Sex-specific foraging strategies and resource partitioning in the southern elephant seal (Mirounga leonina)

The evolution of resource specializations is poorly understood, especially in marine systems. The southern elephant seal (Mirounga leonina) is the largest of the phocid seals, sexually dimorphic, and thought to prey predominantly on fish and squid. We collected vibrissae from male and female southern elephant seals, and assessed stable C and N isotope ratios along the length of the vibrissae. Given that whiskers grow slowly, this sampling strategy reflects any variation in feeding behaviour over a period of time. We found that isotopic variation among females was relatively small, and that the apparent prey choice and trophic level of females was different from that for males. Further, males showed a very broad range of trophic/prey choice positions, grouped into several clusters, and this included isotopic values too low to match a broad range of potential fish and cephalopod prey tested. One of these clusters overlapped with data for South American sea lions (Otaria flavescens), which were measured for comparison. Both male southern elephant seals and southern sea lions forage over the continental shelf, providing the potential for competition. We discuss the possibility that individual southern elephant seals are pursuing specialist foraging strategies to avoid competition, both with one another, and with the South American sea lions that breed nearby.     

The chromosomes of the southern elephant seal, Mirounga leonina (Phocidae: Mammalia).

Chromosome analyses were carried out on bone marrow cells from three elephant seal pups (Mirounga leonina), a species characterized by a 2n = 34 chromosome number. Arm ratios for the different chromosomes were calculated to facilitate the classification of chromosomes into groups. The karyotype of the southern elephant seal differs from other reported 34-chromosome phocids in the morphology of the Y chromosome and in that of one of the autosomal chromosome pairs.     

Past and present status of the southern elephant seal (Mirounga leonina) at Macquarie Island

The southern elephant seal population at Macquarie Island has undergone a serious decline since regular surveys were commenced in 1949. Approximately 2,900 cows and 250 bulls were counted in the isthmus study area during the 1985 breeding season. Comparisons with 20 other counts made in the same area between 1949 and 1984 showed that, although there was considerable fluctuation between the years, the number of seals ashore has dropped at an average rate of 2.1% per year, resulting in a net decrease of approximately 50% for both males and females. This is similar to the ratio of decline of elephant seal populations in the southern Indian Ocean. The census information from all major elephant seal populations was reviewed and it was concluded that there may be a common factor, or group of factors, acting to reduce the Macquarie Island and Indian Ocean populations, while the populations of the South Atlantic seem to be stable. Several potential explanations for these observations are advanced, but it is concluded that a greater knowledge of the ecology of the seals is needed if the declines are to be understood.     

Growth of female southern elephant seals Mirounga leonina at Macquarie Island

Body growth of 137 female southern elephant seals (Mirounga leonina) over 1 year of age was investigated at subantarctic Macquarie Island. An asymptotic straight line, snout–tail body length of 2.57±0.03 m was estimated to be attained at 9 years of age, using a three-parameter Gompertz equation. A significant increase of approximately 0.1 m (5%) in mean body length of females between 1 and 10 years of age was estimated to have occurred between the 1950–1960s and 1990s at Macquarie Island. This is consistent with a reduction in both the rate of population decline and the age of onset of sexual maturity. Age determination using dental cementum layers and the importance of standardised measurements in pinniped growth studies are discussed.     

Southern elephant seal,Mirounga leonina: composition of milk during lactation

An analysis of milk constituents during various stages of lactation in the southern elephant seal Mirounga leonina was carried out. Forty-six milk samples were taken from 30 females throughout lactation during 1985, 1987, 1990 and 1991 on Stranger Point, King George Island, South Shetland Islands. Total nitrogen (TN), non-protein nitrogen (NPN), sugar, fat, ash and water were measured, and from some of these data true protein and energy content were calculated. The results showed a high degree of variation in water and fat concentrations among samples at different stages of lactation. During the first 20 days the fat content of milk increased from about 12 to approximately 52%, while water content fell from 70 to 33%. The composition of milk changes rapidly during the first days post-partum. Protein, minerals and sugar appear to remain stable after the fourth day of lactation. Milk samples contain significant levels of sugars; thin layer chromatography indicates the presence of lactose and glucose together with other unidentified components. There is evidence of a striking change in composition of the milk in the later part of lactation; the progressive increase in the fat:water ratio is abruptly reversed just prior to weaning.     

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.     

