AGING LIVE ANTARCTIC FUR SEALS AND SOUTHERN ELEPHANT SEALS

This study describes a method for extracting post-canine or incisor teeth from live antarctic fur seals ( Arctocephalus gazella ) and southern elephant seals ( Mirounga leonina ) respectively and their use to determine age in a field situation. Dental elevators were used to loosen the teeth from the alveolus and periodental ligament. Most teeth were removed within l-2 min and a total of 214 and 81 teeth were collected by this method from antarctic fur seals and southern elephant seals respectively. No seal recaptured at intervals up to a year after a tooth was extracted showed signs of infection or distress related to removal of the tooth. Teeth were thin-sectioned for the purpose of aging. In both species cementum growth layer groups were a more satisfactory indicator of age than dentinal growth layer groups. Estimates of age from cementum growth layers were confirmed for Antarctic fur seals using seals which had been tagged as pups up to 16 yr before sampling.     

Factors affecting early mortality in the South American fur seal (Arctocephalus australis) in Peru: density-related effects and predation

Early mortality in the South American fur seal ( Arctocephatus australis ) in Peru is considerably higher than that seen in any other population of fur seal; 31–49% in the first month as opposed to a maximum of 20% in other populations. In 1987 and 1988, pup mortality was found to be enhanced by density-related effects and predation on pups by the southern sea-lion ( Olaria byronia ). At a high density beach, where 60% of the fur seals bred, mortality correlated with the number of females ashore as did aggression. Female aggression was driven to high levels by a combination of high density and movement within the colony to thermoregulate. However, even at a low density, beach mortality remained high due to a high level of predation by male southern sea-lions (low density beach 5·0–8·3% of all pups; high density beach 0·2%). Predation rates were lower at the high density beach because male fur seals expelled most attacking sea-lions during the breeding season (low density beach 10·0% of attacking sea-lions expelled; high density beach 58·8%).     

The contemporary condition and some demographic characteristics of the Steller sea lion (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>) reproductive group on Tyuleniy Island,Sea of Okhotsk

In 2010, the largest part of the Steller sea lion breeding community on Tyuleniy Island was located on the harem rookery of northern fur seals, which occupied the eastern beach, as well as on the western side of the island, which was free of fur seals. At the culmination of harem activity on June 29, 26.5% of the animals at the age of 1+ concentrated on the eastern beach and 41.1%, on the western beach in the daytime. However, 52.3% of the pups were born on the eastern beach and only 30.4% were born on the western beach. Pups were also present on the capes: 9.1% of the pups were observed on the northern cape and 8.2% on the southern cape, while the main population on these sites consisted of non-harem bulls, bachelors, and young animals. At the peak of harem activity, the number of females per one harem bull was 13.1 at sites 1 to 3 of the eastern beach and each of them, on average, had 1.05 pups; on sites 7–12 there were, respectively, 9.1 females and 1.42 pups per female, and on the western beach, 21.7 females and 0.64 pups. The resulting abundance of sea lions on Tyuleniy Island in 2010 exceeded 1500, which was almost ten times as many as their number in 1989. A total of about 100 bulls, 60 harem bulls, 1000 females, and 700 pups were recorded there. Half-bulls and young animals amounted to one-third of the entire population. Meanwhile the overall sex ratio at the culmination of harem activity was 11.5 females per one bull and 18.8 per one harem bull. About 75% of the females belonged to the parous group. The mortality rate among newborns reached 5.4%. No mortality was observed in adults. As many as 133 previously branded Steller sea lions were found and 109 of them (81.9%) were immigrants. Among immigrants, 29% were branded individuals of reproductive groups from the Kuril Islands, 54% were from the Iony Islands, 16% were from the Yamsky Islands, and about 1% were from Kamchatka. Four-year-old individuals predominated among the branded immigrants (23.8%). The oldest Steller sea lion (21 years of age) was one that was branded on the Srednego Islands in 1989. The rate of marked animal return from 175 pups that were branded on Tyuleniy Island the year before was 13.8%.     

