BEHAVIORAL EVIDENCE OF THERMAL STRESS FROM OVERHEATING IN UK BREEDING GRAY SEALS

Gray seals ( Halichoerus grypus ) in the UK exhibit clear preferences for pupping close to pools of water on inland breeding sites. The reasons for this are unclear as seals are thought to derive all their water requirements from the metabolism of fat. Likewise, the prospect of seals overheating during the UK's cool, damp, autumnal breeding seasons, has seemed unlikely, but has not been previously explored. Here, we provide preliminary behavioral evidence of thermal stress in female gray seals breeding on the island of North Rona, Scotland. Video footage provided measures of proximity to, and the proportion of animals bathing in, pools in relation to meteorological data (temperature, mean sea-level pressure, rainfall, wind speed and direction) on four dates spread through a single breeding season. The proportion of females close to pools increased with pressure (warmer, drier conditions) and decreased on wetter days. In addition, analyses of colony-wide patterns of seal dispersion showed that females tended to be closer to pools of water at higher pressures and temperatures and at lower wind-speeds. These results provide the first evidence of thermal stress and behavioral thermoregulation through access to pools of water in a phocid breeding in temperate autumnal conditions.     

The behavioural response of Australian fur seals to motor boat noise

Australian fur seals breed on thirteen islands located in the Bass Strait, Australia. Land access to these islands is restricted, minimising human presence but boat access is still permissible with limitations on approach distances. Thirty-two controlled noise exposure experiments were conducted on breeding Australian fur seals to determine their behavioural response to controlled in-air motor boat noise on Kanowna Island (39°10'S, 146°18'E). Our results show there were significant differences in the seals' behaviour at low (64-70 dB) versus high (75-85 dB) sound levels, with seals orientating themselves towards or physically moving away from the louder boat noise at three different sound levels. Furthermore, seals responded more aggressively with one another and were more alert when they heard louder boat noise. Australian fur seals demonstrated plasticity in their vocal responses to boat noise with calls being significantly different between the various sound intensities and barks tending to get faster as the boat noise got louder. These results suggest that Australian fur seals on Kanowna Island show behavioural disturbance to high level boat noise. Consequently, it is recommended that an appropriate level of received boat sound emissions at breeding fur seal colonies be below 74 dB and that these findings be taken into account when evaluating appropriate approach distances and speed limits for boats.     

Human disturbance and stage‐specific habitat requirements influence snowy plover site occupancy during the breeding season

Habitat use has important consequences for avian reproductive success and survival. In coastal areas with recreational activity, human disturbance may limit use of otherwise suitable habitat. Snowy plovers Charadrius nivosus have a patchy breeding distribution along the coastal areas on the Florida Panhandle, USA. Our goal was to determine the relative effects of seasonal human disturbance and habitat requirements on snowy plover habitat use. We surveyed 303 sites for snowy plovers, human disturbance, and habitat features between January and July 2009 and 2010. We made multiple visits during three different sampling periods that corresponded to snowy plover breeding: pre‐breeding, incubation, and brood‐rearing and used multi‐season occupancy models to examine whether human disturbance, habitat features, or both influenced site occupancy, colonization (probability of transition from an unoccupied site to an occupied site), and extinction (probability of transition from an occupied site to an unoccupied site). Snowy plover site occupancy and colonization was negatively associated with human disturbance and site extinction was positively associated with human disturbance. Interdune vegetation had a negative effect on occupancy and colonization, indicating that plovers were less likely to use areas with uniform, dense vegetation among dunes. Also, dune shape, beach debris, and access to low‐energy foraging areas influenced site occupancy, colonization, and extinction. Plovers used habitat based on beach characteristics that provided stage‐specific resource needs; however, human disturbance was the strongest predictor of site occupancy. In addition, vegetation plantings used to enhance dune rehabilitation may negatively impact plover site occupancy. Management actions that decrease human disturbance, such as symbolic fencing and signage, may increase the amount of breeding habitat available to snowy plovers on the Florida Panhandle and in other areas with high human activity. The specific areas that require this protection may vary across snowy plover life history stages.     

