Pregnancy and ovulation rates in Grey seals (Halichoerus grypus) on the British coast

This study determined the pregnancy and ovulation rates in two stocks of Grey seals from Britain and described changes in pregnancy and ovulation rate with age. Seals sampled from the Hebrides during August 1978 and from the Fame Islands between March 1979 and October 1981 were used. Adult female pregnancy rate in the stock from the Fame Islands was estimated to be 94–29%, compared with 82–95% for seals from the Hebrides. These values were not significantly different. Pregnancy rate estimated by direct observation of a conceptus was comparable with an estimate from the presence or absence of corpora albicantia in the ovaries. By the age of four years and over (4 +), i.e. at maturity, 50% of females became pregnant. Pregnancy rate remained constant from age 5 + onwards. Pregnancy rate amongst females which became sexually mature at a younger than average age was lower than in older age classes.     

The relationship between body condition and the timing of implantation in pregnant Grey seals (Halichoerus grypus)

Adult female Grey seals were sampled from the Farne Islands, Northumbria at monthly intervals during 1980 and 1981. The mean date of implantation was 7 April and most females implanted between 11 March and 5 May. Sculp, core and total body weights declined most at the time of lactation in November and were lowest during February. These weights increased most rapidly during March and April; at the same time as implantation occurred. The pectoral blubber depth followed a similar pattern. Females in which the spring increase in body condition occurred earliest also had the earliest implantation. The study demonstrates a correlation between body condition and the time of implantation.     

Perinatal activities in the Grey seal (Halichoerus grypus)

Births and postnatal behaviour were observed at a breeding assembly of Grey seals on North Rona. Birth is extremely rapid and the cow immediately turns to smell the pup. Smelling continues at intervals, particularly during the first 10 min after birth and allows the cow to establish the identity of her pup. She also smells the placenta and defends it against attack by gulls. Disturbance by gulls, neighbouring seals and human intrusion may lead to the cow failing to establish a bond with the pup and consequently abandoning it.     

Movements, diving and foraging behaviour of grey seals (Halichoerus grypus)

This study presents the first direct observations of the movements and behaviour of free-ranging grey seals at sea. Radio and ultrasonic transmitters were attached to three sub-adult male grey seals which were then tracked from a suitable vessel. Behaviour at sea fell into one of three categories: travelling between haul-outs, short duration trips, and resting adjacent to haul-out sites. Travelling was characterized by direct, relatively fast horizontal movement and by V-shaped dive profiles. During short duration trips the seals swam slower and invariably exhibited square-wave dive profiles, spending approximately 60% of total dive duration at the maximum depth. Resting involved shallow dives close to haul-out sites and an absence of directed lateral movement.
The excellent navigational abilities of grey seals are illustrated by the rapid, direct swimming between distant haul-out sites. It is proposed that short duration trips are specifically for foraging because of their association with other piscivores, and because swimming was slow and mostly on or near the sea bed (grey seals are known to feed almost exclusively on demersal and benthic fish). These trips accounted for only 14% of the nine days that seal 1 was tracked. It is also proposed that the habit of diving to the sea bed whilst travelling between distant haul-out sites is to allow opportunistic foraging with only a small increase in total swimming distance.     

Sealpox in captive grey seals (Halichoerus grypus) and their handlers

Histopathologic, ultrastructural, and negative-staining studies indicated that nodular lesions on the flippers, head, and necks of recently weaned, captive grey seals (Halichoerus grypus) were similar to sealpox lesions reported from several other species of seals. Virions associated with the nodules were characteristic of the parapoxvirus subgroup of pox viruses. Two of the three persons handling the seals developed nodular lesions similar to "milker's nodules," the characteristic lesion in persons infected with parapoxvirus. The clinical course of the parapoxvirus infection in both the grey seals and their handlers is described. It was concluded that although sealpox is transmissible to man, the mild clinical manifestations place it in the nuisance category of zoonotic diseases.     

