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.     

Species identification and likely catch time period of whale bones from South Georgia

Skeletal remains of baleen whales killed during the onset of 20th century commercial whaling lie scattered across the shores and abandoned whaling stations of the subantarctic island of South Georgia. Here we report on genetic species identification of whale bones collected from South Georgia using standard historical DNA protocols. We amplified and sequenced short fragments of the mitochondrial DNA (mtDNA) control region from 281 available bone samples. Of these, 231 provided mtDNA sequences of sufficient quality and length (174–194 bp) for species identification: 158 bones were identified as humpback whale (Megaptera novaeangliae), 51 bones were identified as fin whale (Balaenoptera physalus), 18 bones were identified as blue whale (B. musculus), two bones were identified as sei whale (B. borealis), one bone was identified as a southern right whale (Eubalaena australis), and one bone was identified as a southern elephant seal (Mirounga leonina). The prominence of humpback, fin, and blue whale bones in the sample collection corresponds to the catch record of the early years of whaling on the island of South Georgia (pre‐1915), prior to the depletion of these populations.     

RIGHT WHALES (EUBALAENA GLACIALIS) ON JEFFREYS LEDGE: A HABITAT OF UNRECOGNIZED IMPORTANCE?

North Atlantic right whales ( Eubalaena glacialis ) are known to spend the majority of the year between the Great South Channel southeast of Cape Cod, and the Nova Scotian shelf. We examined sightings of right whales on and around Jeffreys Ledge, a 54-km-long glacial deposit off the coast of northern Massachusetts, New Hampshire, and Maine. Sightings on Jeffreys Ledge were extracted from three data sets: (1) a systematic survey of the entire northeastern continental shelf between 1979 and 1982, (2) whale-watch and research-cruise sighting data from 1984 to 1997, and (3) a collaborative database of sightings collected by organizations conducting right whale research and all other available sources. Each database supported two seasonal sighting peaks. During summer (especially July and August) sightings were primarily of mother-calf pairs. Several cow-calf pairs were seen over several days to weeks. Several females were resighted in more than one year, but only when calves were present. During October, November, and December, sightings included all age classes, surface-feeding behavior was frequently observed, and some animals were resighted over several weeks. Given the relatively reduced sighting effort during fall, this number of sightings is surprising. During the 20 yr of observations, 52 of 374 photo-identified North Atlantic right whales (13.9%) were seen at least once on Jeffreys Ledge. We suggest that Jeffreys Ledge may be a more important right whale habitat than previously believed, and that it may play an important role in annual movements and distribution of this population.     

Climate impacts on transocean dispersal and habitat in gray whales from the Pleistocene to 2100

Arctic animals face dramatic habitat alteration due to ongoing climate change. Understanding how such species have responded to past glacial cycles can help us forecast their response to today's changing climate. Gray whales are among those marine species likely to be strongly affected by Arctic climate change, but a thorough analysis of past climate impacts on this species has been complicated by lack of information about an extinct population in the Atlantic. While little is known about the history of Atlantic gray whales or their relationship to the extant Pacific population, the extirpation of the Atlantic population during historical times has been attributed to whaling. We used a combination of ancient and modern DNA, radiocarbon dating and predictive habitat modelling to better understand the distribution of gray whales during the Pleistocene and Holocene. Our results reveal that dispersal between the Pacific and Atlantic was climate dependent and occurred both during the Pleistocene prior to the last glacial period and the early Holocene immediately following the opening of the Bering Strait. Genetic diversity in the Atlantic declined over an extended interval that predates the period of intensive commercial whaling, indicating this decline may have been precipitated by Holocene climate or other ecological causes. These first genetic data for Atlantic gray whales, particularly when combined with predictive habitat models for the year 2100, suggest that two recent sightings of gray whales in the Atlantic may represent the beginning of the expansion of this species' habitat beyond its currently realized range.     

