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

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

Development of the blood and muscle oxygen stores in gray seals (Halichoerus grypus): implications for juvenile diving capacity and the necessity of a terrestrial postweaning fast

To successfully transition from nursing to foraging, phocid seal pups must develop adequate diving physiology within the limited time between birth and their first independent foraging trip to sea. We studied the postpartum development of oxygen stores in gray seals (Halichoerus grypus, n=40) to better understand the ontogeny of diving capacity in phocids. Hemoglobin (Hb), hematocrit (Hct), blood volume (BV), and myoglobin (Mb) levels in newborn (3 d postpartum [DPP]) and newly weaned (17+/-0.4 DPP) pups were among the lowest measured across age classes. During the pups' terrestrial postweaning fast (PWF), Hb, Hct, mass-specific BV, and Mb increased by 28%, 21%, 13%, and 29%, respectively, resulting in a 35% increase in total body mass-specific oxygen stores and a 23% increase in calculated aerobic dive limit (CADL). Although Hb and Hct levels at the end of the PWF were nearly identical to those of yearlings, total body mass-specific oxygen stores and CADL of weaned pups departing for sea were only 66%-67% and 32%-62%, respectively, of those for yearlings and adult females. The PWF represents an integral component of the physiological development of diving capacity in phocids; however, newly independent phocids still appear to have limited diving capabilities at the onset of foraging.     

Morphometrics,body condition,and growth of the ringed seal (Pusa hispida saimensis) in Lake Saimaa: Implications for conservation

Morphometrics and growth of the critically endangered Saimaa ringed seal (Pusa hispida saimensis), which inhabits a freshwater lake in Finland were studied using data from 344 carcasses. This study presents the first detailed information on ringed seal pup growth and body condition from birth to the age of one year. Newborn pups were on average 68 cm long and weighed 5 kg. Pups attained the majority of their first year growth during the nursing period. Body condition and growth declined after weaning, but pups recovered from postweaning nutritional deprivation at the age of 8 mo. The seals achieved their maximum body length, girth, and mandible size around the age of 4 yr, and asymptotic body mass two years later. Baculum growth indicated that males reached sexual maturity at age 5–6 yr. The Saimaa ringed seals' asymptotic body length and mass were 132 cm and 59 kg, respectively, which is similar to medium sized marine ringed seals. Saimaa ringed seals' survival to adulthood is extremely low due to high bycatch mortality; furthermore climate change may hamper pup growth and elevate mortality. Therefore we recommend continuation of fishing closures to improve pup survival.     

Movements and diving behavior of weaned Weddell seal (Leptonychotes weddellii ) pups

Between 1993 and 1995, the diving behavior and movement patterns of 23 weaned Weddell seal pups (Leptonychotes weddellii) were tracked in the Ross Sea. Antarctica, using satellite-linked time-depth recorders. Regression analyses revealed that for seals of between 8 and 27 weeks old, age was poorly correlated with the dive depth, duration, or frequency. However, changes in dive parameters suggested that Weddell seal pups were attempting to maximize dive time, but the manner in which this was done depended on age and time of day. Movement patterns indicated that most Weddell seal pups left their natal area by the end of February, and traveled north along the Antarctic continent coastline. Several individuals returned to McMurdo Sound, but others were last located more than 400 km from McMurdo. Routes followed suggest that pups can use the pack ice habitat, but prefer to remain closer to the coastline than do adults. Accepted: 21 July 1998     

Diet composition and seasonal feeding patterns of a freshwater ringed seal (Pusa hispida saimensis)

The Saimaa ringed seal (Pusa hispida saimensis) is one of the few freshwater seal populations worldwide. The major conservation issue of this critically endangered population is bycatch mortality. We used digestive tract content and stable isotopes (δ¹³C and δ¹⁵N) to estimate the diet and seasonal feeding patterns for gaining better understanding of the seals feeding habits and potential conservation implications. The diet was similar across age groups. Altogether 15 fish species were identified and the most important were smelt (Osmerus eperlanus), ruff (Gymnocephalus cernuus), perch (Perca fluviatilis), vendace (Coregonus albula) and cyprinids. The high δ¹⁵N values suggested that the seals lose weight during winter and spring. Additionally the drop in δ¹⁵N values indicated that pups start to recover from postweaning stress and gain weight around the age of 6 mo. The isotope values differed regionally, which emphasizes that samples from consumers and prey should be collected from the same regions to improve interpretation of the stable isotopic results. Overall, diet composition suggests minimal to nonexistent competition with commercial or recreational fishing. However, observed weight loss of pups during summer may be related to higher risk of bycatch and this should be taken into account when planning temporal fishing closures.     

