Finescale ecological niche modeling provides evidence that lactating gray seals (Halichoerus grypus) prefer access to fresh water in order to drink

Many phocids are capital breeders, relying on stored reserves to sustain energetic requirements while on land. Their large body size, high energy expenditure during lactation, and the insulative effects of the blubber layer can lead to thermal stress from overheating, especially in warm and temperate climates. Thermal stress can influence fine‐scale site choice on breeding colonies, and behavioral thermoregulation has been proposed as an explanation for the clear preferences shown by breeding female gray seals for proximity to pools of water. However, anecdotal observations suggest that pools of water may also be preferred for drinking, though water intake is difficult to verify without real‐time physiological monitoring. Here, an alternative approach demonstrates that gray seals also require access to water for drinking. Using Ecological Niche Factor Analysis to examine fine‐scale physical determinants of pupping site choice at North Rona, Scotland, we found that lactating mothers showed preference for lower salinity pools. This is most pronounced early in the season, when ambient temperatures and presumably thermal stress are greatest. Given that the cooling effect of fresh and salt water should be equivalent, the most parsimonious explanation for this preference for fresh water pools is that lactating females use these pools for drinking.     

Finescale topographical correlates of behavioural investment in offspring by female grey seals, Halichoerus grypus

Grey seals breed colonially on substrates ranging from ice to rocky or sandy beaches. Clear differences in seal behaviour patterns exist among such broad classes of breeding habitat. However, finer scale topographical variation is likely to influence individual behaviour with consequences for pupping success. We examined topographical influences on the behaviour of breeding female grey seals by quantifying topography at a subseal size resolution. Using submetre resolution digital terrain models of two sites within a rocky breeding colony, we compared site topography in relation to observed differences in female behaviour at these sites. Females at both sites preferred breeding close to water (standing pools or sea) and frequently commuted between their pups and water. Topographical models indicated that one site was more costly for seals in terms of their locations and movements within the site. This was due to a lack of low-elevation land adjacent to the main access points from the sea and the reduced availability of pools. Females at this site showed reduced pup attendance and an increase in energetically costly behaviours, whilst females at the lower-cost site spent more time interacting with their pups and resting. These topographically induced behavioural differences are likely to affect the quantity and quality of pup provisioning by mothers and influence individual pupping site selection. Less costly sites are likely to be colonized preferentially and by larger, older and more dominant females, potentially generating finescale spatial heterogeneity in female quality within the breeding colony. Copyright 2000 The Association for the Study of Animal Behaviour.     

Control and amount of heat dissipation through thermal windows in harbor seals (Phoca vitulina)

Pinnipeds have to cope with the thermoregulatory demands of their amphibious lifestyle. As they are effectively insulated against heat loss in water by their blubber, they have to bypass the blubber for heat dissipation while staying ashore. In previous studies thermal windows on the body of captive phocid seals have been described as areas of heat dissipation in air. In this study we used infrared thermography (IRT) to examine thermal windows in seals hauling out as well as in training situations, where the first refers to a voluntary and the latter to an induced stay on shore. Furthermore we provide an IRT-based estimate of heat loss through thermal windows in pinnipeds. Hauling out seals developed thermal windows within a few minutes irrespective of environmental conditions. By contrast, seals staying on shore on a trainer's command did not develop thermal windows at all. The calculation of heat loss through thermal windows resulted in considerable values in air, but above all in water, which is energetically disadvantageous as it takes up to four minutes to close thermal windows in water.     

