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.     

How few whales were there after whaling? Inference from contemporary mtDNA diversity

Reconstructing the history of exploited populations of whales requires fitting a trajectory through at least three points in time: (i) prior to exploitation, when abundance is assumed to be at the maximum allowed by environmental carrying capacity; (ii) the point of minimum abundance or 'bottleneck', usually near the time of protection or the abandonment of the hunt; and (iii) near the present, when protected populations are assumed to have undergone some recovery. As historical abundance is usually unknown, this trajectory must be extrapolated according to a population dynamic model using catch records, an assumed rate of increase and an estimate of current abundance, all of which have received considerable attention by the International Whaling Commission (IWC). Relatively little attention has been given to estimating minimum abundance (N(min)), although it is clear that genetic and demographic forces at this point are critical to the potential for recovery or extinction of a local population. We present a general analytical framework to improve estimates of N(min) using the number of mtDNA haplotypes (maternal lineages) surviving in a contemporary population of whales or other exploited species. We demonstrate the informative potential of this parameter as an a posteriori constraint on Bayesian logistic population dynamic models based on the IWC Comprehensive Assessment of the intensively exploited southern right whales (Eubalaena australis) and published surveys of mtDNA diversity for this species. Estimated historical trajectories from all demographic scenarios suggested a substantial loss of mtDNA haplotype richness as a result of 19th century commercial whaling and 20th century illegal whaling by the Soviet Union. However, the relatively high rates of population increase used by the IWC assessment predicted a bottleneck that was implausibly narrow (median, 67 mature females), given our corrected estimates of N(min). Further, high levels of remnant sequence diversity (theta) suggested that pre-exploitation abundance was larger than predicted by the logistic model given the catch record, which is known to be incomplete. Our results point to a need to better integrate evolutionary processes into population dynamic models to account for uncertainty in catch records, the influence of maternal fidelity on metapopulation dynamics, and the potential for inverse density dependence (an 'Allee effect') in severely depleted populations.     

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 Search for a New Compliance Mechanism Within the International Whaling Commission

The final area of work that needs to be completed by the International Whaling Commission (IWC) before the moratorium on commercial whaling may be lifted is the Revised Management Scheme (RMS). The key issue stalling conclusion of the RMS is the failure to agree on the details of a suitable inspection, observation, and compliance regime. This contribution looks at the negotiations respecting the development of a suitable inspection, observation, and compliance regime.­     

The world's smallest whale population?

The North Pacific right whale (Eubalaena japonica) was heavily exploited by both nineteenth century whaling and recent (1960s) illegal Soviet catches. Today, the species remains extremely rare especially in the eastern North Pacific. Here, we use photographic and genotype data to calculate the first mark–recapture estimates of abundance for right whales in the Bering Sea and Aleutian Islands. The estimates were very similar: photographic = 31 (95% CL 23–54), genotyping = 28 (95% CL 24–42). We also estimated the population contains eight females (95% CL 7–18) and 20 males (95% CL 17–37). Although these estimates may relate to a Bering Sea subpopulation, other data suggest that the total eastern North Pacific population is unlikely to be much larger. Its precarious status today—the world''s smallest whale population for which an abundance estimate exists—is a direct consequence of uncontrolled and illegal whaling, and highlights the past failure of international management to prevent such abuses.     

Locomotor patterns, territory, and tunnel utilization in the mole-rat Spalax ehrenbergi

