Observations of cetaceans in the south-east Atlantic sector of the Southern Ocean,during summer 2008

Knowledge of cetacean species composition and their distribution in the south-east Atlantic sector of the Southern Ocean is scarce. During a survey in February–March 2008, systematic whale sightings were carried out along transect lines following the 5° and 15° E meridians between 35° and 67° S. In total, 67 toothed whales and 126 baleen whales were observed. Both fin whales (four animals) and Antarctic minke whales Balaenoptera bonaerenses (three animals) in addition to 16 individuals of unidentified species were among the observed baleen whales. The dominating baleen whale species in our study was humpback whales Megaptera novaeangliae with 108 individuals observed. They occurred single or in groups up to seven individuals (N _mean = 2.5 ind) and eight of the counts were of calves. The relationship between humpback whale occurrence and environmental variables including Antarctic krill (Euphausia superba) abundance from acoustic recordings, hydrography, bathymetry and production was tested using general additive models. Only temperature increased the predictive power of the model with whale occurrence increasing with the decreasing temperature in more southern areas.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

A surplus no more? Variation in krill availability impacts reproductive rates of Antarctic baleen whales

The krill surplus hypothesis of unlimited prey resources available for Antarctic predators due to commercial whaling in the 20th century has remained largely untested since the 1970s. Rapid warming of the Western Antarctic Peninsula (WAP) over the past 50 years has resulted in decreased seasonal ice cover and a reduction of krill. The latter is being exacerbated by a commercial krill fishery in the region. Despite this, humpback whale populations have increased but may be at a threshold for growth based on these human-induced changes. Understanding how climate-mediated variation in prey availability influences humpback whale population dynamics is critical for focused management and conservation actions. Using an 8-year dataset (2013–2020), we show that inter-annual humpback whale pregnancy rates, as determined from skin-blubber biopsy samples (n = 616), are positively correlated with krill availability and fluctuations in ice cover in the previous year. Pregnancy rates showed significant inter-annual variability, between 29% and 86%. Our results indicate that krill availability is in fact limiting and affecting reproductive rates, in contrast to the krill surplus hypothesis. This suggests that this population of humpback whales may be at a threshold for population growth due to prey limitations. As a result, continued warming and increased fishing along the WAP, which continue to reduce krill stocks, will likely impact this humpback whale population and other krill predators in the region. Humpback whales are sentinel species of ecosystem health, and changes in pregnancy rates can provide quantifiable signals of the impact of environmental change at the population level. Our findings must be considered paramount in developing new and more restrictive conservation and management plans for the Antarctic marine ecosystem and minimizing the negative impacts of human activities in the region.     

Evidence of resource partitioning between humpback and minke whales around the western Antarctic Peninsula

For closely related sympatric species to coexist, they must differ to some degree in their ecological requirements or niches ( e.g. , diets) to avoid interspecific competition. Baleen whales in the Antarctic feed primarily on krill, and the large sympatric prewhaling community suggests resource partitioning among these species or a nonlimiting prey resource. In order to examine ecological differences between sympatric humpback and minke whales around the Western Antarctic Peninsula, we made measurements of the physical environment, observations of whale distribution, and concurrent acoustic measurements of krill aggregations. Mantel's tests and classification and regression tree models indicate both similarities and differences in the spatial associations between humpback and minke whales, environmental features, and prey. The data suggest (1) similarities (proximity to shore) and differences (prey abundance versus deep water temperatures) in horizontal spatial distribution patterns, (2) unambiguous vertical resource partitioning with minke whales associating with deeper krill aggregations across a range of spatial scales, and (3) that interference competition between these two species is unlikely. These results add to the paucity of ecological knowledge relating baleen whales and their prey in the Antarctic and should be considered in conservation and management efforts for Southern Ocean cetaceans and ecosystems.     

