Pup Weaning Mass Research Articles

Effects of external tags on maternal postpartum, offspring body mass and breeding frequency in gray seals Halichoerus grypus

AbstractFew studies have examined the impacts of externally fitted data‐loggers and telemetry tags on pinnipeds. We tested for instrument effects on body mass of lactating female gray seals and their offspring and probability of pupping in the next breeding season. Known‐age adult females (n = 216) were fitted with instruments in winter, spring, and fall from 1992 to 2018 at Sable Island, Nova Scotia. Of those tagged in spring and fall, 61 of 135 returning females and 59 of their offspring were weighed within 5 days postpartum and 79 pups were weighed at weaning. Instrumented females were assigned to treatments based on tag frontal area sums, tag mass, deployment duration, and acoustic tag presence compared to control females without instruments using linear mixed‐effects models. None of the treatment effects were included in the preferred models predicting birth mass of offspring or probability of breeding in the following year. The small negative effect (−3% to −7%) on postpartum maternal mass and pup weaning mass (−4.7%) for females instrumented in fall may be an artifact as longer spring deployments showed no effect. Overall, we found that the instruments deployed had no detectable negative effects on the maternal and offspring variables measured.

Fitness correlates of blubber oxidative stress and cellular defences in grey seals (Halichoerus grypus): support for the life-history-oxidative stress theory from an animal model of simultaneous lactation and fasting.

Life-history-oxidative stress theory predicts that elevated energy costs during reproduction reduce allocation to defences and increase cellular stress, with fitness consequences, particularly when resources are limited. As capital breeders, grey seals are a natural system in which to test this theory. We investigated oxidative damage (malondialdehyde (MDA) concentration) and cellular defences (relative mRNA abundance of heat shock proteins (Hsps) and redox enzymes (REs)) in blubber of wild female grey seals during the lactation fast (n = 17) and summer foraging (n = 13). Transcript abundance of Hsc70 increased, and Nox4, a pro-oxidant enzyme, decreased throughout lactation. Foraging females had higher mRNA abundance of some Hsps and lower RE transcript abundance and MDA concentrations, suggesting they experienced lower oxidative stress than lactating mothers, which diverted resources into pup rearing at the expense of blubber tissue damage. Lactation duration and maternal mass loss rate were both positively related to pup weaning mass. Pups whose mothers had higher blubber glutathione-S-transferase (GST) expression at early lactation gained mass more slowly. Higher glutathione peroxidase (GPx) and lower catalase (CAT) were associated with longer lactation but reduced maternal transfer efficiency and lower pup weaning mass. Cellular stress, and the ability to mount effective cellular defences, could proscribe lactation strategy in grey seal mothers and thus affect pup survival probability. These data support the life-history-oxidative stress hypothesis in a capitalbreeding mammal and suggest lactation is a period of heightened vulnerability to environmental factors that exacerbate cellular stress. Fitness consequences of stress may thus be accentuated during periods of rapid environmental change.

Exploring causal components of plasticity in grey seal birthdates: Effects of intrinsic traits, demography, and climate.

Change in breeding phenology is often a response to environmental forcing, but less is known of the mechanism underlying such changes and their fitness consequences. Here, we report on changes in the breeding phenology from a 27‐year longitudinal study (1991–2017) of individually marked, known‐aged grey seals (Halichoerus grypus) on Sable Island, Nova Scotia, Canada. We used generalized linear mixed models and a 3‐step process to develop a model that includes interactions between intrinsic and extrinsic covariates and to test hypotheses about the influence of fixed factors (maternal age, parity, previous reproductive success, pup sex, colony density, Atlantic Multidecal Oscillation (AMO), North Atlantic Oscillation (NAO), and Sea Surface Temperature) and a random factor (female identity) on parturition dates. We also examined the consequences of the shift in birthdates on maternal energy allocation in offspring as measured by pup weaning mass. Birthdates were known for 2,768 pups of 660 known‐age females. For 494 females with ≥2 parturition dates, repeatability as measured by the intraclass correlation was high (mean = 0.66). 87% of the variation in birthdates was explained by a mixed‐effects model that included intrinsic and extrinsic fixed effects. Most of the explained variation was associated with the random effect of female identity. Parity was the most important intrinsic fixed effect, with inexperienced mothers giving birth later in the season than multiparous females. Over almost 3 decades, mean birthdates advanced by 15 days. The mixed model with intrinsic effects and population size, the detrended AMO from the previous year and mean NAO in the previous 3 years explained 80% of the variation with 21% of variation from the fixed effects. Both primiparous and multiparous individuals responded to the climate forcing, and there was strong evidence for heterogeneity in the response. Nevertheless, the shift in birthdates did not impact pup weaning mass.

