Space and resource use by brown hyenas Hyaena brunnea in the Namib Desert

Two clans of brown hyenas were studied on the arid chilly Namib Desert coast where very large seal colonies are present. Carcasses from seals were abundant during summer and autumn following the seal pupping season, and scarce in winter and spring. Hyenas fed predominantly on carcasses. Members of one clan of three hyenas obtained most of their food along 3 km of coastline adjacent to the seal colonies. The other clan of nine individuals obtained food from an area along 6 km of coastline north of the seal colonies. Although carcasses were usually scavenged, seal pups were seen to be killed on occasions. Time spent foraging was not affected by carcass availability. Territories of the two clans overlapped and individual home-range size varied from 31.9 to 220 km². Food availability apparently had no influence on foraging activities but the dispersal of food affected territory and group sizes. Over abundance of food results in the maintenance of territories extending beyond the distribution of food. The shape and area of these territories may result from the cultural inheritance of space.     

Cephalopods in the diet of the South African fur seal (Arctocephalus pusillus pusillus)

The stomach contents of 1522 adult seals ( Arctocephalus pusillus pusillus ) and 673 pups, which were collected around the West and South Coasts of Southern Africa, were examined and cephalopod prey identified. About 20 cephalopod species (some identifications were uncertain) were found in the stomachs, of which only six were significant: Loligo vulgaris reynaudii. Sepia spp., Octopus sp., Todaropsis eblanae, Todarodes angolensis and Ocythoe tuberculata. This list suggests that A. pusillus feeds exclusively on the continental shelf, frequently on or near the bottom. Among adults, cephalopod prey varied in importance around the coast. It was most important on the South Coast of South Africa where it comprised 35.0% of the weight of all prey, of which L. v. veynaudii constituted about 88%. On the West Coast cephalopods comprised about 26.6% by weight of all prey, and Octopus sp. predominated, whereas in Namibian waters cephalopods were least important, constituting only 3.4% by weight of all prey, with Todarodes angolensis and Octopus sp. being most significant. Among pups (aged 8–10 months) in the Lüderitz (Namibia) area, cephalopods were about 16.9% of prey by weight, of which Ocythoe tuberculata , small Sepia spp. and Octopoda were most important.     

Fasting metabolism in Antarctic fur seal (Arctocephalus gazella) pups.

The metabolism of 52-73-day old Antarctic fur seal pups from Bird Island, South Georgia, was investigated during fasting periods of normal duration while their mothers were at sea foraging. Body mass decreased exponentially with pups losing 3.5-3.8% of body mass per day. Resting metabolic rate also decreased exponentially from 172-197 ml (O2) x min(-1) at the beginning of the fast and scaled to M(b)(0.74) at 2.3 times the level predicted for adult terrestrial mammals of similar size. While there was no significant sex difference in RMR, female pups had significantly higher (F(1,18)=6.614, P<0.019) mass-specific RMR than male pups throughout the fasting period. Fasting FMR was also significantly (t(15)=2.37, P<0.035) greater in females (823 kJ x kg(-1) x d(-1)) than males (686 kJ x kg(-1) x d(-1)). Average protein turnover during the study period was 19.3 g x d(-1) and contributed to 5.4% of total energy expenditure, indicating the adoption of a protein-sparing strategy with a reliance on primarily lipid catabolism for metabolic energy. This is supported by observed decreases in plasma BUN, U/C, glucose and triglyceride concentrations, and an increase in beta-HBA concentration, indicating that Antarctic fur seals pups adopt this strategy within 2-3 days of fasting. Mean RQ also decreased from 0.77 to 0.72 within 3 days of fasting, further supporting a rapid commencement of protein-sparing. However, RQ gradually increased thereafter to 0.77, suggesting a resumption of protein catabolism which was not substantiated by changes in plasma metabolites. Female pups had higher TBL (%) than males for any given mass, which is consistent with previous findings in this and other fur seal species, and suggests sex differences in metabolic fuel use. The observed changes in plasma metabolites and protein turnover, however, do not support this.     

