Body mass loss of moulting female southern elephant seals,Mirounga leonina,at Macquarie Island

Thirteen female southern elephant seals moulting at Macquarie Island lost an average of 4.46±0.80 kg/day (10.01±1.20g/kg/day). There was no significant difference between this rate of body mass loss and that reported for moulting female southern elephant seals from South Georgia. Moulting female southern elephant seals however exhibited larger mass specific mass loss than either female northern elephant seals or male southern elephant seals, indicating a higher metabolic cost of moult in these animals.     

Postweaning duration and body composition changes in Southern elephant seal (Mirounga leonina) pups at King George Island.

Weaning mass in southern elephant seals is highly variable, the heaviest pups being three times as heavy as the lightest ones. After weaning, pups undergo an extensive postweaning period in which they draw on their reserves. To quantify the energy expenditure during the postweaning period, changes in mass, body composition, and postweaning duration were measured in southern elephant seals at King George Island, South Shetland Islands, Antarctica. Overall, mean pup weaning mass was 154 +/- 26 kg (n=117) and did not differ between sexes. Mean minimum postweaning duration was 42.5 +/- 7.5 d. Heavier animals at weaning had lower mass-specific mass loss rates than lighter ones, and a faster depletion of body reserves was associated with a shorter postweaning period. The proportion of body mass represented by fat at weaning was 37% +/- 4% (n=47) and did not differ between sexes. Of these pups, 36 were recaptured after a mean period of 36 d after weaning. On average, total mass loss measured in these animals (39 kg) was composed of 39% water, 47% fat, and 12% protein. The composition of mass loss was not significantly different between sexes and was not related to weaning mass or total body energy reserves. However, fatter animals at weaning lost more fat per kilogram lost than thinner ones. Late in the fast, males and females appeared to be in a similar body condition. Nevertheless, the overall proportion of body mass represented by fat at this time was lower than that presented by the same animals at weaning. We estimated that during the postweaning period pups lost, on average, 30% of their mass at weaning. This comprised approximately 35% of the energy and 32% of the fat in the pup's body.     

GROWTH AND DEVELOPMENT IN FREE-RANGING HARBOR SEAL (PHOCA VITULINA) PUPS FROM SOUTHERN BRITISH COLUMBIA, CANADA

Harbor seals ( Phoca vitulina ) are small pinnipeds that are widely distributed throughout the temperate coastal regions of the Atlantic and Pacific oceans. We determined birth mass, neonatal growth rates, weaning age, and weaning mass of NE Pacific harbor seals ( P. v. richardsi ) during a capture-recapture study that spanned the nursing period (Sidney Island, British Columbia, Canada). Of 46 harbor seal pups initially captured, 28 were classified as newborns ( i. e. , < 24 h old). Mean body mass of newborns was 11.2 ± SE 0.31 kg. Pups were individually tagged and recaptured throughout the nursing period. Average daily mass gain during the nursing period was 394 ± 26 g. Mean birth mass of males did not differ significantly from females, although pups found with fetal pelage (lanugo) (21.4% of all newborns) were smaller at birth (9.8 ± 0.44 kg) than non-lanugo pups (11.6 ± 0.33 kg). Mean weaning mass was estimated at 23.6 ± 1.2 kg at a mean weaning age of 32 d ± 1.5 d. While birth and weaning masses differed little from the published data for offshore Sable Island harbor seals ( P. v. concolor ), British Columbia harbor seals are characterized by half the daily mass gain and a longer nursing period.     

Age and sex differences in the timing of moult in the common seal, Phoca vitulina

This study followed the progress of the annual moult within a population of common seals in Orkney, Scotland. Moulting seals were seen over a three-month period, from 7 June until 16 September. Yearlings were first to start moulting. Amongst older seals, females completed their moult an average of seven days earlier than immature males and 19 days earlier than mature males. Differences in the timing of moult appeared to be related to the age or reproductive status of the animals, and may be the result of differential changes in levels of the sex hormones.     