Reaction of Harbor Seals to Cruise Ships

Abstract: The largest aggregations of harbor seals (Phoca vitulina) in Alaska, USA, haul out on floating ice in tidewater glacial fjords. Seals use these fjords in peak numbers during the critical periods of pupping, breeding, and molting when visits by tour ships also peak. Documented and suspected declines of harbor seals in fjords with rising vessel traffic underscore the need to better understand possible impacts, particularly in areas where ship visits have risen substantially in the past 2 decades. We examined the interruption of haul-out bouts of harbor seals due to approaching cruise ships in Disenchantment Bay, Alaska. We conducted observations from cruise ships and focused on disturbance of seals as evidenced by seals flushing into the water from the floating ice on which they rested. We investigated rate of flushing in relation to vessel distance, approach angle, group size, and seal type (mother, pup, or other). Using a survival-regression analysis, we found that the risk of disturbing harbor seals increased when ships approached within 500 m; seals approached as close as 100 m were 25 times more likely to enter the water than seals 500 m from a ship. Seals were 4 times more prone to enter the water when ships were approaching directly rather than passing abeam. Seals responded similarly regardless of group size or seal type. Energetic models indicated a potential to disrupt energy balance and cause thermal stress in disturbed pups if they spent >50% of their time in ice-chilled water. Studies at non-glacial sites suggest that pups spend 40–70% of their time in the water. Voluntary guidelines for approaching seals in Alaska recommend that cruise ships approach ≥91 m (100 yards), a distance at which we show 90% of seals would flush into the water. Our findings indicate a need to develop regulations to maintain a 500-m separation between cruise ships and seals in all Alaskan glacial fjords.     

Effects of environmental variables on surface temperature of breeding adult female northern elephant seals,Mirounga angustirostris,and pups

Pinnipeds spend extended periods of time on shore during breeding, and some temperate species retreat to the water if exposed to high ambient temperatures. However, female northern elephant seals (Mirounga angustirostris) with pups generally avoid the water, presumably to minimize risks to pups or male harassment. Little is known about how ambient temperature affects thermoregulation of well insulated females while on shore. We used a thermographic camera to measure surface temperature (T_s) of 100 adult female elephant seals and their pups during the breeding season at Point Reyes National Seashore, yielding 782 thermograms. Environmental variables were measured by an onsite weather station. Environmental variables, especially solar radiation and ambient temperature, were the main determinants of mean and maximum T_s of both females and pups. An average of 16% of the visible surface of both females and pups was used as thermal windows to facilitate heat loss and, for pups, this area increased with solar radiation. Thermal window area of females increased with mean T_s until approximately 26 °C and then declined. The T_s of both age classes were warmer than ambient temperature and had a large thermal gradient with the environment (female mean 11.2±0.2 °C; pup mean 14.2±0.2 °C). This large gradient suggests that circulatory adjustments to bypass blubber layers were sufficient to allow seals to dissipate heat under most environmental conditions. We observed the previously undescribed behavior of females and pups in the water and determined that solar radiation affected this behavior. This may have been possible due to the calm waters at the study site, which reduced the risk of neonates drowning. These results may predict important breeding habitat features for elephant seals as solar radiation and ambient temperatures change in response to changing climate.     

Behaviour of female and pup grey seals Halichoerus grypus during the breeding period at Froan, Norway

We studied the behaviour of grey seals, Halichoerus grypus , during the breeding period at Froan, Norway, and compared our findings with existing studies on grey seals at breeding sites in Britain and Canada. The pups at Froan spent more time in water than pups at other breeding sites. While on-shore, the pups at Froan spent most of their time resting, behaviour similar to pups at other breeding sites. Lactating females at Froan spent most of their time in the sea, thus differing from females at most other breeding sites which spend most of their time on-shore. While in the sea, the females at Froan spent most time diving, typically interrupted by regular periods of surface swimming.     

SEX AND AGE SEGREGATION BY PHOCA VITULINA CONCOLOR AT HAUL-OUT SITES DURING THE BREEDING SEASON IN THE PASSAMAQUODDY BAY REGION, NEW BRUNSWICK

The size and composition of groups of harbor seals at two haul-out sites were studied during the breeding season of 1989, in the Passamaquoddy Bay region of Atlantic Canada. Evidence of segregation both by age and sex was found in the distinct composition of the two groups. One group contained mainly males and no pups, and the other had a sex ratio not significantly different from one and contained pups. The proportion of females increased at the nursery site with the onset of birthing in the region while the proportion of males increased through the breeding season at the other site. No increase in the number of adults, in total, was detected over the study period, suggesting that sexual segregation and not a change in haul-out frequency was responsible for the disparity in the sex structure of the two groups. The proportion of juveniles was significantly greater at the male dominated site than at the nursery site.     