Detection and characterization of a Cryptosporidium isolate from a southern elephant seal (Mirounga leonina) from the Antarctic peninsula

The presence of Cryptosporidium and Giardia in 221 fecal samples from different species of Antarctic pinnipeds was investigated by immunofluorescence microscopy and PCR. Cryptosporidium, a skunk-like genotype, was detected only in a southern elephant seal. Giardia was not detected. This is the first report of a Cryptosporidium sp. in Antarctic marine mammals.     

Counts of southern elephant seals, Mirounga leonina, at Bouvet Island

Southern elephant seals were counted and classified into subjective sex-age classes on a weekly basis during expeditions to Bouvet Island in the austral summers of 1996/1997 and 1998/1999. The expeditions coincided with the moulting period of elephant seals aged one?year and older. The presence of weaned pups at the principal haulout site, Nyrøysa/Westwindstranda, during the latter expedition, indicates that breeding took place here during 1998. Elephant seal counts from previous expeditions are summarised.     

Population status, trends and a re-examination of the hypotheses explaining the recent declines of the southern elephant seal Mirounga leonina

1. Between the 1950s and 1990s the southern elephant seal Mirounga leonina underwent large decreases in population size throughout most of its breeding range in the Southern Ocean. While current population estimates suggest a recent recovery, some breeding populations have continued to decrease in recent years (Macquarie and Marion Islands), others have either remained stable (South Georgia, Kerguelen and Heard Island) or have increased (Peninsula Valdés, Argentina). 2. Intrinsic hypotheses for patterns of regional decline include factors that are affected by density-dependent mechanisms: (i) paucity of males, (ii) population ‘overshoot’ and (iii) pandemic disease. Extrinsic hypotheses include (iv) predation, (v) competition with fisheries concerns, (vi) interspecific competition, (vii) environmental change and (viii) human disturbance. Of the eight hypotheses proposed and examined here, we conclude that three can be discounted (i, v, viii), three are unlikely, but may require more testing (ii, iii, iv) and two are plausible (vi, vii). 3. The interspecific competition hypothesis is difficult to test because it requires the simultaneous monitoring of species that overlap directly with elephant seals, many of which have not been identified or little is known. However, an analysis of the relationship between log variance and log abundance (Taylor's power law) for populations of southern and northern elephant seals suggests that interspecific competition is not a significant factor in the decline of the southern elephant seal. 4. The hypothesis that decreases in southern elephant seal populations between the 1950s and 1990s were caused by the environmental change is the easiest to test and most plausible of the hypotheses. We propose a framework by which to test this hypothesis to determine how food availability affects individual survival.     

Lipid composition of blood platelets and erythrocytes of southern elephant seal (Mirounga leonina) and antarctic fur seal (Arctocephalus gazella)

Erythrocyte and blood platelet phospholipid compositions were studied in three elephant seals and two fur seals, two species of marine mammals living in the Subantarctic region feeding on preys rich in (n-3) polyunsaturated fatty acids. Results were compared with those reported for related species and humans. In erythrocytes, the phospholipid (PL) and cholesterol (CHOL) contents were lower in pinnipeds than in humans. Phosphatidylcholine (PC) levels were higher in elephant seals than in fur seals, with a reverse trend for phosphatidylethanolamine (PE) and phosphatidylserine (PS). Both species had lower SM/PC ratios and PE plasmalogen concentrations than human. Erythrocytes were richer in (n-3) fatty acids (FA) in pinnipeds than in humans. In platelets, the PL content was lower and the CHOL content higher in elephant seals than in humans or in other phocid seal species studied to date. The SM/PC ratio was much higher than in other seal species or in man. In both species, the proportion of PE plasmalogens was higher in platelets than in erythrocytes. PL were more saturated in elephant seals than in fur seals. These results suggest that the erythrocytes and platelets of wild marine mammals may prove useful models to study the influence of dietary lipids on the structure and hemostatic function of these cells.     

Characterization of postdive recovery using sound recordings and its relationship to dive duration,exertion, and foraging effort of southern elephant seals (Mirounga leonina)

It is notoriously difficult to measure physiological parameters in cryptic free‐ranging marine mammals. However, it is critical to understand how marine mammals manage their energy expenditure and their diving behavior in environments where the predation risks are low and where survival is mainly linked to capacities to maintain physiological homeostasis and energy budget balance. Elephant seals are top marine predators that dive deeply and continuously when at sea. Using acoustic recorders deployed on two postbreeding southern elephant seals (SES) females, we developed methods to automatically estimate breathing frequency at the surface. Using this method, we found that seals took successive identical breaths at high frequency (0.29 Hz) when recovering at the surface and that breath count was strongly related to postdive surfacing time. In addition, dive depth was the main factor explaining surfacing time through the effects of dive duration and total underwater swimming effort exerted. Finally, we found that recovery does not only occur over one dive timescale, but over a multidive time scale for one individual. The way these predators manage their recovery will determine how they respond to the change in oceanic water column structure in the future.     