Future recovery of baleen whales is imperiled by climate change

Historical harvesting pushed many whale species to the brink of extinction. Although most Southern Hemisphere populations are slowly recovering, the influence of future climate change on their recovery remains unknown. We investigate the impacts of two anthropogenic pressures—historical commercial whaling and future climate change—on populations of baleen whales (blue, fin, humpback, Antarctic minke, southern right) and their prey (krill and copepods) in the Southern Ocean. We use a climate–biological coupled “Model of Intermediate Complexity for Ecosystem Assessments” (MICE) that links krill and whale population dynamics with climate change drivers, including changes in ocean temperature, primary productivity and sea ice. Models predict negative future impacts of climate change on krill and all whale species, although the magnitude of impacts on whales differs among populations. Despite initial recovery from historical whaling, models predict concerning declines under climate change, even local extinctions by 2100, for Pacific populations of blue, fin and southern right whales, and Atlantic/Indian fin and humpback whales. Predicted declines were a consequence of reduced prey (copepods/krill) from warming and increasing interspecific competition between whale species. We model whale population recovery under an alternative scenario whereby whales adapt their migratory patterns to accommodate changing sea ice in the Antarctic and a shifting prey base. Plasticity in range size and migration was predicted to improve recovery for ice‐associated blue and minke whales. Our study highlights the need for ongoing protection to help depleted whale populations recover, as well as local management to ensure the krill prey base remains viable, but this may have limited success without immediate action to reduce emissions.     

Decline in energy storage in the Antarctic minke whale (<Emphasis Type="Italic">Balaenoptera bonaerensis</Emphasis>) in the Southern Ocean

The annual trend in energy storage in the Antarctic minke whale was examined using catch data from all 18 survey years in the Japanese Whale Research Program (JARPA). Regression analyses clearly showed that blubber thickness, girth and fat weight have been decreasing for nearly 2 decades. The decrease per year is estimated at approximately 0.02 cm for mid-lateral blubber thickness and 17 kg for fat weight, corresponding to 9% for both measurements over the 18-year period. Furthermore, “date”, “extent of diatom adhesion”, “sex”, “body length”, “fetus length”, “latitude”, “age” and “longitude” were all identified as partially independent predictors of blubber thickness. The direct interpretation of this substantial decline in energy storage in terms of food availability is difficult, since no long-term krill abundance series is available. However, an increase in the abundance of krill feeders other than minke whales and a resulting decrease in the krill population must be considered as a likely explanation.     

Are Antarctic minke whales unusually abundant because of 20th century whaling?

Severe declines in megafauna worldwide illuminate the role of top predators in ecosystem structure. In the Antarctic, the Krill Surplus Hypothesis posits that the killing of more than 2 million large whales led to competitive release for smaller krill‐eating species like the Antarctic minke whale. If true, the current size of the Antarctic minke whale population may be unusually high as an indirect result of whaling. Here, we estimate the long‐term population size of the Antarctic minke whale prior to whaling by sequencing 11 nuclear genetic markers from 52 modern samples purchased in Japanese meat markets. We use coalescent simulations to explore the potential influence of population substructure and find that even though our samples are drawn from a limited geographic area, our estimate reflects ocean‐wide genetic diversity. Using Bayesian estimates of the mutation rate and coalescent‐based analyses of genetic diversity across loci, we calculate the long‐term population size of the Antarctic minke whale to be 670 000 individuals (95% confidence interval: 374 000–1 150 000). Our estimate of long‐term abundance is similar to, or greater than, contemporary abundance estimates, suggesting that managing Antarctic ecosystems under the assumption that Antarctic minke whales are unusually abundant is not warranted.     

Cetacean mitochondrial DNA control region: sequences of all extant baleen whales and two sperm whale species

The sequence of the mitochondrial control region was determined in all 10 extant species commonly assigned to the suborder Mysticeti (baleen or whalebone whales) and to two odontocete (toothed whale) species (the sperm and the pygmy sperm whale). In the mysticetes, both the length and the sequence of the control region were very similar, with differences occurring primarily in the first approximately 160 bp of the 5' end of the L-strand of the region. There were marked differences between the mysticete and sperm whale sequences and also between the two sperm whales. The control region, less its variable portion, was used in a comparison including the 10 mysticete sequences plus the same region of an Antarctic minke whale specimen and the two sperm whales. The difference between the minke whales from the North Atlantic and the Antarctic was greater than that between any acknowledged species belonging to the same genus (Balaenoptera). The difference was similar to that between the families Balaenopteridae (rorquals) and Eschrichtiidae (gray whales). The findings suggest that the Antarctic minke whale should have a full species status, B. bonaerensis. Parsimony analysis separated the bowhead and the right whale (family Balaenidae) from all remaining mysticetes, including the pygmy right whale. The pygmy right whale is usually included in family Balaenidae. The analysis revealed a close relationship between the gray whale (family Eschrichtiidae) sequence and those of the rorquals (family Balaenopteridae). The gray whale was included in a clade together with the sei, Bryde's, fin, blue, and humpback whales. This clade was separated from the two minke whale types, which branched together.      