Remarkable development of diving performance and migrations of hooded seals (Cystophora cristata) during their first year of life

Newborn hooded seals (Cystophora cristata) have smaller weight-specific oxygen stores than adults, but nothing is known about how this affects their diving behaviour. Here, we present data on the diving behaviour and migrations of seven weaned hooded seal pups of the Greenland Sea stock during their first year of life, as collected by use of satellite telemetry. The pups started diving 1–2 days after tagging, and during a tracking period of 25–398 days they dispersed over vast areas of the Greenland and Norwegian Seas in a manner similar to adults. The initial development of diving depths and durations in April–May was rapid, and pups reached depths of >100 m and dived for >15 min within 3 weeks of age. During early summer (May–June) this development was temporarily discontinued, to be resumed throughout autumn and winter, during which time maximum depths and durations of >700 m and >30 min, respectively, were reached. Depths and durations were significantly related to age/season, location and time of day. The dive behaviour in early summer, with relatively shallow and short dives without diurnal variations, resembles that of adults and probably reflects the vertical distribution of prey rather than physiological constraints. Dives of pups were nevertheless shallower and shorter than those of adults, but relative to body mass both hooded seal pups and adults display a remarkable diving capacity which makes the species particularly suited for studies of defence mechanisms against hypoxia insult in mammals.     

Weaning mass affects changes in body composition and food intake in harbour seal pups during the first month of independence

In phocid seals, the transition to nutritional independence is abrupt, with females abandoning their offspring after weaning and returning to sea. We hypothesized that body size at weaning may play an important role in the nature of this transition. We studied the changes in body composition and water flux of newly weaning harbour seals over the first 4-6 wk postweaning. Thirty-three pups were dosed with deuterium oxide to estimate total body water (TBW) and a subset of 24 was dosed twice to estimate changes in body composition and water flux. All pups lost body mass over the study period, but TBW increased during the period of mass loss, indicating continued lean tissue growth. Combined data from this and our early study indicated that heavy (>median mass) pups were relatively fatter (41.0% vs. 37.1%) and had significantly greater total body energy at weaning than did light (< or = median mass) pups. Percentage TBW declined linearly over time in light pups but was constant in heavy pups for the first 19 d postweaning and then declined linearly. Both the temporal pattern and composition of mass loss differed between light and heavy pups. Estimated food intake increased in the second 2 wk of study compared to the first 2 wk, in both light and heavy pups, reflecting increased foraging success but at levels still insufficient to meet daily expenditures of most individuals.     

Effects of age and body mass on development of diving capabilities of gray seal pups: costs and benefits of the postweaning fast

Development of adequate diving capabilities is crucial for survival of seal pups and may depend on age and body size. We tracked the diving behavior of 20 gray seal pups during their first 3 mo at sea using satellite relay data loggers. We employed quantile analysis to track upper limits of dive duration and percentage time spent diving, and lower limits of surface intervals. When pups first left the breeding colony, extreme (ninety-fifth percentile) dive duration and percentage time spent diving were positively correlated with age, but not mass, at departure. Extreme dive durations and percentage time spent diving peaked at [Formula: see text] d of age at values comparable with those of adults, but were not sustained. Greater peaks in extreme percentage time spent diving occurred in pups that had higher initial values, were older at their peak, and were heavier at departure. Pups that were smaller and less capable divers when they left the colony improved extreme dive durations and percentage time spent diving more rapidly, once they were at sea. Minimum survival time correlated positively with departure mass. Pups that were heavier at weaning thus benefitted from being both larger and older at departure, but smaller pups faced a trade-off. While age at departure had a positive effect on early dive performance, departure mass impacted on peak percentage time spent diving and longer-term survival. We speculate that once small pups have attained a minimum degree of physiological development to support diving, they would benefit by leaving the colony when younger but larger to maximize limited fuel reserves, rather than undergoing further maturation on land away from potential food resources, because poor divers may be able to "catch up" once at sea.     