Unmarked individuals in mark–recapture studies: Comparisons of marked and unmarked southern elephant seals at Marion Island

The presence of unmarked individuals is common in mark–recapture study populations; however, their origin and significance in terms of population dynamics remain poorly understood. At Marion Island, southern Indian Ocean, where virtually all southern elephant seal Mirounga leonina pups born annually (1983–2008) were marked in a long‐term mark–resight study, large numbers of unmarked seals occur. Unmarked seals originate either from marker (tag) loss or from immigration. We aimed to identify patterns in the occurrence of marked and unmarked individuals that will allude to the possible origin and significance of the untagged component of the population, predicting that tag loss will add untagged seals to mainly adult age categories whereas migrating untagged individuals will be mostly juveniles. We fitted a generalized linear model using the factors month, year and age‐class to explain the relative abundance of untagged seals (tag ratio) from 1997 to 2009. Site usage of untagged seals relative to tagged seals was assessed using a binomial test. Untagged seals, predominantly juveniles, were present in the highest proportions relative to tagged seals during the winter haulout (tagged seals/total seals less than 0.3) and the lowest proportion (approximately 0.5) during the female breeding haulout, increasing in relative abundance from 1997 to 2009. Untagged seals were distributed evenly across suitable haulout sites while tagged seals displayed high local site fidelity and occurred in greater numbers at or near large breeding beaches. Untagged seals are considered to be mostly migrant seals that disperse from other islands within the southern Indian Ocean and haul out at Marion Island during non‐breeding haulouts in particular. Some of these seals immigrate to the breeding population, which can be a key component of the local population dynamics. We emphasize the need for mark–recapture studies to evaluate the role of the unmarked component of a population, thereby inducing a more confident estimation of demographic parameters from the marked sample.     

DECLINES IN HARBOR SEAL (PHOCA VITULINA) NUMBERS IN GLACIER BAY NATIONAL PARK, ALASKA, 1992–2002

Glacier Bay National Park had one of the largest breeding aggregations of harbor seals in Alaska, and it is functionally the only marine reserve for harbor seals in Alaska; yet, numbers of seals in the Bay are declining rapidly. Understanding why seals in Glacier Bay are declining may clarify their minimal habitat needs. We estimated population trends using models that controlled for environmental and observer‐related factors. In 1992, 6,200 seals were counted on icebergs in a tidewater glacial fjord and at terrestrial sites; by 2002 only 2,550 seals were counted at these same haul‐outs. Numbers of non‐pups in the glacial fjord declined by 6.6%/yr (?39%/8 yr) in June and by 9.6%/yr (?63%/11 yr) in August and at all other haul‐outs by 14.5%/yr (?75%/10 yr) during August. In the glacial fjord the number of pups remained steady from 1994 to 1999 and made up an increasing proportion of seals counted (5.4%/yr), and the proportion of pups peaked at 34%–36%. The rapid declines do not appear to be due to changes in seal behavior or redistribution. The declines reinforce genetic evidence that harbor seals in Glacier Bay are demographically isolated from other populations and indicate that current management stocks need to be redefined. Changes in Glacier Bay's ecosystem and population demographic data from the glacial fjord suggest that interspecific competition and predation are likely factors in the declines.     

Mass of weaned elephant seal pups in areas of low and high human presence

On sub-Antarctic Macquarie Island, we examined pup weaning mass of southern elephant seals in relation to human presence. Pup weaning mass was previously found to be positively associated with 1st-year survivorship. Weaned pups were weighed in a remote area, Middle Beach, and in an area of relatively high human presence, Isthmus East. The areas were reasonably similar in beach topography, wind and surf conditions, numbers of seals present per kilometre of coastline, and numbers of males and females present in harems. For a sub-sample of measured pups, data on the respective maternal size were collected using photogrammetry. Both male and female weaned pups on Middle Beach were significantly heavier than those on Isthmus East. Estimated length of mothers was significantly higher on Middle Beach. In proportion to their own size, mothers in both areas produced weaners of similar mass, indicating no direct effect of human disturbance on the efficiency of lactation. It remained unclear whether the area differences in maternal and pup size were due to natural or human-related factors.     