Mass transfer efficiency between harp seal (Phoca groenlandica) mothers and their pups during lactation

We investigated the efficiency of mass transfer in lactating harp seals through serial measurements on individual mother-pup pairs during the whelping seasons of 1988 and 1989. We also compared the influence of longitudinal versus cross-sectional sampling on estimates of the efficiency of mass transfer. Among longitudinally sampled pairs, pups grew at an average rate of 2·3 ± 0·5 (mean ± S.D.) kg/d (N = 20). The concomitant mass loss by females averaged 3·1 ± 0·8 kg/d (N = 19). The mean efficiency of mass transfer was 77·0 ± 13·6% (N= 19 pairs).
Estimates of pup growth and female mass loss from regressions of cross-sectional data were 2·0 kg/d and 3·1 kg/d, respectively. These values produce an estimate of 65% for the efficiency of mass transfer.
Consistent with the high efficiency of mass transfer, harp seal females contribute less of their total body mass to nursing ( c. 28%) than most other phocids examined. The resulting energy savings may be important for females of an ice-breeding species, which migrate a long distance shortly after weaning their pups     

Intestinal length of three California pinniped species

Forty-eight intestinal tracts, extracted from both sexes of California sea lions, Harbour seals and Northern elephant seals, were measured. The majority of intestinal tracts were removed from stranded animals that died from various causes. The sea lions and elephant seals, approximately equal in size, were larger than the Harbour seals. All species possess a small intestine which is significantly longer than even the entire gastrointestinal tract of herbivores of comparable size. Elephant seal small intestines, averaging approximately 25 times the seal's body length, were considerably longer than the small intestines of either sea lions (averaged more than 18 times the body length) or Harbour seals (averaged nearly 16 times the body length). However, the large intestines of elephant seals were shorter than either of the other two species. Among the sea lions and Harbour seals the large intestines were approximately equal in length. Sea lions and Harbour seals also showed a close correlation between standard length and total intestinal length. Among elephant seals these two parameters showed greater variability. The functional significance of the extremely long small intestine remains unclear. Certainly, the large body mass and high energy requirements of these animals has contributed to the development of a long intestinal tract. It also appears likely that diet and the high motility rate of digesta influenced the intestinal development. Comparatively, the significantly shorter large intestine of elephant seals probably relates to this species' remarkable capabilities in water conservation and metabolic water retention.     

Juvenile southern elephant seals exhibit seasonal differences in energetic requirements and use of lipids and protein stores

Growing juvenile animals undergo many morphological, physiological, and behavioural changes that influence their energetic requirements, patterns of energy use, and ultimately, their survival and reproductive success. We examined changes in mass loss and body composition of juvenile southern elephant seals (1- and 2-yr-olds) during their two annual haul-outs. At the start and end of the midyear and molt haul-outs, we caught, weighed, and measured 41 and 14 seals, respectively. We measured blubber depth using ultrasound to estimate body composition (lean and adipose tissue mass). Using energy densities of the adipose and lean tissue, we calculated total, lean, and adipose mass changes and energy expenditure. While molting, juvenile seals used more energy than during the midyear, which is related to the increased use of lean tissue for hair and skin regeneration. The amount of energy used increases with mass as individuals mature. We found sexual differences in energy use where females retained greater fat reserves than males by utilizing more lean tissue. These differences are most likely related to haul-out function and behavior, growth, and earlier development of females toward sexual maturity.     

SEASONAL CHANGES IN BLUBBER DISTRIBUTION IN ATLANTIC HARBOR SEALS: INDICATIONS OF THERMODYNAMIC CONSIDERATIONS

Among its functions, the hypodermal blubber layer of pinnipeds serves as both an energy reserve and insulation. This study examined seasonal changes in blubber distribution and body morphology in a group of captive changesharbor seals to test whether these changes were designed to maximize insulative effectiveness. Seasonal changes were found in girth, blubber volume, mean blubber depth, and the ratio of blubber depth to body radius (d/r ratio). These changes were more evident in older seals. The d/r ratio demonstrated a smaller relative annual change than mean blubber depth. The d/r ratio also exhibited less variation along the length of the seal than blubber depth at any given time. Similar to reports for ringed seals, and contrary to those for southern elephant seals, the harbor seals preferentially lost blubber from overinsulated areas of the body. These results suggest that core tissue and blubber mass are lost in a manner that maximizes insulative effectiveness.     

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.     