Decadal changes in blood δ13C values, at-sea distribution, and weaning mass of southern elephant seals from Kerguelen Islands.

Changes in the foraging environment and at-sea distribution of southern elephant seals from Kerguelen Islands were investigated over a decade (2004-2018) using tracking, weaning mass, and blood δ13C values. Females showed either a sub-Antarctic or an Antarctic foraging strategy, and no significant shift in their at-sea distribution was detected between 2004 and 2017. The proportion of females foraging in sub-Antarctic versus Antarctic habitats did not change over the 2006-2018 period. Pup weaning mass varied according to the foraging habitat of their mothers. The weaning mass of sub-Antarctic foraging mothers' pups decreased by 11.7 kg over the study period, but they were on average 5.8 kg heavier than pups from Antarctic foraging mothers. Pup blood δ13C values decreased by 1.1‰ over the study period regardless of their sex and the presumed foraging habitat of their mothers. Together, these results suggest an ecological change is occurring within the Indian sector of the Southern Ocean with possible consequences on the foraging performance of southern elephant seals. We hypothesize that this shift in δ13C is related to a change in primary production and/or in the composition of phytoplankton communities, but this requires further multidisciplinary investigations.

Sources of variation in maternal allocation in a long‐lived mammal

Life history theory predicts allocation of energy to reproduction varies with maternal age, but additional maternal features may be important to the allocation of energy to reproduction.We aimed to characterize age‐specific variation in maternal allocation and assess the relationship between maternal allocation and other static and dynamic maternal features.Mass measurements of 531 mothers and pups were used with Bayesian hierarchical models to explain the relationship between diverse maternal attributes and both the proportion of mass allocated by Weddell seal mothers, and the efficiency of mass transfer from mother to pup during lactation as well as the weaning mass of pups.Our results demonstrated that maternal mass was strongly and positively associated with the relative reserves allocated by a mother and a pup's weaning mass but that the efficiency of mass transfer declines with maternal parturition mass. Birthdate was positively associated with proportion mass allocation and pup weaning mass, but mass transfer efficiency was predicted to be highest at the mean birthdate. The relative allocation of maternal reserves declined with maternal age but the efficiency of mass transfer to pups increases, suggestive of selective disappearance of poor‐quality mothers.These findings highlight the importance of considering multiple maternal features when assessing variation in maternal allocation.

Geographic variation in subantarctic fur seal pup growth: linkages with environmental variability and population density

Abstract Marine predator populations are sensitive to temporal variation in prey availability, but prey dynamics are often difficult to quantify. Long-term measures of offspring growth is a useful performance attribute to gauge the potential demographic direction for such predator populations, especially where other metrics (e.g., population size estimates) are lacking. Subantarctic fur seal ( Arctocephalus tropicalis ) females are central place foragers during a protracted lactation period, and their foraging success determines the growth and vitality of their offspring. Using data spanning over 2 decades, we assessed geographic and temporal variation in growth rates and weaning mass of subantarctic fur seal pups at 2 of the species’ principal populations (Gough and Marion islands) and identified environmental conditions that may, through assumed bottom-up mechanisms, affect body mass at weaning. While Marion Island pups grew at an average rate of between 0.040 and 0.067kg/day early in lactation (comparable to conspecific growth at Amsterdam Island), the mean growth rate at Gough Island (approximately 0.030kg/day) was lower than the growth rate represented by the bottom 5% of the body mass distribution at Marion Island. Notwithstanding substantial interannual variability, we found support for a negative trend in weaning mass at both populations, suggesting a rise in limiting factors that is hypothesized to relate to concurrent local population size increases. Weaning mass tended to be higher when sea surface temperatures were warmer (with a stronger positive effect at Gough Island) and during positive phases of the Southern Oscillation Index (La Niña events), with a stronger positive effect in males. Given the low weaning mass of Gough Island fur seal pups, continued population growth here seems unlikely. While density-dependent regulation appears to have increased in strength at Marion Island, terminating rapid population growth, current weaning weights remain above the physiological limits of growth in subantarctic fur seals.