The contrasting hidden consequences of parasitism: Effects of a hematophagous nematode (Uncinaria sp.) in the development of a marine mammal swimming behavior

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Harbor seal pup dispersal and individual morphology,hematology, and contaminant factors affecting survival

Understanding the factors affecting individual harbor seal (Phoca vitulina) survival is essential for determining population level health risks. We estimated postweaning dispersal, and modeled the effects of morphology, hematology, and blubber contaminants on the survival of recently weaned harbor seal pups using a mark recapture framework. We deployed satellite transmitters on apparently healthy pups captured in San Francisco Bay (SFB, n = 19) and Tomales Bay (TB, n = 7), and pups released after rehabilitation that stranded along the central California coast preweaning (n = 21). Dispersal distances were further than previously reported for harbor seal pups (maximum = 802 km) which has implications for understanding risks to this vulnerable age class. We found differences in body condition, serum immunoglobulin and thyroxine (T4) concentrations, white blood cell count, and blubber organohalogen contamination (OH) among the three groups. Overall, increased T4, decreased OH, and increased mass were associated with greater survival probabilities; whereas, among stranded seals, greater mass gain, shorter time in rehabilitation, and admission to rehabilitation earlier in the season were associated with greater survival probabilities. Attention to these latter factors may improve the success of rehabilitation efforts. For wild pups, reduction of legacy contaminants and direct causes of mortality, such as ship strike, may enhance pup survival.     

Body condition at weaning affects the duration of the postweaning fast in gray seal pups (Halichoerus grypus)

Gray seals (Halichoerus grypus) undergo a terrestrial postweaning fast (PWF) that depletes energy reserves acquired during the suckling interval. Plasticity in PWF duration may ensure that pups of variable body condition depart for sea with adequate energy reserves. To test this hypothesis, we examined body condition of 30 gray seal pups at weaning and monitored their PWF duration. On average, fat accounted for 47.3% +/- 0.7% of their 53.2 +/- 1.3-kg weaning mass. Although fasting duration averaged 21 +/- 1.1 d (n = p28), there was considerable variation in fasting duration (9 to > 31 d) and the resulting age when pups departed to sea (26 to > 49 d). Percent fat at weaning(38.6%-54.6%) was positively correlated with fasting duration(n = 28, r = 0.376, P = 0.0489). In contrast, total body gross energy (735.3-1,447.4 MJ) and body mass (39.0-66.0 kg) were not correlated with fasting duration. Thus, body composition,not overall body reserves, predicted fasting duration, but the effect was weak, indicating that other factors also account for the observed variation in fasting duration. We speculate that pups with greater percent fat more effectively utilized lipid and conserved protein while meeting metabolic costs throughout the PWF. As a result, fatter pups extended the PWF duration,which may be critical for development of diving physiology and may have facilitated their survivorship to age 1.     

Diving Related Changes in the Blood Oxygen Stores of Rehabilitating Harbor Seal Pups (Phoca vitulina)

