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.     

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.     

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.     

Seasonal and Geographic Variation of Southern Blue Whale Subspecies in the Indian Ocean

Understanding the seasonal movements and distribution patterns of migratory species over ocean basin scales is vital for appropriate conservation and management measures. However, assessing populations over remote regions is challenging, particularly if they are rare. Blue whales (Balaenoptera musculus spp) are an endangered species found in the Southern and Indian Oceans. Here two recognized subspecies of blue whales and, based on passive acoustic monitoring, four “acoustic populations” occur. Three of these are pygmy blue whale (B.m. brevicauda) populations while the fourth is the Antarctic blue whale (B.m. intermedia). Past whaling catches have dramatically reduced their numbers but recent acoustic recordings show that these oceans are still important habitat for blue whales. Presently little is known about the seasonal movements and degree of overlap of these four populations, particularly in the central Indian Ocean. We examined the geographic and seasonal occurrence of different blue whale acoustic populations using one year of passive acoustic recording from three sites located at different latitudes in the Indian Ocean. The vocalizations of the different blue whale subspecies and acoustic populations were recorded seasonally in different regions. For some call types and locations, there was spatial and temporal overlap, particularly between Antarctic and different pygmy blue whale acoustic populations. Except on the southernmost hydrophone, all three pygmy blue whale acoustic populations were found at different sites or during different seasons, which further suggests that these populations are generally geographically distinct. This unusual blue whale diversity in sub-Antarctic and sub-tropical waters indicates the importance of the area for blue whales in these former whaling grounds.     

Estimation of minke whale abundance from an acoustic line transect survey of the Mariana Islands

The minke whale is one of the most abundant species of baleen whales worldwide, yet is rarely sighted in subtropical waters. In the North Pacific, they produce a distinctive sound known as the “boing,” which can be used to acoustically localize individuals. A vessel‐based survey using both visual and passive acoustic monitoring was conducted during the spring of 2007 in a large (616,000 km²) study area encompassing the Mariana Islands. We applied line transect methods to data collected from a towed hydrophone array to estimate the abundance of calling minke whales in our study area. No minke whales were sighted, but there were hundreds of acoustic detections of boings. Computer algorithms were developed to localize calling minke whales from acoustic recordings, resulting in over 30 independent localizations, a six‐fold increase over those estimated during the survey. The two best estimates of abundance of calling minke whales were determined to be 80 and 91 animals (0.13 and 0.15 animals per 1,000 km2, respectively; CV = 34%). These are the first density and abundance estimates for calling minke whales using towed hydrophone array surveys, and the first estimates for this species in the Mariana Islands region. These are considered minimum estimates of the true number of minke whales in the study area.     

Seasonality of Antarctic minke whale (Balaenoptera bonaerensis) calls off the western Antarctic Peninsula

The Antarctic minke whale (Balaenoptera bonaerensis) is a difficult species to study because of its low visual detectability and preference for living within the sea ice habitat, accessible only by ice‐strengthened vessels. Recent identification of the Antarctic minke whale as the source of the seasonally ubiquitous bio‐duck call has allowed the use of this sound, as well as downsweeps, to investigate seasonality trends and diel patterns in Antarctic minke whale call production, and their relationship to sea ice cover. Passive acoustic data were collected using an autonomous Acoustic Recording Package (ARP) off the western Antarctic Peninsula. Bio‐duck calls were classified into four distinct call variants, with one variant having two subtypes. Bio‐duck calls were detected between April and November, with increasing call duration during the austral winter, indicating a strong seasonality in call production. Downsweeps, which were also attributed to Antarctic minke whales, were present throughout most months during the recording period, with a peak in July, and an absence in March and April. Both bio‐duck and downsweeps were significantly correlated with sea ice cover. No diel patterns were observed in bio‐duck calls or in downsweep call production at this site.     

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.     

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.     