The northern bottlenose whale has been caught in the Faroe Islands for centuries, with written catch records going back to 1584 and unbroken from 1709. A total of 811 whales has been reported in the period 1584-1993. The Faroese bottlenose whaling is an opportunistic drive fishery of pods sighted very close to shore. Natural strandings also occur. Most of the fishery has taken place in two close southern villages of the Faroese archipelago (72% of the catch). The high season is 20 August-20 September. The pod contains 1-7 whales with an average of 2.1 whales. Most of them are immature males or mature females with juveniles, but as many males as females have been caught overall. Females and males at every stage of development have been caught in the Faroes, although it appears that the bottlenose whales approaching the Faroese coast and then driven ashore have not included as large and as small individuals as those shot offshore. A body weight (W in kg) and length (L in cm) relationship has been calculated for both sexes combined: W = 0.0000131 x L^3.07. Females and immature males have a grey and bulbous forehead. As the males mature their forehead becomes flatter and lighter, and only large mature males have a white and flat forehead. The stomach contents of nine whales contained in total at least 13 squid species. A comparison with pilot whaling shows that bottlenose whales arrive 2-4 weeks later than the pilot whales and that the geographical distribution of the catch is very different for both species, suggesting a different pattern of migration through the archipelago.     

Bones as biofuel: a review of whale bone composition with implications for deep-sea biology and palaeoanthropology

Whales are unique among vertebrates because of the enormous oil reserves held in their soft tissue and bone. These ‘biofuel’ stores have been used by humans from prehistoric times to more recent industrial-scale whaling. Deep-sea biologists have now discovered that the oily bones of dead whales on the seabed are also used by specialist and generalist scavenging communities, including many unique organisms recently described as new to science. In the context of both cetacean and deep-sea invertebrate biology, we review scientific knowledge on the oil content of bone from several of the great whale species: Balaenoptera musculus, Balaenoptera physalus, Balaenoptera borealis, Megaptera novaeangliae, Eschrichtius robustus, Physeter macrocephalus and the striped dolphin, Stenella coeruleoalba. We show that data collected by scientists over 50 years ago during the heyday of industrial whaling explain several interesting phenomena with regard to the decay of whale remains. Variations in the lipid content of bones from different parts of a whale correspond closely with recently observed differences in the taphonomy of deep-sea whale carcasses and observed biases in the frequency of whale bones at archaeological sites.     

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.     

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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World whale stocks

The history of whaling is very largely one of repeated over-exploitation of the various whale stocks which became available through discovery or technological advance. Modern whaling has similarly caused considerable reductions in the numbers of some species in the major whaling grounds. Stock assessment methods are based on catch and effort statistics, biological information including age and reproductive status, marking and sightings records. Catch effort data have to be used with caution, because of changes in species preference, shifts in the whaling grounds and national fleet variations. With allowance made for these factors, cumulative catches adjusted for recruitment can be used to estimate the initial stock number. Changes in stock density after known catches also lead to abundance estimates. Logarithmic regression of age composition data are used to find the total mortality rates. The natural mortality can be estimated from early season catches in a fishery or pre-fishery year classes caught more recently; fishing mortality is found by subtraction, which again leads to abundance estimates. Mathematical approaches incorporating recruitment estimates from actual age composition data and theoretical population models have been employed. Additional estimates come from mark release-recapture experiments and direct sightings counts from whaling vessels and research ships. The latter are the only means of estimating the protected species. The yields which the various stocks can sustain are calculated from direct observations and theoretical considerations of the changes in recruitment, largely due to increased pregnancy rates and the lower ages at sexual maturity which occur in exploited stocks. The results of all the available analyses have been compared and combined to produce the population estimates and yields tabulated. The object of whale management is to bring all stocks to the levels providing the maximum or optimum sustainable yields. These are defined in terms of numbers at the moment, but may be expressed as biomass in the future.     

Low genetic diversity in pygmy blue whales is due to climate-induced diversification rather than anthropogenic impacts

Unusually low genetic diversity can be a warning of an urgent need to mitigate causative anthropogenic activities. However, current low levels of genetic diversity in a population could also be due to natural historical events, including recent evolutionary divergence, or long-term persistence at a small population size. Here, we determine whether the relatively low genetic diversity of pygmy blue whales (Balaenoptera musculus brevicauda) in Australia is due to natural causes or overexploitation. We apply recently developed analytical approaches in the largest genetic dataset ever compiled to study blue whales (297 samples collected after whaling and representing lineages from Australia, Antarctica and Chile). We find that low levels of genetic diversity in Australia are due to a natural founder event from Antarctic blue whales (Balaenoptera musculus intermedia) that occurred around the Last Glacial Maximum, followed by evolutionary divergence. Historical climate change has therefore driven the evolution of blue whales into genetically, phenotypically and behaviourally distinct lineages that will likely be influenced by future climate change.     