Harbor seal (Phoca vitulina) pups begin diving within hours of birth, stimulating the development of the blood oxygen (O₂) stores necessary to sustain underwater aerobic metabolism. Since harbor seals experience a brief nursing period, the early-life development of these blood O₂ stores is necessary for successful post-weaning foraging. If mothers and pups become prematurely separated, the pup may be transported to a wildlife rehabilitation center for care. Previous studies suggest that the shallow pools and lack of diving in rehabilitation facilities may lead to under-developed blood O₂ stores, but diving behavior during rehabilitation has not been investigated. This study aimed to simultaneously study the diving behaviors and blood O₂ store development of rehabilitating harbor seal pups. Standard hematology measurements (Hct, Hb, RBC, MCV, MCH, MCHC) were taken to investigate O₂ storage capacity and pups were equipped with time-depth recorders to investigate natural diving behavior while in rehabilitation. Linear mixed models of the data indicate that all measured blood parameters changed with age; however, when compared to literature values for wild harbor seal pups, rehabilitating pups have smaller red blood cells (RBCs) that can store less hemoglobin (Hb) and subsequently, less O₂, potentially limiting their diving capabilities. Wild pups completed longer dives at younger ages (maximum reported <25 days of age: 9 min) in previous studies than the captive pups in this study (maximum <25 days of age: 2.86 min). However, captivity may only affect the rate of development, as long duration dives were observed (maximum during rehabilitation: 13.6 min at 89 days of age). Further, this study suggests that there may be a positive relationship between RBC size and the frequency of long duration dives. Thus, rehabilitating harbor seal pups should be encouraged to make frequent, long duration dives to prepare themselves for post-release foraging.     

Absence of sexual differences in aggregation behavior and association patterns in South American sea lion pups (Otaria flavescens) at Isla de Lobos,Uruguay

In the South American sea lion (Otaria flavescens), a polygynous species with a high degree of sexual dimorphism, pups form groups during maternal foraging trips. The aim of this study was to investigate the existence of sexual differences in aggregation behavior and association patterns in O. flavescens pups at Isla de Lobos, Uruguay, during the first 2 mo of life. Scan samplings of marked pups were conducted every hour from 0700 to 1800 to register behavior. Association patterns were analyzed in SOCPROG (using the Half-Weight Index of Association). We did not find significant sex differences in any variable (time spent in groups, average group size, time interacting with other pups, and time in groups at sea). Each pup showed preference for at least one partner. Associations between individuals of the same and different sex were not significantly different. As expected, during the first month, pups associated more strongly with pups born in the same zone than with those born in a different zone. This research provides new evidence on the development of social behavior in otariids and serves as a basis for future studies focusing on sexual differences in pup behavior and association patterns among individuals (e.g., related with kinship).     

Mass changes of grey seal Halichoerus grypus pups on Sable Island: differential maternal investment reconsidered

Theory predicts that where variance in reproductive success differs between sexes, as in many sexually dimorphic species, mothers should invest more in male than in female offspring. In pinnipeds, parental investment is the sole responsibility of the mother. Both size at birth and subsequent mass gain during lactation are measures of that investment. We studied mass gain of grey seal pups, a sexually dimorphic species, during lactation and mass loss during the first 13 days of the postweaning fast using longitudinal data obtained on Sable Island, Canada. A mixed-effects repeated measures model was used to account for the serial correlation in these longitudinal data. We found no significant difference in the birth mass of male (16.6 ± 0.45 kg) and female (15.6 ±0.74 kg) pups. Rate of mass gain by males (3.0 ± 0.13 kg/d) and females (2.7 ± 0.12 kg/d) also did not differ significantly during the lactation period. Grey seal pups on Sable Island tended to be larger at birth and gain mass more rapidly (70%–100%) during lactation than those in the United Kingdom. Mass loss of weaned pups was curvilinear over time. Over the first 10 days of the fast, pups lost 22% of their weaning mass. In both males and females, large pups lost mass more slowly than small ones during the postweaning fast. Based on our results and a review of previous studies, we found little evidence of differential maternal investment in male and female grey seal pups.     

Diving behavior in a free‐living,semi‐aquatic herbivore,the Eurasian beaver Castor fiber

Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.     

