The analysis of blubber fatty acids (FAs) is a useful tool to infer diet of mammals that live in remote regions where year-round studies are difficult. The FA may not be distributed uniformly within the blubber, which can have implications for dietary predictive studies. The aim of this study was to determine the FA composition in the blubber core of the Antarctic leopard seal, Hydrurga leptonyx, and evaluate the potential implications of FA stratification for dietary analysis. The blubber cores of 24 seals were sub-sectioned into outer, middle and inner layers and their FA were compared to those of their potential prey species. A vertical variation in FA composition was found across the whole blubber core of the leopard seal. 17 FAs were found at greater than trace amounts (>0.5%) across all samples and the most abundant were: C18:1ω9, C16:1, C22:6ω3, C16:0 and C18:1ω7, which accounted for approximately 70% of the total FA. Almost all FAs had a continuous gradient through the blubber. Principal Component Analysis confirmed separation between inner and outer layers while the middle layer was a transition. The stratification of the leopard seal blubber was similar to the general pattern observed in a variety of marine species: monounsaturated FA (MUFA) dominated the three layers being more abundant in the outer layer, polyunsaturated (PUFA) and saturated FA (SFA) were more abundant in the inner layer. Polyunsaturated FAs are of dietary origin and SFAs are chemically inert so they can be used as a long-term reserve, which suggest that the inner layer is the site of deposition of the FA obtained from diet. The influence of prey on the composition of the leopard seals' blubber was clearer in the inner layer, although neither outer nor inner layers exactly matched the FA of the potential prey. This suggests that there are other components influencing the FA composition of this predator; therefore, in order to carry out dietary analysis it is important to consider the stratification of blubber and to use the inner layer, where the influence of diet is more evident. This has significant implications for sampling methods in the field.
Southern elephant seals (Mirounga leonina) range widely throughout the Southern Ocean and are associated with important habitats (e.g., ice edges, shelf) where they accumulate energy to fuel their reproductive efforts on land. Knowledge of the fine scale foraging behaviour used to garner this energy, however, is limited. For the first time, acceleration loggers were deployed on three adult southern elephant seals during a translocation study at Kerguelen Island. The aims of the study were to (1) identify prey capture attempts using 2-D accelerometer tags deployed on the head of southern elephant seals, (2) compare the number of foraging dives identified by simple dive depth profiles and accelerometer profiles and (3) compare dive characteristics between prey encounter and non-prey encounter dives. The 2-D loggers recorded depth every second, surge and heave accelerations at 8 or 16 Hz and were carried for periods between 23 and 121 h. Rapid head movements were interpreted to be associated with prey encounter events. Acceleration data detected possible prey encounter events in 39–52% of dives whilst 67–80% of dives were classified as foraging dives when using dive depth profiles alone. Prey encounters occurred in successive dives during days and nights and lasted between tenths of a second and 7.6 min. Binomial linear mixed effect models showed that seals were diving significantly deeper and increased both descent rate and bottom duration when encountering prey. Dive duration, however, did not significantly increase during dives with prey encounters. These results are in accordance with optimal foraging theory, which predicts that deep divers should increase both their transit rates and the time spent at depth when a profitable prey patch is encountered. These findings indicate that this technique is promising as it more accurately detects possible prey encounter events compared with dive depth profiles alone and thus provides a better understanding of seal foraging strategies.