Equal investment in male and female offspring in southern elephant seals

Sex ratio theory predictions concerning differential parental investment in offspring by sex were tested on southern elephant seals, Mirounga leonina , breeding at Península Valdés, Argentina. Females invested equally in sons and daughters, as reflected by the similar mass at birth (mean ± 1 S.D.) of 14 males (44.1 ± 6.5 kg) and 14 females (43.4 ± 3.8 kg), and similar mass at weaning of 52 males (131.5 ± 22.4 kg) and 38 females (131.4 ± 18.3 kg). There were also no sex differences in the rate of mass gain during nursing (males = 4.0 ± 0.9 kg/day; females = 3.9 ± 0.8 kg/day), rate of mass loss during the first month of post-weaning fast (males = 0.85 ± 0.19 kg/day; females = 0.92 ± 0.15 kg/day), mean age at weaning (males = 22.3 ± 1.6 days; females = 22.7 ± 1.7 days), and female nursing behaviour. Mother's size accounted for most of the variation in mass of pups at weaning. Mothers ranked as small, medium and large, weaned pups with a mean mass of 102, 130 and 145 kg, respectively. The sex ratio of weanlings did not differ from unity. These data are consistent with Fisher's (1930) sex ratio theory.     

Phenotypic flexibility of a southern African duck Alopochen aegyptiaca during moult: do northern hemisphere paradigms apply?

Phenotypic flexibility during moult has never been explored in austral nomadic ducks. We investigated whether the body condition, organ (pectoral muscle, gizzard, liver and heart) mass and flight‐feather growth Egyptian geese Alopochen aegyptiaca in southern Africa show phenotypic flexibility over their 53‐day period of flightless moult. Changes in body mass and condition were examined in Egyptian geese caught at Barberspan and Strandfontein in South Africa. Mean daily change in primary feather length was calculated for moulting geese and birds were dissected for pectoral muscle and internal organ assessment. Mean body mass and condition varied significantly during moult. Body mass and condition started to decrease soon after flight feathers were dropped and continued to do so until the new feathers were at least two‐thirds grown, after which birds started to regain body mass and condition. Non‐moulting geese had large pectoral muscles, accounting for at least 26% of total body mass. Once moult started, pectoral muscle mass decreased and continued to do so until the flight feathers were at least one‐third grown, after which pectoral muscle mass started to increase. The regeneration of pectoral muscles during moult started before birds started to gain overall body mass. Gizzard mass started to increase soon after the onset of moult, reaching a maximum when the flight feathers were two‐thirds grown, after which gizzard mass again decreased. Liver mass increased significantly as moult progressed, but heart mass remained constant throughout moult. Flight feather growth was initially rapid, but slowed towards the completion of moult. Our results show that Egyptian geese exhibit a significant level of phenotypic flexibility when they moult. We interpret the phenotypic changes that we observed as an adaptive strategy to minimize the duration of the flightless period. Moulting Egyptian geese in South Africa undergo more substantial phenotypic changes than those reported for ducks in the northern hemisphere.     

Changes in body mass and feeding behaviour in male harbour seals, Phoca vitulina, in relation to female reproductive status

Loss of body mass has been used as an index of the cost of reproductive effort in mammals. We studied changes in body mass of male harbour seals, Phoca vitulina , during the breeding season on Sable Island, Nova Scotia, Canada. Individually marked subadult ( n = 21) and adult ( n = 22) males were captured at approximately six-day intervals throughout the breeding season. Adult males weighed an average of 108 kg ± 5·6 (S.E.) at initial capture, whereas subadults weighed an average of 76 ± 6·6 kg. The rate of mass loss by adult males did not differ from zero during the pre-mating period (i.e. the period without receptive females), but differed significantly from zero (-0·91 ± 0·007 kg/day) during the mating period (i.e. the period with receptive females). By contrast, the rate of mass change of subadults did not differ from zero in either period, indicating that food availability was unlikely to be responsible for the observed changes in adults. Adult males lost up to 24% of body mass during the breeding season. Examination of sera for the presence of chylomicrons (i.e. evidence of recent feeding) also suggested that adults stopped feeding during the mating period, while subadults did not. These results suggest that reproduction represents a significant energetic cost to adult male harbour seals.     