Age-related reproductive variation in a wild marine mammal population

Life history theory predicts a change in reproduction success with age as energy resources are limited and must be allocated effectively to maximize reproduction and survival. In this study, we use three reproductive performance measures, maternal expenditure, offspring weaning mass, and first-year survival, to investigate the role that maternal age plays in successful reproduction. Long-term uninterrupted life history data available for Marion Island’s southern elephant seals and mass change estimates from photogrammetry data allow for assessment of age-related reproduction performance and trade-offs. Known-aged adult females were photographed for photogrammetric mass estimation (n = 29) and their pups weighed at weaning during the 2009 breeding season. Maternal age and proportional mass loss positively influenced pup weaning mass. In turn, first-year pup return rates (as a proxy for survival) were assessed through the intensive mark–recapture program. Pup survival increased with female age and weaning mass. Pups of young females aged 3–6 years have a lower first-year survival probability compared with pups of older and larger females.

O' mother where wert thou? Maternal strategies in the southern elephant seal: a stable isotope investigation

Maternal effects are widespread in ecology and can alter the dynamics of a population. We investigated the impact of maternal foraging strategies on offspring weaning mass--a proxy of maternal foraging success and of offspring survival--in southern elephant seals on îles Kerguelen. Using 4 years of data, we modelled pup weaning mass as a two-component mixture and used blood stable isotope values to discriminate between maternal foraging strategies previously identified from bio-logging studies. Carbon isotope ratio was a strong predictor of weaning mass, but the relationship was non-monotonic in contrast to a priori expectations. Females foraging in the interfrontal zone weaned pups with a smaller mass compared with females foraging in Antarctic waters. Pup mass was positively correlated with a proxy of global primary production in the interfrontal zone for small weanlings. Maternal effects, via a poor foraging efficiency in the 1970s, may help explain the large population decrease observed at that time on îles Kerguelen because of an overall decrease in pup weaning mass, survival and subsequent recruitment.

Repeatability in lactation performance and the consequences for maternal reproductive success in gray seals

In mammals, the most significant maternal effect on offspring growth during lactation is the ability of females to efficiently transfer milk energy to their neonates. However, despite the importance of the transfer of milk energy to both maternal and offspring fitness, nothing is known about the extent to which variation among females may be attributed to differences in individual quality or environmental variation in natural populations. We measured repeatability over multiple lactation periods in components of lactation performance in free-ranging, multiparous gray seal (Halichoerus grypus) females to examine to what extent variation among females in pup weaning mass may be attributed to inherent differences in their physiological capacity to deliver milk energy. Levels of repeatability were high for milk composition (r = 0.38-0.50), daily milk output (r = 0.46), and the duration of lactation (r = 0.57), demonstrating that there are consistent differences among females in these characters across lactations and that the overall capacity of gray seal females to deliver milk energy to their pups is characteristic of individuals. The repeatability in pup weaning mass (r = 0.48) was consistent with the values for the components of total milk energy output and suggests that, over a large proportion of their reproductive life, individual gray seal females will consistently wean pups with greater or lesser probabilities of survival. Our results suggest that inherent differences among females in their physiological capacity to deliver milk energy may be an important component of variation in individual quality and, thus, lifetime reproductive success in mammals. High levels of repeatability also suggest that components of milk energy transfer may have a significant heritable genetic basis.

