Provisioning Strategies of Antarctic Fur Seals and Chinstrap Penguins Produce Different Responses to Distribution of Common Prey and Habitat

Central-place foragers that must return to a breeding site to deliver food to offspring are faced with trade-offs between prey patch quality and distance from the colony. Among colonial animals, pinnipeds and seabirds may have different provisioning strategies, due to differences in their ability to travel and store energy. We compared the foraging areas of lactating Antarctic fur seals and chinstrap penguins breeding at Seal Island, Antarctica, to investigate whether they responded differently to the distribution of their prey (Antarctic krill and myctophid fish) and spatial heterogeneity in their habitat. Dense krill concentrations occurred in the shelf region near the colony. However, only brooding penguins, which are expected to be time-minimizers because they must return frequently with whole food for their chicks, foraged mainly in this proximal shelf region. Lactating fur seals and incubating penguins, which can make longer trips to increase energy gain per trip, and so are expected to be energy-maximizers, foraged in the more distant (>20 km from the island) slope and oceanic regions. The shelf region was characterized by more abundant, but lower-energy-content immature krill, whereas the slope and oceanic regions had less abundant but higher-energy-content gravid krill, as well as high-energy-content myctophids. Furthermore, krill in the shelf region undertook diurnal vertical migration, whereas those in the slope and oceanic regions stayed near the surface throughout the day, which may enhance the capture rate for visual predators. Therefore, we sug- gest that the energy-maximizers foraged in distant, but potentially more profitable feeding regions, while the time-minimizers foraged in closer, but potentially less profitable regions. Thus, time and energy constraints derived from different provisioning strategies may result in sympatric colonial predator species using different foraging areas, and as a result, some central-place foragers use sub- optimal foraging habitats, in terms of the quality or quantity of available prey.

Hunting and social behaviour of leopard seals (Hydrurga leptonyx) at Seal Island, South Shetland Islands, Antarctica

The hunting behavior of leopard seals Hydrurga leptonyx was monitored opportunistically at Seal Island, South Shetland Islands, during the austral summers from 1986/87 to 1994/95. Leopard seals used several methods to catch Antarctic fur seal pups Arctocephalus gazella and chinstrap penguins Pygoscelis antarctica, and individuals showed different hunting styles and hunting success. One to two leopard seals per year were responsible for an average of 60% of observed captures of fur seal pups. Leopard seals preyed on penguins throughout the summer, but preyed on fur seal pups only between late December and mid-February. Hunting behavior differed significantly between different locations on the island; fur seals were hunted only at one colony, and penguins were hunted in several areas. The relative abundance of prey types, size of prey in relation to predator, and specialization of individual leopard seals to hunt fur seal prey probably influence individual prey preferences among leopard seals. On five occasions, two leopard seals were seen together on Seal Island. Possible interpretations of the relationship between the interacting leopard seals included a mother-offspring relationship, a consorting male-female pair, and an adult leopard seal followed by an unrelated juvenile. In two incidents at Seal Island, two leopard seals were observed interacting while hunting: one seal captured fur seal pups and appeared to release them to the other seal. Observations of leopard seals interacting during hunting sessions were difficult to confirm as co-operative hunting, but they strongly implied that the two seals were not agonistic toward one another. The hunting success of individual leopard seals pursuing penguins or fur seals is probably high enough for co-operative hunting not to become a common hunting strategy; however, it may occur infrequently when it increases the hunting productivity of the seals.

Taking account of dependent species in management of the Southern Ocean krill fishery: estimating crabeater seal abundance off east Antarctica

1. The crabeater seal Lobodon carcinophaga is considered to be a key species in the krill-based food web of the Southern Ocean. Reliable estimates of the abundance of this species are necessary to allow the development of multispecies, predator–prey models as a basis for management of the krill fishery in the Southern Ocean. 2. A survey of crabeater seal abundance was undertaken in 1500 000 km2 of pack-ice off east Antarctica between longitudes 64–150° E during the austral summer of 1999/2000. Sighting surveys, using double observer line transect methods, were conducted from an icebreaker and two helicopters to estimate the density of seals hauled out on the ice in survey strips. Satellite-linked dive recorders were deployed on a sample of seals to estimate the probability of seals being hauled out on the ice at the times of day when sighting surveys were conducted. Model-based inference, involving fitting a density surface, was used to infer densities in the entire survey region from estimates in the surveyed areas. 3. Crabeater seal abundance was estimated to be between 0.7 and 1.4 million animals (with 95% confidence), with the most likely estimate slightly less than 1 million. 4. Synthesis and applications. The estimation of crabeater seal abundance in Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR) management areas off east Antarctic where krill biomass has also been estimated recently provides the data necessary to begin extending from single-species to multispecies management of the krill fishery. Incorporation of all major sources of uncertainty allows a precautionary interpretation of crabeater abundance and demand for krill in keeping with CCAMLR’s precautionary approach to management. While this study focuses on the crabeater seal and management of living resources in the Southern Ocean, it has also led to technical and theoretical developments in survey methodology that have widespread potential application in ecological and resource management studies, and will contribute to a more fundamental understanding of the structure and function of the Southern Ocean ecosystem.

