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.     

Dispersion and site fidelity of breeding female grey seals (Halichoerus grypus) on North Rona, Scotland

Female grey seals ( Halichoerus grypus ) formed breeding aggregations on the island of North Rona, Scotland. Aggregations of females were associated particularly with gullies leading from the sea, leaving large areas of available space unoccupied. Changes in the degree of aggregation of females during the breeding season were similar in 1987, 1988 and 1989. Pronounced aggregations occurred in the early and late parts of each breeding season.
Of 67 breeding females marked in 1985, 62 (93%)) returned to N. Rona to breed in at least one season up to 1989, but 18 (27%) were present in all five years. Females came ashore up to 14 days before giving birth and 82% were observed first in the vicinity of their subsequent pupping site. Between 1985 and 1989, marked females which returned were faithful to their previous pupping sites, even when the previous pup had died. There was no evidence of a gradual change in the location of individual pupping sites over time. This pupping site fidelity may generate aggregations whose location, timing and composition is predictable.     

Distribution, pup production and mortality of New Zealand sea lion

Surveys were undertaken at Campbell Island / Motu Ihupuku during January and February 2008, to determine the distribution and pup production of New Zealand sea lions (NZ sea lion; Phocarctos hookeri). In addition, necropsies were performed at the main breeding site of Davis Point to determine the principal causes of early mortality for NZ sea lion pups. In total, 397 pups were tagged and 186 untagged pups were found dead, giving a minimum pup production of 583 pups and a one month of age mortality estimate of 40%. This represents a higher pup production than previous estimates from Campbell Island (although survey techniques are not comparable), and equates to 21% of the total pup production for NZ sea lions in the 2007/08 season. Early pup mortality was high (40%) at Campbell Island, with trauma, starvation, and drowning in rock pools and peat mires the major causes of death. Pups were concentrated in two colonial breeding sites: Davis Point on the north shore of Perseverance Harbour (76%) and a newly recorded breeding site (Paradise Point) on the southern shore of Perseverance Harbour (21%). Non-colonial breeding or single pups occurred around the southern parts of the island from sea level to 400 m; however, these only contributed 3% of the known pup production.     

Grey seal (Halichoerus grypus) pup production at North Rona: A study of birth and survival statistics collected in 1972

The Grey seal breeding assembly on North Rona was studied throughout autumn 1972. Weekly censuses provided data on birth rate and mortality. It is estimated that 1736 pups were born in 1972. This is about 500 less than the mean production estimate for this assembly during the period 1959–71, possibly a result of disturbance of the breeding site by the expedition. About 600 of the pups died before putting to sea. The mean date of birth and the duration of the pupping season agree closely with earlier work on North Rona, although the mean daily birth rate curve is somewhat irregular, perhaps because there was an unusually dry spell of weather at a critical time in the pupping season. An improved estimate of mortality at this assembly showed a relationship with pup density similar to that observed at the Fame Islands. Techniques for estimating pup production (and consequently total population), though not reliable for determining absolute values, are useful for indicating trends and they suggest a stable population of 8000–9000 animals at North Rona. Such estimates might be improved by the use of better methods of aerial photography.     

Timing of births and juvenile mortality in the South American fur seal in Peru

In most species, synchronous, seasonal reproduction is usually associated with higher offspring survival in animals giving birth around the peak, relative to those breeding at the extremes of the reproductive season. In contrast, in the South American fur seal ( Arctocephalus australis ) at Punta San Juan, Peru, females pupping around the peak of births had a greater probability of losing their pups and/or tended to lose them at an earlier age than females breeding early or late in the season. However, the timing of breeding in this population varies little between years and is consistently synchronous. Individual females maintained their relative breeding times in consecutive years, regardless of whether or not they lost their pup the previous year. Thus, pup mortality seems to have no effect on the timing of reproduction in this population.
High breeding densities and the consequent high pup mortality in the S. American fur seal in Peru may have resulted from intense poaching outside protected areas and are of recent origin. The reproductive synchrony in this population could have originally evolved as a response to seasonal variations in food availability and weather conditions, differences in female or pup body condition, predation pressure, sexual selection and/or harassment avoidance, but at the present high density levels has become maladaptive.     