Iron defecation by sperm whales stimulates carbon export in the Southern Ocean

The iron-limited Southern Ocean plays an important role in regulating atmospheric CO₂ levels. Marine mammal respiration has been proposed to decrease the efficiency of the Southern Ocean biological pump by returning photosynthetically fixed carbon to the atmosphere. Here, we show that by consuming prey at depth and defecating iron-rich liquid faeces into the photic zone, sperm whales (Physeter macrocephalus) instead stimulate new primary production and carbon export to the deep ocean. We estimate that Southern Ocean sperm whales defecate 50 tonnes of iron into the photic zone each year. Molar ratios of C_export ∶Fe_added determined during natural ocean fertilization events are used to estimate the amount of carbon exported to the deep ocean in response to the iron defecated by sperm whales. We find that Southern Ocean sperm whales stimulate the export of 4 × 10⁵ tonnes of carbon per year to the deep ocean and respire only 2 × 10⁵ tonnes of carbon per year. By enhancing new primary production, the populations of 12 000 sperm whales in the Southern Ocean act as a carbon sink, removing 2 × 10⁵ tonnes more carbon from the atmosphere than they add during respiration. The ability of the Southern Ocean to act as a carbon sink may have been diminished by large-scale removal of sperm whales during industrial whaling.     

Decrease in stomach contents in the Antarctic minke whale (Balaenoptera bonaerensis) in the Southern Ocean

The Antarctic minke whale (Balaenoptera bonaerensis) is one of the major krill predators in Antarctic waters. A reported decline in energy storage over almost two decades indicates that food availability for the whales may also have declined recently. To test this hypothesis, catch data from 20 survey years in the Japanese Whale Research Program in the Antarctic (JARPA) and its second phase (JARPA II) (1990/91–2009/10), which covered the longitudinal sector between 35°E and 145°W south of 58°S, were used to investigate whether there was any annual trend in the stomach contents weight of Antarctic minke whales. A linear mixed-effects analysis showed a 31 % (95 % CI 12.6–45.3 %) decrease in the weight of stomach contents over the 20 years since 1990/1991. A similar pattern of decrease was found in both males and females, except in the case of females sampled at higher latitude in the Ross Sea. These results suggest a decrease in the availability of krill for Antarctic minke whales in the lower latitudinal range of the research area. The results are consistent with the decline in energy storage reported previously. The decrease in krill availability could be due to environmental changes or to an increase in the abundance of other krill-feeding predators. The latter appears somewhat more likely, given the recent rapid recovery of humpback whale. Furthermore, humpback whales are not found in the Ross Sea, where both Antarctic krill and ice krill (Euphausia crystallorophias) are available, and where no change in prey availability for Antarctic minke whales is indicated.     

Observations of blue whales feeding in Antarctic waters

There are no published accounts of blue whales (Balaenoptera musculus) feeding in Antarctic waters. This note describes the behaviour of two groups of blue whales feeding in Antarctic pelagic waters. Whales were observed during the 18th IWC/IDCR southern hemisphere minke whale assessment cruise. Feeding behaviour in both cases resembled those described previously for both northern hemisphere blue whales and fin whales (B. physalus). These observations suggest that a programme of comparative behavioural observations could be developed to test the “feeding competition” hypothesis, which suggests that recovery of populations of blue whales will be impeded by feeding competition with sympatric minke whales. Accepted: 29 April 1999     

Super-aggregations of krill and humpback whales in Wilhelmina Bay, Antarctic Peninsula

Ecological relationships of krill and whales have not been explored in the Western Antarctic Peninsula (WAP), and have only rarely been studied elsewhere in the Southern Ocean. In the austral autumn we observed an extremely high density (5.1 whales per km(2)) of humpback whales (Megaptera novaeangliae) feeding on a super-aggregation of Antarctic krill (Euphausia superba) in Wilhelmina Bay. The krill biomass was approximately 2 million tons, distributed over an area of 100 km(2) at densities of up to 2000 individuals m(-3); reports of such 'super-aggregations' of krill have been absent in the scientific literature for >20 years. Retentive circulation patterns in the Bay entrained phytoplankton and meso-zooplankton that were grazed by the krill. Tagged whales rested during daylight hours and fed intensively throughout the night as krill migrated toward the surface. We infer that the previously unstudied WAP embayments are important foraging areas for whales during autumn and, furthermore, that meso-scale variation in the distribution of whales and their prey are important features of this system. Recent decreases in the abundance of Antarctic krill around the WAP have been linked to reductions in sea ice, mediated by rapid climate change in this area. At the same time, baleen whale populations in the Southern Ocean, which feed primarily on krill, are recovering from past exploitation. Consideration of these features and the effects of climate change on krill dynamics are critical to managing both krill harvests and the recovery of baleen whales in the Southern Ocean.     