Does disappearance mean extirpation? The case of right whales off Namibia

Right whales off Namibia were severely depleted by early 19th century whaling, and rarely featured in modern whaling catches in the 1920s. Aerial surveys of the Namibian coastline from 1978 and onwards revealed increasing numbers of right whales, but few cow‐calf pairs. Aerial surveys off South Africa since 2009 showed a major decline in the availability of animals without calves. Twenty individual matches were made between 94 whales photographed off Namibia/Northern Cape in 2003–2012 and 1,677 photographed off South Africa in 1979–2012. Eight were adult females that calved in South African waters, but only one was also seen with a calf off Namibia. Twelve out of 13 individuals off Namibia with distinctive dorsal pigmentation were first seen as calves off South Africa. These results strongly indicate connectivity between the two regions, while the presence off Namibia of three adult females from the South African population in the season in which they are believed to conceive suggests that there is unlikely to be any genetic differentiation between the two areas. We conclude that the reappearance of right whales off Namibia represents range expansion from South Africa rather than the survival of a few remnants of an originally separate stock.     

From the southern right whale hunting decline to the humpback whaling expansion: a review of whale catch records in the tropical western South Atlantic Ocean

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SEXUAL COMPOSITION and ANALYSIS OF REPRODUCTIVE FEMALES IN THE NORTH ATLANTIC RIGHT WHALE, EUBALAENA GLACIALIS, POPULATION

To test hypotheses involving reproduction and demographics, the sex of individuals must be established, but many species of Cetacea are not obviously dimorphic. In the North Atlantic right whale, Eubalaena glacialis , population, the sex of 61 males and 55 females had been determined previously by observation of the urogenital region, and the sex of 43 more females had been inferred from repeated sightings with a calf. To confirm the sex of some of these animals and to identify the sex of mote animals, genomic DNA was isolated from skin samples of 95 individual right whales (54 from among those described above and 41 additional recognizable individuals). The DNA was surveyed using the human Y-chromosome probe pDP1007. With Eco RI-digested DNA, a clear, sex-discriminating banding pattern was apparent. This method verified the sex of all 54 animals whose sex was previously known or inferred and identified the sex of an additional 41 recognizable individuals. A total of 89 male and 111 female right whales was identified in the population. The most unbiased estimate of sex ratio available is the 36 male and 34 female calves identified by genital morphology and DNA techniques. The sex ratio of this sample does not differ significantly from unity (P = 0.811). Only 38% (58/152) of the females in the North Atlantic population are known to have been reproductively successful compared with 54% in the population of right whales in the western South Atlantic. The population growth rate reported for the North Atlantic population is only 33% of that reported for right whales in the South Atlantic. Thirteen adult North Atlantic females have been identified that have not been known to calve during the past 11 yr. These data suggest that the absence of measurable recovery may be due to a combination of fewer actively reproducing females and lower reproductive rates of some females.     

Historic and current habitat use by North Pacific right whales Eubalaena japonica in the Bering Sea and Gulf of Alaska

1. To help define areas and ecological parameters critical to the survival and recovery of the remnant population of North Pacific right whales, habitat use was investigated by examining all available sighting and catch records in the south-eastern Bering Sea (SEBS) and Gulf of Alaska (GOA) over the past two centuries. 2. Based on re-analyses of commercial whaling records, search effort, and resultant catches and sightings, waters of the: (i) SEBS slope and shelf, (ii) eastern Aleutian Islands and (iii) GOA slope and abyssal plain were important habitat for North Pacific right whales through the late 1960s. 3. Since 1980, the only area where right whales have been seen consistently is on the SEBS middle shelf. However, acoustic detections and single sightings have been reported in all other regions except the SEBS slope and oceanic GOA (areas where little, if any, acoustic and visual effort has occurred). 4. Sightings since 1979 were in waters < 200 m deep which may simply reflect the paucity of search effort elsewhere. From the commercial whaling era to the late 1960s, right whales were commonly seen in waters > 2000 m deep, indicating that their distribution is not restricted to shallow continental shelves. 5. North Pacific right whale sightings through the centuries have been associated with a variety of oceanic features, and there is little in common in the bathymetry of these regions. These whales appear to have a greater pelagic distribution than that observed in the North Atlantic, which may be related to the availability of larger copepods across the SEBS and GOA.     