The effects of vessel approaches on the New Zealand fur seal (Arctocephalus forsteri) in the Bay of Plenty,New Zealand

Animals that establish new sites near the edge of the species' range may be vulnerable to disturbance as they are low in numbers and are not tied to the sites. Pinniped distributions world‐wide are changing as many species are recolonizing areas of their former ranges and establishing new colonies. Little research is available on the impact that vessel presence may pose on pinnipeds at such sites. This study documents responses of New Zealand fur seals to vessels in the Bay of Plenty, New Zealand, at a recently established breeding colony. Fur seal behavior at the breeding location was recorded in the presence of vessels. GLMM and GAM analyses revealed that fur seal responses varied with month, time of day, duration of vessel exposure, and the distance to the vessel. Age and sex of the seals, and the number of seals present also influenced fur seal response. Fur seals at this site became disturbed when vessels approached to the 10–20 m distance category, and a precautionary minimum approach distance of 50 m has been suggested. This research provides direction for monitoring and minimizing impacts of vessels on fur seals, especially where new sites are being colonized.     

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.     

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.     

Does shade cover availability limit nest-site choice in two populations of a turtle with temperature-dependent sex determination?

Shifts in resource use may be an important mechanism by which organisms can adjust to novel environmental conditions, such as those imposed by climate change. However, for such shifts to be possible, environmental space must exist into which organisms can move. Habitat that ensures successful survival and reproduction is one such critical resource. We studied resource selection of shade cover over nest sites by painted turtles in populations in Illinois (center of range) and New Mexico (southern edge of range). We targeted this habitat feature because shade can influence hatching success and offspring phenotype (including sex in the study species) by affecting nest microenvironments. We found that while turtles in both populations selected nest sites that were shadier than average available sites, overall resource selection differed between the populations. This disparity may have been due to differences in structure of vegetation that provides shade at each site, because areas with high shade cover in New Mexico (low dense thickets) were much more difficult for turtles to access than those in Illinois (dense tree canopy cover). Further, shade cover predicted different parameters of incubation regime at each site, suggesting that turtles must assess dissimilar components of shade cover in order to choose nest sites and predict their future incubation regimes. Our results suggest that shade cover within nesting areas is a key component of painted turtle habitat, and that accessible, highly-shaded nest sites may be limited at the New Mexico site. Maintaining a range of shade cover from which turtles can select nest sites would permit plasticity in nest-site choice to be expressed, which may be important in preventing sex ratio skews due to climate change.     

Holocene changes in the ecology of northern fur seals: insights from stable isotopes and archaeofauna

The remains of northern fur seals (Callorhinus ursinus) are among the most abundant of pinniped elements recovered from mainland coastal archaeological sites in both California and Oregon. This is surprising as all contemporary northern fur seals breed exclusively on offshore islands, primarily at high latitudes, and the species is otherwise pelagic. The vulnerability of these animals to human predation suggests that either humans were foraging much further offshore than has been presumed or alternatively that the ecology of these animals has shifted during the late Holocene. We used isotopic and archaeofaunal analysis of the remains of pinnipeds from the middle to late Holocene of central and northern California to clarify the breeding and foraging behavior, and migration patterns of these ancient animals. The carbon and nitrogen isotope compositions of ancient northern fur seals reveal that these animals fed as far offshore as they do today, and that they remained at middle latitudes throughout the year. From an archaeological site at Moss Landing, California, we identified 16 skeletal elements from at least 12 very small northern fur seal pups. From another site near Mendocino, California, we identified the remains of at least 6 pups. We estimate the size and age of 5 of the young animals using sex-specific regressions of body length on the short dentary length derived from measurements of modern specimens. Our estimates indicate these ancient pups were substantially smaller, and therefore younger, than modern 3-month-old northern fur seal pups from similar latitudes and their nitrogen isotope compositions suggest they had not been weaned. As present-day northern fur seals do not leave their rookeries until they are at least 4 months old, we consider it highly unlikely that these ancient pups swam to these mainland locations from some distant island rookery. While there are numerous nearshore rocky outcrops along the Mendocino Coast, which may have supported small breeding colonies, the Moss Landing site is centered on a 40-km-long sandy beach, and is more than 120 km from what at the time were the nearest offshore islands. We conclude that northern fur seal adult females, subadults, and pups whose remains were recovered at the Moss Landing archaeological site must have been taken at a mainland rookery. Evidence that northern fur seals once bred on the mainland at this central California location suggests that the abundant remains of these animals at numerous other archaeological sites along the California coast also reflect the presence of nearby mainland rookeries. Based on the relative abundance of their remains in ancient human occupation sites and the widespread distribution of sites where their remains have been found, it appears that northern fur seals were once the predominant pinniped throughout a region where they now only rarely occur. Furthermore, their presence along the central and northern California coasts appears to have once severely limited the distribution of other pinnipeds, which are now common to the region.     