Maternal influences on reproduction in two populations of Columbian ground squirrels

In this study, we examined influences of maternal traits on offspring birth mass, growth rate, and weaning mass for two populations of Columbian ground squirrels (Spermophilus columbianus). We tested relationships between maternal body condition, structural size, change in mass (during gestation, during lactation, and during the entire reproductive period), timing of reproduction, and litter size on offspring traits using path analyses. To assess whether maternal investment in offspring traits extended beyond the period of direct maternal care, we examined associations between offspring traits and overwinter survival of pups. In general, females in better condition raised pups that were heavier at weaning and that had faster growth rates during lactation. Litter size had a negative effect on mass and growth rate, and only litter size had a significant effect on birth mass. For both populations, the average weaning mass of pups within a litter had a positive effect on the number of pups that survived to yearling age. In a population for which birth masses and growth rates were available, pups with faster growth rates survived better to yearling age, whereas birth mass had no effect on the number of surviving offspring in litters. We found substantial maternal influences on offspring growth and size, and evidence that these influences may extend beyond the juvenile period and constitute influences on fitness. The key to arriving at these conclusions was to take the number of offspring into account before testing for maternal effects.

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (Mirounga leonina) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ13C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ13C and δ15N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ13C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ13C and δ15N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.

The effects of El Niño–La Niña on reproductive parameters of elephant seals feeding in the Bellingshausen Sea

AbstractAim To assess the impacts of El Niño–La Niña events on the pup weaning mass and diet of female southern elephant seals (Mirounga leonina) feeding in the Bellingshausen Sea, Antarctica, and to understand the ecological processes that drive these impacts.Location Atlantic southern elephant seal weaning mass and diet were measured at King George Island (62º14′ S, 58º30′ W). Feeding areas for pregnant female seals from King George Island are located west of Alexander Island in the Bellingshausen Sea.Methods Data on weaning mass were collected between 1985 and 1994 during the breeding season (September–November). Moulting females were anaesthetized and cephalopod beaks were isolated and identified from stomach contents obtained from stomach lavages. Sea‐surface temperature anomaly (SSTA) data for the ‘El Niño 3.4’ geographical region (5º N–5º S, 120º W–170º W) were used to define El Niño–Southern Oscillation (ENSO) event years (grouped as El Niño, La Niña and Neutral) as well as the strength of each ENSO event year. Using data from the US National Center for Environmental Prediction, temperature, sea ice concentration and atmospheric pressure anomalies in the Bellingshausen Sea were calculated from March to August, corresponding to the feeding period of pregnant female seals.Results Positive temperature anomalies and negative pressure anomalies in the Bellingshausen Sea were observed during La Niña years and negative temperature anomalies and positive pressure anomalies during El Niño years. These data correlate with sea ice concentration anomalies, which are highly negative during La Niña years and highly positive during El Niño years. Warm temperature conditions in the Bellingshausen Sea during La Niña years are strongly related to both higher weaning mass in elephant seals and to an increase in squid beaks in the stomach contents of females.Main conclusions It is possible that higher elephant seal weaning masses in La Niña years correlate with warmer waters in the Bellingshausen Sea leading to the rapid growth of squid and their more frequent descents to depths frequented by elephant seals. This results in increased predation by pregnant females, leading to a greater mass among weaned pups. This hypothesis may guide future research about interactions between climate and the marine biosphere.

THE IMPORTANCE OF CONSIDERING PREDICTION VARIANCE IN ANALYSES USING PHOTOGRAMMETRIC MASS ESTIMATES

AbstractDevelopment and application of photogrammetric mass‐estimation techniques in marine mammal studies is becoming increasingly common. When a photogrammetrically estimated mass is used as a covariate in regression modeling, the error associated with estimating mass induces bias in regression statistics and decreases model explanatory power. Thus, it is important to understand and account for prediction variance when addressing ecological questions that require use of estimated mass values. In a simulation study based on data collected from Weddell seals, we developed regression models of pup weaning mass as a function of maternal postparturition mass where maternal mass was directly measured and second where maternal mass was photogrammetrically estimated. We demonstrate that when estimated mass was used, the regression coefficient was biased toward zero and the coefficient of determination was 30% less than the value obtained when using maternal postparturition mass obtained from direct measurement. After applying bias correction procedures, however, the regression coefficient and coefficient of determination were within 2% of their true values. To effectively use photogrammetrically estimated masses, prediction variance should be understood and accounted for in all analyses. The methods presented in this paper are effective and simple techniques to explore and account for prediction variance.