What saved the whales? An economic analysis of 20th century whaling

Catches of whales show a historically cyclical pattern, with catches declining as stocks of the financially most attractive species fell, but expanding as substitute species were caught. Total combined catch peaked in the early 1960s and fell thereafter to the current regulated levels. While it is widely thought that international whaling agreements account for the current stable stock levels, economic analysis reveals that market forces leading to reduced catch were already in place well before the agreements took hold. To some extent, therefore, catches were destined to decline as whale products ceased to be commercially attractive on a large scale. Using econometric analysis, the paper shows the various forces at work: declining stocks, the rise of substitute products, internationally increasing environmentalism, and rising incomes. Of these forces, stock decreases, which resulted in high unit catch costs, and income growth, which reduced rather than increased demand, were the most important factors, with regulation following, rather than leading, catch changes.     

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.     

Legal aspects of the IWC decision on the southern ocean sanctuary

This comment is based on, and largely incorporates, a memorandum by the author submitted by Japan to the International Whaling Commission (IWC) at its forty‐eighth meeting in June 1996, in Aberdeen, Scotland. It reviews relevant international law principles of interpretation—including ordinary meaning, objectives and purposes, subsequent conduct, and evolutionary interpretation—in their application to the question of the validity of the IWC action in adopting a Southern Ocean Sanctuary, which forbids the commercial harvest of whales in that area regardless of their conservation status. The review concludes that the IWC acted contrary to the provisions of its charter, the 1946 International Convention for the Regulation of Whaling. An epilogue summarizes the IWC discussion of the memorandum, and the general conclusion that an action by the commission ipso facto determines that it complies with the convention, and suggests that this latest instance of ignoring treaty obligations raises the question of how long the IWC can survive when the majority of its members repudiate the objectives and purposes of the convention.     

Increasing abundance of bowhead whales in West Greenland

In April 2006, a dedicated survey of bowhead whales (Balaena mysticetus) was conducted on the former whaling ground in West Greenland to determine the current wintering population abundance. This effort included a double platform aerial survey design, satellite tracking of the movements of nine whales, and estimation of high-resolution surface time from 14 whales instrumented with time-depth recorders. Bowhead whales were estimated to spend an average of 24% (cv=0.03) of the time at or above 2m depth, the maximum depth at which they can be seen on the trackline. This resulted in a fully corrected abundance estimate of 1229 (95% CI: 495-2939) bowhead whales when the availability factor was applied and sightings missed by observers were corrected. This surprisingly large population estimate is puzzling given that the change in abundance cannot be explained by a recent or rapid growth in population size. One possible explanation is that the population, which demonstrates high age and sex segregation, has recently attained a certain threshold size elsewhere, and a higher abundance of mature females appears on the winter and spring feeding ground in West Greenland. This in combination with the latest severe reduction in sea ice facilitating access to coastal areas might explain the surprising increase in bowhead whale abundance in West Greenland.     

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.     

Modern whaling in Britain and the north-east Atlantic Ocean

Modern whaling, using an explosive harpoon fired from a steam catcher-boat to kill the fast-swimming rorquals, began from shore whaling stations in northern Norway in the 1860s. It spread to Iceland, the Faeroe Islands and Spitsbergen, before reaching the British Isles in 1903.
Whaling took place from four stations in the Shetland Islands, one in the Outer Hebrides, and two in Ireland, before the First World War. Fin whales were the main species caught but Blue, Humpback, Sei, Right, Sperm and Bottle-nosed whales were also taken. Four stations re-opened in 1920, but from 1923 onwards only two continued to operate and whaling ceased in 1929, though the Hebridean station worked again for two seasons in 1950–1951.
The species composition of catches at the Hebridean and Irish stations was very similar and different from that of the stations in the Shetland Islands where few Blue whales and Right whales were taken. There is evidence that Fin whales were being overfished on the Shetland Islands whaling grounds at an early date, and that Blue whales and Right whales, but not Fin whales, declined in numbers on the Hebridean grounds.
The history of modern whaling in the north-east North Atlantic region as a whole indicates that the stocks of Blue, Humpback and Right whales were not large enough to support continuous whaling on the scale which took place there. The development of whaling since 1945 supports the view that there are separate populations of Fin whales in the region. The numbers of this species have declined on the whaling grounds of the Faeroe Islands and western Norway, and possibly also of north Norway, but not on the Icelandic grounds where there is no evidence of overfishing.     