Population status and breeding season chronology of Heard Island fur seals

Fur seals were eliminated by sealers at Heard Island soon after its discovery in the 1850s. The first recorded breeding of Antarctic fur seals (Arctocephalus gazella) since sealing was reported in early 1963 (two pups). The most recent survey of the Heard Island fur-seal population was undertaken between November 2000 and March 2001, when 1,012 Antarctic fur-seal pups were born. This represents a fourfold increase since the last complete census in 1987/1988 (13 years), when 248 births were recorded. Pup estimates and counts available for eight breeding seasons since 1962/1963 suggest the population has been increasing at between 12 and 20% per year. Based on pup production, the breeding population is estimated to number approximately 4,100 seals. The number of fur seals on Heard Island peaked in late February/early March at 29,256 indicating that, in addition to the breeding population, a significant number of seals born elsewhere haul out on the island. Most of these are moulting sub-adult and adult males. As in 1987/1988, only one subantarctic fur-seal pup (A. tropicalis) was observed, suggesting this species is not colonising the island, as has been speculated.     

Harbor seal use of glacier ice and terrestrial haul‐outs in the Kenai Fjords,Alaska

Harbor seals, Phoca vitulina, use diverse haul‐out substrates including ice calved by tidewater glaciers. Numbers of seals at glacial and terrestrial haul‐outs on the southeastern Kenai Peninsula, Alaska, were assessed using aerial, vessel, and video surveys. Mean annual abundance at glacial and terrestrial haul‐outs differed temporally. From 2004 to 2011, numbers of seals counted during the molt increased 5.4%/yr at glacial haul‐outs and 9%/yr at terrestrial haul‐outs while numbers of pups increased 5.0%/yr at glacial sites and 1.5%/yr at terrestrial sites. Numbers of seals without pups counted during pupping increased 7.96%/yr at glacial sites and 5.1%/yr at terrestrial sites. Results indicate that pupping and molting locations are not equivalent and population monitoring during the molt does not necessarily reflect habitat association of pupping seals. Ratios of pups to total seals counted during pupping and the subsequent molt were used to contrast habitat use. Low proportions of pups at terrestrial haul‐outs, relative to most glacial haul‐outs, indicate an overall preference for pupping in glacial haul‐outs. High proportions of pups at most glacial sites (during pupping and molting) suggest reduced use of tidewater glacier habitats by nonbreeders and molting seals. Results suggest more seals associate with glacial haul‐outs than currently estimated.     

Prey selection of subadult male northern fur seals (Callorhinus ursinus) and evidence of dietary niche overlap with adult females during the breeding season

During the breeding season northern fur seals (Callorhinus ursinus) congregate on the Pribilof Islands in large numbers creating the potential for intraspecific competition. Due to the declining trend in the Pribilof Islands population of fur seals, it is important to understand how prey resources are partitioned among the population. Fur seals exhibit a high degree of sexual dimorphism resulting in energetic differences among age and sex classes. Therefore, we hypothesized that subadult male and adult female fur seals would differ in the type and size of prey consumed. We examined the diets of subadult male (age 2–8; mean mass 28–176 kg) and adult female (age ≥ 3 yr; mean mass 13–50 kg) seals on St. Paul Island from 1992 to 2000. Prey remains found in fecal samples were compared using niche overlap indices. There was nearly complete dietary niche overlap between subadult male and adult female fur seals. Walleye pollock (Theragra chalcogramma), Pacific salmon (Oncorhynchus spp.), Pacific herring (Clupeia pallasi), and cephalopods were common prey items found in the diets of both groups. We found differences in the size of pollock consumed and that geographic location of sample collection may be important in determining diet differences. Our results indicate high levels of dietary overlap among subadult male and adult female fur seals.     

SURVIVAL RATES FOR THE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

Abstract: Endangered Hawaiian monk seal ( Monachs schauinslandi ) pups at all the major breeding islands in the Northwestern Hawaiian Islands have been tagged since the early 1980s. Pups were double flipper tagged as soon as possible post-weaning. With few exceptions, an extensive tag resighting effort was conducted annually at the same islands. These resighting data were used to estimate seal survival rates from the time of tagging to age one at all locations using the ratio of seals alive in the second year to number of pups tagged. These survival rates among the islands, from weaning to age one, averaged over the years of the study, ranged from 0.80 to 0.90. For young seals over age one, capture-recapture methods were used to calculate survival pooled through several years, and these rates ranged from 0.85 to 0.98. At French Frigate Shoals and Laysan Island, the higher numbers of tagged pups allowed separate estimates of male and female survival to be calculated. These rates suggested that survival of immature females was better than males. Beginning in 1989, survival of immature seals at French Frigate Shoals declined sharply.     