Population Growth of Antarctic Fur Seals: Limitation by a Top Predator, The Leopard Seal?

Antarctic fur seals (Arctocephalus gazella) in the South Shetland Islands are recovering from 19th-century exploitation more slowly than the main population at South Georgia. To document demographic changes associated with the recovery in the South Shetlands, we monitored fur seal abundance and reproduction in the vicinity of Elephant Island during austral summers from 1986/1987 through 1994/1995. Total births, mean and variance of birth dates, and average daily mortality rates were estimated from daily live pup counts at North Cove (NC) and North Annex (NA) colonies on Seal Island. Sightings of leopard seals (Hydrurga leptonyx) and incidents of leopard seal predation on fur seal pups were recorded opportunistically during daily fur seal research at both sites. High mortality of fur seal pups, attributed to predation by leopard seals frequently observed at NC, caused pup numbers to decline rapidly between January and March (i.e., prior to weaning) each year and probably caused a long-term decline in the size of that colony. The NA colony, where leopard seals were never observed, increased in size during the study. Pup mortality from causes other than leopard seal predation appeared to be similar at the two sites. The number of pups counted at four locations in the Elephant Island vicinity increased slowly, at an annual rate of 3.8%, compared to rates as high as 11% at other locations in the South Shetland Islands. Several lines of circumstantial evidence are consistent with the hypothesis that leopard seal predators limit the growth of the fur seal population in the Elephant Island area and perhaps in the broader population in the South Shetland Islands. The sustained growth of this fur seal population over many decades rules out certain predator–prey models, allowing inference about the interaction between leopard seals and fur seals even though it is less thoroughly studied than predator–prey systems of terrestrial vertebrates of the northern hemisphere. Top-down forces should be included in hypotheses for future research on the factors shaping the recovery of the fur seal population in the South Shetland Islands.

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups.

Social cohesion among kin, gene flow without dispersal and the evolution of population genetic structure in the killer whale (Orcinus orca)

In social species, breeding system and gregarious behavior are key factors influencing the evolution of large-scale population genetic structure. The killer whale is a highly social apex predator showing genetic differentiation in sympatry between populations of foraging specialists (ecotypes), and low levels of genetic diversity overall. Our comparative assessments of kinship, parentage and dispersal reveal high levels of kinship within local populations and ongoing male-mediated gene flow among them, including among ecotypes that are maximally divergent within the mtDNA phylogeny. Dispersal from natal populations was rare, implying that gene flow occurs without dispersal, as a result of reproduction during temporary interactions. Discordance between nuclear and mitochondrial phylogenies was consistent with earlier studies suggesting a stochastic basis for the magnitude of mtDNA differentiation between matrilines. Taken together our results show how the killer whale breeding system, coupled with social, dispersal and foraging behaviour, contributes to the evolution of population genetic structure.

Cetaceans of Southeast Alaska: Distribution and Seasonal Occurrence

Aim To assess the distribution, group size, seasonal occurrence and annual trends of cetaceans. Location The study area included all major inland waters of Southeast Alaska. Methods Between 1991 and 2007, cetacean surveys were conducted by observers who kept a constant watch when the vessel was underway and recorded all cetaceans encountered. For each species, we examined distributional patterns, group size, seasonal occurrence and annual trends. Analysis of variance (anova F) was used to test for differences in group sizes between multiple means, and Student’s t-test was used to detect differences between pairwise means. Cetacean seasonal occurrence and annual trends were investigated using a generalized linear model framework. Results Humpback whales (Megaptera novaeangliae) were seen throughout the region, with numbers lowest in spring and highest in the fall. Fin whale (Balaenoptera physalus) and minke whale (Balaenoptera acutorostrata) distributions were more restricted than that reported for humpback whales, and the low number of sightings precluded evaluating seasonal trends. Three killer whale (Orcinus orca) eco-types were documented with distributions occurring throughout inland waters. Seasonal patterns were not detected or could not be evaluated for resident and offshore killer whales, respectively; however, the transient eco-type was more abundant in the summer. Dall’s porpoise (Phocoenoides dalli) were distributed throughout the region, with more sightings in spring and summer than in fall. Harbour porpoise (Phocoena phocoena) distribution was clumped, with concentrations occurring in the Icy Strait/Glacier Bay and Wrangell areas and with no evidence of seasonality. Pacific white-sided dolphins (Lagenorhynchus obliquidens) were observed only occasionally, with more sightings in the spring. For most species, group size varied on both an annual and seasonal basis. Main conclusions Seven cetacean species occupy the inland waters of Southeast Alaska, with distribution, group size, seasonal occurrence and annual trends varying by species. Future studies that compare spatial and temporal patterns with other features (e.g. oceanography, prey resources) may help in identifying the key factors that support the high density and biodiversity of cetaceans found in this region. An increased understanding of the region’s marine ecology is an essential step towards ensuring the long-term conservation of cetaceans in Southeast Alaska.