MIGRATION AND POPULATION STRUCTURE OF NORTHEASTERN PACIFIC WHALES OFF COASTAL BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS FROM 1908-1967

Data recorded from 24,862 whales killed by British Columbia coastal whaling stations between 1908 and 1967 revealed trends in the abundance, sex ratios, age structure, and distribution of sperm ( Physeter macrocephalus ), fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), humpback ( Megaptera novaeangliae ), and blue ( Balaenoptera musculus ) whales. The catch data were analyzed using annual and monthly mean values. Monthly and annual variation in whaling effort was deduced from accounts of the history of British Columbia coastal whaling, and biases arising from changes in effort were considered in the interpretation of the results. During the later years of whaling (1948-1967), the mean lengths of captured whales declined significantly and pregnancy rates dropped to near zero in fin, sei, and blue whales. Monthly patterns in numbers killed revealed a summer migration of sei and blue whales past Vancouver Island, and confirms anecdotal suggestions that local populations of fin and humpback whales once spent extended periods in the coastal waters of British Columbia. Furthermore, the data strongly suggest that sperm whales mated (April-May) and calved (July-August) in British Columbia's offshore waters. The historic whaling records reveal much about the migratory behavior and distribution of the large whales species as they once were, and may continue to be, in the northeastern Pacific. Verifying the persistence of these trends in the remnant populations is a necessary and logical next step.     

A citizen science approach to long-term monitoring of humpback whales (Megaptera novaeangliae) off Sydney,Australia

The Cape Solander Whale Migration Study is a citizen science project that annually counts northward migrating humpback whales (Megaptera novaeangliae) off Cape Solander, Sydney, Australia. Dedicated observers have compiled a 20-year data set (1997–2017) of shore-based observations from Cape Solander's high vantage point. Using this long-term data set collected by citizen scientists, we sought to estimate the humpback whale population trend as it continues to recover postexploitation. We estimated an exponential growth rate of 0.099 (95% CI = 0.079–0.119) using a generalized linear model, based on observer effort (number of observation days) and number of whales observed, equating to 10% per annum growth in sightings since 1997. We found that favorable weather conditions for spotting whales off Cape Solander consisted of winds <30 km/hr from a southerly through a north westerly direction. Incidental observations of other cetacean species included the endangered blue whale (Balaenoptera musculus) and data deficient species such as killer whales (Orcinus orca) and false killer whales (Pseudorca crassidens). Citizen science-based studies can provide a cost-effective approach to monitoring wildlife over the time necessary to detect change in a population. Information obtained from citizen science projects like this may help inform policy makers responsible for State and Federal protection of cetaceans in Australian waters and beyond.     

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.     

Photo-identification and satellite telemetry connect southern right whales from South Georgia Island (Islas Georgias del Sur) with multiple feeding and calving grounds in the southwest Atlantic

The sub-Antarctic waters of South Georgia Island (Islas Georgias del Sur, SG/IG) are a regularly visited feeding ground for southern right whales (Eubalaena australis, SRW) in the southwest Atlantic. Satellite telemetry and photo-identification records were compared to better understand the role of SG/IG in the SRW migratory network. We present the first insights from SRW satellite-tracked from the SG/IG feeding ground, habitat use patterns in the Scotia Arc, and movements to Antarctic habitats. Photo-identification comparisons to calving and feeding areas across the South Atlantic and a review of sightings of cetaceans reported from Bird Island (west of SG/IG) since 1979 illuminate long-term habitat use patterns in SG/IG. We present the first recorded migratory movement between SG/IG and multiple countries: Argentina, Uruguay, and Brazil. Photo-identification (1) linked SG/IG to a female SRW with a long-term sighting history in Brazil, and (2) provided the first match between SG/IG and the western Antarctic Peninsula, suggesting the latter could extend the feeding area for southwest Atlantic SRW. Satellite tracking and opportunistic sightings suggest that shelf and coastal waters west of SG/IG represent an important multi-season SRW feeding habitat and add to our overall understanding of habitats and ranges occupied by recovering southwest Atlantic SRW.     