Distribution of post-weaning Antarctic fur seal Arctocephalus gazella pups at South Georgia

Potentially some of the biggest gaps in our knowledge about the ecology of Antarctic fur seals (Arctocephalus gazella) relate to juvenile animals. We investigated the at-sea distribution of five male and five female fur seal pups post-weaning. The study was carried out at Bird Island, South Georgia during two successive winters using satellite-linked platform transmitter terminals (PTTs). Our results are analysed in relation to pup sex and the physical environment and productivity of those 2 years, as well as in the context of our present knowledge of where post-breeding females and males forage. The available physical and biological data during both of the winters of this study suggest that both years were not unusual. We report marked differences between the sexes with male pups foraging significantly further away from land and their birth site than do females. The pups foraged in areas to the East of Bird Island seldom reported as foraging areas for the adult population. Also as winter progressed they showed a more oceanic distribution leaving the continental shelf, possibly to exploit a different prey source that was more readily available in the upper water column.     

Growth rates of Antarctic fur seal Arctocephalus gazella pups at South Georgia

Large numbers of pups of the Antarctic fur seal were weighed at regular intervals through their rearing period in six seasons between 1972 and 1981 at South Georgia. Growth rates and birth and weaning weights were estimated from linear regressions of mean weight on mean age. Female pups had significantly lower growth rates and weaning weights than males, but higher variances for all growth parameters. This may reflect the importance to males of polygynous species of attaining high body weight. Except for an inverse relation between growth rate and birth weight there were no systematic relationships between growth parameters. Seasonal differences in growth rates were substantial and there was no indication that the continuing rapid population increase was affecting pup growth rates. The use of these data in detecting seasonal variations in local food availability is discussed. Preliminary estimates of the energy costs of pup rearing indicate that pup existence metabolism consumes c. 80% and mass gain to weaning only c. 15% of the energy provided by the female.     

Temperature regulation and microhabitat choice by free-ranging Galapagos fur seal pups (Arctocephalus galapagoensis)

Summary Behavioural activity, and core and surface temperatures of 4 unrestrained Galapagos fur seals were recorded in the natural habitat during their first weeks of life. Climatic variables were registered simultaneously. Pup behaviours were divided into bouts of resting (55% of total time), sucking (23%) and other activities (22%). Pups maintained a constant body temperature from their first day. Core temperature (T _e ) was 37.7° C±0.3° C (x ± SD) over 39 pup-days and 8 pup-nights. Skin temperature was correlated with T _c , but flipper temperature was not. No daily T _c rhythm was detected. Microclimate data were used to calculate operative temperature T _e . Environmental temperatures can be very high, with T _e above T _c 6–9 h a day for animals exposed to the sun, but below it in the shade. T _c is about 22° C at night. Pups avoid overheating mainly by withdrawing into the shade and reducing activity to a minimum during the hot hours of the day. Sun-exposed pups could be active at any time during the day if they had access to water, which was usually around high tide.     

The composition of Australian fur seal (Arctocephalus pusillus doriferus) milk throughout lactation.

The proximate chemical composition of milk was determined throughout lactation in the Australian fur seal (Arctocephalus pusillus doriferus Wood Jones), a temperate species with an 11-mo lactation period typical of most temperate otariids. Average lipid and protein contents were 42% and 10%, respectively, which is similar to that recorded for the polar otariid species but higher than reported in other temperate, long-lactation species. Milk composition, however, varied substantially throughout lactation, with lipid content rising from 30% soon after parturition to 50% at 230 d postpartum before decreasing to 45% toward weaning. The trend in water content was opposite to that of lipid, while protein content increased only slightly during lactation from 10% to 12%. The peak in lipid content occurred at approximately 70% of the way through lactation, similar to that found in the Australian sea lion and several phocid seal species. Significant positive relationships were found between milk lipid content and both the duration of the preceding foraging trip and body mass. A highly significant and close correlation was found between milk water and lipid contents such that gross energy content may be accurately predicted from water content alone. Similar predictive equations have been found for several other otariid and phocid species, but these appear to differ between phylogenetic groups.     