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.     

Hematology and serum chemistry of harp (Phoca groenlandica) and hooded seals (Cystophora cristata) during the breeding season, in the Gulf of St. Lawrence, Canada

Standard hematologic and serum chemistry parameters were determined from 28 harp seals (Phoca groenlandica) and 20 hooded seals (Cystophora cristata) sampled from 6 March 2001 to 13 March 2001 during the breeding season. Whole blood was collected immediately postmortem from harp seal mother-pup pairs and from six hooded seal pups, and from live-captured adult hooded seals and three hooded seal pups; blood was analyzed within 24 hr at a local human hospital. A certified veterinary laboratory validated subsamples of whole blood and analyzed all serum chemistry parameters. Significant interlaboratory differences in mean values of packed cell volume (PCV) and mean cell volume (MCV) were found. Significant differences were found between samples from the five seal groups (adult male hooded seals, lactating female hooded seals, unweaned hooded seal pups; lactating female harp seals, and unweaned harp seal pups) for hematology and most serum chemistry parameters. In general, age-class influenced mean values of PCV, hemoglobin (HB), red blood cell (RBC) counts, MCV, mean cell hemoglobin (MCH), mean cell hemoglobin concentration (MCHC), and nucleated red blood cell (NRBC) counts per 100 leucocytes, but most age-related variations were species specific. Harp seal pups had significantly lower mean values of HB, PCV, MCH, and MCHC than did other seal groups, and significantly lower mean RBC counts than did hooded seal pups. Mean NRBC counts per 100 leukocytes were more than three times higher in harp seal pups than in hooded seal pups, but this difference was not statistically significant. Mean MCV were significantly lower in harp and hooded seal pups compared to those of adult harp and hooded seals. Differences in hemograms between pup species were likely because of the precocious development of hooded seal pups, which are weaned within 4 days, compared to 12 days for harp seal pups. Among adult seal groups, male hooded seals had significantly higher mean values of PCV and HB than did female harp and hooded seals, and significantly higher mean RBC counts than did adult female hooded seals. Among adult females, mean values of MCH and MCHC were statistically higher in hooded seals than in harp seals. Adult female harp and hooded seals did not differ significantly in other RBC parameters and mean leukocyte counts. Mean values of glucose, blood urea nitrogen, total bilirubin, alanine aminotransferase (ALT), alkaline phosphatase (ALP), total protein, and albumin showed species-specific variations between adults and pups. Except for ALP, few significant differences in mean enzyme activities of aspartate aminotransferase (AST), ALT, creatine kinase and gamma-glutamyltransferase were found between seal groups. Mean concentrations of electrolytes (calcium, phosphorus, sodium, potassium, chloride, magnesium, and total carbon dioxide) varied with age class, but variations in potassium and magnesium were species specific. Harp seal pups had significantly higher mean phosphorus and potassium levels compared to other seal groups.     

Holocene changes in the ecology of northern fur seals: insights from stable isotopes and archaeofauna