Resource-defense Polygyny and Male Territory Quality in the New Zealand Fur Seal

The polygynous New Zealand fur seal, Arctocephalus forsteri, breeds in densely-packed colonies with males defending exclusive territories. The distribution of individuals within a colony of these seals was monitored over three consecutive breeding seasons and behavioural thermoregulation was found to be an important factor influencing the site selection of both males and females. Shaded areas and pools of water were used by both sexes for cooling and use of these cooling substrates increased as rock surface temperature increased, resulting in a clumped distribution on hot days. Substrate preferences were tested experimentally by manipulating the availability of cooling substrates during two seasons. The number of females using an area increased significantly after shade or pools of water were added to that area. For males, pool additions resulted in increased use of those areas, while adding shade had no effect. Cooling substrates had a patchy distribution and could therefore be monopolised by territorial males. Using the number of females on a territory as a measure of male mating success, it was found that female numbers were most strongly positively correlated with the area of shaded substrate on a territory. Total territory area was also positively correlated with number of females. It is argued that thermoregulatory constraints are a major factor affecting female site choice, and that this in turn affects male mating success.     

Philopatry, Site Fidelity and Local Kin Associations within Grey Seal Breeding Colonies

Abstract Colonially breeding animals such as grey seals typically have a discrete number of breeding sites. The extent to which offspring are philopatric (return to breed at their natal site) and the prevalence of philopatry by sex of pup has fundamental consequences for the social and genetic structure within and between each colony.
Grey seals born and marked by tagging and cohort brands at colonies on North Rona, Outer Hebrides since 1960 and at the Isle of May, Firth of Forth since 1981 have shown philopatry. Overall re-sight rates of 1620 pups tagged at North Rona and 1667 pups at the Isle of May ranged from 0 to 17%. Although most evidence of philopatry relates to females, males at each colony displayed philopatry. In addition, females born at N. Rona pupped closer to their natal sites than would be expected by chance (p = 0.005), providing evidence of fine scale natal site fidelity. Females born at the Isle of May did not show the same degree of fine scale natal site fidelity, but there was considerable individual variation. The spatial scale of philopatry shown by seals differed according to the local area within each island. Fidelity of known adult females to pupping sites at both colonies was high. In addition, we found an instance of mother and daughter pupping together away from the daughter's natal site.
These data: (i) provide evidence of philopatry in both sexes at each site; (ii) describe occurrences of fine scale philopatry at both sites; (iii) show that mothers and offspring occupied the same areas on the colony, suggesting the possibility of highly related groups within colonies, which would provide conditions for kin-specific altruistic behaviours; (iv) suggest the possibility of individual or kin recognition, even though contact between mothers and offspring has been thought to cease after the 18 d lactation period.     

Factors influencing and consequences of breeding dispersal and habitat choice in female grey seals (<Emphasis Type="Italic">Halichoerus grypus</Emphasis>) on Sable Island,Nova Scotia

Selection of breeding location can influence reproductive success and fitness. Breeding dispersal links habitat use and reproduction. This study investigated factors affecting breeding dispersal and its reproductive consequences in grey seals (Halichoerus grypus) on Sable Island, Nova Scotia. Breeding dispersal distance was determined in 692 individually marked, known-age female grey seals observed from 2004 to 2014. We used generalized linear mixed-effects models to test hypotheses concerning environmental and demographic factors influencing breeding dispersal distance and the consequences of dispersal distance on offspring weaning mass. Grey seal females rarely exhibited fidelity to previous breeding sites. Median dispersal distance between years was 5.1 km. Only 2.9% of females returned to a previous breeding site. Breeding dispersal distance was affected by parity and density, but effects were small and are presumably of no biological significance. Variation in dispersal distance among adult females was large. Dispersal distance had no significant influence on offspring weaning mass; however, as previously found, pup sex and maternal age did. Although breeding location was not important, heavier pups were born in habitats with no tidal or storm-surge influence indicating that breeding habitat type did influence offspring size at weaning. The lack of site fidelity in grey seals on Sable Island is associated with an unpredictable and changing landscape (sand dunes) that could make it difficult for females to locate previous breeding locations. Although breeding location within habitat type had small consequences on offspring weaning mass, we detected no evidence that breeding site selection within the habitat had consequences to females.     