The Southern elephant seal (Mirounga leonina) is a major consumer from the Southern Ocean. This species is highly sexually dimorphic and frequently exhibits resource partitioning according to sex and/or age classes. This study analysed the feeding habits of the M. leonina population from Isla 25 de mayo (King George Island) in the spring/summer seasons of 1995/1996–2002/2003. A total of 232 individuals from different sex-age groups were stomach lavaged. The analysis of stomach samples showed that cephalopods were the main prey followed by fish, their frequency of occurrence being 98.1 and 17.9 % respectively. Cephalopods were dominated by the Antarctic glacial squid, Psychroteuthis glacialis, which occurred in 83 % of samples, constituting 57.2 % in numbers and 61.3 % in mass. Octopods were of lesser relevance, occurring in 18 % of samples, but became more important in the diet of male individuals. Juvenile seals fed on a lower variety of cephalopod prey than older ones. This would coincide with the diving pattern characteristic of the different sex-age categories of seals. The predominance of P. glacialis might be associated with the more southerly location of the foraging areas of this population compared to others. Fish were largely represented by the myctophid Gymnoscopelus nicholsi, which occurred in 81.3 % of samples containing otoliths and constituted 76.4 % in numbers and 66.4 % in mass. However, while myctophids may be the dominant fish prey of elephant seals in areas close to the South Shetlands, they would be probably replaced by P. antarcticum as seals migrate towards higher latitudes.
Seventy-seven immobilizations were carried out on adult male southern elephant seals at Stranger Point, Isla 25 de Mayo (King George Island) using a combination of Zoletil ® (tiletamine and zolazepam) and ketamine in order to obtain biological samples. During 2006/2007, 22 males were immobilized at the beginning of their breeding period (EB), 19 of which were recaptured at the end of breeding (LB). Four were given only once at an unknown stage of breeding (USB) and 18 males were immobilized at the beginning of molting (BM). During 2007/2008, 14 adult males were immobilized at an USB. Zoletil ® was administered using an automatic discharge device, whereas ketamine was injected directly with a syringe, and was used only when the initial sedation was not enough to carry out the programmed sampling. The initial mean dose of Zoletil ® was 1,387 ± 304 mg, which represented 0.60 ± 0.14 mg/kg, range 0.36–1.05, n = 77. In 47 procedures, an average dose of 1.04 ± 0.66 mg/kg of ketamine was added. Mean immobilization time was 34 ± 14 min. In 25 out of the 77 procedures, males showed apnea, which lasted 8 ± 4 min (range 2–15 min). The necessary doses of Zoletil ® and ketamine to attain immobilization differed between stages. For animals taken twice, doses (mg/kg) of Zoletil ® and ketamine were significantly higher at the beginning than at the end of breeding. During molting, the doses of Zoletil ® given were significantly lower than those used during breeding, although the proportion of animals that required ketamine during molting was significantly higher than during breeding. Zoletil ® proved to be a safe immobilizing agent for field work on adult males of this species, given the wide range of doses used without any serious consequences. Furthermore, the addition of ketamine was useful when the initial sedation was not satisfactory or for prolonging the immobilization period in a practical and reliable way.
Serum protein values in Southern elephant seals (Mirounga leonina) were analyzed during the breeding and molting periods at the 25 de Mayo Island (King George Island), Antarctica, during the 1999/2000 and 2000/2001 field seasons, in order to study the reaction of adults to fasting, and of pups to nursing and fasting. The following analyses were carried out: Total Proteins (TP) and Albumin (Alb) were analyzed by a colorimetric technique, and Apolipoprotein B (Apo-B) was determined by the immunodiffusion technique on agarose gel plates. The general ranges of average values for total proteins and specific fractions during the breeding period were: TP (g/dl) = 5.12–9.83; Alb (g/dl) = 1.72–5.71 and Apo-B (mg/dl) = 10–266, and during the post-breeding period: TP = 4.85–9.45; Alb = 2.06–4.20 and Apo-B = 13–232. The essential obtained data were: (a) fasting does not impact adult males, except for a significant decrease of Apo-B during breeding; (b) fasting does not impact adult females, except for a significant decrease in TP during molting; (c) suckling increases significantly TP and Apo-B in pups; (d) post-weaning fast decreases significantly all measured serum components in pups. We can conclude that the adults are adapted to long term fasting, without any metabolic downside they maintain homeostasis during this period, as shown by the serum data. The pups clearly react to both suckling and the post-weaning fast, which are periods driven almost exclusively by lipid chemistry. These impacts may be seen in the chemistry of serum proteins.