Effects of maternal age and condition on parturition and the perinatal period of Antarctic fur seals

The effect of maternal age and condition on the date of parturition and the duration of the perinatal period of Antarctic fur seals at Bird Island, South Georgia, were investigated over three consecutive breeding seasons. Females rear young during a four-month lactation period in a highly seasonal but predictable environment. Although females may first pup at three years of age, they did not attain full adult size until six years of age; older females (≥ 6 years) tended to be heavier, longer, and in better condition than younger females (3–5 years). Older females returned to breeding beaches earlier and could occupy the most suitable pupping sites, and gave birth when densities of animals on the beaches were low (i.e. more favourable for pup survival). Females that arrived earlier were able to remain ashore longer with their pups prior to departing on their first foraging trips but this was unrelated to either maternal age or condition. Younger females returned later in the pupping season, possibly as a result of late implantation due to smaller energy reserves than older and larger females. In 1990 all females arrived late, were in poorer condition, gave birth to lighter pups, and had shorter perinatal periods. This suggests that not only was implantation late but that females returned to an area of low food availability prior to parturition.     

Influence of the physical environment and conspecific aggression on the spatial arrangement of breeding grey seals

Understanding the habitat requirements of a species for breeding is essential for its conservation, particularly if the availability of suitable habitat is a limiting factor for population growth. This is postulated to be the case for grey seals, one of the more abundant marine apex predators in northern European waters. In common with similar studies that have investigated the habitat preferences of breeding grey seals, we use abiotic (topographical, climatological) attributes but, unlike previous work, we also incorporate behavioural variables, particularly the occurrence of aggressive interactions between females and the presence of neighbouring seals. We used two Generalized Additive Models (GAM) in a sequential and iterative fashion. The first model links the occurrence of aggression at particular points in the colony to local topography derived from a Geographical Information System (GIS), presence of neighbouring seal pups and the day of the breeding season. The output of this GAM is used as one of the explanatory variables in a GAM of daily variation in the spatial distribution of births. Although proximity of a birth site to a water source and the presence of neighbouring seal pups both had significant influences on the distribution of newborn pups over time and space, at the scale of the study site it was found that simple rules could predict pup distribution more efficiently than a complex individual-based simulation model.     

Alloparenting in Steller sea lions (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>): correlations with misdirected care and other observations

Alloparental care is rarely observed in Steller sea lions (Eumetopias jubatus) where maternal care is extended to a single pup for up to 1 year or more. However, we observed 28 allonursing events and one case of adoption at a small breeding rookery in the western Gulf of Alaska between the years 2001 and 2005. Multiparous and primiparous females were observed nursing nonfilial individuals with equal frequency, but primiparous females spent significantly more time nursing nonfilial individuals. Multiparous females allowed allonursing only while sleeping and unaware while most primiparous females were aware that they were allonursing. These results are consistent with the misdirected-care hypothesis suggesting that primiparous (presumably younger) females nurse nonfilial pups due to inexperience, whereas multiparous (presumably older) females are victims of milk stealing during times of inattentiveness. Nonfilial pups were aggressively tossed most often during the pupping season and only by multiparous females, while allonursing events occurred more frequently after the pupping season. Starveling pups were not cared for by any female, but two were attended by a single bull during separate autumn seasons.     

Foraging locations of female New Zealand sea lions (

Figure of Eight Island is located in the southern end of the Auckland Islands and hosts the smallest breeding colony of New Zealand (NZ) sea lions (Phocarctos hookeri). Between 1995/96 and 2005/06, pup production in this colony decreased by 57% (from 144 to 62 pups). In contrast, there was a 30% decrease in pup production in the largest colony in the north-east of the Auckland Islands over the same period. NZ sea lions in the Auckland Islands area are subject to by-catch deaths and resource competition from subantarctic trawl fisheries. The present study investigated where four lactating females from Figure of Eight Island foraged during the austral summer of 2007/08 and compared their foraging areas with female NZ sea lions from the northern Auckland Islands breeding locations (Enderby and Dundas islands) and with fisheries activities. Females foraged south of Adams Island (the southernmost Auckland Island), predominantly at the edge of the Auckland Islands shelf, but those from Figure of Eight Island made shorter foraging trips within more concentrated areas than females from Enderby or Dundas islands. The 59 female NZ sea lions satellite-tracked to date from Figure of Eight, Enderby and Dundas islands foraged over the entire area of the Auckland Islands shelf and many (including three of the four females from Figure of Eight Island) had extensive overlap with subantarctic trawl fisheries. Further research is needed to determine whether the foraging behaviour of females from Figure of Eight Island is linked to their greater decline in pup production.     

