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.     

SEASONAL OCCURRENCE OF NORTH ATLANTIC RIGHT WHALE (EUBALAENA GLACIALIS) VOCALIZATIONS AT TWO SITES ON THE SCOTIAN SHELF

A year-long acoustic survey for critically endangered North Atlantic right whales was conducted at two sites on the central and western Scotian Shelf. Autonomous hydrophones recorded sound continuously from July 2004 to August 2005. Right whale contact calls (upcalls) were identified using automatic recognition software, and the resulting detections were checked manually. Substantial numbers of hours with upcalls were observed at both sites, with approximately four times as many hours with calls at the western site as the central one. Calls occurred mainly from August through October, with the earliest calls in late June and the latest at the end of December at both sites. In addition to this seasonal trend, there was a significant diel pattern in calling at the central site but not at the more westerly site. Results are analyzed in light of feeding ecology and broad-scale movements of right whales.     

Red Shift,Blue Shift: Investigating Doppler Shifts,Blubber Thickness,and Migration as Explanations of Seasonal Variation in the Tonality of Antarctic Blue Whale Song

The song of Antarctic blue whales (Balaenoptera musculus intermedia) comprises repeated, stereotyped, low-frequency calls. Measurements of these calls from recordings spanning many years have revealed a long-term linear decline as well as an intra-annual pattern in tonal frequency. While a number of hypotheses for this long-term decline have been investigated, including changes in population structure, changes in the physical environment, and changes in the behaviour of the whales, there have been relatively few attempts to explain the intra-annual pattern. An additional hypothesis that has not yet been investigated is that differences in the observed frequency from each call are due to the Doppler effect. The assumptions and implications of the Doppler effect on whale song are investigated using 1) vessel-based acoustic recordings of Antarctic blue whales with simultaneous observation of whale movement and 2) long-term acoustic recordings from both the subtropics and Antarctic. Results from vessel-based recordings of Antarctic blue whales indicate that variation in peak-frequency between calls produced by an individual whale was greater than would be expected by the movement of the whale alone. Furthermore, analysis of intra-annual frequency shift at Antarctic recording stations indicates that the Doppler effect is unlikely to fully explain the observations of intra-annual pattern in the frequency of Antarctic blue whale song. However, data do show cyclical changes in frequency in conjunction with season, thus suggesting that there might be a relationship among tonal frequency, body condition, and migration to and from Antarctic feeding grounds.     

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.     

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.     

Blue and fin whale acoustic presence around Antarctica during 2003 and 2004

Seasonal and spatial variations of blue ( Balaenoptera musculus ) and fin whale ( B. physalus ) calls were analyzed from recordings collected with Acoustic Recording Packages (ARPs) deployed between January 2003 and July 2004 at four circumpolar locations: the Western Antarctic Peninsula (WAP), the Scotia Sea (SS), Eastern Antarctica (EA), and the Ross Sea (RS). Call characteristics were compared among sites using the average pressure spectrum levels from 1 month of data at each location. Presence of calls was analyzed using automatic call detection and acoustic power analysis methods. Blue whale calls were recorded year-round, with the highest detections in February–May and November. This suggests that the blue whale population may not migrate synchronously, and may indicate long duration calls are more common during migrations. Fin whale calls were detected only during February–July. Two distinct fin whale call types were recorded, suggesting a possible separation into two populations. The calls at the EA site had a secondary frequency peak in the pressure spectrum at 99 Hz and the calls at the WAP and the SS sites had a peak at 89 Hz. No fin whale calls were detected at the RS site. Acoustics are a good tool to monitor large whales in the Southern Ocean.     

Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean

• 1 Blue whale locations in the Southern Hemisphere and northern Indian Ocean were obtained from catches (303 239), sightings (4383 records of ≥8058 whales), strandings (103), Discovery marks (2191) and recoveries (95), and acoustic recordings.
• 2 Sighting surveys included 7 480 450 km of effort plus 14 676 days with unmeasured effort. Groups usually consisted of solitary whales (65.2%) or pairs (24.6%); larger feeding aggregations of unassociated individuals were only rarely observed. Sighting rates (groups per 1000 km from many platform types) varied by four orders of magnitude and were lowest in the waters of Brazil, South Africa, the eastern tropical Pacific, Antarctica and South Georgia; higher in the Subantarctic and Peru; and highest around Indonesia, Sri Lanka, Chile, southern Australia and south of Madagascar.
• 3 Blue whales avoid the oligotrophic central gyres of the Indian, Pacific and Atlantic Oceans, but are more common where phytoplankton densities are high, and where there are dynamic oceanographic processes like upwelling and frontal meandering.
• 4 Compared with historical catches, the Antarctic (‘true’) subspecies is exceedingly rare and usually concentrated closer to the summer pack ice. In summer they are found throughout the Antarctic; in winter they migrate to southern Africa (although recent sightings there are rare) and to other northerly locations (based on acoustics), although some overwinter in the Antarctic.
• 5 Pygmy blue whales are found around the Indian Ocean and from southern Australia to New Zealand. At least four groupings are evident: northern Indian Ocean, from Madagascar to the Subantarctic, Indonesia to western and southern Australia, and from New Zealand northwards to the equator. Sighting rates are typically much higher than for Antarctic blue whales.
• 6 South‐east Pacific blue whales have a discrete distribution and high sighting rates compared with the Antarctic. Further work is needed to clarify their subspecific status given their distinctive genetics, acoustics and length frequencies.
• 7 Antarctic blue whales numbered 1700 (95% Bayesian interval 860–2900) in 1996 (less than 1% of original levels), but are increasing at 7.3% per annum (95% Bayesian interval 1.4–11.6%). The status of other populations in the Southern Hemisphere and northern Indian Ocean is unknown because few abundance estimates are available, but higher recent sighting rates suggest that they are less depleted than Antarctic blue whales.