The remains of northern fur seals (Callorhinus ursinus) are among the most abundant of pinniped elements recovered from mainland coastal archaeological sites in both California and Oregon. This is surprising as all contemporary northern fur seals breed exclusively on offshore islands, primarily at high latitudes, and the species is otherwise pelagic. The vulnerability of these animals to human predation suggests that either humans were foraging much further offshore than has been presumed or alternatively that the ecology of these animals has shifted during the late Holocene. We used isotopic and archaeofaunal analysis of the remains of pinnipeds from the middle to late Holocene of central and northern California to clarify the breeding and foraging behavior, and migration patterns of these ancient animals. The carbon and nitrogen isotope compositions of ancient northern fur seals reveal that these animals fed as far offshore as they do today, and that they remained at middle latitudes throughout the year. From an archaeological site at Moss Landing, California, we identified 16 skeletal elements from at least 12 very small northern fur seal pups. From another site near Mendocino, California, we identified the remains of at least 6 pups. We estimate the size and age of 5 of the young animals using sex-specific regressions of body length on the short dentary length derived from measurements of modern specimens. Our estimates indicate these ancient pups were substantially smaller, and therefore younger, than modern 3-month-old northern fur seal pups from similar latitudes and their nitrogen isotope compositions suggest they had not been weaned. As present-day northern fur seals do not leave their rookeries until they are at least 4 months old, we consider it highly unlikely that these ancient pups swam to these mainland locations from some distant island rookery. While there are numerous nearshore rocky outcrops along the Mendocino Coast, which may have supported small breeding colonies, the Moss Landing site is centered on a 40-km-long sandy beach, and is more than 120 km from what at the time were the nearest offshore islands. We conclude that northern fur seal adult females, subadults, and pups whose remains were recovered at the Moss Landing archaeological site must have been taken at a mainland rookery. Evidence that northern fur seals once bred on the mainland at this central California location suggests that the abundant remains of these animals at numerous other archaeological sites along the California coast also reflect the presence of nearby mainland rookeries. Based on the relative abundance of their remains in ancient human occupation sites and the widespread distribution of sites where their remains have been found, it appears that northern fur seals were once the predominant pinniped throughout a region where they now only rarely occur. Furthermore, their presence along the central and northern California coasts appears to have once severely limited the distribution of other pinnipeds, which are now common to the region.     

Multiple mating strategies explain unexpected genetic mixing of New Zealand fur seals with two congenerics in a recently recolonized population

Human impacts on natural systems can cause local population extinctions, which may promote redistribution of taxa and secondary contact between divergent lineages. In mammalian populations that have mating systems shaped by polygyny and sexual selection, the potential for hybridization to ensue and persist depends on individual and demographic factors. At Macquarie Island, a recently formed fur seal population is comprised of both sexes of breeding Antarctic (Arctocephalus gazella) and subantarctic (A. tropicalis) fur seals, and an itinerant collection of male New Zealand fur seals (A. forsteri), presumed to be non-breeders due to their absence from principle breeding areas. The mating system of the three species is described as resource-defence polygyny: males defend beach territories containing breeding females for exclusive mating rights. A recent genetic study identified a high level of hybridization in the population (17-30%), unexpectedly involving all three species. This study examined the source of involvement in breeding by A. forsteri with respect to mating strategies operating in the population. Ninety-five (10%) pups born from 1992 to 2003 were genetically identified as New Zealand hybrids. Most resulted from reproduction within territories by New Zealand hybrids of both sexes, although some were conceived extra-territorially, indicating that males successfully utilize strategies other than territory holding to achieve paternities. Female reproductive status influenced mating partner and mating location, and females without pups were more likely to conceive extra-territorially and with A. forsteri males. This study illustrates an important consequence of low heterospecific discrimination in a sympatric population of long-lived mammals.     

Mother-young behaviour at Grey seal breeding beaches

A comparative study of mother-young relations at Grey seal breeding beaches was made and results from two sites in Orkney are described to illustrate the different situations. It was found that at the minimally disturbed beach the cows consistently and exclusively suckled their own pups. To achieve this a bond is formed between the mother and pup immediately after parturition. The cow uses a combination of senses to locate and identify her pup; she recognises and responds to her pup's call, she returns to where she last suckled and she identifies the pup by smell. The pup remains on the beach where it was last suckled, it probably does not recognise its mother and does not discriminate between cows when soliciting suckling. These results are considered to be representative of completely undisturbed behaviour in the wild. At the other beach there were many examples of inconsistent suckling; these are described and attributed to disturbed conditions created by the method of study which led to a break-down of the mother-young bond between many of the seals present. The situation was accentuated by the topography of the beach and the density of seals present.     

Does kin selection influence fostering behaviour in Antarctic fur seals (Arctocephalus gazella)?

