Acquisition of gliding skills by Weddell seal (Leptonychotes weddellii) pups during lactation

Pinnipeds give birth to their pups ashore or on ice and forage in water. Therefore, neonates initially lack the adaptations to sustain prolonged underwater diving activity. Although the physiological development for breath-holding during diving has been investigated in seal pups, little is known about the concurrent development of behavioral adaptations during lactation. In this study, multisensor data loggers were used to record diving behavior and swimming gaits of pre-weaned Weddell seal (Leptonychotes weddellii) pups. Experiments were conducted in 16 pups at the Syowa Station, Antarctica, from November to December 1999 and 2004. Swimming speeds, dive depths and flipper stroking rates were recorded for each individual for about 24 h. We found that the glide index during ascending was increased with body length, whereas the dominant stroke cycle frequency were not affected by body length, dive depth and descent or ascent phase. All pups had significantly higher stroke rates in descent than in ascent, but there was no difference between swimming speed. As we found a positive relationship between the body length and age, we considered body length as an index of growth. Therefore, we conclude that pups gradually acquire the ability to glide with utilizing positive buoyancy during ascending toward the end of lactation.     

Microscopic anatomy of the eye of the deep-diving Antarctic Weddell seal (Leptonychotes weddellii)

The microscopic anatomy of the eye of the Weddell seal was studied with various light and electron microscopic methods with a view to correlating morphological findings with the biology of this seal which is adapted to the extremes of the Antarctic environment and to extreme diving excursions into the lightless depths of the sea. In the retina an area centralis was found but no fovea centralis. The densely packed photoreceptors consist exclusively of highly differentiated rods, which in primates detect light at low intensity but have rather poor image discrimination. The ganglion cells are relatively scarce, suggesting a high degree of convergence of the light-sensitive cells on the ganglion cells. The pigment epithelium is almost devoid of pigment granules. The extensive tapetum lucidum is about 400-500 microm thick and is composed of about 30 layers of specialized cells. The cornea is 650 (center) to 800-900 (periphery) microm thick. Its structure and glycosaminoglycan histochemistry correspond to that of other mammals. The iridocorneal angle is unusually deep and pervaded by an elaborate trabecular meshwork, which together with a complex canal of Schlemm can be correlated with the ability to absorb large amounts of fluid. The ciliary muscle and its antagonist, the membrane of Bruch, are poorly developed, suggesting relatively poor abilities of accommodation. The combination of a well-developed tapetum lucidum, an unpigmented pigment epithelium, well-developed rods, and a high number of rods converging on only few ganglion cells is obviously an adaptation to an extreme light sensitivity, enabling the animals to make use of the little light available in the deep sea.     

Microgeographic variation and songs in the underwater vocalisation repertoire of the Weddell seal (Leptonychotes weddellii) from the Vestfold Hills,Antarctica

Recordings of underwater vocalisations of the Weddell seal (Leptonychotes weddellii) were made in fjords of the Vestfold Hills near Davis Station, Antarctica. The repertoire was examined for the presence of microgeographic variation. Recordings covered the period from the end of mating to the beginning of the moult in the 1989/1990 breeding season. The repertoire was classified into 44 vocalisation types using a classification method based on objective parameters. Unique vocalisations were identified at each site, with 48% exclusive to Long Fjord and 41% to Tryne Fjord. A small percentage (11%) of the repertoire was common to both fjords reflecting the limited, observed interchange of individuals between these populations due to their strong breeding site fidelity. Seven male songs are described in the present study and possibly function to maintain underwater territories.     

Physiological and behavioral determinants of the aerobic dive limit in Weddell seal (Leptonychotes weddellii) Pups