Reproductive performance in grey seals: age‐related improvement and senescence in a capital breeder

1. Three hypotheses have been advanced to account for age-related improvement in performance: the selection hypothesis predicts improved due to the loss of lower quality phenotypes, the constraint hypothesis predicts individuals improve function, and the restraint hypothesis predicts younger individuals forego or reduce effort because of mortality risks. A decline in age-related performance (i.e. senescence) is predicted by mutation accumulation, antagonistic pleiotropy and disposable soma (wear and tear) hypotheses. 2. Using five measures of performance - birth rate, maternal and pup birth mass, pup weaning mass, weaning success and lactation length - we tested these hypotheses concerning age-related change in reproduction in 279 female grey seals (Halichoerus grypus), ages 4-42 years, over a 23-year period between 1983 and 2005 on Sable Island, Nova Scotia. These females produced 2071 pups. 3. Although body mass of primiparous females increased with age (4-7 years) birth mass of their pups did not, but pup weaning mass did. Second- and third-parity females of the same age as primiparous females gave birth to and weaned heavier pups. However, parity and age were dropped from models when maternal body mass was included. 4. The proportion of females giving birth varied significantly with maternal age, increasing in young females and then declining late in life. Weaning success rate also increased rapidly to about 8 years and subsequently declined in females > 32 years. 5. Generalized additive models indicated nonlinear changes in 3 day body mass (i.e. approximately birth mass) and weaning mass of pups as a function of maternal age, after accounting statistically for the effects of maternal body mass. Mixed-effects, repeated-measures models fitted to longitudinal data further supported the conclusion that pup birth mass and weaning mass vary nonlinearly with maternal age and indicated nonlinear changes in lactation duration. 6. We found some support for the constraint hypothesis, but our findings were not consistent with the selection hypothesis or the restraint hypothesis as the basis for improvement in reproductive performance. 7. Senescence was evident in multiple female and offspring traits, indicating the degeneration in function of several physiological systems as predicted by the disposable soma hypothesis.

Ocean climate and seal condition

BackgroundThe condition of many marine mammals varies with fluctuations in productivity and food supply in the ocean basin where they forage. Prey is impacted by physical environmental variables such as cyclic warming trends. The weaning weight of northern elephant seal pups, Mirounga angustirostris, being closely linked to maternal condition, indirectly reflects prey availability and foraging success of pregnant females in deep waters of the northeastern Pacific. The aim of this study was to examine the effect of ocean climate on foraging success in this deep-diving marine mammal over the course of three decades, using cohort weaning weight as the principal metric of successful resource accrual.ResultsThe mean annual weaning weight of pups declined from 1975 to the late 1990s, a period characterized by a large-scale, basin-wide warm decadal regime that included multiple strong or long-duration El Niños; and increased with a return to a cool decadal regime from about 1999 to 2004. Increased foraging effort and decreased mass gain of adult females, indicative of reduced foraging success and nutritional stress, were associated with high ocean temperatures.ConclusionDespite ranging widely and foraging deeply in cold waters beyond coastal thermoclines in the northeastern Pacific, elephant seals are impacted significantly by ocean thermal dynamics. Ocean warming redistributes prey decreasing foraging success of females, which in turn leads to lower weaning mass of pups. Annual fluctuations in weaning mass, in turn, reflect the foraging success of females during the year prior to giving birth and signals changes in ocean temperature cycles.