     

Seasonal detection of three types of “pygmy” blue whale calls in the Indian Ocean

The Indian Ocean is an area in which a rich suite of cetacean fauna, including at least two subspecies of blue whale, is found; yet little information beyond stranding data and short‐term surveys for this species is available. Pygmy blue whale (Balaenoptera musculus spp.) call data are presented that provide novel information on the seasonal and geographic distribution of these animals. Acoustic data were recorded from January 2002 to December 2003 by hydrophones at three stations of the International Monitoring System, including two near the subequatorial Diego Garcia Atoll and a third southwest of Cape Leeuwin, Australia. Automated spectrogram correlation methods were used to scan for call types attributed to pygmy blue whales. Sri Lanka calls were the most common and were detected year‐round off Diego Garcia. Madagascar calls were only recorded on the northern Diego Garcia hydrophone during May and July, whereas Australia calls were only recorded at Cape Leeuwin, between December and June. Differences in geographic and seasonal patterns of these three distinct call types suggest that they may represent separate acoustic populations of pygmy blue whales and that these “acoustic populations” should be considered when assessing conservation needs of blue whales in the Indian Ocean.     

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.     

Regional differences in length at sexual maturity for female blue whales based on recovered Soviet whaling data

New blue whale ovarian corpora data from illegal Soviet catches in the Southern Hemisphere and northern Indian Ocean were recovered from the original logbooks. Catches north of 52°S were assumed to be pygmy blue whales ( Balaenoptera musculus brevicauda , n = 1,272); those south of 56°S were assumed to be Antarctic (true) blue whales ( B. m. intermedia , n = 153). Three probable Antarctic blue whales north of 52°S were excluded. Lengths at which 50% and 95% of females become sexually mature ( L ₅₀ and L ₉₅) were estimated from a Bayesian logistic model. These estimates are more precise than previous Japanese estimates because Soviet catches below the legal minimum of 70 ft (21.3 m) were 32 times greater. For pygmy blue whales L ₅₀ was 19.2 m (95% interval 19.1–19.3 m) and L ₉₅ was 20.5 m (95% interval 20.4–20.7 m). Antarctic L ₅₀ (23.4 m, 95% interval 22.9–23.9 m) was much longer than L ₅₀ for pygmy blue whale regions (18.4–19.9 m). The median L ₅₀ for the northern Indian Ocean was 0.5–0.6 m shorter than for pygmy blue whales from other regions; although statistically significant, these small length differences provide little support for northern Indian Ocean blue whales being a separate subspecies, B. m. indica .     

Blue whales respond to anthropogenic noise

Anthropogenic noise may significantly impact exposed marine mammals. This work studied the vocalization response of endangered blue whales to anthropogenic noise sources in the mid-frequency range using passive acoustic monitoring in the Southern California Bight. Blue whales were less likely to produce calls when mid-frequency active sonar was present. This reduction was more pronounced when the sonar source was closer to the animal, at higher sound levels. The animals were equally likely to stop calling at any time of day, showing no diel pattern in their sensitivity to sonar. Conversely, the likelihood of whales emitting calls increased when ship sounds were nearby. Whales did not show a differential response to ship noise as a function of the time of the day either. These results demonstrate that anthropogenic noise, even at frequencies well above the blue whales' sound production range, has a strong probability of eliciting changes in vocal behavior. The long-term implications of disruption in call production to blue whale foraging and other behaviors are currently not well understood.     