Impact of body reserves on energy expenditure, water flux, and mating success in breeding male northern elephant seals

In capital breeders, individual differences in body size and condition can impact mating effort and success. In addition to the collateral advantages of large body size in competition, large nutrient reserves may offer advantages in endurance rivalry and enable the high rates of energy expenditure associated with mating success. We examined the impacts of body reserves and dominance rank on energy expenditure, water flux, mating success, and breeding tenure in the adult male northern elephant seal, a polygynous, capital breeder. Adult males expended energy at a rate of 159 ± 49 MJ d (-1), which is equivalent to 3.1 times the standard metabolic rate predicted by Kleiber's equation. Despite high rates of energy expenditure and a long fasting duration, males spared lean tissue effectively, deriving a mean of 7% of their metabolism from protein catabolism. Body composition had a strong impact on the ability to spare lean tissue during breeding. When controlling for body size, energy expenditure, depletion of blubber reserves, and water efflux were significantly greater in alpha males than in subordinate males. Large body size was associated with increased reproductive effort, tenure on shore, dominance rank, and reproductive success. Terrestrial locomotion and topography appeared to strongly influence energy expenditure. Comparisons with conspecific females suggest greater total seasonal reproductive effort in male northern elephant seals when controlling for the effects of body mass. In polygynous capital breeding systems, male effort may be strongly influenced by physiological state and exceed that of females.     

Plasma melatonin concentration in neonatal northern elephant seals,Mirounga angustirostris

The development of pineal function in northern elephant seals was examined in an attempt to understand the physiological basis for previously observed high daytime levels of melatonin in neonatal southern elephant seals. Pineal glands from four northern elephant seal pups, estimated age less than 1 week, weighed 3.0 ± 0.80 g, which was significantly less than that previously found in southern elephant seals (4.6 ± 0.35 g). Midday concentrations of plasma melatonin in pups averaged more than 3000 pmol/l in the first 5 days post-partum, but declined rapidly to less than 400pmol/l after day 9. Daytime melatonin levels in northern elephant seals tended to be lower than in southern elephant seals, although they were very high compared with other species. A circadian cycle of plasma melatonin concentration was observed in newborn northern elephant seals, with levels of 3000–5000 pmol/1 during the day, rising to more than 10,000 pmol/1 late in the dark phase. Soon after weaning at 4 weeks of age, daytime and night-time levels were in the range 60–100 pmol/1 and 100–400 pmol/1, respectively. When approximately 10 weeks old, most samples were in the range 100–400 pmol/1 with no discernible difference between day and night levels. The results do not support the hypothesis that the pineal gland is involved in thermogenesis in new-born southern elephant seals. Instead, the very active pineal gland may contribute to energy conservation, by lowering body temperature, particularly at night. As physical insulation is acquired by the deposition of blubber, the mechanism is not required and melatonin falls to adult levels.     

Diving behaviour and foraging location of female southern elephant seals from Patagonia

Our aim was to describe the free-ranging diving pattern and to determine the location of foraging of pregnant female southern elephant seals, Mirounga leonina , from Peninsula Valdes, Argentina. This colony is unusual in two respects: it is removed from deep water by a broad shallow shelf (345–630 km wide), and colony numbers have been increasing in recent years in contrast to numbers from other southern hemisphere colonies that are stable or in decline. Microprocessor controlled, geolocation-time-depth recorders were deployed on four females, recording a total of 15,836 dives (270 dive days) during the period February to April, 1992. Departing seals crossed the continental shelf quickly (54–5–62–1 h) and did not show signs of foraging until reaching deep water, due east of the colony in the South Atlantic Ocean. Diving was virtually continuous (93% of the time underwater) with overall mean (±S.D.) rates of 2.5±0.2 dives/h, mean dive durations of 22.8 ± 7.1 min (maximum dive duration = 79 min) with 1.6±0.6min surface intervals between dives, and dive depths of 431±193m (maximum dive depth = 1,072 m). The diving pattern of females from Patagonia is similar to that of seals from colonies where numbers are decreasing (Macquarie stock) or are stable (South Georgia Island). Our subjects did not, however, feed in or south of the Antarctic Polar Front, or in cold waters along the Antarctic coast, where seals from declining or stable colonies forage.     

