Characterization of 25 microsatellite loci in bowhead whales (Balaena mysticetus)

Bowhead whales (Balaena mysticetus) experienced a severe demographic population bottleneck caused by commercial whaling that ceased in 1914. Aboriginal subsistence whale harvests have continued and are managed by the International Whaling Commission. In an effort to provide management advice for bowhead whales, 25 microsatellite loci were isolated from genomic DNA libraries. This panel of markers will be utilized to analyse stock structure hypotheses of current bowhead whale populations.     

OPEN-BOAT WHALING ON THE STRAITS OF GIBRALTAR GROUND AND ADJACENT WATERS

Logbooks ( n = 317) from whaling expeditions made in the North Atlantic during the 19th century were examined to investigate activity in the Gibraltar Straits grounds. At least forty expeditions of whaling vessels from European and American ports visited the area. In all cases the main target was the sperm whale, but pilot whales, dolphins, sea turtles, and even a blue whale were also taken. Whaling effort concentrated on the Atlantic side of the Straits; only two expeditions ventured into the Mediterranean Sea, obtaining negligible catches. The whaling season extended during spring and summer and peaked in June–July. This seasonality appeared not to be governed by changes in whale density but by the trade winds necessary to sail southward or westward to cross the Atlantic. Searching effort continued while trying out, but the rate of sighting cetaceans was about half that of searching periods. However, the rate of sighting or capturing a sperm whale remained unchanged during processing, probably because the gregarious habits of the species produced clumping of catches. For every whale secured, 1.31 whales were struck. After correcting for struck but lost whales and for "gammed" vessels, the minimum number of removals of sperm whales during 1862–1889 is estimated at 237.     

The origins and character of 'aboriginal subsistence' whaling: a global review

The International Whaling Commission (IWC) recognizes aboriginal subsistence whaling to be distinct from commercial whaling, and these two broad categories of whaling are subject to different management approaches. This paper describes recent, ongoing and likely future whale hunts that qualify, or may qualify, for aboriginal subsistence status within the IWC’s management framework. To ensure conservation of the whale populations, a forthright exposition of the origins, development and character of these hunts is needed in addition to stock assessment, a risk‐averse catch limit algorithm and appropriate mechanisms within the whaling communities to ensure compliance. The hunts for Bowhead Whales (Balaena mysticetus) and Gray Whales (Eschrichtius robustus) in the Arctic and North Pacific, respectively, and Sperm Whales (Physeter macrocephalus) in Indonesia have long histories and local origins. Those for Humpback Whales (Megaptera novaeangliae) in the Lesser Antilles and at Tonga in the South Pacific were introduced by foreign commercial whalers. Whale hunting in the Philippines appears to have originated both locally and as a result of foreign influence. The relatively recent initiation of whaling for Fin Whales (Balaenoptera physalus) and Minke Whales (Balaenoptera acutorostrata) in Greenland required introduced technology but can be viewed as a modern adaptation of an ancient tradition. Consensus in deciding how to classify and manage non‐industrial whaling has been, and will remain, elusive. Even with common definitions of key terms such as ‘subsistence’, ‘commercial’ and ‘aboriginal’, interpretations will depend on whether one’s priorities are whale‐centred or human‐centred.     

Molecular genetic identification of whale and dolphin products from commercial markets in Korea and Japan