Characterization of Australian fur seal vocalizations during the breeding season

The vocal repertoire, structure, and behavioral context of airborne vocalizations produced by Australian fur seals (Arctocephalus pusillus doriferus) are described using recordings made at a breeding colony on Kanowna Island, Bass Strait, Australia. The study identified six different call types: three produced by males (bark, guttural threat, and submissive call); five produced by females (bark, guttural threat, submissive call, growl, and pup attraction call) and the female attraction call produced by pups and yearlings. Vocalizations were compared according to age and sex classes. The overall structure and function of the pup attraction and female attraction call produced by females, yearlings, and pups, was similar. However, while similar in their overall appearance, certain call types have a lower fundamental frequency when compared with other fur seals. In addition, the male bark call alters in rate of production according to the context used, where calls are slower when males are stationary and advertising their territorial status and faster when males are involved in confrontations with other males or actively herding females. Further research is required to investigate changes in environmental conditions and their effects on shaping the call structure and communication in Australian fur seals.     

Spatial variation in age-specific probabilities of first reproduction for Weddell seals

Spatial variation in vital rates can affect the dynamics and persistence of a population. We evaluated the prediction that age-specific probabilities of survival and first reproduction for Weddell seals would vary as a function of birth location in Erebus Bay, Antarctica. We used multi-state mark–resight models and 25 years of data to estimate demographic rates for female seals. We predicted that probabilities of survival and first reproduction would be higher for seals born at near-shore colonies or more southerly-located colonies with consistent ice conditions. Contrary to predictions, results revealed higher age-specific probabilities of first reproduction at offshore colonies relative to near-shore colonies and no spatial variation in survival rates. For 7-year old females (average age at 1st reproduction=7.6 years old) born at offshore colonies to mothers aged 10.8 years (average maternal age), probability of first reproduction was 0.43 (SE=0.07), whereas probability of first reproduction for females born at near-shore colonies was 0.30 (SE=0.05) based on estimates from our top-ranked model. Breeding probabilities following first reproduction were also higher at offshore colonies. Thus, our results (1) provide evidence of spatial variation in breeding probabilities, (2) reveal the importance of birth location on a female's vital rates, and (3) suggest that the effect persisted for many years. Birth-colony effects may be attributed to spatial variation in prey availability, or to heterogeneity in female quality in this population. If females who are superior competitors consistently chose offshore colonies for pupping, pups born at these locations may have inherited those superior qualities and displayed higher probabilities of first reproduction, relative to seals born at other colonies. Further research into physical or food-related differences among colonies may offer insight into spatial variation in breeding probabilities documented in this paper.     

Influences of habitat features and human disturbance on use of breeding sites by a declining population of southern fur seals (Arctocephalus australis)

Southern fur seals Arctocephalus australis in Peru have declined gradually over the past decade, and declined dramatically (72%) as a result of low food availability during the severe El Niño in 1997–98. In 1999, seals abandoned some historically important breeding sites. This is particularly alarming because new sites were not colonized. Our objective was to examine how habitat features and human disturbance influenced whether sites were currently used, abandoned or apparently not used in the past by fur seals for breeding. Data were collected on 14 variables at 70 potential breeding sites at three guano reserves in Peru. Discriminant analysis revealed significant multivariate differences among sites currently used for breeding, abandoned sites and unused sites ( F =5.97, P <0.00001), and the model classified 74% of sites correctly. Currently used sites were less likely to have human disturbance and more likely to have offshore islands, stacked rocks, tide pools and abundant shade. Separate discriminant analyses for each reserve produced similar results. Habitat associated with thermoregulation (e.g. shade or pools) may be more important to fur seals in Peru, which breed at lower latitudes and are at greater risk of overheating on land than other populations. Habitat with minimized human access may be especially important to seals in small populations in which individuals may perceive themselves as more vulnerable because of decreased vigilance and dilution effects. Seals in our study selected breeding habitat with stacked rocks, which create shade and tide pools for thermoregulation and make human access difficult; but pups might suffer higher mortality in this habitat. We hypothesize that fur seals in Peru may exhibit an Allee effect, whereby suitability of habitat varies with population abundance.     