Signals from the south; humpback whales carry messages of Antarctic sea‐ice ecosystem variability

Southern hemisphere humpback whales (Megaptera novaeangliae) rely on summer prey abundance of Antarctic krill (Euphausia superba) to fuel one of the longest‐known mammalian migrations on the planet. It is hypothesized that this species, already adapted to endure metabolic extremes, will be one of the first Antarctic consumers to show measurable physiological change in response to fluctuating prey availability in a changing climate; and as such, a powerful sentinel candidate for the Antarctic sea‐ice ecosystem. Here, we targeted the sentinel parameters of humpback whale adiposity and diet, using novel, as well as established, chemical and biochemical markers, and assembled a time trend spanning 8 years. We show the synchronous, inter‐annual oscillation of two measures of humpback whale adiposity with Southern Ocean environmental variables and climate indices. Furthermore, bulk stable isotope signatures provide clear indication of dietary compensation strategies, or a lower trophic level isotopic change, following years indicated as leaner years for the whales. The observed synchronicity of humpback whale adiposity and dietary markers, with climate patterns in the Southern Ocean, lends strength to the role of humpback whales as powerful Antarctic sea‐ice ecosystem sentinels. The work carries significant potential to reform current ecosystem surveillance in the Antarctic region.     

A Southern Ocean archipelago enhances feeding opportunities for a krill predator

Productivity in the oceans is heightened around oceanographic and bathymetric features such as fronts and islands. This can have a flow-on effect, providing increased food availability for higher trophic level species. Using data from a 5-day combined visual and acoustic survey, we examined the hypothesis that higher Antarctic krill (Euphausia superba) density provides a lucrative resource for humpback whales (Megaptera novaeangliae) at a remote Antarctic feeding area, the Balleny Islands (67^oS, 164°E). We assessed whale presence at the feeding area in relation to prey (krill), productivity and environmental variables using density surface modeling. We found stark differences between krill swarms near the islands and those in adjacent open water. Swarms were twice as dense and three times more numerous near the Balleny Islands compared to an open water pelagic environment, suggesting that the islands offered a profitable feeding opportunity. At the feeding area, whales were found in deeper and more productive waters with medium krill densities. These relationships, along with the high krill availability around the islands, may be the result of the Island Mass Effect.     

Interspecific relationships in density among the whale community in the Antarctic

Interspecific relationships in density among a whale community in Antarctic feeding grounds were examined using the sightings data derived from the systematic surveys conducted between 1978/1979 and 1987/1988. A clear difference in densities against the physiographic variables (the sea floor-slope type) was identified between baleen whales and toothed whales. Densities of sperm whales and ziphiids were low in the waters over the continental shelf where minke whales' densities were highest. This led to an apparent negative correlation in the density between minke and sperm whales, and minke whale and ziphiids. A significant positive correlation in density between minke and blue whales was identified. No association in density between minke and humpback whales was observed. Distribution of killer whales shows strong positive correlation with that of minke whales. The positive correlation existed between minke and blue whales, and minke and killer whales even when the effect of environmental variables was excluded. Analysis also revealed that the environmental variables, including physiographic variables, are major factors affecting the distributions and density of whales, especially between baleen whales and toothed whales. Accepted: 8 December 1999     

Occurrence of dwarf minke whales (Balaenoptera acutorostrata subsp.) around the Antarctic Peninsula

The occurrence of dwarf minke whales (Balaenoptera acutorostrata subsp.) around the Antarctic Peninsula was examined based on 406 sightings of minke whales recorded during the Chilean Antarctic Scientific Expeditions and other opportunistic cetacean surveys. Identification of the species was made only for the whales sighted in the proximity of the vessels when the specific diagnostic characters could be confirmed. Of the 406 sightings, 296 were assigned to Antarctic (519 individuals), nine (11 individuals) to dwarf and 101 to unidentified minke whales (149 individuals). Dwarf minke whales were identified by the reported external diagnostic characters for this species. Seven animals occurred around the South Shetland Island and four in the Gerlache Strait. In addition, another two animals were identified as dwarf minke whales in the Bellinghausen Sea in winter 1993, being these the most southern records for this species. These results confirm the occurrence of dwarf minke whales around the Antarctic Peninsula during the summer seasons, as well as in the Bellinghausen Sea in winter. The geographical range of these sightings was comprised between 61°03′ and 69°25′S and between 55°29′ and 86°53′W. These results also suggest that some dwarf minke whales remain in the Antarctic during the austral winter.     