Hormonal responses to extreme fasting in subantarctic fur seal (Arctocephalus tropicalis) pups

Surviving prolonged fasting implies closely regulated alterations in fuel provisioning to meet metabolic requirements, while preserving homeostasis. Little is known, however, of the endocrine regulations governing such metabolic adaptations in naturally fasting free-ranging animals. The hormonal responses to natural prolonged fasting and how they correlate to the metabolic adaptations observed, were investigated in subantarctic fur seal (Arctocephalus tropicalis) pups, which, because of the intermittent pattern of maternal attendance, repeatedly endure exceptionally long fasting episodes throughout their development (1-3 mo). Phase I fasting was characterized by a dramatic decrease in plasma insulin, glucagon, leptin, and total l-thyroxine (T(4)) associated with reductions in mass-specific resting metabolic rate (RMR), plasma triglycerides, glycerol, and urea-to-creatine ratio, while nonesterified fatty acids (NEFA) and β-OHB increased. In contrast, the metabolic steady-state of phase II fasting reached within 6 days was associated with minimal concentrations of insulin, glucagon, and leptin; unchanged cortisol and triiodothyronine (T(3)); and moderately increased T(4). The early fall in insulin and leptin may mediate the shift to the strategy of energy conservation, protein sparing, and primary reliance on body lipids observed in response to the cessation of feeding. In contrast to the typical mammalian starvation response, nonelevated cortisol and minimal glucagon levels may contribute to body protein preservation and downregulation of catabolic pathways, in general. Furthermore, thyroid hormones may be involved in a process of energy conservation, independent of pups' nutritional state. These original hormonal settings might reflect an adaptation to the otariid repeated fasting pattern and emphasize the crucial importance of a tight physiological control over metabolism to survive extreme energetic constraints.     

Chemical characterisation of the oligosaccharides in hooded seal (Cystophora cristata) and Australian fur seal (Arctocephalus pusillus doriferus) milk

Carbohydrates were extracted from hooded seal milk, Crystophora cristata (family Phocidae). Free oligosaccharides were separated by gel filtration and then purified by ion exchange chromatography, gel filtration and preparative thin layer or paper chromatography and their structures determined by 1H-NMR. The hooded seal milk was found to contain inositol and at least nine oligosaccharides, most of which had lacto-N-neotetraose or lacto-N-neohexaose as core units, similar to those in milk of other species of Carnivora such as bears (Ursidae). Their structures were as follows: Gal(beta1-4)Glc (lactose); Fuc(alpha1-2)Gal(beta1-4)Glc (2'-fucosyllactose); Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (lacto-N-neotetraose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (lacto-N-fucopentaose IV); Gal(beta1-4)GlcNAc(beta1-3)[Gal(beta1-4)GlcNAc(beta1-6)]Gal(1-4)Glc (lacto-N-neohexaose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)[Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (monofucosyl lacto-N-neohexaose a); Gal(beta1-4)GlcNAc(beta1-3)[Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (monofucosyl lacto-N-neohexaose b); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)[Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc (difucosyl lacto-N-neohexaose); Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (para lacto-N-neohexaose); Fuc(alpha1-2)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (monofucosyl para lacto-N-neohexaose). Milk of the Australian fur seal, Arctophalus pusillus doriferus (family Otariidae) contained inositol but no lactose or free oligosaccharides. These results, therefore, support the hypothesis that the milk of otariids, unlike that of phocids, contains no free reducing saccharides.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in response to spatial variation in white shark (Carcharodon carcharias) predation risk

Foraging and predation risk are often separated at rookeries of marine central place foragers, thus offering an opportunity to gain insight into how predator‐avoidance shapes the behavior of prey. Here we compare the behavior of Cape fur seals (Arctocephalus pusillus pusillus) at two island rookeries with and without white shark (Carcharodon carcharias) predations, and assess seal behavior in relation to marked spatiotemporal variation in risk at the high‐risk site (Seal Island, South Africa). Our results show that seal behavior at the two sites is comparatively similar in summer, when predation risk is low at both sites, but not in winter. Compared to seals at the “low‐risk” site, seals at Seal Island avoided deep‐water habitat around the island at high risk times and restricted their use of this habitat in favor of safe, shallow waters when engaging in social and thermoregulatory behaviors. Seals increased their frequency of jostling, porpoising, and diving when moving through the danger zone and seals in groups were safer than single individuals. Overall, our results suggest that seal behavior around the high‐risk site is strongly affected by predation risk, and show this rookery to be an excellent predator‐prey system at which to evaluate long‐standing ecological hypotheses.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in relation to temporal variation in predation risk by white sharks (Carcharodon carcharias) around a seal rookery in False Bay,South Africa