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.      

Separating pygmy and Antarctic blue whales using long-forgotten ovarian data

Pygmy blue whales ( Balaenoptera musculus brevicauda ) are ≤24.1 m and are generally found north of 52°S in summer, whereas the more southerly Antarctic blue whales ( B. m. intermedia ) may exceed 30 m. Previous assessments have assumed that catches and recent surveys south of 60°S recorded Antarctic blue whales, but these may have included pygmy blue whales. Here, we use ovarian corpora, which accumulate with ovulations and hence with length, to separate these subspecies. The resulting Bayesian mixture model, applied to 1,380 Northern Region (north of 52°S and 35°–180°E) and 3,844 Southern Ocean (south of 52°S) blue whales, estimated that only 0.1% (95% credibility intervals 0.0%–0.4%) of the Antarctic region blue whales were pygmy blue whales and, unexpectedly, found significantly lower lifetime ovulation counts for pygmy blue whales than for Antarctic blue whales (7.6 vs . 13.6). Over four decades, despite substantial depletion of Antarctic blue whales, there was no trend in the estimated proportion of pygmy blue whales in the Antarctic. Several lines of investigation found no evidence for sizeable numbers of pygmy blue whales in ovarian corpora data collected in the 1930s, as was previously hypothesized.     

Seasonal and Diel Vocalization Patterns of Antarctic Blue Whale (Balaenoptera musculus intermedia) in the Southern Indian Ocean: A Multi-Year and Multi-Site Study

Passive acoustic monitoring is an efficient way to provide insights on the ecology of large whales. This approach allows for long-term and species-specific monitoring over large areas. In this study, we examined six years (2010 to 2015) of continuous acoustic recordings at up to seven different locations in the Central and Southern Indian Basin to assess the peak periods of presence, seasonality and migration movements of Antarctic blue whales (Balaenoptera musculus intermedia). An automated method is used to detect the Antarctic blue whale stereotyped call, known as Z-call. Detection results are analyzed in terms of distribution, seasonal presence and diel pattern of emission at each site. Z-calls are detected year-round at each site, except for one located in the equatorial Indian Ocean, and display highly seasonal distribution. This seasonality is stable across years for every site, but varies between sites. Z-calls are mainly detected during autumn and spring at the subantarctic locations, suggesting that these sites are on the Antarctic blue whale migration routes, and mostly during winter at the subtropical sites. In addition to these seasonal trends, there is a significant diel pattern in Z-call emission, with more Z-calls in daytime than in nighttime. This diel pattern may be related to the blue whale feeding ecology.     

Using calls as an indicator for Antarctic blue whale occurrence and distribution across the southwest Pacific and southeast Indian Oceans

Understanding species distribution and behavior is essential for conservation programs of migratory species with recovering populations. The critically endangered Antarctic blue whale (Balaenoptera musculus intermedia) was heavily exploited during the whaling era. Because of their low numbers, highly migratory behavior, and occurrence in remote areas, their distribution and range are not fully understood, particularly in the southwest Pacific Ocean. This is the first Antarctic blue whale study covering the southwest Pacific Ocean region from temperate to tropical waters (32°S to 15°S). Passive acoustic data were recorded between 2010 and 2011 across the southwest Pacific (SWPO) and southeast Indian (SEIO) oceans. We detected Antarctic blue whale calls in previously undocumented SWPO locations off eastern Australia (32°S, 152°E) and within the Lau Basin (20°S, 176°W and 15°S, 173°W), and SEIO off northwest Australia (19°S, 115°E).In temperate waters, adjacent ocean basins had similar seasonal occurrence, in that calling Antarctic blue whales were present for long periods, almost year round in some areas. In northern tropical waters, calling whales were mostly present during the austral winter. Clarifying the occurrence and distribution of critically endangered species is fundamental for monitoring population recovery, marine protected area planning, and in mitigating anthropogenic threats.     