The aerobic dive limit, as defined by an increase in plasma lactate levels following dives, has to date only been determined in adult and juvenile Weddell seals (Leptonychotes weddellii). However, theoretical aerobic dive limits based on calculated total body oxygen stores, estimated metabolic rates, and dive duration frequencies have been published for several species. Using data collected over the past 3 years in McMurdo Sound. Antarctica, the aerobic dive limit of Weddell seal pups was determined by both the physiological and modeling methods. Time-depth diving recorders deployed on 36 pups between 2 and 14 weeks of age allowed the aerobic dive limit to be predicted from duration-frequency histograms. The aerobic dive limit was also calculated from estimates of total body oxygen stores and predicted diving metabolic rates. Finally, these two estimates were compared with aerobic dive limits determined from post-dive lactate levels in three pups between 5 and 7 weeks old. The aerobic dive limits of pups increased with age, but pup aerobic dive limits were still significantly shorter than those of yearlings and adults. In addition, the aerobic dive limits determined by the three methods were not equivalent for pups, yearlings, or adults, and indicate that care should be taken when modeling methods are used to estimate the aerobic dive limit in other species. Changes in hematocrit, plasma glucose, and plasma lactate levels during and between rest, diving, and recovery in pups were compared to known values for juveniles and adults. Plasma metabolite levels were more highly regulated in older pups, and together with the increasing aerobic dive limit, suggest that Weddell seal pups are not refined divers until after they are weaned, and that their diving ability continues to develop over several years.     

Histological,histochemical, and ultrastructural investigations on the gastrointestinal system of antarctic seals: Weddell seal (Leptonychotes weddellii) and crabeater seal (Lobodon carcinophagus)

The morphology of the principal sections of the gastrointestinal system of two Antarctic seals with different dietary habits, namely, the Weddell seal (Leptonychotes weddellii) and the crabeater seal (Lobodon carcinophagus), has been investigated. Histologically examined by light microscopy, the tissue layers of the gastrointestinal tract of both seals are almost identical to those observed in most other mammals and no major differences in principle organization could be found between the two seal species. The ultrastructure of the gastric and intestinal epithelial cells has been examined and is also closely comparable to that of these cells in other mammals; however, Paneth cells have not been found in our material. In general, therefore, adaptations of the gastrointestinal tract to the aquatic environment or the diet are not obvious at the morphological levels of organization studied. Histochemical differences are found between the two closely related species; mucins of the surface epithelium in the stomach of Weddell seals are highly sulfated, while those in the crabeater seal are not. Mucous neck cells in Weddell seals contain acid mucosubstances, while those of crabeater seals contain neutral ones. Goblet cells in the small and large intestine in Weddell seals contain both neutral and acid mucosubstances. Both mucin types are detected in the crabeater seal; however, the mucins of the colon in the crabeater seal are more highly sulfated than those in the Weddell seal. The ratio of globet cells to enterocytes in the large intestine of crabeater seals is higher than that in Weddell seals. © 1995 Wiley-Liss, Inc.     

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.     

Chemical restraint of Weddell seals (Leptonychotes weddellii) with a combination of tiletamine and zolazepam.

A 1:1 combination by weight of tiletamine hydrochloride and zolazepam hydrochloride was administered to 30 adult Weddell seals (Leptonychotes weddellii) in doses varying from 100 to 300 mg. Full immobilization was achieved in 16 seals, moderate sedation in seven and light sedation in seven. Three animals died; two were fully immobilized and one was moderately sedated prior to death. The drug combination was considered satisfactory, although its usefulness was limited by the lack of chemical antagonists when complications were encountered in immobilized animals.     

Sex‐related differences in the postmolt distribution of Weddell seals (Leptonychotes weddellii) in the southern Weddell Sea

The population of Weddell seals (Leptonychotes weddellii) in the southern Weddell Sea is in a unique position on the continental shelf edge, with vast shelf waters to the south, and deep Southern Ocean to the north. We describe sex‐related differences in the winter distribution of this population, from data collected by 20 conductivity‐temperature‐depth satellite relay data loggers deployed in February 2011 at the end of the annual molt. The regional daily speed was calculated, and a state‐space model was used to estimate behavioral states to positions along individuals’ tracks. GLMMs estimated that males and smaller individuals, diving in shallower water, traveled less far per day of deployment (males 14.6 ± 2.26 km/d, females 18.9 ± 2.42 km/d), and males were estimated to dive in shallower water (males 604 ± 382 m, females 1,875 ± 1,458 m). Males and smaller individuals were also estimated to be more resident; males spent an average 83.4% ± 7.7% of their time in a resident behavioral state, compared to females at 74.1% ± 7.1%. This evidence that male and female Weddell seals in the southern Weddell Sea are adopting different strategies has not been shown elsewhere along their circumpolar distribution.     