Relationship between the distribution of euphausiids and baleen whales in the Antarctic (35°E – 145°W)

Simultaneous whale sighting and hydroacoustic surveys were conducted from a research vessel in the Antarctic to examine the relationship between the distribution of euphausiids and baleen whales. High densities of minke whales and large aggregations of euphausiids were observed along the ice edge over the continental slope in the southeast region of area IV and in the southwest region of area V. The results suggest that the continental slope zone that coincides with the ice edge would be an important minke whale feeding area. Minke whales were rarely sighted in the offshore region even if euphausiids were abundant. Distributions of humpback whales were correlated with high euphausiid density zones, regardless of the bottom topographic features. Several groups of blue whales were sighted in the small area along the ice edge where euphausiids were abundant, but sightings were too few to draw any general conclusion about the relationship between blue whales and euphausiids. Both baleen whales and euphausiids were scarce in the area east of 170°W where sea ice covered the continental shelf and slope zone.     

Horizontal niche partitioning of humpback and fin whales around the West Antarctic Peninsula: evidence from a concurrent whale and krill survey

A dedicated aerial cetacean survey was conducted concurrently to a standardised net trawl survey for krill in order to investigate distribution patterns of large whales and different krill species and to investigate relationships of these. Distance sampling data were used to produce density surface models for humpback (Megaptera novaeangliae) and fin whales (Balaenoptera physalus) around the West Antarctic Peninsula (WAP). Abundance for both species was estimated over two strata in the Bransfield Strait and Drake Passage. Distinct distribution patterns suggest horizontal niche partitioning of the two whale species around the WAP, with fin whales aggregating at the shelf edge of the South Shetland Islands in the Drake Passage and humpback whales in the Bransfield Strait. Krill biomass estimated from the concurrent krill survey was used along with CTD data from the same expedition, bathymetric parameters and satellite data on chlorophyll-a and ice concentration to model krill distribution. Comparisons of the predicted distributions of both whale species with the predicted distributions of Euphausia superba, Euphausia crystallorophias and Thysanoessa macrura suggest a complex relationship rather than a straightforward correlation between krill and whales. However, results indicate that fin whales were feeding in an area dominated by T. macrura, while humpback whales were found in areas of higher E. superba biomass. Our results provide abundance estimates for humpback whales and, for the first time, fin whales in the WAP and contribute important information on feeding ecology and habitat use of these two species in the Southern Ocean.     

Intra‐season variations in distribution and abundance of humpback whales in the West Antarctic Peninsula using cruise vessels as opportunistic platforms

Fine‐scale knowledge of spatiotemporal dynamics in cetacean distribution and abundance throughout the Western Antarctic Peninsula (WAP) is sparse yet essential for effective ecosystem‐based management (EBM). Cruise vessels were used as platforms of opportunity to collect data on the distribution and abundance of humpback whales (Megaptera novaeangliae) during the austral summer of 2019/2020 in a region that is also important for the Antarctic krill (Euphausia superba) fishery, to assess potential spatiotemporal interactions for future use in EBM. Data were analyzed using traditional design‐based line transect methodology and spatial density surface hurdle models fitted using a set of physical environmental covariates to estimate the abundance and distribution of whales in the area, and to describe their temporal dynamics. Our results indicate a rapid increase in humpback whale abundance in the Bransfield and Gerlache Straits through December, reaching a stable abundance by mid‐January. The distribution of humpback whales appeared to change from a patchier distribution in the northern Gerlache Strait to a significantly concentrated presence in the central Gerlache and southern Bransfield Straits, followed by a subsequent dispersion throughout the area. Abundance estimates agreed well with previous literature, increasing from approximately 7000 individuals in 2000 to a peak of 19,107 in 2020. Based on these estimates, we project a total krill consumption of between 1.4 and 3.7 million tons based on traditional and contemporary literature on per capita krill consumption of whales, respectively. When taken in the context of krill fishery catch data in the study area, we conclude that there is minimal spatiotemporal overlap between humpback whales and fishery activity during our study period of November–January. However, there is potential for significant interaction between the two later in the feeding season, but cetacean survey efforts need to be extended into late season in order to fully characterize this potential overlap.     

Fine‐scale spatial differences in humpback whale diet composition near Kodiak,Alaska

On the North Pacific feeding grounds, humpback whales (Megaptera novaeangliae) are recovering from commercial whaling at a rapid rate (6.8%). The potential effect that this recovery will have on trophic dynamics involving these predators is currently unknown. To better elucidate complex trophic dynamics, variability in diet composition of apex predators on their respective feeding grounds needs to be understood. Thus, we explored the diet composition of two defined subaggregations of humpback whales of the Kodiak Archipelago population (“North,” “South”) using stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin and regional prey samples in Bayesian dietary mixing models. Humpback whales in the “North” region consumed proportionally more fish, dominated by capelin (Mallotus villosus), whereas, whales in the “South” region consumed predominantly krill. The difference in diet composition appears to reflect regional differences in prey availability. Thus, regional variability in diet composition by humpback whales may have disproportionate impacts on prey resources of sympatric predators depending on available prey biomass. As a result, we suggest fine‐scale studies of apex predator diets are needed to better model trophic dynamics with accuracy.     