Killer whale predation on penguins in Antarctica

We report here the first published observations of killer whales (Orcinus orca) feeding on penguins in Antarctica. The sightings took place in the Gerlache Strait off the western Antarctic Peninsula during February 2010. Two species of pygoscelid penguins were taken—gentoo (Pygoscelis papua, at least four individuals) and chinstrap (P. antarctica, 2). From remains left at the surface, it was clear that the killer whales fed mainly on the breast muscles, although some penguins may have been swallowed whole. The killer whales were ecotype B, which are purported seal specialists, but we also saw ecotype A, prey specialists on Antarctic minke whales Balaenoptera bonaerensis, chase, but not catch penguins. Because of their small relative size, if penguins are regularly targeted by killer whales in Antarctica, the impact on their populations could be significant.     

Relative changes in krill abundance inferred from Antarctic fur seal

Antarctic krill Euphausia superba is a predominant species in the Southern Ocean, it is very sensitive to climate change, and it supports large stocks of fishes, seabirds, seals and whales in Antarctic marine ecosystems. Modern krill stocks have been estimated directly by net hauls and acoustic surveys; the historical krill density especially the long-term one in the Southern Ocean, however, is unknown. Here we inferred the relative krill population changes along the West Antarctic Peninsula (WAP) over the 20th century from the trophic level change of Antarctic fur seal Arctocephalus gazella using stable carbon (δ(13)C) and nitrogen (δ(15)N) isotopes of archival seal hairs. Since Antarctic fur seals feed preferentially on krill, the variation of δ(15)N in seal hair indicates a change in the proportion of krill in the seal's diets and thus the krill availability in local seawater. For the past century, enriching fur seal δ(15)N values indicated decreasing krill availability. This is agreement with direct observation for the past ～30 years and suggests that the recently documented decline in krill populations began in the early parts of the 20th century. This novel method makes it possible to infer past krill population changes from ancient tissues of krill predators.     

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.     

Aerial surveys for Antarctic minke whales (Balaenoptera bonaerensis) reveal sea ice dependent distribution patterns

This study investigates the distribution of Antarctic minke whales (AMW) in relation to sea ice concentration and variations therein. Information on AMW densities in the sea ice‐covered parts of the Southern Ocean is required to contextualize abundance estimates obtained from circumpolar shipboard surveys in open waters, suggesting a 30% decline in AMW abundance. Conventional line‐transect shipboard surveys for density estimation are impossible in ice‐covered regions, therefore we used icebreaker‐supported helicopter surveys to obtain information on AMW densities along gradients of 0%–100% of ice concentration. We conducted five helicopter surveys in the Southern Ocean, between 2006 and 2013. Distance sampling data, satellite‐derived sea‐ice data, and bathymetric parameters were used in generalized additive models (GAMs) to produce predictions on how the density of AMWs varied over space and time, and with environmental covariates. Ice concentration, distance to the ice edge and distance from the shelf break were found to describe the distribution of AMWs. Highest densities were predicted at the ice edge and through to medium ice concentrations. Medium densities were found up to 500 km into the ice edge in all concentrations of ice. Very low numbers of AMWs were found in the ice‐free waters of the West Antarctic Peninsula (WAP). A consistent relationship between AMW distribution and sea ice concentration weakens the support for the hypothesis that varying numbers of AMWs in ice‐covered waters were responsible for observed changes in estimated abundance. The potential decline in AMW abundance stresses the need for conservation measures and further studies into the AMW population status. Very low numbers of AMWs recorded in the ice‐free waters along the WAP support the hypothesis that this species is strongly dependent on sea ice and that forecasted sea ice changes have the potential of heavily impacting AMWs.