Hydrographic influences on the summer dive behaviour of Weddell seals (Leptonychotes weddellii) in Atka Bay, Antarctica

In order to gain insights into species-level behavioural responses to the physical environment, it is necessary to obtain information from various populations and at all times of year. We analysed the influences of physical environmental parameters on the mid-summer dive behaviour of Weddell seals (Leptonychotes weddellii) from a little-known population at Atka Bay, Antarctica. Dive depth distributions followed a typical bimodal pattern also exhibited by seals from other populations and seals targeted both shallow water layers of <50 m and depths near the seafloor. Increased stratification of temperature layers within the water column resulted in increased forage efforts by the seals through relatively high numbers of dives to the seafloor, as well as forage effort associated with shallow dives. We interpret these behavioural responses to be due to increased water temperature stratification resulting in the concentration of prey species in particular depth layers.     

A theoretical analysis of diving performance in the Weddell seal (Leptonychotes weddelli)

Marine mammals are constrained in their foraging behaviour because, as obligate air breathers, they must undertake regular trips to the water surface to satisfy the need for respiratory gas exchange. Maximum underwater endurance time is determined by O2 supply and demand, but this does not necessarily imply that O2 is the main factor regulating individual dive and surface times. This study presents a theoretical analysis of diving performance that emphasizes a key role for CO2 in the proximate control of diving behaviour. Computer simulations, based on a mathematical model of the mammalian cardiorespiratory control system, are used to investigate the influence of swimming to depth and other energetic stresses (feeding, thermogenesis, sleep) on predicted diving behaviour in an average adult Weddell seal. The plausibility of the proposed model is supported by the study, which replicated published observations of natural diving behaviour in this species. It is suggested that diving behaviour is tuned to oscillations in respiratory drive and that behavioural and physiological factors can alter the dynamic characteristics of the system to achieve a highly adaptable reciprocal interaction that blurs the boundary between physiology and behaviour.     

Individual variation in pup vocalizations and absence of behavioral signs of maternal vocal recognition in Weddell seals (Leptonychotes weddellii)

Individually stereotyped vocalizations often play an important role in relocation of offspring in gregarious breeders. In phocids, mothers often alternate between foraging at sea and attending their pup. Pup calls are individually distinctive in various phocid species. However, experimental evidence for maternal recognition is rare. In this study, we recorded Weddell seal (Leptonychotes weddellii) pup vocalizations at two whelping patches in Atka Bay, Antarctica, and explored individual vocal variation based on eight vocal parameters. Overall, 58% of calls were correctly classified according to individual. For males (n= 12) and females (n= 9), respectively, nine and seven individuals were correctly identified based on vocal parameters. To investigate whether mothers respond differently to calls of familiar vs. unfamiliar pups, we conducted playback experiments with 21 mothers. Maternal responses did not differ between playbacks of own, familiar, and unfamiliar pup calls. We suggest that Weddell seal pup calls may need to contain only a critical amount of individually distinct information because mothers and pups use a combination of sensory modalities for identification. However, it cannot be excluded that pup developmental factors and differing environmental factors between colonies affect pup acoustic behavior and the role of acoustic cues in the relocation process.     

Satellite imagery can be used to detect variation in abundance of Weddell seals (Leptonychotes weddellii) in Erebus Bay, Antarctica

The Weddell seal population in Erebus Bay, Antarctica, represents one of the best-studied marine mammal populations in the world, providing an ideal test for the efficacy of satellite imagery to inform about seal abundance and population trends. Using high-resolution (0.6 m) satellite imagery, we compared counts from imagery to ground counts of adult Weddell seals and determined temporal trends in Erebus Bay during November 2004–2006 and 2009, and December 2007. Seals were counted from QuickBird-2 and WorldView-1 images, and these counts were compared with ground counts at overlapping locations within Erebus Bay during the same time. Counts were compared across years and within individual haul-out locations. We counted a total of 1,000 adult Weddell seals from five images across all years (for a total of 21 satellite-to-ground count comparisons), approximately 72% of the total counted on the ground at overlapping locations. We accurately detected an increase in abundance during 2004–2009. There was a strong, positive correlation (r = 0.98, df = 3, P < 0.003) between ground counts and counts derived from the imagery. The correlation between counts at individual haul-out locations was also strong (r = 0.80, df = 19, P < 0.001). Detection rates ranged from 30 to 88%. Overall, our results showed the utility of high-resolution imagery to provide an accurate way to detect the presence and variation in abundance of Weddell seals. Our methods may be applied to other species in polar regions, such as walruses or polar bears, particularly in areas where little is known about population status.     