The marked differences in predation risk posed by white sharks (Carcarodon carcarias) at island rookeries of Cape fur seals (Arctocephalus pusillus pusillus) offer a quasi‐experimental design within a natural system for exploring how prey adjust their behavior in response to temporal variation in predation risk. Here we compare movement of juvenile and adult Cape fur seals at a high risk (Seal Island) and low risk (Egg Island) rookery. We further compare juveniles and adults at Seal Island in low and high risk seasons and at low and high risk times of day within those seasons. Adult fur seals at Seal Island avoided traversing the zone of high white shark predation risk during the high risk period (0700–0959) in the season of high risk (winter), but not during the low risk season (summer). By contrast, adult fur seals at Egg Island showed no temporal discretion in either season. Unlike juvenile fur seals at Egg Island, juveniles at Seal Island adjusted their temporal movement patterns to more closely mimic adult seal movement patterns. This suggests that exposure to predators is the primary driver of temporal adjustments to movement by prey species commuting from a central place.     

Species-specific cell-matrix interactions are essential for differentiation of alveoli like structures and milk gene expression in primary mammary cells of the Cape fur seal (Arctocephalus pusillus pusillus).

Few models are in place for analysis of extreme lactation patterns such as that of the fur seals which are capable of extended down regulation of milk production in the absence of involution. During a 10-12 month lactation period, female fur seals suckle pups on shore for 2-3 days, and then undertake long foraging trips at sea for up to 28 days, resulting in the longest intersuckling bouts recorded. During this time the mammary gland down regulates milk production. We have induced Cape fur seal (Arctocephalus pusillus pusillus) mammary cells in vitro to form mammospheres up to 900 microm in diameter, larger than any of their mammalian counterparts. Mammosphere lumens were shown to form via apoptosis and cells comprising the cellular boundary stained vimentin positive. The Cape fur seal GAPDH gene was cloned and used in RT-PCR as a normalization tool to examine comparative expression of milk protein genes (alphaS2-casein, beta-lactoglobulin and lysozyme C) which were prolactin responsive. Cape fur seal mammary cells were found to be unique; they did not require Matrigel for rapid mammosphere formation and instead deposited their own matrix within 2 days of culture. When grown on Matrigel, cells exhibited branching/stellate morphogenesis highlighting the species-specific nature of cell-matrix interactions during morphological differentiation. Matrix produced in vitro by cells did not support formation of human breast cancer cell line, PMC42 mammospheres. This novel model system will help define the molecular pathways controlling the regulation of milk protein expression and species specific requirements of the extracellular matrix in the cape fur seal.     

Ontogeny of aquatic behaviours in Antarctic fur seal (Arctocephalus gazella) pups in relation to growth performances at Kerguelen Islands

In diving marine predators, such as pinnipeds, the development of diving and foraging skills prior to weaning might be critical to post-weaning survival. Here, we examined the effect of pup mass growth on the amount of time devoted to aquatic activities and the dive performance of Antarctic fur seal, Arctocephalus gazella, pups on Kerguelen Island. Maternal attendance and mass-specific growth rate were assessed for 85 pups. Two types of monitoring were applied: visual observations of behaviours for 60 pups and the deployment of time-depth recorders (TDRs) on 19 female pups. At approximately 2 months of age, pups demonstrated minimal diving behaviour, but displayed considerable aquatic activity. While mothers were foraging at sea, pups fasted on land (6.0 ± 1.3 d). As the mass-specific growth rate was different between sexes, only data on female pups were analysed (n = 31). Mass-specific growth rate was related to maternal attendance patterns and impacted the amount of time allocated by pups to aquatic activities. The time spent in the water by pups was quadratically related to fasting progress. This study shows the importance of growth and fasting progress on the quantity of time pups devoted to aquatic activities. Our results suggest that greater post-weaning survival of heavier pups may be due not only to their greater body reserves, as reported in several studies, but also possibly to from their greater aquatic skills and physiological adaptations developed during the suckling period.