CATCHES OF HUMPBACK AND OTHER WHALES FROM SHORE STATIONS AT MOSS LANDING AND TRINIDAD, CALIFORNIA, 1919–1926

Logbook data from California shore whaling stations at Moss Landing (1919–1922 and 1924) and Trinidad (1920 and 1922–1926) are analyzed. The logs for the two stations record the taking of 2,111 whales, including 1,871 humpbacks, 177 fin whales, 26 sei whales, 3 blue whales, 12 sperm whales, 7 gray whales, 1 right whale, 1 Baird's beaked whale, and 13 whales of unspecified type (probably humpbacks). Most whales were taken from spring to autumn, but catches were made in all months of some years. The sex ratios of humpback, fin, and sei whales (the three species with sufficient sample sizes to test) did not differ from parity. Primary prey, determined from stomach contents, included sardines and euphausiids for both humpback and fin whales, and 'plankton' (probably euphausiids) for sei whales. The prevalence of pregnancy was 0.46 among mature female humpbacks and 0.43 among mature female fin whales, although these values are reported with caution. Information on length distribution for all species is summarized. Analysis of the catch data for this and other areas supports the current view that humpback whales along the west coast of the continental United States comprise a single feeding stock and also suggests that the present population is well below pre-exploitation levels.     

Bowhead whales Balaena mysticetus in the Okhotsk Sea

• 1 Little is known about the endangered population of bowhead whales Balaena mysticetus in the Okhotsk Sea (OS). Here, we review existing information about this stock, including much material published in Russian.
• 2 Whaling for OS bowheads began around 1846, was pursued intensively for two decades and continued sporadically until about 1913. Beginning in 1967, whalers from the USSR killed bowheads illegally, although the number of whales taken remains unknown. Estimates of the pre‐exploitation population size have ranged from 3000 to 20000 whales, but all such estimates are based upon untested assumptions and incomplete data.
• 3 Information on historical and current distribution of bowheads comes from whaling records (notably Townsend 1935 ) and from modern (notably Russian/Soviet) marine mammal surveys. Little is known about winter distribution. During spring and summer, known bowhead concentrations occur in Shelikhov Bay and at Shantar. Although historical whaling data show bowheads in Shelikhov Bay during summer and early autumn, there have been no recent sightings later than June. However, extensive 19th century catches were made over much of the northern OS, and the present range and habitat use of the population is probably broader than existing data suggest. There is evidence for age or maturational class segregation between Shantar and Shelikhov Bay; the former hosts immature whales and lactating females, and the latter hosts adults.
• 4 Genetic data indicate that the OS bowhead stock is separate from the Bering‐Chukchi‐Beaufort population, but that the two populations share a common ancestry. There is no evidence that bowheads ever leave the OS.
• 5 Russian observers have put the current size of the OS stock in the low hundreds, but this is not based on quantitative analysis. Overall, the OS bowhead population is very likely to be relatively small; it did not recover from the intensive whaling in the 19th century, and the illegal Soviet catches of the 1960s have further set back its recovery. Dedicated surveys and other research are required to assess the status and conservation needs of the 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.     

Genetic diversity and structure of blue whales (Balaenoptera musculus) in Australian feeding aggregations

The worldwide distribution of blue whales (Balaenoptera musculus) has not prevented this species from becoming endangered due to twentieth century whaling. In Australia there are two known feeding aggregations of blue whales, which most likely are the pygmy subspecies (B. m. brevicauda). It is unknown whether individuals from these feeding aggregations belong to one breeding stock, or multiple breeding stocks that either share or occupy separate feeding grounds. This was investigated using ten microsatellite loci and mitochondrial DNA control region sequences (N = 110). Both sets of markers revealed no significant genetic structure, suggesting that these whales are likely to belong to the same breeding stock.     