Maternal and newborn life‐history traits during periods of contrasting population trends: implications for explaining the decline of harbour seals (Phoca vitulina), on Sable Island

AbstractAnnual censuses of the number of harbour seal Phoca vitulina pups born on Sable Island Canada showed an increasing trend during the 1980s, but a rapid decline through the 1990s from 625 pups in 1989 to only 32 by 1997. Weekly surveys of the North Beach of the island during the 1991–98 breeding seasons showed that the number of adults and juveniles also declined during the 1990s. Despite the dramatic demographic changes, maternal postpartum mass, pup birth mass, relative birth mass, lactation duration, pup weaning mass and relative weaning mass showed no significant trends during 1987–96. However, two traits did change. The age structure of parturient females increased significantly, indicating reduced recruitment to the breeding population. Mean birth date increased by 7 days during the early 1990s, suggesting nutritional stress of females and later implantation dates. This nutritional stress may in turn have been caused by increased competition from the rapidly increasing grey seal population on Sable Island. Although minimum estimates of shark‐inflicted mortality can account for much of the decline, evidence suggests that food shortage arising from interspecific competition may have also played a role in causing the decline of the population through effects on fecundity and juvenile survival.

Human disturbance, nursing behaviour, and lactational pup growth in a declining southern elephant seal (Mirounga leonina) population

We studied lactation behaviour in relation to pup growth in southern elephant seals (Mirounga leonina) at Macquarie Island, and compared harems in areas of high and low human presence to determine if there is an effect attributable to human activities, including scientific research. Pup weaning mass, a known correlate of first–year survival, was positively influenced by suckle bout durations during early and middle lactation and by maternal aggression during late lactation; no other behavioural variables were associated with weaning mass. In the area of high human presence, we observed from a distance the behaviour of mother–pup pairs directly before, during, and after visits to harems by other researchers. Alertness was raised threefold in the presence of people but quickly returned to predisturbance levels after their departure; there were no significant short-term effects on other behavioural variables. In the areas of high and low human presence, we observed the undisturbed behaviour of the seals in the absence of other people. No significant differences in any behavioural variables examined were found, indicating no long-term changes in behaviour resulting from human presence. Human disturbance therefore appears not to have significantly contributed to the population decline observed at Macquarie Island, but the conclusion requires caution given the fairly low power of our analyses.

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.

Maternal effects on offspring growth rate and weaning mass in harbour seals

We studied maternal effects on offspring traits during lactation in harbour seals (Phoca vitulina) on Sable Island, Nova Scotia, from 1988 to 1996. Duration of lactation was correlated with rate of pup mass gain (r = %#150;0.34, n = 116) and weaning mass (r = 0.29, n = 154). Pups that grew faster had shorter nursing periods, whereas those that attained higher weaning masses nursed for a greater number of days. Pup sex did not affect patterns of maternal effects. The pups of young females (4%#150;6 years old) gained mass at a constant but lower rate (0.56 kg/d) than the pups of older females through midlactation (0.74%#150;0.78 kg/d; n = 75). In older females, rates of pup mass gain decelerated between mid and late lactation. Although maternal age did not directly affect weaning mass of pups, path analysis showed that maternal age acted on weaning mass through intermediary traits. Lighter females gave birth to smaller and slower growing pups, but invested relatively more than heavier females (n = 153). Effects of maternal postpartum mass on weaning mass (n = 100) were weaker in harbour seals than in phocids that fast during lactation, but apparently stronger than in otariids that forage during lactation, suggesting that the strength of maternal effects is influenced by lactation strategy.

Mass of weaned elephant seal pups in areas of low and high human presence

On sub-Antarctic Macquarie Island, we examined pup weaning mass of southern elephant seals in relation to human presence. Pup weaning mass was previously found to be positively associated with 1st-year survivorship. Weaned pups were weighed in a remote area, Middle Beach, and in an area of relatively high human presence, Isthmus East. The areas were reasonably similar in beach topography, wind and surf conditions, numbers of seals present per kilometre of coastline, and numbers of males and females present in harems. For a sub-sample of measured pups, data on the respective maternal size were collected using photogrammetry. Both male and female weaned pups on Middle Beach were significantly heavier than those on Isthmus East. Estimated length of mothers was significantly higher on Middle Beach. In proportion to their own size, mothers in both areas produced weaners of similar mass, indicating no direct effect of human disturbance on the efficiency of lactation. It remained unclear whether the area differences in maternal and pup size were due to natural or human-related factors.