Assessing changes in numbers and distribution of large whale entanglements in Newfoundland and Labrador,Canada1

Entanglements of large whales in commercial fisheries in Newfoundland and Labrador, Canada, have been consistently recorded since 1979, as part of a program aimed at releasing captured animals and reducing costs to fishermen. This data set represented an opportunity to identify fisheries posing particular entanglement risks to local whale populations. Data were assessed over the periods 1979–1992 and 1993–2008, corresponding to distinct phases in fisheries distribution and intensity. Between 1979 and 2008, 1,209 large whale entanglements were recorded in Newfoundland and Labrador. These were mostly humpback whales (Megaptera novaeangliae; 80%) and minke whales (Balaenoptera acutorostrata; 15%). Dramatic declines in reported inshore whale entanglement rates were observed following the 1992 moratorium on Atlantic cod (Gadus morhua) fisheries. Recently, more entanglements have been reported further offshore, largely due to expansion of fisheries targeting snow crab (Chionoecetes opilio). For all whale species, entanglement rates and associated mortality rates varied considerably in different fishing gear. Fractions of humpback and minke whales found dead in different fishing gear differed substantially, with minke whales far more likely to be found dead than humpback whales.     

Modeling the relationship of Antarctic minke whales to major ocean boundaries

The population size of Antarctic minke whales Balaenoptera bonaerensis has been changing simultaneously with profound changes in the physics, i.e., mesopredator habitat features, of the Southern Ocean. Although the two trends may not be related, distinguishing among the factors responsible requires a better understanding of minke whale habitat preferences. For the first time at a large geographic scale, i.e., between 140° E and 35° W, we use data not constrained by vessels needing to avoid sea ice to model the habitat affinities of this pagophilic mesopredator. Using Maxent, we modeled minke whale proximity to the Antarctic Shelf Break Front (ASBF) and the southern boundary of Antarctic Circumpolar Current (sbACC), as well as association with sea ice, given that global climate change is altering the positions or intensity of these features. We also included water depth and chlorophyll (proxy for productivity) as variables. Minke whale presence data were gathered using strip and line census on 55 cruises on board icebreakers during late spring and summer, 1976–2005. The most important variable was distance to ASBF, followed by water depth and sea-ice concentration. That is, found principally in waters south of the sbACC during summer, minke whales were most abundant near the outer edge of the continental shelf (shallow depth), including areas heavily covered by sea ice. We propose that as the sbACC moves south and sea ice disappears, as projected by global climate models, minke whale habitat will shrink, and likely intra- and inter-specific competition will increase.     

CONSEQUENCES OF ALTERNATIVE FUNCTIONAL RESPONSE FORMULATIONS IN MODELS EXPLORING WHALE-FISHERY INTERACTIONS

We evaluated the utility of Ecosim for exploring interactions between cetacean predators, their prey, and fisheries. We formulated six Ecosim parameterizations, representing alternative hypotheses of feeding interactions (functional response) between cetaceans and their main fish prey, and examined differences in the predicted responses to simulated harvesting regimes for minke whales and their prey. Regardless of the type of function response formulated, intense fishing on the main fish prey of minke whales had a longer-lasting negative impact on minke whales than when minke whale biomass was removed directly by harvesting. Consumption rate, biomass, feeding time and mortality of minke whales were all sensitive to the type of functional response specified. Inclusion of "handling time" limited minke whales consumption at high prey densities and predicted higher consumption at low prey densities; features characteristic of a type II functional response. Predicted decline and recovery rates of minke whales were slower than when consumption rates were not limited. Addition of "foraging time" adjustments resulted in more conservative estimates of decline and recovery. However, when "other mortality" was linked to time spent foraging, exposure to higher mortality at low prey densities, and reduced mortality at high prey densities resulted in dramatic differences in predicted biomass trajectory. Sensitivity to the "other mortality" assumption is important for cetaceans whose predation mortality is only a small proportion of total mortality. Differences in the feeding and biomass dynamics were also observed when prey availability to predators was represented by changes in prey vulnerability, confirming earlier reports that Ecosim predictions are sensitive to this parameter.