HAUL-OUT SELECTION BY PACIFIC HARBOR SEALS (PHOCA VITULINA RICHARDII): ISOLATION AND PERCEIVED PREDATION RISK

The potential for non-aquatic predators to influence habitat use by harbor seals ( Phoca vitulina ) in a nearshore marine environment was studied by examining haul-out site use and through an experimental approach. Distance from shore, distance to possible foraging depths, peripheral water depth, and haul-out areas were quantified for each haul-out. There was a positive relationship between the number of seals hauled out and the distance from shore for eight known haul-out sites. The hypothesis that harbor seals increasingly hauled out farther offshore to reduce predation risk was tested experimentally by measuring their response to a model of a potential terrestrial predator in comparison to a control object, and to disturbance by a human at one of the study sites. Harbor seals abandoned the haul-out in the presence of the predator model, but showed little response to the controls, suggesting they possess a threat image for terrestrial predators and avoid hauling out when it is perceived. These results support the hypothesis that harbor seals select isolated sites to reduce exposure to terrestrial carnivores.     

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.     

Topographic spatial characterisation of grey seal Halichoerus grypus breeding habitat at a sub-seal size spatial grain

Expansion within breeding colonies may critically depend upon the availability of suitable breeding habitat. Here we use topographic modelling in a GIS to characterise suitable pupping habitat and accurately predict the pattern of colonisation in an expanding grey seal breeding colony – the Isle of May (Scotland). We use high resolution images from large format aerial photographs of the colony to generate sub‐metre accurate Digital Terrain Models (DTMs). GIS modelling with these DTMs provides topographic measures of elevation, slope and ease of access to sea and freshwater pools at a 2 m grid cell size. Seal locations during the 1994 breeding season, with sex‐age class, were also digitised from the same images. We examine how the physical attributes of cells (locations) with and without pups differ and identify areas suitable for pupping but remaining unoccupied during 1994. We predict patterns of future colonisation by characterising areas differentiated by the densities of pups within 5 m grid cells and identifying areas, both occupied or unoccupied, with a potential for increased future pupping densities. Our predictions were tested by examining pup distributions observed in the 1998 breeding season. Occupied sites were significantly closer to freshwater pools and access to the sea (p<0.001) than unoccupied sites suggesting that proximity to water may restrict colony expansion before all areas of suitably flat terrain are occupied. All pup density classes occurred in sites with similar slope values and distance to pools. However, higher pupping densities occurred closer to access points (p=0.014). Pup densities observed in 1998 revealed that our 1994 predictions were accurate (p<0.0001). Only 12% of 466 grid cells had higher densities in 1998 than predicted, of which 88% differed by only 1 pup. These incorrectly classified cells occurred at the expanding edge of the colony (in a more topographically homogenous area) and at the main access points from the sea (major traffic zones). These results demonstrate the value of the accurate quantification of topographic parameters at the appropriate spatial grain (in this case below the size of the individual) for use in habitat classification and predictions of habitat utilisation.     

