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.     

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.     

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.     

Past and present distributions of southern right whales(Eubalaena australis)

Abstract

During the 19th century, sail whalers hunted right whales throughout the southern oceans north of 50 degrees. This review seeks to document the recovery of southern right whales by comparing the northernmost recent sightings with older sightings that survive “fossilised” in historical records of the sail whaling era. Despite a steady expansion northwards in recent years, the present distribution of right whales is still far short of that recorded by sail whalers over a century ago.     

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.

     

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.     

Wal und Mensch

Whale and Man The history of the interaction between man and whale is changeful. As in the past, also today, these gentle giants provide enough material for tales and modern mysteries. The excessive whaling of the past century almost led to extinction of several baleen whale species. Whaling distinguishs between commercial, aboriginal subsistence and scientific whaling. Since the whaling moratorium of the International Whaling Commission came into effect 1986, the stocks of the great whales developed differently. We deliver insight into the history of whaling and demonstrate the current threats to cetaceans, bycatch, entanglement, ship strike, accidents, marine pollution and climate change. Whale‐watching is a fast growing section of tourism industry, however, this form of wildlife watching may have negative consequences for the whales.     

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.     

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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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.     

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.     

KILLER WHALES AND MARINE MAMMAL TRENDS IN THE NORTH PACIFIC—A RE-EXAMINATION OF EVIDENCE FOR SEQUENTIAL MEGAFAUNA COLLAPSE AND THE PREY-SWITCHING HYPOTHESIS

Springer et al . (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling's removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al ., and much of the reduction occurred 50–100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al . suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al .) were likely abundant throughout the period. Overall, we suggest that the Springer et al . hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem.     

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.     

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.     

Whales sustain fisheries: Blue whales stimulate primary production in the Southern Ocean

It has previously been asserted that baleen whales compete with fisheries by consuming potentially harvestable marine resources. The regularly applied “surplus‐yield model” suggests that whale prey becomes available to fisheries if whales are removed, and has been presented as a justification for whaling. However, recent findings indicate that whales enhance ecosystem productivity by defecating iron that stimulates primary productivity in iron‐limited waters. While juvenile whales and whales that are pregnant or lactating retain iron for growth and milk production, nonbreeding adult whales defecate most of the iron they consume. Here, we modify the surplus‐yield model to incorporate iron defecation. After modeling a simplistic trajectory of blue whale recovery to historical abundances, the traditional surplus‐yield model predicts that 10¹¹ kg of carbon yr^?1 would become unavailable to fisheries. However, this ignores the nutrient recycling role of whales. Our model suggests the population of blue whales would defecate 3 × 10⁶ kg of iron yr^?1, which would stimulate primary production equivalent to that required to support prey consumption by the blue whale population. Thus, modifying the surplus‐yield model to include iron defecation indicates that blue whales do not render marine resources unavailable to fisheries. By defecating iron‐rich feces, blue whales promote Southern Ocean productivity, rather than reducing fishery yields.     

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

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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.     

PATTERNS OF BOWHEAD WHALE DISTRIBUTION AND ABUNDANCE NEAR BARROW, ALASKA, IN FALL 1982-1989

Although the distribution and relative abundance of bowhead whales varied annually within the fall whaling area near Barrow, Alaska, the distance of whales from shore was not significantly different among years 1982-1989 (ANOVA, F = 0.5, P > 0.5). The minimum detectable distance for the ANOVA was 12 km (α= 0.05, β= 0.1). Annual median distance of random bowhead sightings from shore ranged from 23 to 39 km, with an eight-year median of 32 km. Highest annual bowhead sighting rates were positively associated with the proportion of feeding whales, indicating that whale feeding opportunities may affect the availability of whales within hunting range each fall.     

Killer whales and whaling: the scavenging hypothesis

Killer whales (Orcinus orca) frequently scavenged from the carcasses produced by whalers. This practice became especially prominent with large-scale mechanical whaling in the twentieth century, which provided temporally and spatially clustered floating carcasses associated with loud acoustic signals. The carcasses were often of species of large whale preferred by killer whales but that normally sink beyond their diving range. In the middle years of the twentieth century floating whaled carcasses were much more abundant than those resulting from natural mortality of whales, and we propose that scavenging killer whales multiplied through diet shifts and reproduction. During the 1970s the numbers of available carcasses fell dramatically with the cessation of most whaling (in contrast to a reasonably stable abundance of living whales), and the scavenging killer whales needed an alternative source of nutrition. Diet shifts may have triggered declines in other prey species, potentially affecting ecosystems, as well as increasing direct predation on living whales.