We report the methods and results of molecular genetic identification of the species and, in some cases, geographical origins of whale and dolphin products purchased from retail markets and restaurants in Japan and South Korea. As reported previously (Baker & Palumbi 1994), we used the polymerase chain reaction (PCR) and a portable laboratory to amplify, purify and later sequence a portion of the mitochondrial DNA control region from 16 commercial products purchased in Japan. This ‘spot check’ revealed a surprising variety of species for sale, including minke, fin and humpback whales and one or two species of dolphins sold as ‘kujira’ or whale. In the Korean survey, DNA amplifications were conducted by two of us (C.S.B. and F.C.) working with independent equipment and reagents. The two sets of DNA amplifications were returned to our respective laboratories and sequenced independently for cross-validation. Among the total of 17 species-specific sequences we found a dolphin, a beaked whale, 13 Northern Hemisphere minke whales (representing at least seven distinct individuals) and two whales which are closely related to the recognized sei and Bryde's whales but could not be identified as either using available type sequences. We suggest that these two specimens represent a currently unrecognized species or subspecies of Bryde's whale, possibly the so-called ‘small-form’ reported from the tropical waters of the Indo-Pacific. We conclude that molecular systematic analyses of DNA sequences have tremendous utility for the identification of whale and dolphin products. However, there are certain constraints on the application of these techniques for monitoring whaling or trade in whale products. First, PCR and DNA sequencing can generate misleading artefacts. These can generally be recognized or eliminated through experimental controls. Second, phylogenetic reconstructions of DNA sequences can be misinterpreted if the database of type sequences is inadequate or the taxonomy of the group is incomplete. This constraint is, at present, a more serious obstacle to molecular monitoring of whaling. Our results highlight uncertainties about the taxonomic status of oceanic populations and morphological forms of two species (or species complexes) targeted by legal and illegal hunting, the minke and Bryde's whales. Despite these uncertainties, it is difficult to reconcile some of the species available in Japanese and Korean commercial markets with recent catch records made available to the International Whaling Commission. It is particularly disturbing that two specimens of an unrecognized species or subspecies of baleen whale were for sale in a restaurant in South Korea in October, 1994, 8 years after the acceptance of an international moratorium on commercial whaling.     

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.     

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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Two Intense Decades of 19th Century Whaling Precipitated Rapid Decline of Right Whales around New Zealand and East Australia

Right whales (Eubalaena spp.) were the focus of worldwide whaling activities from the 16^th to the 20^th century. During the first part of the 19^th century, the southern right whale (E. australis) was heavily exploited on whaling grounds around New Zealand (NZ) and east Australia (EA). Here we build upon previous estimates of the total catch of NZ and EA right whales by improving and combining estimates from four different fisheries. Two fisheries have previously been considered: shore-based whaling in bays and ship-based whaling offshore. These were both improved by comparison with primary sources and the American offshore whaling catch record was improved by using a sample of logbooks to produce a more accurate catch record in terms of location and species composition. Two fisheries had not been previously integrated into the NZ and EA catch series: ship-based whaling in bays and whaling in the 20^th century. To investigate the previously unaddressed problem of offshore whalers operating in bays, we identified a subset of vessels likely to be operating in bays and read available extant logbooks. This allowed us to estimate the total likely catch from bay-whaling by offshore whalers from the number of vessels seasons and whales killed per season: it ranged from 2,989 to 4,652 whales. The revised total estimate of 53,000 to 58,000 southern right whales killed is a considerable increase on the previous estimate of 26,000, partly because it applies fishery-specific estimates of struck and loss rates. Over 80% of kills were taken between 1830 and 1849, indicating a brief and intensive fishery that resulted in the commercial extinction of southern right whales in NZ and EA in just two decades. This conforms to the global trend of increasingly intense and destructive southern right whale fisheries over time.     

The Relationship between Vessel Traffic and Noise Levels Received by Killer Whales (Orcinus orca)

Whale watching has become increasingly popular as an ecotourism activity around the globe and is beneficial for environmental education and local economies. Southern Resident killer whales (Orcinus orca) comprise an endangered population that is frequently observed by a large whale watching fleet in the inland waters of Washington state and British Columbia. One of the factors identified as a risk to recovery for the population is the effect of vessels and associated noise. An examination of the effects of vessels and associated noise on whale behavior utilized novel equipment to address limitations of previous studies. Digital acoustic recording tags (DTAGs) measured the noise levels the tagged whales received while laser positioning systems allowed collection of geo-referenced data for tagged whales and all vessels within 1000 m of the tagged whale. The objective of the current study was to compare vessel data and DTAG recordings to relate vessel traffic to the ambient noise received by tagged whales. Two analyses were conducted, one including all recording intervals, and one that excluded intervals when only the research vessel was present. For all data, significant predictors of noise levels were length (inverse relationship), number of propellers, and vessel speed, but only 15% of the variation in noise was explained by this model. When research-vessel-only intervals were excluded, vessel speed was the only significant predictor of noise levels, and explained 42% of the variation. Simple linear regressions (ignoring covariates) found that average vessel speed and number of propellers were the only significant correlates with noise levels. We conclude that vessel speed is the most important predictor of noise levels received by whales in this study. Thus, measures that reduce vessel speed in the vicinity of killer whales would reduce noise exposure in this population.     