Long-term monitoring of Antarctic pinnipeds in Admiralty Bay (South Shetlands, Antarctica)

Year-round monitoring of five Antarctic pinnipeds was conducted in Admiralty Bay from 1988 up to 2000. Two breeding species: southern elephant sealsMirounga leonina (Linnaeus, 1758) and Weddell sealsLeptonychotes weddellii (Lesson, 1826), were present throughout the year. Three other species: crabeater seals Lobodon carcinophagus (Hobron and Jacquinot, 1842), leopard sealsHydrurga leptonyx (Blainville, 1820), and Antarctic fur sealsArctocephalus gazella (Peters, 1875) visited the area only for short periods. During this study, the abundance of elephant seals was stable, whereas those of Weddell and crabeater seals declined. Leopard seals numbers fluctuated irregularly. We detected a possible immigration from South Georgia: of a stable magnitude for elephant seals, and of variable magnitude, depending on food accessibility, for Antarctic fur seals. We found a strong recurrence of the spatial distributions of elephant, Weddell, and Antarctic fur seals in the 13 oases on the shore of Admiralty Bay. Annual distribution patterns were characteristic for each species. The innermost beaches were used predominantly by the animals during their annual fasts: the breeding and the moulting seasons.     

Habitat selection of the globally threatened Aquatic Warbler Acrocephalus paludicola at the western margin of its breeding range and implications for management

The globally threatened Aquatic Warbler Acrocephalus paludicola is an umbrella species for fen mires and is at risk of extinction in its westernmost breeding population due to severe habitat loss. We used boosted regression trees to model Aquatic Warbler habitat selection in order to make recommendations for effective management of the last remnant habitats. Habitat data were collected in the years 2004–2006 in all remaining breeding sites in Pomerania (eastern Germany and western Poland) as well as in recently abandoned sites. Models were validated using data from similar Aquatic Warbler habitats in Lithuania. The probability of occurrence of Aquatic Warblers in late May/early June was positively associated with low isolation from other occupied sites, less eutrophic conditions, a high proportion of area mown early in the preceding year, high availability of vegetation 60–70 cm high, high prey abundance and high habitat heterogeneity. Early summer land management is needed in the more productive sites to prevent habitat deterioration by succession to higher and denser vegetation. As this also poses a serious threat to broods, management that creates a mosaic of early and late used patches is recommended to preserve and restore productive Aquatic Warbler sites. In less productive sites, winter mowing can maintain suitable habitat conditions. Aquatic Warbler‐friendly land use supports a variety of other threatened plant and animal species typical of fens and sedge meadows and can meet the economic interests of local land users.     

VOCAL INDIVIDUALITY IN MOTHER AND PUP SOUTH AMERICAN FUR SEALS, ARCTOCEPHALUS AUSTRALIS

South American fur seals breeding in Peru are subjected to levels of maternal aggression, and subsequent pup mortality, that are higher than has been reported for any other otariid species. For mothers and pups to maintain contact with each other, a mutual recognition system should exist to facilitate reunion and avoid misdirection of maternal effort. We recorded vocalizations of mothers and pups at Punta San Juan, Peru, during the 1994 and 1995 breeding seasons. Sixteen acoustic variables were measured from a total of 560 calls from 15 mothers and 13 pups. Multivariate analysis showed that calls were variable in several acoustic dimensions. While calls of both mothers and pups showed low variability within and high variability among individuals, mothers' calls were more individualistic. On average, discriminant-function analysis correctly assigned 60% of pup calls and 70% of mother calls to the individual that produced them. Characteristics of the fundamental frequency were most important for distinguishing among mothers, while pup calls, which typically contained less harmonic structure, could be differentiated by formant-like frequency ranges. Thus, calls of mother and pup South American fur seals appear to exhibit sufficient stereotypy to allow for recognition and discrimination among individuals.     