Synchronous Seasonal Change in Fin Whale Song in the North Pacific

Fin whale (Balaenoptera physalus) song consists of down-swept pulses arranged into stereotypic sequences that can be characterized according to the interval between successive pulses. As in blue (B. musculus) and humpback whales (Megaptera novaeangliae), these song sequences may be geographically distinct and may correlate with population boundaries in some regions. We measured inter-pulse intervals of fin whale songs within year-round acoustic datasets collected between 2000 and 2006 in three regions of the eastern North Pacific: Southern California, the Bering Sea, and Hawaii. A distinctive song type that was recorded in all three regions is characterized by singlet and doublet inter-pulse intervals that increase seasonally, then annually reset to the same shorter intervals at the beginning of each season. This song type was recorded in the Bering Sea and off Southern California from September through May and off Hawaii from December through April, with the song interval generally synchronized across all monitoring locations. The broad geographic and seasonal occurrence of this particular fin whale song type may represent a single population broadly distributed throughout the eastern Pacific with no clear seasonal migratory pattern. Previous studies attempting to infer population structure of fin whales in the North Pacific using synchronous individual song samples have been unsuccessful, likely because they did not account for the seasonal lengthening in song intervals observed here.     

Seasonal acoustic behavior of blue whales (Balaenoptera musculus) in the Gulf of California,Mexico

Six years of passive acoustic monitoring data from the Gulf of California reveal seasonality and movements for the northeastern Pacific blue whales. Three sites were studied, one from the southern (Punta Pescadero) and two from the northern (Isla Tiburon and Canal de Ballenas) regions. A total of 4,953 h were analyzed, and 15,539 blue whale calls were detected, of which 2,723 (18%) were A calls, 11,249 were B calls (72%), and 1,567 were D calls (10%). A and B calls were produced both as song units (2,524) or AB singular calls (2,026). The high rate of songs and their seasonality suggest that the GC is a winter‐breeding ground. A shift from AB call predominance in winter, to D calls in spring and early summer, especially at the entrance of the GC, suggests the importance of this area for reproduction and foraging. Analysis of calling frequency suggests a clear movement of blue whales from the southern region (Punta Pescadero) to the northern regions (Canal de Ballenas and Isla Tiburon), with subsequent southern movement in March. The seasonality and mobility of blue whales in the Gulf of California, inferred from their calling, contributes to the ecological understanding of this population.     

Hybridization of Southern Hemisphere blue whale subspecies and a sympatric area off Antarctica: impacts of whaling or climate change?

Understanding the degree of genetic exchange between subspecies and populations is vital for the appropriate management of endangered species. Blue whales (Balaenoptera musculus) have two recognized Southern Hemisphere subspecies that show differences in geographic distribution, morphology, vocalizations and genetics. During the austral summer feeding season, the Antarctic blue whale (B. m. intermedia) is found in polar waters and the pygmy blue whale (B. m. brevicauda) in temperate waters. Here, we genetically analyzed samples collected during the feeding season to report on several cases of hybridization between the two recognized blue whale Southern Hemisphere subspecies in a previously unconfirmed sympatric area off Antarctica. This means the pygmy blue whales using waters off Antarctica may migrate and then breed during the austral winter with the Antarctic subspecies. Alternatively, the subspecies may interbreed off Antarctica outside the expected austral winter breeding season. The genetically estimated recent migration rates from the pygmy to Antarctic subspecies were greater than estimates of evolutionary migration rates and previous estimates based on morphology of whaling catches. This discrepancy may be due to differences in the methods or an increase in the proportion of pygmy blue whales off Antarctica within the last four decades. Potential causes for the latter are whaling, anthropogenic climate change or a combination of these and may have led to hybridization between the subspecies. Our findings challenge the current knowledge about the breeding behaviour of the world's largest animal and provide key information that can be incorporated into management and conservation practices for this endangered species.     

A Simulation Study of Acoustic-Assisted Tracking of Whales for Mark-Recapture Surveys

Collecting enough data to obtain reasonable abundance estimates of whales is often difficult, particularly when studying rare species. Passive acoustics can be used to detect whale sounds and are increasingly used to estimate whale abundance. Much of the existing effort centres on the use of acoustics to estimate abundance directly, e.g. analysing detections in a distance sampling framework. Here, we focus on acoustics as a tool incorporated within mark-recapture surveys. In this context, acoustic tools are used to detect and track whales, which are then photographed or biopsied to provide data for mark-recapture analyses. The purpose of incorporating acoustics is to increase the encounter rate beyond using visual searching only. While this general approach is not new, its utility is rarely quantified. This paper predicts the “acoustically-assisted” encounter rate using a discrete-time individual-based simulation of whales and survey vessel. We validate the simulation framework using existing data from studies of sperm whales. We then use the framework to predict potential encounter rates in a study of Antarctic blue whales. We also investigate the effects of a number of the key parameters on encounter rate. Mean encounter rates from the simulation of sperm whales matched well with empirical data. Variance of encounter rate, however, was underestimated. The simulation of Antarctic blue whales found that passive acoustics should provide a 1.7–3.0 fold increase in encounter rate over visual-only methods. Encounter rate was most sensitive to acoustic detection range, followed by vocalisation rate. During survey planning and design, some indication of the relationship between expected sample size and effort is paramount; this simulation framework can be used to predict encounter rates and establish this relationship. For a case in point, the simulation framework indicates unequivocally that real-time acoustic tracking should be considered for quantifying the abundance of Antarctic blue whales via mark-recapture methods.     