Changes in mass and body composition, measured with labelled water, were used to quantify the energy expenditure during lactation and energy replenishment during the post-breeding aquatic phase in Southern elephant seal females at Stranger Point, King George Island. During lactation females spent a mean of 6,021±1,365 MJ, which resulted in a loss of 35% of the initial mass, comprising 63% of initial body fat and 20% of initial body protein. During the 58±5.4 day post-breeding foraging period, females gained 135±39 Kg, which allowed them to recover an average of 55% of the mass, including 46% of the fat, 71% of the protein and 47% of the energy lost during lactation. Neither the mass nor the energy lost during lactation were related to those replenished while at sea. However, protein loss expressed in absolute terms or as a proportion of that present at the beginning of lactation explained about 50% of the variation in the protein gained during the post-breeding phase. This might indicate the presence of a mechanism favouring an increase in lean tissue during post-breeding. Daily energy requirements for an average sized female, during the post-breeding aquatic phase were estimated at 96 MJ. Estimation of prey consumption varies according to assumptions about diet composition. On a basis of 450 females, the total biomass of fish and squid consumed by the breeding group, assuming a diet composed of 75% cephalopods and 25% fish, was estimated to be 521 and 174 metric tonnes, respectively, for the period examined.
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 mother’s 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 pup’s 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.
We quantified immunoglobulins (Ig) in mammary secretions of 12 female southern elephant seals ( Mirounga leonina) at King George Island, Antarctica, using single radial immunodiffusion on agarose plates. Seals were chemically immobilized for milk sample collection at four time points during the 3- to 4-week suckling period. All three major mammalian immunoglobulins (IgG, IgM, IgA) were detected in southern-elephant seal milk. Total immunoglobulin levels and the ratio of Ig subclasses varied throughout the suckling period. Total immunoglobulin levels were highest on the 1st day of lactation, when they represented over 57% of the mean total protein concentration of the milk, and declined steadily throughout lactation, representing approximately 40% of the mean total protein concentration at the end of the suckling period. IgG was the most abundant immunoglobulin class (85.93–93.78% of total milk immunoglobulins), followed by IgM (6.06–13.93%), and IgA (0.14–0.23%). There was no significant difference in IgA levels throughout the suckling period. IgG levels were significantly lower during the second stage (3–6 days post-parturition) than during the first, third or fourth stages (1, 13–15, and 19–25 days post-parturition, respectively). IgM levels were highest during the first stage of lactation; these values were significantly higher than levels measured during the second, third and fourth stages of lactation. Transfer of passive immunity from female to offspring in other mammalian species is correlated with the subclass of immunoglobulin secreted in the milk; species acquiring passive immunity in utero, via the placenta, secrete a preponderance of IgA, whereas species acquiring immunity post-partum, via lacteal secretions and gut resorption, secrete a preponderance of IgG. The Ig patterns and concentrations observed in our study of southern elephant seals are consistent with an important role of post-partum transmission of passive immunity during the pinniped lactation period.
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.
Mass changes in female southern elephant seals, sampled sequentially at different points through their annual cycle, were measured at King George Island, South Shetland Islands, during the 1995/1996 and 1996/1997 field seasons. Females weighed after they had given birth showed an increase of 37 ± 36 kg (mean ± SD), which represented 6.2 ± 6.4% in relation to their mass in the first breeding season. During the first aquatic phase, between the end of lactation and the beginning of moult, females gained a mean of 128 ± 35 kg, (n = 18) (2.19 ± 0.65 kg day−1), which represented between 27 and 83% of the mass they had lost during lactation. Nine females followed during moulting showed a mass loss rate of 5.0 ± 0.4 kg day−1, which was half the rate during lactation. Total mass loss during moulting (129 ± 22 kg) was not significantly different from mass gain for the same females between lactation and moult (135 ± 37 kg). Furthermore, at the end of moulting, female mass was not significantly different from the mass at the end of lactation. These masses represented 65 ± 5% and 64 ± 5%, respectively, of their initial mass after parturition. During the second period at sea, from the end of the moult until females hauled out to give birth in the following breeding season, the estimated mass gain was 1.45 ± 0.24 kg day−1 (n = 5), which was not significantly different to the rate of mass gain shown by the same females during the first period at sea (2.26 ± 0.70 kg day−1). Total mass gain during the second aquatic phase (364 ± 63 kg) was not correlated with the mass at the end of moulting, but it was positively related to the mass loss experienced by females from parturition until the end of the moulting period in the first breeding season.