An in‐field experimental test of pinniped behavioral types

There has been extensive recent interest in the concepts of behavioral types, behavioral syndromes, and personalities in nonhuman animal species. Evidence for behavioral types now exists from a wide range of taxa, from mollusks to mammals. However, marine mammals are poorly represented in this literature. Here, we describe an in‐field experimental test of behavioral types in breeding gray seals, using a remotely controlled vehicle to deliver a standardized test stimulus to target individuals. We report on the design and implementation of this test and on the behavioral responses of individuals. Analysis of behavioral responses from both males and females revealed consistent individual differences across tests, suggesting that this is a practical and viable technique for determining individual variation in behavioral type in the field. Despite extensive literature on behavioral types, studies of behavioral types in wild populations remain rare. It is, therefore, important to develop ways to identify and quantify the existence of behavioral types in natural populations, because only by doing this, can we hope to ascertain the ecological and evolutionary relevance of behavioral types.     

Successful foraging zones of southern elephant seals from the Kerguelen Islands in relation to oceanographic conditions

Southern elephant seals, Mirounga leonina, undertake large-scale oceanic movements to access favourable foraging areas. Successful foraging areas of elephant seals from the Kerguelen Islands are investigated here in relation to oceanographic parameters. Movements and diving activity of the seals as well as oceanographic data were collected through a new generation of satellite relayed devices measuring and transmitting locations, pressure, temperature and salinity. For the first time, we have associated foraging behaviour, determined by high increased sinuosity in tracks, and dive density (i.e. number of dives performed per kilometre covered), and changes in body condition, determined by variations in drift rate obtained from drift dives, to identify the oceanographic conditions of successful foraging zones for this species. Two main sectors, one close to the Antarctic continent and the other along the Polar Front (PF), where both foraging activity and body condition increase, seem to be of particular interest for the seals. Within these regions, some seals tended to focus their foraging activity on zones with particular temperature signatures. Along the Antarctic continent, some seals targeted colder waters on the sea bottom during benthic dives, while at the PF the favourable zones tended to be warmer. The possible negative effect of colder waters in Antarctic on the swimming performances of potential fish or squid prey could explain the behaviour of elephant seals in these zones, while warmer waters within the PF could correspond to the optimal conditions for potential myctophid prey of elephant seals.     

Hot spring frogs (Buergeria japonica) prefer cooler water to hot water

“Hot spring frog” is an informal name used for the Japanese stream tree frog (Buergeria japonica), which is widely distributed in Taiwan and the Ryukyu Archipelago in Japan. Some populations of the species are known to inhabit hot springs. However, water temperature can be extremely high around the sources of hot springs. Thus, it is questionable whether B. japonica selectively inhabits such dangerous environments. To address this question, we conducted a series of observations of water temperature preferences of a hot spring population of B. japonica in Kuchinoshima Island in Japan: (a) a field observation of tadpole density in water pools of different temperatures, (b) a field observation of water temperatures where adult males appear for breeding, and (c) an indoor observation of water temperatures selected by adult females for oviposition. As a result, tadpoles showed a higher density in cooler water. Adult males avoided water pools hotter than 37°C, and adult females selected cooler pools for oviposition. Camera records also showed that adult individuals tend to appear around cooler pools. Thus, we did not find any support for the hypothesis that hot spring frogs prefer hot water. Conversely, they apparently tended to prefer cooler water if it was available. Water temperatures around the sources of the hot spring exceed thermal tolerances of the species and could be a strong selective pressure on the population. Thus, the ability to sense and avoid lethal temperatures may be a key ecological and physiological characteristic for the species that inhabit hot springs.     

Individual heterogeneity in life-history trade-offs with age at first reproduction in capital breeding elephant seals