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.     

BLUE WHALE (BALAENOPTERA MUSCULUS)DISTRIBUTION IN THE EASTERN TROPICAL PACIFIC

The distribution of blue whales, Balaenoptera musculus , in the eastern tropical Pacific (ETP) was analyzed from 211 sightings of 355 whales recorded during research vessel sighting surveys or by biologists aboard fishing vessels. Over 90% of the sightings were made in just two areas: along Baja California, and in the vicinity of the Costa Rica Dome (a large, stationary eddy centered near 9°N, 89°W), with the rest made along the equator near the Galapagos islands, the coasts of Ecuador and northern Peru. All sightings occurred in relatively cool, upwelling-modified waters. Because these areas are the most productive parts of the ETP, and have relatively large standing stocks of euphausiids, it seems possible that blue whales select low latitude habitats which permit foraging. The waters off western Baja California were occupied seasonally, with a peak in sightings coinciding with the spring peak in upwelling and biological production. The Costa Rica Dome area was occupied year round, suggesting either a resident population, or that both northern and southern hemisphere whales visit, with temporal overlap. The modal group size was one for all areas and seasons, but the frequency of groups with two or more whales was significantly higher in sightings made near the Galapagos Islands and the coast of Ecuador and northern Peru.     

Do ship strikes threaten the recovery of endangered eastern North Pacific blue whales?

Blue whales were targeted in the North Pacific from 1905–1971 and are listed as endangered by the IUCN. Despite decades without whaling, abundance estimates for eastern North Pacific (ENP) blue whales (Balaenoptera musculus) suggest little evidence for a recent increase. One possible reason is fatal strikes by large ships, which have affected populations of other cetaceans and resulted in mitigation. We used a population dynamics model to assess the trends and status of ENP blue whales, and the effects of ship strikes. We estimate the population likely never dropped below 460 individuals, and is at 97% of carrying capacity (95% interval 62%–99%). These results suggest density dependence, not ship strikes, is the key reason for the observed lack of increase. We also estimate future strikes will likely have a minimal impact; for example, an 11‐fold increase in vessels would lead to a 50% chance the long‐term population would be considered depleted. Although we estimate ship strike mitigation would have minimal impacts on population trends and status, current levels of ship strikes are likely above legal limits set by the U.S. The recovery of ENP blue whales from whaling demonstrates the ability of blue whale populations to rebuild under careful management.     

Distribution and migratory destinations of humpback whales off the Pacific coast of Central America during the boreal winters of 1996–2003

Here, we examine the distribution, habitat use, and migratory destinations of North Pacific humpback whales wintering off Central America. Coastal boat surveys were conducted off Costa Rica and Panama between 1996 and 2003. In 1999, a broader survey was conducted along most of Central America. Over 23,000 km were surveyed, with the greatest effort off southern Costa Rica. We made 191 sightings of 320 individual humpback whales. Whales were seen between 14°N and 8°N, making this the most southerly of the North Pacific wintering areas. Encounters included singles, adult pairs, singers, and mother/calf pairs. Mother/calf pairs accounted for 27% of all groups sighted, which is one of the highest sighting rates reported among North Pacific wintering areas. Sixty percent of sightings occurred in depths <50 m. Average sea surface temperature was 28.6°C (±1.0 SD). Ninety percent of the 77 unique whales photo‐identified were also seen in the California–Oregon–Washington feeding aggregation. The 1999 survey showed that humpback whales were widely distributed along the Central American coast at relatively low densities. The extensive distribution of animals, the higher proportion of calves, and the almost exclusive migration to a single feeding area contrast with observations in other regions.