Mass loss by females and changes in the body composition of their pups were measured during lactation in southern elephant seals, Mirounga leonina, at King George Island, South Shetland Islands. Mothers lost 10.78±2.07 kg/day (n=6), while theor pups gained 5.18±1.5 kg/day. The period measured (19±0.6 days) represented 86% of the whole lactation period. During the period measured females lost an average 31% of their weight after parturition. Of the total mass gained by pups (98.7±30.5 kg), 45% was water, 12% was protein, and 41% was fat. Fat reserves of pups, which represented less than 2% of their body weight at birth, increased to 31% at weaning. This constituted a mean gain of 45 kg of fat during the experimental period. At weaning, pups at King George Island showed greater fat reserves than those at other breeding sites, a fact which could contribute to their survival when they start to feed at sea.
This paper reports Immunoglobulin M (IgM) levels in serum samples from eight female-pup pairs of southern elephant seals (Mirounga leonina), at King George Island, Antarctica. IgM levels were determined on sera obtained from sequential sampling throughout the suckling period (approximately 23 days). The IgM concentration in southern elephant seal serum was measured by single radial immunodiffusion on agarose plates. Female IgM levels (123.5–613.0 mg/dL, n = 8) were significantly higher than pup levels (5.9–123.6 mg/dL, n = 8). Both groups showed an increasing trend throughout the entire suckling period, with significant differences in relation to stages of lactation. Pup IgM levels on the first day of life (mean ± SD, 7.6 ± 2.9 mg/dL, n = 3) suggest that endogenous synthesis takes place before birth.
Mass transfer from mother to pup during the lactation period, and mass recovery for the same females during the foraging period were measured in the southern elephant seal at King George Island, Antarctica. During the 19.2 ± 0.9-day lactation period measured (which represented 87% of the entire nursing), females lost a mean mass of 10.56 ± 1.76 kg/day (n = 27), while their pups gained a mean mass of 5.27 ± 1.1 kg/day. There was a correlation between daily body weight gain in pups and daily weight loss by their mothers. Pup weaning mass was positively related to maternal post-partum mass. Serial samples showed that weight losses by females and gains by their pups were not linear over lactation, but showed lower values at the beginning and at the end of lactation. During the 60.5 ± 6.2-day foraging phase between the end of lactation and molt, females gained 2.21 ± 0.65 kg/day (n = 12), or 54% of the mass lost during nursing. Growth rates reported here are higher than those reported in other breeding sites. However, the ratio of body mass loss by females to gain by their pups was similar, suggesting that higher growth rates and greater weaning mass at South Shetland are due to a higher mean weight of females on arrival at this breeding site. The foraging period was shorter and the mass gained greater than those measured at South Georgia; this could be related to relatively shorter distances to foraging areas.
Milk protein fractions during various stages of lactation in the southern elephant seal Mirounga leonina were analysed. Twelve milk samples were taken from ten females throughout the lactation period during 1990 and 1991 at Stranger Point, King George Island, South Shetland Islands. Milk samples were subjected to polyacrylamide gel electrophoresis (PAGE). Samples from different days of lactation gave similar qualitative electrophoretic patterns. True protein content was significantly higher (P<0.05) at the beginning of lactation, and then remained constant until weaning. Caseins and whey proteins each consisted of several protein entities (four and five distinct bands respectively). Casein constituted only about 30% of the protein nitrogen, the remaining 70% being derived from whey proteins. There was some variation in concentration of casein and whey proteins as a function of time (P<0.0.5).
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.