Recruitment age plays a key role in life-history evolution. Because individuals allocate limited resources among competing life-history functions, theory predicts trade-offs between current reproduction and future growth, survival and/or reproduction. Reproductive costs tend to vary with recruitment age, but may also be overridden by fixed individual differences leading to persistent demographic heterogeneity and positive covariation among demographic traits at the population level. We tested for evidence of intra- and inter-generational trade-offs and individual heterogeneity relating to age at first reproduction using three decades of detailed individual life-history data of 6,439 capital breeding female southern elephant seals. Contrary to the predictions from trade-off hypotheses, we found that recruitment at an early age was associated with higher population level survival and subsequent breeding probabilities. Nonetheless, a survival cost of first reproduction was evident at the population level, as first-time breeders always had lower survival probabilities than prebreeders and experienced breeders of the same age. However, models accounting for hidden persistent demographic heterogeneity revealed that the trade-off between first reproduction and survival was only expressed in “low quality” individuals, comprising 35% of the population. The short-term somatic costs associated with breeding at an early age had no effect on the ability of females to allocate resources to offspring in the next breeding season. Our results provide strong evidence for individual heterogeneity in the life-history trajectories of female elephant seals. By explicitly modeling hidden persistent demographic heterogeneity we show that individual heterogeneity governs the expression of trade-offs with first reproduction in elephant seals.     

Ontogenetic niche partitioning in southern elephant seals from Argentine Patagonia

Elephant seals, Mirounga spp., are highly dimorphic, having different energetic requirements according to age and sex, and foraging in various ecological and oceanographic contexts. Resource partitioning has been shown for the sub-Antarctic populations of southern elephant seals, M. leonina, where colonies are surrounded by narrow shelves that deepen abruptly. In contrast, seals from Península Valdés (Argentina), in the northernmost extent of the breeding range, face an extended, shallow, temperate, and productive continental shelf. We integrated tracking data from 98 animals (juveniles and adults, males and females) gathered over more than two decades, and found that although all available habitats were used, individuals segregated by age and sex. Juvenile males favored shelf habitats, whereas subadult and adult males also used the shelf break. Juvenile females preferred the shelf and the more distant Argentine Basin used by postbreeding and postmolt adult females. Males showed the highest proportion of area-restricted search locations, suggesting more spatially concentrated feeding activity, and likely reflecting a preference for foraging habitat and prey. Our results are consistent with those from other populations, implying that elephant seals show remarkable similarities in habitat use by age and sex classes, despite broad differences in the offshore habitats between sub-Antarctic and temperate ecosystems.     

Seasonal variation in the metabolic rate and body composition of female grey seals: fat conservation prior to high-cost reproduction in a capital breeder?

Many animals rely on stored energy through periods of high energy demand or low energy availability or both. A variety of mechanisms may be employed to attain and conserve energy for such periods. Wild grey seals demonstrate seasonal patterns of energy storage and foraging behaviour that appear to maximize the allocation of energy to reproduction—a period characterized by both high energy demand and low food availability. We examined seasonal patterns in resting rates of oxygen consumption as a proxy for metabolic rate (RMR) and body composition in female grey seals (four adults and six juveniles), testing the hypothesis that adults would show seasonal changes in RMR related to the reproductive cycle but that juveniles would not. There was significant seasonal variation in rates of resting oxygen consumption of adult females, with rates being highest in the spring and declining through the summer months into autumn. This variation was not related to changes in water temperature. Adults increased in total body mass and in fat content during the same spring to autumn period that RMR declined. RMR of juveniles showed no clear seasonal patterns, but did increase with increasing mass. These data support the hypothesis that seasonal variation in RMR in female grey seals is related to the high costs of breeding.     

South American sea lions in Peru have a lek‐like mating system

Five years of behavioral observations revealed significant effects of high air temperatures and breeding site topography on the mating system of South American sea lions in Peru. Unlike most polygynous mammals that defend females or fixed territories, male sea lions in Peru maintained positions along the shoreline where females passed each day to thermoregulate, and where most copulations occurred. Sex ratios (1 male per 17 females) and male mating success were extremely skewed (14% of males achieved 50% of the copulations, and 25% of them did not copulate at all). The mass daily movements of females toward the water and cool substrate of the shoreline, along with a highly skewed sex ratio, accentuated the difficulty for males to monopolize and restrict female movements. Females moved freely and chose their mates, unlike in temperate regions of their range where male South American sea lions control groups of females or access to tide pools. Our observations indicate that the South American sea lion in Peru has a lek‐like breeding system. This is a rare alternative to the common male strategies of defending females and resources, and is likely an evolutionary product of their highly skewed sex ratio, protracted breeding season, and the extreme subtropical climate where they breed.