Morphometric analysis of Chilean blue whales and implications for their taxonomy

In the Southern Hemisphere, blue whales are currently divided into two subspecies, Antarctic blue whales (Balaenoptera musculus intermedia) and pygmy blue whales (B. m. brevicauda), but there is some debate about whether Chilean blue whales should also be considered as a separate subspecies. Here, we provide novel morphometric data to directly address this taxonomic question from a biological survey of 60 blue whales taken during the 1965/1966 Chilean whaling season. The data show that maximum body length and mean body length of both sexually mature females and males for Chilean blue whales are intermediate between pygmy and Antarctic blue whales; and that fluke-anus lengths of Chilean blue whales are significantly different from pygmy blue whales, but not necessarily from Antarctic blue whales. There is also some support from the data that snout-eye measurements are different among all three groups. These data provide further confirmation that Chilean blue whales are a distinct population requiring separate management from other blue whale populations, and are also consistent with suggestions that Chilean blue whales are not the same subspecies as pygmy blue whales.     

Insights into the population structure of blue whales in the Eastern North Pacific from recent sightings and photographic identification

Blue whales were widely distributed in the North Pacific prior to the primary period of modern commercial whaling in the early 1900s. Despite concentrations of blue whale catches off British Columbia and in the Gulf of Alaska, there had been few documented sightings in these areas since whaling for blue whales ended in 1965. In contrast, large concentrations of blue whales have been documented off California and Baja California and in the eastern tropical Pacific since the 1970s, but it was not known if these animals were part of the same population that previously ranged into Alaskan waters. We document 15 blue whale sightings off British Columbia and in the Gulf of Alaska made since 1997, and use identification photographs to show that whales in these areas are currently part of the California feeding population. We speculate that this may represent a return to a migration pattern that has existed for earlier periods for eastern North Pacific blue whale population. One possible explanation for a shift in blue whale use is changes in prey driven by changes in oceanographic conditions, including the Pacific Decadal Oscillation (PDO), which coincides with some of the observed shifts in blue whale occurrence.     

Seasonal occurrence of southeast Pacific blue whale songs in southern Chile and the eastern tropical Pacific

Passive acoustic data were collected January 2012 to April 2013 at four sites in the Chiloense Ecoregion (CER) in southern Chile (≈43°S–44°S, 71°W–73°W) and 1996–2002 from one site in the eastern tropical Pacific (ETP) (8°S, 95°W). Automatic detectors were used to detect the two songs (SEP1 and SEP2) described for southeast Pacific (SEP) blue whales. There was a strong seasonal pattern of occurrence of SEP songs in the CER from December to August, peaking March to May. In the ETP, the occurrence of songs was an order of magnitude lower but songs were present year‐round, with a peak around June. These findings support austral summer/autumn seasonal residency in the CER and a seasonal movement of blue whales towards the ETP during June/July, returning in December. Interannual differences in the ETP were possibly linked to the 1997–1998 El Niño event. At both study sites, SEP2 was significantly more common than SEP1; both songs largely followed the same temporal trends. These findings contribute to our understanding of the seasonal movements of endangered SEP blue whales and can inform conservation strategies, particularly in the CER coastal feeding ground. We recommend future year‐round passive acoustic studies in the CER and the ETP (e.g., near the Galapagos Islands), ideally coupled with oceanographic data.     

Humpback Whale Song and Foraging Behavior on an Antarctic Feeding Ground

Reports of humpback whale (Megaptera novaeangliae) song chorusing occurring outside the breeding grounds are becoming more common, but song structure and underwater behavior of individual singers on feeding grounds and migration routes remain unknown. Here, ten humpback whales in the Western Antarctic Peninsula were tagged in May 2010 with non-invasive, suction-cup attached tags to study foraging ecology and acoustic behavior. Background song was identified on all ten records, but additionally, acoustic records of two whales showed intense and continuous singing, with a level of organization and structure approaching that of typical breeding ground song. The songs, produced either by the tagged animals or close associates, shared phrase types and theme structure with one another, and some song bouts lasted close to an hour. Dive behavior of tagged animals during the time of sound production showed song occurring during periods of active diving, sometimes to depths greater than 100 m. One tag record also contained song in the presence of feeding lunges identified from the behavioral sensors, indicating that mating displays occur in areas worthy of foraging. These data show behavioral flexibility as the humpbacks manage competing needs to continue to feed and to prepare for the breeding season during late fall. This may also signify an ability to engage in breeding activities outside of the traditional, warm water breeding ground locations.