Medieval and Early Modern Whaling in Portugal

ABSTRACT

Mainland Portugal is not renowned for having been a whaling nation of significance. However, preliminary studies have brought to light enough historical references to suggest that whaling occurred from at least the 13th century, and the present work identifies 38 historical sources documenting whale use or whaling on the Portuguese coast between 1201 and 1728. A peak of whale-related sources occurred during the 13th and 14th centuries, and almost all Portuguese accounts are contemporary to those found from the French and Spanish Basque countries, such that the beginning of the whaling activity seems to be coeval. No geographical cluster of whaling activities can be established—they seem to have been unevenly scattered along the entire coastline. Nor can a chronological north–south movement of coastal whaling activities be discerned. The geographical and chronological patterns give support to the assumption that whaling was not introduced to Portugal by the Basques, who are known to have spread westward from the French Labourd (11th century), via Golf of Biscay, to Asturias, and southward to Galicia (14th century). Rather, Portuguese whale use seems to have originated independently of Basque influence. Several of the sources specify “black whales” as the target species. This is consistent with modern knowledge about the distribution and migration patterns of North Atlantic right whales during Basque medieval and early modern whaling. The Portuguese sources are not clear as to numbers of whales taken, nor to the whaling technology used, but the activity was sufficiently well organized and developed to warrant the levying of tithes in the feudal system of 13th-century Portugal.     

ICJ's Decision in Australia v. Japan: Giving up the Spear or Refining the Scientific Design?

The case Whaling in the Antarctic (Australia v. Japan) decided by the International Court of Justice is a landmark that introduces new parameters for measuring the “reasonableness” of scientific research by permit under the International Convention on Whaling. However, aspects of these parameters and how they may be applied in future cases remain uncertain. Because the Court's interpretation of the language “for purposes of scientific research” avoids defining scientific research, the Court's decision provides only a limited degree of clarification for States that intend to operate scientific whaling programs under Article VIII of the Convention. The Court's reasonableness test is unlikely to prevent scientific whaling. States who no longer support the dual object and purpose of the Whaling Convention may want to consider negotiating a new international instrument that would be more protective of whales and their habitat.     

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.     

Global distribution of fin whales Balaenoptera physalus in the post‐whaling era (1980–2012)

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Baleen whales: conservation issues and the status of the most endangered populations

Most species of baleen whales were subject to intensive overexploitation by commercial whaling in this and previous centuries, and many populations were reduced to small fractions of their original sizes. Here, we review the status of baleen whale stocks, with an emphasis on those that are known or thought to be critically endangered. Current data suggest that, of the various threats potentially affecting baleen whales, only entanglement in fishing gear and ship strikes may be significant at the population level, and then only in those populations which are already at critically low abundance. The impact of some problems (vessel harassment, and commercial or aboriginal whaling) is at present probably minor. For others (contaminants, habitat degradation, disease), existing data either indicate no immediate cause for concern, or are insufficient to permit an assessment. While the prospect for many baleen whales appears good, there are notable exceptions; populations that are of greatest concern are those suffering from low abundance and associated problems, including (in some cases) anthropogenic mortality. These include: all Northern Right Whales Eubalaena glacialis, Bowhead Whales Balaena mysticetus of the Okhotsk Sea and various eastern Arctic populations, western Gray Whales Eschrichtius robustus, and probably many Blue Whale Balaenoptera musculus populations. We review the status of these populations and, where known, the issues potentially affecting their recovery. Although Humpback Whales Megaptera novaeangliae and Southern Right Whales Eubalaena australis were also heavily exploited by whaling, existing data indicate strong recovery in most studied populations of these species.     

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.     

Using pre- and postexploitation samples to assess the impact of commercial whaling on the genetic characteristics of eastern North Pacific gray and humpback whales and to compare methods used to infer historic demography

Many species of whales went through recent bottlenecks due to commercial whaling. These declines were rapid and recent relative to the life spans and generation times of these species, raising questions regarding to what degree commercial whaling influenced the genetic characteristics of these populations. We analyzed mitochondrial and nuclear DNA from pre- and postwhaling samples from two populations that have arguably shown the greatest degree of recovery: eastern North Pacific gray and humpback whales. We also compare the performance of different methods to test for historic bottlenecks and infer past demography based on genetic data. We found substantially higher levels of genetic diversity in gray than in humpback whales (for both time periods), likely due to recent connectivity between Atlantic and Pacific gray whale populations. Other than mitochondrial diversity in humpback whales, levels of diversity were not lower in contemporary samples relative to prewhaling samples, indicating that commercial whaling had a minimal impact on metrics of genetic diversity themselves. However, it did have large impacts on the patterns of diversity, as evidenced by all coalescent-based methods showing clear evidence of a bottleneck for both populations, whereas all but one method not based on the coalescent failed to detect a bottleneck.     