Finescale topographical correlates of behavioural investment in offspring by female grey seals, Halichoerus grypus

Grey seals breed colonially on substrates ranging from ice to rocky or sandy beaches. Clear differences in seal behaviour patterns exist among such broad classes of breeding habitat. However, finer scale topographical variation is likely to influence individual behaviour with consequences for pupping success. We examined topographical influences on the behaviour of breeding female grey seals by quantifying topography at a subseal size resolution. Using submetre resolution digital terrain models of two sites within a rocky breeding colony, we compared site topography in relation to observed differences in female behaviour at these sites. Females at both sites preferred breeding close to water (standing pools or sea) and frequently commuted between their pups and water. Topographical models indicated that one site was more costly for seals in terms of their locations and movements within the site. This was due to a lack of low-elevation land adjacent to the main access points from the sea and the reduced availability of pools. Females at this site showed reduced pup attendance and an increase in energetically costly behaviours, whilst females at the lower-cost site spent more time interacting with their pups and resting. These topographically induced behavioural differences are likely to affect the quantity and quality of pup provisioning by mothers and influence individual pupping site selection. Less costly sites are likely to be colonized preferentially and by larger, older and more dominant females, potentially generating finescale spatial heterogeneity in female quality within the breeding colony. Copyright 2000 The Association for the Study of Animal Behaviour.     

Grey seal, Halichoerus grypus, energetics: females invest more in male offspring

Grey seal females transfer large amounts of energy to their pups during the brief lactation period. The costs of lactation have been measured using weight changes of mother and pup pairs. Large females come ashore to give birth earlier in the season and lose weight more rapidly than smaller females. The sex ratio of Grey seal pups born is skewed towards males in the early part of the breeding season. Male pups are larger at birth and gain weight more rapidly than female pups, and their mothers show a correspondingly faster rate of weight loss than mothers of female pups. The energy costs of gestation and lactation to a Grey seal mother are 31 GJ for male pups and2–8 GJ for female pups. Males are therefore 10% more costly in energy terms to raise to weaning. Because, on average, large females arrive at breeding sites before smaller animals, biased results on weight changes would be obtained from methods which do not use repeated weighings. We suggest that the high efficiencies of lactation estimated for Harp seals compared with other phocid seals could be accounted for by such a bias.     

Microsatellite variation in grey seals (Halichoerus grypus) shows evidence of genetic differentiation between two British breeding colonies

Eight highly variable microsatellite loci were used to examine the genetic variability and differentiation of grey seals (Halichoerus grypus) at two widely spaced British breeding colonies. Samples were collected from adults and pups on the island of North Rona, off the north-west coast of Scotland, and on the Isle of May, situated at the mouth of the Firth of Forth on the east coast Highly significant differences in allele frequencies between these two sites were found for all eight loci, indicating considerable genetic differentiation. Thus, although grey seals are known to range over very large areas outside the breeding season, site fidelity of adults and philopatry of pups for these breeding colonies must be sufficiently common to have effects, through genetic drift, at the sub-population level. Migration rate was estimated using Wrighf's fixation index (F_ST), Slatkin's private alleles model and the new statistic, R_ST, which is analogous to (F_ST) but which takes into account the process of microsatellite mutation. An almost 8-fold discrepancy between the values we obtained provides cautionary evidence that microsatellite loci may contravene one or more of the assumptions on which these methods are based.     

Reproductive behaviour of the Grey seal (Halichoerus grypus) on Sable Island, Nova Scotia

We describe an unusual mating system, observed in a land-breeding colony of Grey seal, Halichoerus grypus , in the western Atlantic. Males and females begin to visit the breeding beach about a week before the season begins, but none stay ashore for long until the first pup is born. The cows are gregarious, probably return to the same part of the beach to give birth from one year to the next, and tend to remain in the general vicinity of the birth site during their two and a half week sojourn ashore. Within these limits, however, they are quite mobile, and the size, location and composition of the temporary aggregations which they form vary from one day to the next. The cows become thinner and more sedentary as oestrus approaches, but otherwise they give no overt signs of receptivity. The males do not defend territories, nor do they form dominance hierarchies. Instead, they compete for tenure, the right to remain within the shifting population of females. Tenured bulls directly test the receptivity of nearby cows from time to time, and they continually manoeuvre in ways which maximize their chances of being next to cows which are either in oestrus or likely to become so in the near future. Bulls which fail to establish themselves amongst females try to intercept cows as they are leaving for the sea at the end of their season, but their reproductive success, estimated in several ways, is significantly lower than that of bulls with tenure.
The system would be adaptive for seals which breed on the pack ice, but it is unique amongst land-breeding pinnipeds. If, as some circumstantial evidence suggests, Grey seals were originally pack-ice breeders, the persistence of such a system in a land-breeding colony raises some interesting questions about the plasticity of mating behaviour.     