Important marine habitat off east Antarctica revealed by two decades of multi‐species predator tracking

Satellite telemetry data are a key source of animal distribution information for marine ecosystem management and conservation activities. We used two decades of telemetry data from the East Antarctic sector of the Southern Ocean. Habitat utilization models for the spring/summer period were developed for six highly abundant, wide‐ranging meso‐ and top‐predator species: Adélie Pygoscelis adeliae and emperor Aptenodytes forsteri penguins, light‐mantled albatross Phoebetria palpebrata, Antarctic fur seals Arctocephalus gazella, southern elephant seals Mirounga leonina, and Weddell seals Leptonychotes weddellii. The regional predictions from these models were combined to identify areas utilized by multiple species, and therefore likely to be of particular ecological significance. These areas were distributed across the longitudinal breadth of the East Antarctic sector, and were characterized by proximity to breeding colonies, both on the Antarctic continent and on subantarctic islands to the north, and by sea‐ice dynamics, particularly locations of winter polynyas. These areas of important habitat were also congruent with many of the areas reported to be showing the strongest regional trends in sea ice seasonality. The results emphasize the importance of on‐shore and sea‐ice processes to Antarctic marine ecosystems. Our study provides ocean‐basin‐scale predictions of predator habitat utilization, an assessment of contemporary habitat use against which future changes can be assessed, and is of direct relevance to current conservation planning and spatial management efforts.     

Man's impact on seals

The major direct effects of man on seal populations are caused by hunting. Subsistence hunting using traditional methods has a long history and generally resulted in a balance between the hunters and the seals. More recently commercial hunting, using sophisticated methods, has reduced many seal stocks to low levels. In some cases no balance was achieved and either the seals or the industry became extinct. Minor direct effects of man on seals include sport hunting and the destruction of seals for fisheries protection. These are generally of local importance only. Indirect effects may be either detrimental or beneficial to seal stocks. Detrimental effects include fishing operations which may result in the unintentional destruction of seals or compete with seals for available limited fish stocks, habitat destruction, pollution, and anthropogenic disturbance. Beneficial indirect effects include increase of available food stocks resulting from the reduction of another predator, increase of available breeding habitat as a result of human population shifts, and changes in ethical attitudes to seals.     

Migratory Disturbance Thresholds with Mule Deer and Energy Development

Fine-scale movement data has transformed our knowledge of ungulate migration ecology and now provides accurate, spatially explicit maps of migratory routes that can inform planning and management at local, state, and federal levels. Among the most challenging land use planning issues has been developing energy resources on public lands that overlap with important ungulate habitat, including the migratory routes of mule deer (Odocoileus hemionus). We generally know that less development is better for minimizing negative effects and maintaining habitat function, but we lack information on the amount of disturbance that animals can tolerate before reducing use of or abandoning migratory habitat. We used global positioning system data from 56 deer across 15 years to evaluate how surface disturbance from natural gas well pads and access roads in western Wyoming, USA, affected habitat selection of mule deer during migration and whether any disturbance threshold(s) existed beyond which use of migratory habitat declined. We used resource and step selection functions to examine disturbance thresholds at 3 different spatial scales. Overall, migratory use by mule deer declined as surface disturbance increased. Based on the weight of evidence from our 3 independent but complementary metrics, declines in migratory use related to surface disturbance were non-linear, where migratory use sharply declined when surface disturbance from energy development exceeded 3%. Disturbance thresholds may vary across regions, species, or migratory habitats (e.g., stopover sites). Such information can help with management and land use decisions related to mineral leasing and energy development that overlap with the migratory routes of ungulates. © 2020 The Wildlife Society.     

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.     

Factors affecting early mortality in the South American fur seal (Arctocephalus australis) in Peru: density-related effects and predation

Early mortality in the South American fur seal ( Arctocephatus australis ) in Peru is considerably higher than that seen in any other population of fur seal; 31–49% in the first month as opposed to a maximum of 20% in other populations. In 1987 and 1988, pup mortality was found to be enhanced by density-related effects and predation on pups by the southern sea-lion ( Olaria byronia ). At a high density beach, where 60% of the fur seals bred, mortality correlated with the number of females ashore as did aggression. Female aggression was driven to high levels by a combination of high density and movement within the colony to thermoregulate. However, even at a low density, beach mortality remained high due to a high level of predation by male southern sea-lions (low density beach 5·0–8·3% of all pups; high density beach 0·2%). Predation rates were lower at the high density beach because male fur seals expelled most attacking sea-lions during the breeding season (low density beach 10·0% of attacking sea-lions expelled; high density beach 58·8%).