Geographic variation of Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>) airborne mother–pup vocalisations

Geographic variation of vocal behaviour may be reflective of genetic, cultural and ecological differences between locations. Such differences have been previously reported in six pinnipeds, including Weddell seals (Leptonychotes weddellii). This study investigated whether the acoustic characteristics of airborne Weddell seal mother and pup ‘contact’ calls vary geographically between the Vestfold Hills and McMurdo Sound. From each location, 72 mother and 120 pup calls were analysed, with 11 acoustic characteristics measured. MANOVA and Discriminant function analysis (DFA) identified significant differences in several fundamental frequency characteristics. The cross-validated DFA classified greater than 60% of mother and pup vocalisations to the correct location. For novel calls, the probabilities of correctly classifying these percentages by chance are low. Additionally, ordinal logistic regression identified significant differences in the maximum energy distribution of mother and pup vocalisations. For pups, this variation is likely the result of genetic differences between the two populations. Nevertheless, results support suggestions that, when compared with underwater repertoire differences, airborne within-call differences are less efficient at identifying Weddell seal breeding populations.     

Architecture and functional aspects of the distal airways of Antarctic seals with different habits,the Weddell Seal (Leptonychotes weddellii) and the Crabeater Seal (Lobodon carcinophagus)

Summary The lung of the deep diving Weddell Seal is characterized by an unusually well developed periacinar dense collagenous connective tissue, and a thick coat of smooth musculature particularly in the distal bronchioli. Both, collagen and smooth musculature appear to be functionally interrelated, the first serving presumably as site of origin or attachment for the latter. The orientation of the bronchiolar smooth muscle cells is complex: there exists a basic pattern of two crisscrossing helical bundles that wind in opposite direction. In addition, longitudinal bundles are frequent both at the inside and the outside of the muscular coat. Furthermore, more or less complete ringshaped bundles occur as well as groups of muscle fibres running radially into the collagenous tissue of the surroundings of a bronchiolus. This complex architecture presumably allows active adjustment to various physiological needs of the Weddell Seal including as extremes both closing and widening of the bronchiolar lumen. Isometric contractions of the smooth musculature may stiffen the wall of the distal airways while diving. In the Crabeater Seal which dives for shorter durations and by far less deeply than the Weddell Seal, both periacinar collagen and bronchiolar smooth musculature are of similar arrangement, however, occur in considerably reduced amounts. A rich supply of autonomie nerve fibres with abundant varicosities controls the smooth muscle cells, which are interconnected by gap junctions and receive their innervation par distance (visceral type of smooth musculature). The majority of varicosities contains small clear vesicles, as is typical for cholinergic nerves, suggesting a strong parasympathetic influence. Other varicosities are presumably of peptidergic type. Mast cells and epithelial endocrine cells may exert additional influence on the musculature.     

Variation of annual apparent survival and detection rates with age,year and individual identity in male Weddell seals (Leptonychotes weddellii) from long-term mark-recapture data

Exploring age- and sex-specific survival rates provides insight regarding population behavior and life-history trait evolution. However, our understanding of how age-specific patterns of survival, including actuarial senescence, compare between the sexes remains inadequate. Using 36 years of mark-recapture data for 7,516 male Weddell seals (Leptonychotes weddellii) born in Erebus Bay, Antarctica, we estimated age-specific annual survival rates using a hierarchical model for mark-recapture data in a Bayesian framework. Our male survival estimates were moderate for pups and yearlings, highest for 2-year-olds, and gradually declined with age thereafter such that the oldest animals observed had the lowest rates of any age. Reports of senescence in other wildlife populations of species with similar longevity occurred at older ages than those presented here. When compared to recently published estimates for reproductive Weddell seal females, we found that peak survival rates were similar (males: 0.94, 95% CI = 0.92–0.96; females: 0.92, 95% CI = 0.93–0.95), but survival rates at older ages were lower in males. Age-specific male Weddell seal survival rates varied across years and individuals, with greater variation occurring across years. Similar studies on a broad range of species are needed to contextualize these results for a better understanding of the variation in senescence patterns between the sexes of the same species, but our study adds information for a marine mammal species to a research topic dominated by avian and ungulate species.