PUP PRODUCTION AND POPULATION TRENDS OF THE AUSTRALIAN FUR SEAL (ARCTOCEPHALUS PUSILLUS DORIFERUS)

We estimated the number of live Australian fur seal pups using capture-markresights, direct ground counts, or aerial photography at all breeding sites following the pupping season of November-December 2002. Pups were recorded at 17 locations; nine previously known colony sites, one newly recognized colony and seven haul-out sites where pups are occasionally born. In order of size, the colonies were Lady Julia Percy Island (5,899 pups), Seal Rocks (4,882), The Skerries (2,486), Judgment Rocks (2,427), Kanowna Island (2,301), Moriarty Rocks (1,007), Reid Rocks (384), West Moncoeur Island (257), and Tenth Island (124). The newly recognized site was Rag Island, in the Cliffy Group, where we recorded 30 pups. We also recorded pups at the following haul-out sites: Cape Bridge-water (7 pups), Bull Rock (7), Wright Rock (5), Twin Islet (1), The Friars (1), He des Phoques (1), and Montague Island (1). In total, we estimate there were 19,819 (SE = 163) live pups at the time of the counts. We discuss trends in pup numbers and derive current population estimates for the Australian fur seal.     

Philopatry, Site Fidelity and Local Kin Associations within Grey Seal Breeding Colonies

Abstract Colonially breeding animals such as grey seals typically have a discrete number of breeding sites. The extent to which offspring are philopatric (return to breed at their natal site) and the prevalence of philopatry by sex of pup has fundamental consequences for the social and genetic structure within and between each colony.
Grey seals born and marked by tagging and cohort brands at colonies on North Rona, Outer Hebrides since 1960 and at the Isle of May, Firth of Forth since 1981 have shown philopatry. Overall re-sight rates of 1620 pups tagged at North Rona and 1667 pups at the Isle of May ranged from 0 to 17%. Although most evidence of philopatry relates to females, males at each colony displayed philopatry. In addition, females born at N. Rona pupped closer to their natal sites than would be expected by chance (p = 0.005), providing evidence of fine scale natal site fidelity. Females born at the Isle of May did not show the same degree of fine scale natal site fidelity, but there was considerable individual variation. The spatial scale of philopatry shown by seals differed according to the local area within each island. Fidelity of known adult females to pupping sites at both colonies was high. In addition, we found an instance of mother and daughter pupping together away from the daughter's natal site.
These data: (i) provide evidence of philopatry in both sexes at each site; (ii) describe occurrences of fine scale philopatry at both sites; (iii) show that mothers and offspring occupied the same areas on the colony, suggesting the possibility of highly related groups within colonies, which would provide conditions for kin-specific altruistic behaviours; (iv) suggest the possibility of individual or kin recognition, even though contact between mothers and offspring has been thought to cease after the 18 d lactation period.     

How mothers find their pups in a colony of Antarctic fur seals

In Antarctic fur seals, Arctocephalus gazella, mothers must identify their own young among hundreds or even thousands of pups, if they are to invest in their own offspring and avoid misdirecting their parental care. When returning to their breeding colony from a foraging trip of several days at sea, mothers have to find and identify their young before suckling can occur. There appears to be little confusion about which pup belongs to a mother, and adoption is absent or rare. Using behavioral observations, we investigated the means by which female Antarctic fur seals identified their pups in a breeding colony of about 750 mother-pup pairs on Kerguelen Island. We evaluated the importance of vision, scent communication, vocalizations, and rendezvous locations as possible explanations of how mothers find their pups. Every pup that a mother examined, whether her own or not, exchanged naso-nasal inspection with her, suggesting a strong role for olfactory communication in individual recognition. Both mothers and pups called to each other, and mothers that searched for pups over a longer period gave more calls and encountered more pups. Thus, vocalizations may have been used to attract pups that might be offspring. Nursing usually occurred in the same place from the end of one maternal visit to the colony and the arrival at the beginning of the next visit, suggesting that nursing locations may serve as a meeting place, or rendezvous, for mothers and pups. These results suggest that finding pups is a two-stage process for females, in which pups for sampling are attracted by calls or examined at the previous nursing location, and then individual identification is made by olfactory cues.     