Energy transfer and female condition in nursing harp seals Phoca groenlandica

Lactating female harp seals, most with their pups, were collected on the "whelping" ice in the Gulf of St. Lawrence in 1976 and from 1978 to 1980. During lactation females lost weight at an average rate of 3.17±0.52 (SE_b) kg d⁻¹, for a total energy loss of approximately 250000 kcal. Pups grew at a mean rate of 2.78±0.19 kg d⁻¹ for a gain in production energy of about 194000 kcal. Compared to 1976, adult females sampled in 1978 to 1980 had lower energy reserves at the onset of lactation. Coincidentally there has been a decrease in newborn lengths and girths. Although compensatory growth in weight appears to occur, between-year differences in newborn lengths persist to weaning. Various reproductive strategies of female harp seals faced with reduced energy stores are discussed.     

Lactation costs in southern elephant seals at King George Island,South Shetland Islands

Labelled-water methodology was used to quantify energy costs and energy transfer efficiency in 18 mother-pup pairs of southern elephant seals (Mirounga leonina) during lactation. During the lactation period, mothers lost a mean mass of 227±47 kg. Mass loss included 22% of the protein, 60% of the fat, and 51% of the energy in the mothers body upon arrival. Total body-energy reserves at parturition explained 69% of the variation in the total lactation costs and 50% of the variation in the pups body energy at weaning. On average, pups retained 48% of the mass, 49% of protein, 53% of fat and 51% of energy lost by their mothers. Greater, fatter females showed a decrease in the efficiency of energy and fat transfer and, at the same time, an increase in the efficiency of protein transfer. This may be due to an increased use of protein as metabolic fuel, as fat demands for milk production increase. There was no evidence that greater total lactation costs influence the ability of mothers to produce a pup in the next breeding season.     

Metabolic limits on dive duration and swimming speed in the southern elephant seal Mirounga leonina

The ability of air-breathing marine predators to forage successfully depends on their ability to remain submerged. This is in turn related to their total O(2) stores and the rate at which these stores are used up while submerged. Body size was positively related to dive duration in a sample of 34 adult female southern elephant seals from Macquarie Island. However, there was no relationship between body size and dive depth. This indicates that smaller seals, with smaller total O(2) stores, make shorter dives than larger individuals but operate at similar depths, resulting in less time being spent at depth. Nine adult female elephant seals were also equipped with velocity time depth recorders. In eight of these seals, a plot of swimming speed against dive duration revealed a cloud of points with a clear upper boundary. This boundary could be described using regression analysis and gave a significant negative relationship in most cases. These results indicate that metabolic rate varies with activity levels, as indicated by swimming speed, and that there are quantifiable limits to the distance that a seal can travel on a dive of a given swimming speed. However, the seals rarely dive to these physiological limits, and the majority of their dives are well within their aerobic capacity. Elephant seals therefore appear to dive in a way that ensures that they have a reserve of O(2) available.     

Dispersion during the moult haulout of southern elephant seals at the Courbet Peninsula, Iles Kerguelen

We studied the dispersion of 4-year-old southern elephant seals (Mirounga leonina) along a 75.5 km coastal area at the Courbet Peninsula, Iles Kerguelen, relative to their birth site when they were ashore to moult in early 1984. The seals were mostly faithful to their natal sites, but availability of suitable moulting habitat (e.g. wallows, vegetated areas) influenced seal dispersion. As moult progressed, the seals moved farther away from their initial moult sites and natal sites, but remained largely on the easterly beaches of the Courbet Peninsula. This behaviour would facilitate mark-recapture estimates of age and sex specific survival.     

SHORT NOTES

Laycock, H. T. 1982. Moulting and plumage changes in the Thickbilled Weaver. Ostrich 53:91-101.

Thickoilled Weavers were studied in captivity, in the wild and as museum specimens. Moulting follows the normal passerine pattern, but a difference from related species is that there is no post-fledging moult of the flight feathers. Methods were devised for identifying isolated feathers and for aging trapped birds, this being easier in the male. After the breeding season the male undergoes eclipse, which has apparently not been described before, and loses his white forehead patches. Adult males and females moult about the same time, but second-year males moult six or eight weeks earlier. The duration of post-nuptial moult is about four months and is timed to occur during the season when there is maximum food availability. The use of a “moult score” is avoided in this account and the timing of feather loss substituted as having more real meaning.