A recent observation that female Antarctic fur seals foster pups born to related females raises the fascinating possibility that kin selection may promote altruistic behaviour even on a crowded breeding beach, where individual interactions are frequent and complex. However, the use of genetic markers to identify small numbers of unusually highly related individuals is fraught with difficulty due to the likely presence of genotyping errors and related problems. Consequently, we examined an enlarged dataset where errors had been reduced to an absolute minimum by a combination of close scrutiny and repeat genotyping. We find no support for the idea that females preferentially suckle pups born to female relatives. Instead, the previously reported pattern can be explained by a combination of genotyping errors and de novo mutations. Our study emphasizes the need for caution when interpreting rare events that occur at a rate approaching that expected for normal genotyping errors.     

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.     

BREEDING BIOLOGY OF SOUTHERN ELEPHANT SEALS IN PATAGONIA

Abstract: Elephant seals breed in Patagonia (Península Valdés, Argentina) from late August to early November, reaching peak numbers during the first week in October. Observations of this population over the past ten years yielded similar results. Eighty percent of the pups were born by 2 October. Most (96%) of 663 females marked during three breeding seasons gave birth to a pup. Females stayed on land a mean of 28 d, gave birth 6 d after arrival, nursed their pups for 22 d, and copulated a mean of 2.5 times 20 d after parturition and 2 d before departure. Copulations peaked during the third week in October. Males spent 57–80 d on land fasting and defending harems of up to 134 females (median 11–13 females, depending on year). Most (96%) marked females that gave birth ( n = 636) also weaned their pups successfully. Pup sex ratio was unity. Harems were smaller and breeding occurred about three weeks earlier in Patagonia than in other colonies. Thermal conditions, day length and food availability may explain clines in the timing of breeding events between populations, Other parameters of the breeding season for the expanding Patagonia colony are similar to those for declining southern elephant seal populations elsewhere.     

BODY TEMPERATURE IN THE NEWBORN SOUTHERN ELEPHANT SEAL, MIROUNGA LEONINA, AT MACQUARIE ISLAND

Newborn southern elephant seal pups were reported by Laws (1953) to be "to some extent poikilothermic at birth." Rectal temperatures of known age southern elephant seal pups were recorded during the 1985 pupping season at Macquarie Island. The mean pup rectal temperature was found to be 381°C ± 0.1°C SEM ( n = 131, range = 36.5°-39.1°C). Pups at two hours, six hours, and one day after birth had significantly higher rectal temperatures than pups two, three, or four days of age. Rectal temperatures of neonatal southern elephant seals were within the range observed for other pinnipeds, (but never as low as the 31°C previously observed for southern elephant seals at Signy Island in 1953). A significant though weak positive correlation was found between pup temperature and body weight. However, no correlation between pup temperature and age or any environmental factor was found. These observations demonstrated that southern elephant seal pups at Macquarie Island are homeothermic, rather than heterothermic from birth.     

Spatial and temporal distribution of Antarctic fur seals (Arctocephalus gazella) on the breeding grounds at Bird Island,South Georgia

Summary The number of Antarctic fur seals (Arctocephalus gazella) in the area of the Scotia Sea has been increasing. Observation of their distribution on the breeding grounds are important to help design and interpret censuses. These seals are highly polygynous. In the study area at Bird Island, South Georgia, females give birth and are mated in densely packed colonies located along the shore close to the tideline. Males establish territories in this area, but are also found further inland. This study examined the density-dependent processes regulating the instantaneous size of the male breeding population; the distribution of males in relation to space and the number of females available to be mated and the effect of gregarious behaviour of females on male dispersion. Males were only territorial on the beaches in the areas where most females gave birth and subsequently had their post-partum oestrus. There was an apparent lower (19–20 m²) and upper (40 m²) limit to territory size. Males were excluded from the beach areas when the average density on the beaches was greater than 5 males per 200 m². An asymptotic density of 10–11 males per 200 m² was reached on the beaches and 4–5 males per 200 m² elsewhere. These two asymptotic densities may represent the upper and lower limits of density for a territorial system of dispersion. A model of the temporal changes in female numbers suggested that the total number of females occupying an area of beach during the mating period was approximately twice the number at the peak of the season. There was no indication that males compensated for low female density by increasing territory size. Females and pups became more dispersed after the end of the mating period. It is suggested that one function of gregariousness in females is as a mechanism for mate selection. This study also has implications for methods used to measure population size.