On‐land habitat preferences of female New Zealand sea lions at Sandy Bay,Auckland Islands

Terrestrial habitat is important for breeding in most pinnipeds. On land, most species remain near the shore, but New Zealand (NZ) sea lions, Phocarctos hookeri, often rest inland up to 1.5 km from the sea. Only three breeding areas of NZ sea lions exist today after the species was extirpated from its historical range (NZ mainland). The study was conducted at the Sandy Bay breeding colony, Auckland Islands, between December 2002 and March 2003. We used daily Global Positioning System locations of breeding females with pups and mapping in a Geographic Information System to determine terrestrial habitat use and preferences. Slopes less than 20° were preferred throughout the study. Females chose nursing sites with a seasonal change, preferentially based on the distance from the sea and habitat type. Comparisons with the other breeding colonies of NZ sea lions are presented and data are discussed in the context of the recolonization of the NZ mainland. Overall, the most suitable terrestrial habitat configuration for a breeding aggregation of NZ sea lions appears to be a sandy beach, with a wide area above high tide and moderate intertidal zone (for breeding), backed with vegetated sand dunes and forest on primarily flat terrain (for later dispersion).     

Reproductive performance links to fine-scale spatial patterns of female grey seal relatedness

Fine-scale spatial patterns of female relatedness throughout the established grey seal breeding colony of North Rona, Scotland, were investigated by accurate mapping and spatially explicit analyses of a large sample (n = 262) of mothers using variation at nine microsatellite DNA loci. Local spatial autocorrelation analyses identified locations where seals were more highly related to the colony than average. These locations were also areas where the more successful females bred, were occupied first during each breeding season, were centrally placed locations of preferred habitat types and were likely to be the locations which were the first to be colonized historically. Mothers occupying such sites achieved higher than average pup growth rates, suggesting a founder fitness benefit.     

Distribution, abundance and breeding cycle of the Australian sea lion Neophoca cinerea (Mammalia: Pinnipedia)

Surveys of the Australian sea lion Neophoca cinerea were conducted throughout its range in Western and South Australia between December 1987 and February 1992. Almost every island was visited between Houtman Abrolhos and The Pages ( n = 255), many of them more than once.
Sea lions breed on at least 50 islands, 27 in Western Australia and 23 in South Australia. Of the 50 breeding sites, 31 have not been reported previously. A further 19 islands may also support breeding colonies. A total of 1,941 pups was counted and pup production was estimated at 2,432. Only five colonies produced more than 100 pups each and they accounted for almost half of the pup production. Most of these colonies are near Kangaroo Island, South Australia. A breeding cycle of 17–18 months has been reported for N. cinerea at Kangaroo Island and on the west coast of Western Australia; this was also noted at another 11 islands where repeated visits coincided with breeding. No evidence was found for breeding seasons shorter or longer than 17–18 months. The breeding season was not synchronized between islands, as it is in other pinnipeds. A predictive model is developed to estimate the population size from pup production figures. It indicates that pup numbers should be multiplied by between 3.81 and 4.81 to estimate the total population size just before the pupping season begins. This leads to estimates of 9,300–11,700 for the total population, considerably greater than earlier estimates.
Causes of the unique reproductive cycle of N. cinerea are unknown, but we hypothesize that it results from living in a temperate climate in some of the most biologically depauperate waters of the world. It is also clear that day length and water temperature cannot act as exogenous cues for implantation of the blastocyst; the physiological events of gestation must, rather, be cued endogenously.     

SUMMER HOME RANGE AND HABITAT SELECTION OF HARBOR SEAL (PHOCA VITULINA) PUPS

We studied the summer home range and habitat selection of harbor seal pups tracked using VHF radio telemetry along the coast of central Norway in 1997 and 1998. Median fixed kernel home range of six pups tracked in 1998 was 10.4 km² and the median core area was 1.2 km². One particular deep basin (>100 m deep) was highly selected, and extensively used by four of the pups, probably for feeding. An area adjacent to the deep basin also was selected, probably for haul outs. One pup spent most of the time in inshore, kelp forest areas. Shallow areas without kelp forest and exposed land were used least frequently, possibly because they did not provide suitable areas for feeding and to haul out. The seals hauled out more during calm wind and low tide than during rough wind and high tide. Seals seemed to haul out on intertidal rocks associated with islands, and alternated between several haul-out sites. We found no pattern in haul-out activity associated with age, time of day, air/sea temperature, cloud cover, or rainfall. In 1997, seven pups were tracked manually. These data were associated with methodological limitations, and they were used only to illustrate that these seals stayed mainly within the shallow, inshore, kelp forest area, indicating a possible difference in habitat use between years.     