Territorial behaviour in the Western Atlantic Grey seal (Halichoerus grypus)

Virtually all published accounts of territorial behaviour in the Grey seal apply to breeding males only. Long-range field studies reveal, however, that lactating cows also maintain territories for a period of several weeks. During the summer months, adults and sub-adults have specific resting places to which they return each day at low water and which they defend against other individuals of their own species.     

Behaviour of female and pup grey seals Halichoerus grypus during the breeding period at Froan, Norway

We studied the behaviour of grey seals, Halichoerus grypus , during the breeding period at Froan, Norway, and compared our findings with existing studies on grey seals at breeding sites in Britain and Canada. The pups at Froan spent more time in water than pups at other breeding sites. While on-shore, the pups at Froan spent most of their time resting, behaviour similar to pups at other breeding sites. Lactating females at Froan spent most of their time in the sea, thus differing from females at most other breeding sites which spend most of their time on-shore. While in the sea, the females at Froan spent most time diving, typically interrupted by regular periods of surface swimming.     

Seasonal movements and diurnal activity rhythms of the Grey seal (Halichoerus grypus)

The seasonal movements and diurnal activity rhythms of the Grey seals occurring along the Atlantic Coast of Nova Scotia and in the southern part of the Gulf of St Lawrence have been studied over a period of three and a half years. The seasonal movements of the adults can be divided into five periods or phases: (1) prepupping, (2) pupping and mating, (3) postmating, (4) spring concentration and (5) summer dispersal. The diurnal activity rhythm is attuned to the tides; the effects of the hours of daylight appear to be minimal. During the breeding season the cows go to sea as the tide is falling and return when it is rising. Adults of both sexes have a similar rhythm at other times of the year, except in mid-summer when the pattern is reversed.     

Trend in the pup production of grey seals Halichoerus grypus at Froan, Norway, from 1974 to 1987

Pup counts of grey seals at Froan, Norway, during the period 1974 to 1987 are given. The counts indicate an increase in pup production from 1974 to 1983. The counts from 1985 to 1987 indicate a stagnation and possibly a decrease in production. This is probably due to culls at the breeding ground, particularly during the 1984 breeding season.     

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.     

Finding fathers: spatio-temporal analysis of paternity assignment in grey seals (Halichoerus grypus)

Molecular studies of pinniped breeding systems exhibit a broad range of agreement and disagreement with observational indices of male breeding success. Grey seal studies have reported considerable discrepancies between genetic and behavioural paternity measures that have been interpreted as evidence of previously unidentified male strategies and/or tactics. Therefore, these studies have the power to fundamentally alter our perceptions of mating systems. However, other pinniped studies exhibit no such disagreements, and one possible explanation for disparities may be sampling biases in space and time. Therefore, it is essential that potential sampling biases are examined to evaluate the likelihood of previously unidentified male strategies. We examined paternities assigned at the North Rona grey seal colony between 1999 and 2002 in relation to concurrent detailed behavioural and locational data for males and females. We found that (i) for females observed in sexual interaction(s) during their oestrus period, it was highly probable that one of the interacting males fathered their next pup; (ii) over 80% of assigned paternities agreed with observations of the in-colony behaviour and spatio-temporal proximity of the males and females involved; and (iii) a minority of females exhibit mate choice and seek sires outside their local male's home range, although evidence suggests that these females mate on the colony rather than at sea. In conclusion, nearly all paternities assigned agreed with expectation based upon detailed knowledge of the spatio-temporal patterns of individuals during the breeding season. We found little evidence of unidentified male strategies at North Rona, Scotland, whereas further examination of mechanisms of female choice may be productive.     

The relative influence of age and weight on the reproductive behaviour of male grey seals Halichoerus grypus

The breeding behaviour of male grey seals Halichoerus grypus of known age and weight was studied on Sable Island, Nova Scotia. Branded males ranged in age from 8 to 16 years and although there was a positive relationship between age and weight, there was a large overlap between age classes. In general, there was an increase in length of tenure and rates of copulation with increasing male age but this was less marked among males between the ages of 13 and 16 years. Compared to older ones, males of 8 and 12 years were observed at more sites, travelled more extensively around the breeding colony, were transient more frequently and were unable to lie as close to females. There was no correlation between male body weight and either length of tenure or rates of copulation when males of age 8 (the smallest males) were excluded from the analysis. Most agonistic behaviour was made by larger, older males towards smaller, younger ones and appeared to be responsible for the short length of tenure and low reproductive rates of young bulls.