easy to use in the field even by inexperienced researchers.

Stage-structured transmission of phocine distemper virus in the Dutch 2002 outbreak.

Heterogeneities in transmission among hosts can be very important in shaping infectious disease dynamics. In mammals with strong social organization, such heterogeneities are often structured by functional stage: juveniles, subadults and adults. We investigate the importance of such stage-related heterogeneities in shaping the 2002 phocine distemper virus (PDV) outbreak in the Dutch Wadden Sea, when more than 40 per cent of the harbour seals were killed. We do this by comparing the statistical fit of a hierarchy of models with varying transmission complexity: homogeneous versus heterogeneous mixing and density- versus frequency-dependent transmission. We use the stranding data as a proxy for incidence and use Poisson likelihoods to estimate the 'who acquires infection from whom' (WAIFW) matrix. Statistically, the model with strong heterogeneous mixing and density-dependent transmission was found to best describe the transmission dynamics. However, patterns of incidence support a model of frequency-dependent transmission among adults and juveniles. Based on the maximum-likelihood WAIFW matrix estimates, we use the next-generation formalism to calculate an R(0) between 2 and 2.5 for the Dutch 2002 PDV epidemic.

Pup production and breeding distribution of the Caspian seal (Phoca caspica) in relation to human impacts.

Aerial surveys of Caspian seals on the winter ice field in Kazakhstan territorial waters were carried out in February 2005 and 2006 to assess the annual pup production for the species and natural predation on newborn pups. Estimated pup production was 21063 in 2005 and 16905 in 2006 (including an estimated figure for pups born in Russian territory in each year). The breeding population size of approximately 20 000 females is much less than published estimates from the late 1980s. Eagles were the principal natural predators of pups. Commercial icebreaker routes passed through areas of dense pup concentrations in 2006, although not in 2005. Our findings have important implications for the development of conservation strategies for the species. Natural mortality, loss to predators, and, more important, the current hunting quota substantially exceed the recruitment of the Caspian seal population. Anthropogenic sources of mortality should be managed to avoid further declines in the species.

[Structural organization of mesenteric lymph nodes in postnatal development of the Baikal seal, Pusa sibirica Gmel].

Structural organization of mesenteric lymph nodes in the Baikal seal has been studied with regard to its age-dependent changes. It has been shown that the relative area of connective tissue structures (the capsule and trabeculae) increases during postnatal development, while the areas of the cortex and medulla decrease. The proportions of secondary lymph nodes and paracortical zone in the cortex become smaller, with the corticomedullary index tending to decrease with age. These phenomena indicate that mesenteric lymph nodes undergo regression during postnatal ontogeny, with their lymphopoietic function being attenuated. Lower values of the corticomedullary index in Baikal seal pups aged 1 month to 4 years are apparently explained by a decrease in the proportion of cortex substance related to the enhancement of the motor function of lymph nodes.

Quasi-Poisson vs. negative binomial regression: how should we model overdispersed count data?

Quasi-Poisson and negative binomial regression models have equal numbers of parameters, and either could be used for overdispersed count data. While they often give similar results, there can be striking differences in estimating the effects of covariates. We explain when and why such differences occur. The variance of a quasi-Poisson model is a linear function of the mean while the variance of a negative binomial model is a quadratic function of the mean. These variance relationships affect the weights in the iteratively weighted least-squares algorithm of fitting models to data. Because the variance is a function of the mean, large and small counts get weighted differently in quasi-Poisson and negative binomial regression. We provide an example using harbor seal counts from aerial surveys. These counts are affected by date, time of day, and time relative to low tide. We present results on a data set that showed a dramatic difference on estimating abundance of harbor seals when using quasi-Poisson vs. negative binomial regression. This difference is described and explained in light of the different weighting used in each regression method. A general understanding of weighting can help ecologists choose between these two methods.

A longitudinal and cross-sectional analysis of total body oxygen store development in nursing harbor seals (Phoca vitulina).

This study compared the efficacy of longitudinal and cross-sectional sampling regimes for detecting developmental changes in total body oxygen (TBO(2)) stores that accompany behavioral development in free-ranging harbor seal pups. TBO(2) stores were estimated for pup (n = 146) and adult female (n = 20) harbor seals. Age related changes were compared between pups captured repeatedly during the lactation period (longitudinal dataset) and a second group of pups handled only once (cross-sectional dataset). At each handling, hematocrit, hemoglobin, red blood cell count, total plasma volume, blood volume, muscle myoglobin concentration, and blood and muscle oxygen stores were determined. Comparisons across age categories revealed newborn blood oxygen stores were initially elevated, declined to low values by early lactation, and increased through post-weaning. Muscle oxygen stores remained low and constant throughout lactation and only increased significantly post-weaning. Overall TBO(2)stores increased 17% during lactation, and weaned pups had TBO(2)stores that were 55% as large as those of adults. Thus, significant increases in TBO(2) stores must occur after weaning, as pups begin to forage independently. Results from the two sampling schemes did not differ, indicating that the logistically simpler cross-sectional design can be used to monitor physiological development in harbor seals.

Vocal development in captive harbor seal pups, Phoca vitulina richardii: age, sex, and individual differences.

The goal of this research was to determine when harbor seal pup vocalizations become sufficiently distinctive to allow individual recognition. A total of 4593 calls were analyzed from 15 captive pups. Nineteen were harsh, broadband, staccato calls used in an aggressive context. The rest were tonal "mother attraction calls," having an inverted "v"- or "u"-shaped spectrogram with harmonics and a fundamental frequency around 200-600 Hz. Calls were individually distinctive even in pups less than 2 weeks old, suggesting that mothers may be able to recognize pup vocalizations at this early age. Classification rates from discriminant function analysis were generally comparable to those of other phocids and less than in otariids, supporting the theory that recognition is more highly developed in otariids. Significant differences were found between male and female pup calls, and there were significant interactions between pup sex and age. The results of this study should be interpreted with caution until the findings are verified in wild harbor seal pups.

Erythropoietin concentration in developing harbor seals (Phoca vitulina).

Tissue hypoxia elicits the production of erythropoietin (Epo), a hormone that stimulates red blood cell production. In young diving mammals, oxygen is stored primarily in the blood, and blood oxygen stores increase significantly during the first weeks of life. In an effort to establish the role of Epo during this period of blood development, this study measured Epo concentration in plasma of 134 harbor seal (Phoca vitulina) pups and adults. Concurrent measurements of hematocrit (Hct), hemoglobin concentration [Hb], and red blood cell (RBC) counts allowed the evaluation of the effect of Epo on blood oxygen store capacity. Erythropoietin and most blood parameters varied with age. At birth, neonatal [Hb], Hct, and RBC were elevated, possibly due to the rapid expansion of plasma volume associated with growth rates of 0.5 kg/day. In contrast, Epo concentration increased from 6.64 +/- 0.83 mU/ml in newborns to 9.53 +/- 0.86 mU/ml in early nursing pups. Erythropoietin concentration remained elevated above newborn and adult concentration (5.71 +/- 0.79 mU/ml) through weaning, suggesting that Epo was responding to tissue hypoxia brought on by early anemia. Since similar changes in erythropoietin have been documented in terrestrial mammals, it appears that Epo plays a similar role in the blood development of harbor seals.