Adenohypophysis of the elephant seal (Mirounga leonina): morphology and seasonal histological changes

Changes in adenohypophyseal cell populations over a 12-month period were studied in the seasonally breeding elephant seal (24 adult males, 3 adult females, and 5 neonates) at Macquarie Island. The glands were weighed and fixed in formol sublimate. Selected sections were stained with the oxidation-alcian blue-periodic acid Schiff-orange G technique. Gonadotropic, thyrotropic, lactotropic, and somatotropic cells were readily identifiable; whilst corticotropes, inactive secretory cells of all types, and stellate cells were not stained and were counted collectively as chromophobic cells. Hypophyseal weight was low throughout autumn and winter, but increased significantly during the spring breeding season and the summer. Thyrotropes were distributed evenly throughout the pars distalis, but the other secretory cells showed areas of concentration. Acidophils were common peripherally, particularly lactotropes, while gonadotropes were largely confined to the 'basophilic wedge,' a narrow, central superior zone. In males, lactotropic and gonadotropic cells showed significant seasonal changes in number. Gonadotropes were more common in sexually active males than sexually quiescent ones, while lactotrope numbers were much greater at midsummer than midwinter. This lactotrope cycle appeared to be related to photoperiod but unrelated to breeding.     

Energy gain and loss during lactation and postweaning in southern elephant seal pups (Mirounga leonina) at King George Island

Deuterium-labeled water was used to measure changes in the proximate body composition during the lactation period and after weaning in southern elephant seal pups at King George Island, South Shetland Islands, Antarctica. During the lactation period (23.0 ± 1.4 days) pups gained a mean of 4.9 ± 0.5 kg/day (n=7). Of the total mass gain (112 ± 8 kg), 38% was water, 48% was fat, and 11% was protein. This represented an increase in total body gross energy of 2437 ± 145 MJ. The proportion of body mass represented by fat was less than 2% at birth, increasing to 35 ± 2% at weaning. We followed the pups during a mean period of 36 ± 3 days after weaning. During this period, pups had a mean loss of 1.21 ± 0.10 kg/day (n=7) comprising 39% water, 48% fat, and 12% protein. The energy cost over this period was 952 ± 168 MJ, which represented, on average, 39% of the total energy gained during the suckling period. Accepted: 3 January 2000     

Climate change and seal survival: evidence for environmentally mediated changes in elephant seal, Mirounga leonina, pup survival

Maternal and physical factors play a significant role in animal life-history variability, which means that large scale climate change has the potential to affect the size and dynamics of animal populations indirectly through maternal investment and directly through conditions that animals are exposed to. However, little is known about the effects of large-scale oceanographic events such as the El-Ni?o southern oscillation (ENSO) that influence productivity in the Southern Ocean and the abundance, quality and distribution of prey. The possible mechanisms by which physical factors and primary productivity could influence life-history traits, such as survival of apex predators, includes direct influences such as food availability and foraging success and indirect influences such as stored maternal investment and resource transfer during lactation. Here, we quantify the relative contribution of maternal investment and climate conditions at remote foraging sites to survival in the first year of life for southern elephant seals. We present evidence linking climate (ENSO) and variations in a key demographic parameter--first-year survival--and demonstrate that survival was highest during ENSO events and that the ability of mothers to store and acquire resources, which is typically related to ocean productivity, is the most important determinant of survival in the first year. This functional link provides valuable insights that can be used to model the responses of the seal populations to climate change scenarios.     

The annual cycle of the epididymis of the elephant seal (Mirounga leonina) at Macquarie Island

The tubules of the head and body of the elephant seal epididymis undergo large increases in diameter and principal cell height for the duration of the breeding season, and regress at the conclusion of breeding. The tubules of the tail show a lesser development, and only towards the end of breeding. Spermatozoa, first produced in the testis in August, reach the tail of the epididymis in September, and are present in large numbers throughout the epididymis in October and November. In December, the first month of sexual quiescence following the breeding season, sperm are almost totally absent from the epididymis.