Historical Observations of Humpback And Blue Whales in the North Atlantic Ocean: Clues to Migratory Routes and Possibly Additional Feeding Grounds

The seasonal distributions of humpback and blue whales (Megaptera novaeangliae and Balaenoptera musculus, respectively) in the North Atlantic Ocean are not fully understood. Although humpbacks have been studied intensively in nearshore or coastal feeding and breeding areas, their migratory movements between these areas have been largely inferred. Blue whales have only been studied intensively along the north shore of the Gulf of St. Lawrence, and their seasonal occurrence and movements elsewhere in the North Atlantic are poorly known. We investigated the historical seasonal distributions of these two species using sighting and catch data extracted from American 18th and 19th century whaling logbooks. These data suggest that humpback whales migrated seasonally from low-latitude calving/ breeding grounds over a protracted period, and that some of them traveled far offshore rather than following coastal routes. Also, at least some humpbacks apparently fed early in the summer west of the Mid-Atlantic Ridge, well south of their known present-day feeding grounds. In assessing the present status of the North Atlantic humpback population, it will be important to determine whether such offshore feeding does in fact occur. Blue whales were present across the southern half of the North Atlantic during the autumn and winter months, and farther north in spring and summer, but we had too few data points to support inferences about these whales’ migratory timing and routes.

Acoustic Detections of Singing Humpback Whales in Deep Waters Off The British Isles

From October 1996 through September 1998, we used bottom-mounted hydrophone arrays to monitor deep-water areas north and west of the British Isles for songs of humpback whales (Megaptera novaeangliae). Singing humpbacks were consistently detected between October and March from the Shetland- Faroe Islands south to waters west of the English Channel. Temporal and geographic patterns of song detections, and movements of individually tracked whales, exhibited a southwesterly trend over this period, but with no corresponding northward trend between April and September. These results, together with a review of historical data from this area, suggest that the offshore waters of the British Isles represent a migration corridor for humpbacks, at least some of which summer in Norwegian (and possibly eastern Icelandic) waters. The migratory destination of the detected animals remains unknown, but the limited data suggest that these whales are bound primarily for the West Indies rather than historical breeding areas off the northwestern coast of Africa. Humpbacks detected in British waters after early to mid- March probably do not undertake a full migration to the tropics. These data provide further evidence that singing is not confined to tropical waters in winter, but occurs commonly on migration even in high latitudes.

Segregation of migration by feeding ground origin in North Atlantic humpback whales (Megaptera novaeangliae)

Results from a large-scale, capture–recapture study of humpback whales Megaptera novaeangliae in the North Atlantic show that migration timing is influenced by feeding ground origin. No significant differences were observed in the number of individuals from any feeding area that were re-sighted in the common breeding area in the West Indies. However, there was a relationship between the proportion (logit transformed) of West Indies sightings and longitude (r2 = 0.97, F1,3 = 98.27, P = 0.0022) suggesting that individuals feeding farther to the east are less likely to winter in the West Indies. A relationship was also detected between sighting date in the West Indies and feeding area. Mean sighting dates in the West Indies for individuals identified in the Gulf of Maine and eastern Canada were significantly earlier than those for animals identified in Greenland, Iceland and Norway (9.97 days, t179 = 3.53, P = 0.00054). There was also evidence for sexual segregation in migration; males were seen earlier on the breeding ground than were females (6.63 days, t105 = 1.98, P = 0.050). This pattern was consistently observed for animals from all feeding areas; a combined model showed a significant effect for both sex (F1 = 5.942, P = 0.017) and feeding area (F3 =4.756, P=0.0038). The temporal difference in occupancy of the West Indies between individuals from different feeding areas, coupled with sexual differences in migratory patterns, presents the possibility that there are reduced mating opportunities between individuals from different high latitude areas.