PUP SURVIVAL IN THE MEDITERRANEAN MONK SEAL (MONACHUS MONACHUS) COLONY AT CABO BLANCO PENINSULA (WESTERN SAHARA-MAURITANIA)

We surveyed pup survival in Mediterranean monk seals ( Monachus monachus ), at Cabo Blanco Peninsula (Western Sahara-Mauritania) colony from May 1993 to December 1997. This species breeds and hauls out on beaches inside two main caves. During the study period we detected a total of 93 pups that died or disappeared. The survival rate of 84 pups through the age of first moult (approximately two months) was 0.47. This value is similar to those reported for other pinnipeds breeding in caves but lower than for those breeding on open beaches. Mortality varied seasonally and appeared to increase as a result of storms, large ocean swells, and high tides. Mother-pup pair separation (and resulting pup starvation) and physical injury caused by impact against the rock walls of the cave and cliffs were established as the causes of most deaths. Beach surface area inside the caves also appeared to be a mediating factor in the effects of sea conditions. High pup mortality may be a limiting growth factor in this population, although cave dwelling protects the population from predators and human disturbance.     

REPRODUCTIVE CHARACTERISTICS OF FEMALE SEA OTTERS

Abstract: Several important aspects of reproduction in the female sea otter, such as gestation, pupping frequency, period of pup dependency, and annual pupping rate, were unclear when this study was begun. We present data from 75 tagged adult females that indicate gestation is variable, but on average is about 6 months, the length of pup dependency is 6 mo, thus the pupping interval is usually 12–13 mo. Most females breed for the first time in their fifth year of life. About 85–90% of adult females pup in a given year.     

Survival estimates for the Australian sea lion: Negative correlation of sea surface temperature with cohort survival to weaning

The Australian sea lion (Neophoca cinerea) population at Seal Bay Conservation Park, South Australia, is estimated to be declining at a rate of 1.14% per breeding season. To better understand the potential causes of this decline, survival rates were examined to 14 yr of age for eight cohorts marked as pups (aged 0.17 yr) between 1991 and 2002. Apparent yearly survival rates (Φ) varied by cohort for pups from marking to weaning at 1.5 yr (Φ= 0.30–0.67). Postweaning juvenile survival (1.5–3 yr) was 0.89 and survival from 3 to 14 yr was constant (Φ female:male = 0.96:0.89). Φ of pup cohorts was negatively correlated to local sea surface temperature where the sea lions forage (SST) and was especially low for cohort 7 in 2000 (0.30). It is possible that periods of unusually warm oceanographic conditions may be limiting primary production and inhibiting maternal provisioning to pups. Pup survival to weaning is relatively low compared to other otariid species, is likely to limit recruitment, and may be contributing to the decline in pup abundance observed in the colony.     

Variation in Female Grey Seal (Halichoerus grypus) Reproductive Performance Correlates to Proactive-Reactive Behavioural Types

Consistent individual differences (CIDs) in behaviour, indicative of behavioural types or personalities, have been shown in taxa ranging from Cnidaria to Mammalia. However, despite numerous theoretical explanations there remains limited empirical evidence for selective mechanisms that maintain such variation within natural populations. We examined behavioural types and fitness proxies in wild female grey seals at the North Rona breeding colony. Experiments in 2009 and 2010 employed a remotely-controlled vehicle to deliver a novel auditory stimulus to females to elicit changes in pup-checking behaviour. Mothers tested twice during lactation exhibited highly repeatable individual pup-checking rates within and across breeding seasons. Observations of undisturbed mothers (i.e. experiencing no disturbance from conspecifics or experimental test) also revealed CIDs in pup-checking behaviour. However, there was no correlation between an individuals’ pup-checking rate during undisturbed observations with the rate in response to the auditory test, indicating plasticity across situations. The extent to which individuals changed rates of pup-checking from undisturbed to disturbed conditions revealed a continuum of behavioural types from proactive females, who maintained a similar rate throughout, to reactive females, who increased pup-checking markedly in response to the test. Variation in maternal expenditure (daily mass loss rate) was greater among more reactive mothers than proactive mothers. Consequently pups of more reactive mothers had more varied growth rates centred around the long-term population mean. These patterns could not be accounted for by other measured covariates as behavioural type was unrelated to a mother’s prior experience, degree of inter-annual site fidelity, physical characteristics of their pupping habitat, pup sex or pup activity. These findings are consistent with the hypothesis that variation in behavioural types is maintained by spatial and temporal environmental variation combined with limits to phenotype-environment matching.