Soil water availability for plants as quantified by conventional available water, least limiting water range and integral water capacity

There are different approaches to define the soil available water (SAW) for plants. The objectives of this study are to evaluate the SAW values of 12 arable soils from Hamadan province (western Iran) calculated by plant available water (PAW), least limiting water range (LLWR) and integral water capacity (IWC) approaches and to explore their relations with Dexter’s index of soil physical quality (i.e., S-value). Soil water retention and mechanical resistance were determined on the intact samples which were taken from the 5–10 cm layer. For calculation of LLWR and IWC, the van Genuchten-Mualem model was fitted to the observed soil water retention data. Two matric suctions (h) of 100 and 330 cm were used for the field capacity (FC). There were significant differences (P?<?0.01) between the SAW values calculated by PAW₁₀₀, PAW₃₃₀, LLWR₁₀₀, LLWR₃₃₀ and IWC. The highest (i.e., 0.210 cm³ cm^?3) and the lowest (i.e., 0.129 cm³ cm^?3) means of SAW were calculated for the IWC and LLWR₃₃₀, respectively. The upper limit of LLWR₃₃₀ for all of the soils was h of 330 cm, and that of LLWR₁₀₀ (except for one soil that was air-filled porosity of 0.1 cm³ cm^?3) was h of 100 cm. The lower limit of LLWR₃₃₀ and LLWR₁₀₀ for five soils was h of 15,000 cm and for seven soils was mechanical resistance of 2 MPa. The IWC values were smaller than those of LLWR₁₀₀ for two soils, equal to those of LLWR₁₀₀ for three soils and greater than those of LLWR₁₀₀ for the rest. There is, therefore, a tendency to predict more SAW using the IWC approach than with the LLWR approach. This is due to the chosen critical soil limits and gradual changes of soil limitations vs. water content in the IWC calculation procedure. Significant relationships of SAW with bulk density or relative bulk density were found but not with the clay and organic matter contents. Linear relations between IWC and LLWR₁₀₀ or LLWR₃₃₀ were found as: IWC?=??0.0514 + 1.4438LLWR₁₀₀, R ²?=?0.83; and IWC?=??0.0405 + 2.0465LLWR₃₃₀, R ²?=?0.84, respectively (both significant at P?<?0.01). Significant relationships were obtained between the SAW values and S indicating the suitability of the index S to explain the availability of soil water for plants even when complicated approaches like IWC are considered. Overall, the results demonstrate the importance of the choice of the approach to be used and its critical limits in the estimation of the soil available water to plants.     

Modelling the past and future of whales and whaling

Historical reconstruction of the population dynamics of whales before, during and after exploitation is crucial to marine ecological restoration and for the consideration of future commercial whaling. Population dynamic models used by the International Whaling Commission require historical catch records, estimates of intrinsic rates of increase and current abundance, all of which are subject to considerable uncertainty. Population genetic parameters can be used for independent estimates of historical demography, but also have large uncertainty, particularly for rates of mutational substitution and gene flow. At present, demographic and genetic estimates of pre-exploitation abundance differ by an order of magnitude and, consequently, suggest vastly different baselines for judging recovery. Here, we review these two approaches and suggest the need for a synthetic analytical framework to evaluate uncertainty in key parameters. Such a framework could have broad application to modelling both historical and contemporary population dynamics in other exploited species.     

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.