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.     

RELATIVE ABUNDANCE OF LARGE WHALES AROUND SOUTH GEORGIA (1979–1998)1

To assess large-whale stocks following the cessation of land-based South Georgia whaling in 1965, we report three independent sighting databases: a cruise in 1997, observations from Bird Island (NW of South Georgia) between 1979 and 1998, and mariner sightings between 1992 and 1997. All species were rare, with sightings of southern right whales being the most common event. Two right whales photographed off South Georgia matched animals known from Peninsula Valdés, Argentina, a population known to be growing at 7%per annum. In contrast, blue and fin whales appeared to be less abundant. A single blue whale mother-calf pair was observed off the Shag Rocks in February 1997. Extirpation of animals from this particular feeding ground is the most likely reason for ongoing low numbers of all species. Other factors may include competition for krill by traditional predators such as penguins and seals and more recently by humans, an unusually high rate of natural mortality, habitat change such as alteration in sea ice coverage, and/or the impact of ongoing whaling. The history of this critical area of large-whale habitat and this report demonstrate the need for improved, consistent longterm monitoring of population trends for these depleted stocks.     

Pre-whaling genetic diversity and population ecology in eastern Pacific gray whales: insights from ancient DNA and stable isotopes

Commercial whaling decimated many whale populations, including the eastern Pacific gray whale, but little is known about how population dynamics or ecology differed prior to these removals. Of particular interest is the possibility of a large population decline prior to whaling, as such a decline could explain the ~5-fold difference between genetic estimates of prior abundance and estimates based on historical records. We analyzed genetic (mitochondrial control region) and isotopic information from modern and prehistoric gray whales using serial coalescent simulations and Bayesian skyline analyses to test for a pre-whaling decline and to examine prehistoric genetic diversity, population dynamics and ecology. Simulations demonstrate that significant genetic differences observed between ancient and modern samples could be caused by a large, recent population bottleneck, roughly concurrent with commercial whaling. Stable isotopes show minimal differences between modern and ancient gray whale foraging ecology. Using rejection-based Approximate Bayesian Computation, we estimate the size of the population bottleneck at its minimum abundance and the pre-bottleneck abundance. Our results agree with previous genetic studies suggesting the historical size of the eastern gray whale population was roughly three to five times its current size.     

Fin whales in the NE Atlantic; relationships between abundance and distribution

It has been a widely held opinion that several fin whale populations inhabit the NE Atlantic. In the present communication the existence of one versus several fin whale populations in this area was considered. A survey of catch figures indicated considerable similarities among the various whaling grounds. Thus the figures did not support the belief that the various grounds were inhabited by different populations. The low productivity of the whales and their good mobility in an environment where the effects of physical barriers are limited were among the factors discussed as acting against the origination of separate populations. A model, ("the funnel analogy"), was presented according to which decrease in density would first be expected at the periphery of distribution. The area W of Iceland was suggested to constitute the present distributional core of the fin whale in the NE Atlantic. An account was given of the catch of fin whales in Iceland during the past 32 yr. Catch figures calibrated for Increased catcher efficiencies showed a falling tendency.     

DNA profile of a sixteenth century western North Atlantic right whale (Eubalaena glacialis)

Low levels of genetic variability identified within the North Atlantic right whale (Eubalaena glacialis), when compared to the Southern right whale (E. australis) and other large whales, have been suggested to result from population reductions due to whaling. Previous genetic analysis of 218 whale bones from sixteenth century Basque whaling sites in the western North Atlantic revealed only a single right whale bone. We determined the genotypes of 27 microsatellite loci using DNA isolated from this bone. All alleles from the historic specimen occur in the extant western North Atlantic population and both the probability of identity of the specimen and the number of heterozygous loci are similar to that in the extant population. Assessments of how genetically different the historical population might have been suggest genetic characteristics have not changed substantially over four centuries of whaling.