The influence of temperature on diving behaviour in the alpine newt, Triturus alpestris

An aquatic lifestyle poses serious restriction to air-breathing animals in terms of time and energy spent during a dive cycle. The diving frequency increases with water temperature, therefore an ectotherm's time budget greatly depends on the thermal characteristics of the aquatic environment. Available data suggests that time costs caused by temperature-dependent dive frequency can be partially compensated for by adjusting the swimming speed and diving angle during dive cycle. We tested this prediction by examining the influence of temperature on the diving behaviour of the alpine newt, Triturus alpestris. The ascending speed and angle showed disparate patterns of temperature dependency, with a minor influence on travel duration. Surprisingly, at higher temperatures, the diving newts saved most of their time by restricting swimming activity in the water column during their return to the bottom and not by adjusting their ascending duration. Hence, aquatic newts have the capacity to reduce temperature-dependent time costs of aerial breathing primarily by behavioural modifications during the descending phase of the dive cycle.     

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.     

The physiological and behavioural development of diving in Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>) pups

The physiological and behavioural development of diving was examined in Australian fur seal (Arctocephalus pusillus doriferus) pups to assess whether animals at weaning are capable of exploiting the same resources as adult females. Haematocrit, haemoglobin and myoglobin contents all increased throughout pup development though total body oxygen stores reached only 71% of adult female levels just prior to weaning. Oxygen storage components, however, did not develop at the same pace. Whereas blood oxygen stores had reached adult female levels by 9 months of age, muscle oxygen stores were slower to develop, reaching only 23% of adult levels by this age. Increases in diving behaviour corresponded to the physiological changes observed. Pups spent little time (<8%) in the water prior to moulting (age 1–2 months) whereas following the moult, they spent >27% of time in the water and made mid-water dives (maximum depth 35.7 ± 2.9 m) with durations of 0.35 ± 0.03 min. By 9 months (just prior to weaning), 30.5 ± 9.3% of all dives performed were U-shaped benthic dives (maximum depth 65.0 ± 6.0 m) with mean durations of 0.87 ± 0.25 min, significantly shorter than those of adult females. These results suggest that while Australian fur seal pups approaching the age of weaning are able to reach similar depths as adult females, they do not have the physiological capacity to remain at these depths for sufficient durations to exploit them to the same efficiency.     

Age-related changes in hematocrit in the Galápagos sea lion (Zalophus wollebaeki) and the Weddell seal (Leptonychotes weddellii)

In mammals, hematocrit (Hct) is optimized between the divergent requirements of blood flow characteristics and oxygen transport and storage capacity. This trade‐off plays a particularly major role in marine mammals, in which oxygen demand during sustained diving requires high Hct levels. Galápagos sea lions (Zalophus wollebaeki) need a long time after birth to develop from terrestrial life to the state of an independent forager at sea. We here show that pups were born with high Hct of 45%, then reduced Hct during the first 40 d of life to 31% while they remain constantly ashore, and increased Hct again until the adult level (57%) is reached at 1 yr of age when they begin to dive for foraging at sea. A similar, but much more rapidly changing pattern is seen in Weddell seal pups, but not in northern elephant seals, where no reduction in Hct is seen after birth. These and similarly changing patterns in terrestrial mammals likely reflect species specific functional adjustments during development due to a trade‐off between the costs of circulation and the changing need to store and transport oxygen.     

Ontogeny of Time Allocation in Black-tailed Prairie Dogs

The ontogeny of time allocation was studied in a field population of black-tailed prairie dogs, Cynomys ludovicianus. Pups first emerged from their natal burrows in late May. All individuals in the population, including all pups, were observed for 12 weeks following this emergence. Both cross-sectional and longitudinal analyses of ontogeny revealed considerable age differences. Upon first emergence, pups were extremely wary and spent most of their time vigilant and little time feeding. As pups aged, they increased time spent feeding and decreased time spent vigilant. Male and female pups behaved similarly. Pups differed from adults in their time allocation patterns and these differences changed as pups aged. Pups were initially more vigilant and fed less than adults, but became less vigilant than adults as the summer progressed. Yearlings behaved similarly to adults. Possible intrinsic (e.g., pup age and weight) and extrinsic (e.g., weather conditions, microhabitat location and coterie composition) influences on time allocation by pups were also examined. In general, extrinsic influences appeared to have more impact on pup behavior than intrinsic ones, suggesting that time allocation by pups may be largely context-dependent. Overall, the considerable number of age differences argues for more consideration of ontogeny in models of antipredator behavior.