Movements of satellite-monitored humpback whales from New Caledonia

Knowledge of the local and migratory movements of humpback whales (Megaptera novaeangliae) from New Caledonia is very limited. To investigate this topic, we attached satellite-monitored tags to 12 whales off southern New Caledonia. Tag longevity ranged from 1 to 52 days (X = 22.5 days). Tagged whales generally moved to the south or southeast, with several spending time in a previously unknown seamount habitat named Antigonia before resuming movement, generally toward Norfolk Island or New Zealand. However, 1 female with a calf traveled the entire length of the western coast of New Caledonia (~450 km) and then west in the direction of the Chesterfield Reefs, a 19th century American (“Yankee”) whaling ground. None of the New Caledonia whales traveled to or toward eastern Australia, which is broadly consistent with the low rate of interchange observed from photo-identification comparisons between these 2 areas. The connections between New Caledonia and New Zealand, together with the relatively low numbers of whales seen in these places generally, support the idea that whales from these 2 areas constitute a single population that remains small and unrecovered.

A New Birth-Interval Approach to Estimating Demographic Parameters of Humpback Whales

A demographic model is developed based on interbirth intervals and is applied to estimate the population growth rate of humpback whales (Megaptera novaeangliae) in the Gulf of Maine. Fecundity rates in this model are based on the probabilities of giving birth at time t after a previous birth and on the probabilities of giving birth first at age x. Maximum likelihood methods are used to estimate these probabilities using sighting data collected for individually identified whales. Female survival rates are estimated from these same sighting data using a modified Jolly–Seber method. The youngest age at first parturition is 5 yr, the estimated mean birth interval is 2.38 yr (SE = 0.10 yr), the estimated noncalf survival rate is 0.960 (SE = 0.008), and the estimated calf survival rate is 0.875 (SE = 0.047). The population growth rate (l) is estimated to be 1.065; its standard error is estimated as 0.012 using a Monte Carlo approach, which simulated sampling from a hypothetical population of whales. The simulation is also used to investigate the bias in estimating birth intervals by previous methods. The approach developed here is applicable to studies of other populations for which individual interbirth intervals can be measured.

Killer Whales and Marine Mammal Trends in The North Pacific—A Re-Examination of Evidence for Sequential Megafauna Collapse and the Prey-Switching Hypothesis

Springer et al. (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling’s removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al., and much of the reduction occurred 50–100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al. suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al.) were likely abundant throughout the period. Overall, we suggest that the Springer et al. hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem.

Evidence of a Feeding Aggregation of Humpback Whales (Megaptera novaeangliae) Around Kodiak Island, Alaska

The known summer feeding range of the North Pacific humpback whale (Megaptera novaeangliae) extends from California, along the coasts of Oregon, Washington, and Alaska, into the Bering Sea, along the Aleutian Islands, the Sea of Okhotsk (Tomilin 1957), and to northern Japan (Rice 1977). In feeding areas of the northeastern Pacific Ocean, humpback whale photoidentification research has been concentrated off California (Calambokidis et al. 1993), southeastern Alaska (Darling and McSweeney 1985, Baker et al. 1986, 1992; Perry et al. 1990), Prince William Sound in Alaska (von Ziegesar 1992), the Oregon and Washington coasts (Calambokidis et al. 1993), and British Columbia (Darling and McSweeney 1985; Graerne Ellis, unpublished data). Results of these photoidentification studies have documented that individual whales tend to return to the same general areas in subsequent years (Darling and McSweeney 1985, Baker et al. 1986, Calambokidis et a(. 1996, von Ziegesar et al. 1994).

Seasonal Variation in Reception of Fin Whale Calls at Five Geographic Areas in the North Pacific

In late August 1991 scientists at the National Oceanic and Atmospheric Administration’s (NOAA) National Marine Mammal Laboratory (NMML) and Pacific Marine Environmental Laboratory (PMEL) began a pilot study to investigate the capability of hydrophones from the US. Navy’s fixed array system to detect large whales in the North Pacific by passive reception of their calls. PMEL had previously established a direct data link from five bottom-mounted arrays of the Navy SOSUS (Sound Surveillance System), via the Naval Oceanographic Processing Facility (NOPF) at Whidbey Island, Washington, to study low-level seafloor seismicity (Fox et al. 1994). PMEL subsequently provided NMML tapes of SOSUS hydrophone data from which whale calls were analyzed. As in an analogous study conducted in the North Atlantic (Nishimura and Conlon 1994, Clark 1995, Mellinger and Clark 1995), calls attributable to whales were received at each SOSUS site at rates that varied seasonally (Anonymous 1996).