Human-induced ecological cascades: Extinction, restoration, and rewilding in the Galápagos highlands.

Oceanic islands support unique biotas but often lack ecological redundancy, so that the removal of a species can have a large effect on the ecosystem. The larger islands of the Galápagos Archipelago once had one or two species of giant tortoise that were the dominant herbivore. Using paleoecological techniques, we investigate the ecological cascade on highland ecosystems that resulted from whalers removing many thousands of tortoises from the lowlands. We hypothesize that the seasonal migration of a now-extinct tortoise species to the highlands was curtailed by decreased intraspecific competition. We find the trajectory of plant community dynamics changed within a decade of the first whaling vessels visiting the islands. Novel communities established, with a previously uncommon shrub, Miconia, replacing other shrubs of the genera Alternanthera and Acalypha. It was, however, the introduction of cattle and horses that caused the local extirpation of plant species, with the most extreme impacts being evident after c. 1930. This modified ecology is considered the natural state of the islands and has shaped subsequent conservation policy and practice. Restoration of El Junco Crater should emphasize exclusion of livestock, rewilding with tortoises, and expanding the ongoing plantings of Miconia to also include Acalypha and Alternanthera.

Genomic Impact of Whaling in North Atlantic Fin Whales.

It is generally recognized that large-scale whaling in the 19th and 20th century led to a substantial reduction of the size of many cetacean populations, particularly those of the baleen whales (Mysticeti). The impact of these operations on genomic diversity of one of the most hunted whales, the fin whale (Balaenoptera physalus), has remained largely unaddressed because of the paucity of adequate samples and the limitation of applicable techniques. Here, we have examined the effect of whaling on the North Atlantic fin whale based on genomes of 51 individuals from Icelandic waters, representing three temporally separated intervals, 1989, 2009 and 2018 and provide a reference genome for the species. Demographic models suggest a noticeable drop of the effective population size of the North Atlantic fin whale around a century ago. The present results suggest that the genome-wide heterozygosity is not markedly reduced and has remained comparable with other baleen whale species. Similarly, there are no signs of apparent inbreeding, as measured by the proportion of long runs of homozygosity, or of a distinctively increased mutational load, as measured by the amount of putative deleterious mutations. Compared with other baleen whales, the North Atlantic fin whale appears to be less affected by anthropogenic influences than other whales such as the North Atlantic right whale, consistent with the presence of long runs of homozygosity and higher levels of mutational load in an otherwise more heterozygous genome. Thus, genome-wide assessments of other species and populations are essential for future, more specific, conservation efforts.

Ancestry testing of "Old Tom," a killer whale central to mutualistic interactions with human whalers.

Cooperative hunting between humans and killer whales (Orcinus orca) targeting baleen whales was reported in Eden, New South Wales, Australia, for almost a century. By 1928, whaling operations had ceased, and local killer whale sightings became scarce. A killer whale from the group, known as "Old Tom," washed up dead in 1930 and his skeleton was preserved. How these killer whales from Eden relate to other populations globally and whether their genetic descendants persist today remains unknown. We extracted and sequenced DNA from Old Tom using ancient DNA techniques. Genomic sequences were then compared with a global dataset of mitochondrial and nuclear genomes. Old Tom shared a most recent common ancestor with killer whales from Australasia, the North Atlantic, and the North Pacific, having the highest genetic similarity with contemporary New Zealand killer whales. However, much of the variation found in Old Tom's genome was not shared with these widespread populations, suggesting ancestral rather than ongoing gene flow. Our genetic comparisons also failed to find any clear descendants of Tom, raising the possibility of local extinction of this group. We integrated Traditional Custodian knowledge to recapture the events in Eden and recognize that Indigenous Australians initiated the relationship with the killer whales before European colonization and the advent of commercial whaling locally. This study rectifies discrepancies in local records and provides new insight into the origins of the killer whales in Eden and the history of Australasian killer whales.

Diversity of mitochondrial DNA in 3 species of great whales before and after modern whaling.

The 20th century commercial whaling industry severely reduced populations of great whales throughout the Southern Hemisphere. The effect of this exploitation on genetic diversity and population structure remains largely undescribed. Here, we compare pre- and post-whaling diversity of mitochondrial DNA (mtDNA) control region sequences for 3 great whales in the South Atlantic, such as the blue, humpback, and fin whale. Pre-whaling diversity is described from mtDNA extracted from bones collected near abandoned whaling stations, primarily from the South Atlantic island of South Georgia. These bones are known to represent the first stage of 20th century whaling and thus pre-whaling diversity of these populations. Post-whaling diversity is described from previously published studies reporting large-scale sampling of living whales in the Southern Hemisphere. Despite relatively high levels of surviving genetic diversity in the post-whaling populations, we found evidence of a probable loss of mtDNA lineages in all 3 species. This is evidenced by the detection of a large number of haplotypes found in the pre-whaling samples that are not present in the post-whaling samples. A rarefaction analysis further supports a loss of haplotypes in the South Atlantic humpback and Antarctic blue whale populations. The bones from former whaling stations in the South Atlantic represent a remarkable molecular archive for further investigation of the decline and ongoing recovery in the great whales of the Southern Hemisphere.

Recovery of carbon benefits by overharvested baleen whale populations is threatened by climate change.

Despite the importance of marine megafauna on ecosystem functioning, their contribution to the oceanic carbon cycle is still poorly known. Here, we explored the role of baleen whales in the biological carbon pump across the southern hemisphere based on the historical and forecasted abundance of five baleen whale species. We modelled whale-mediated carbon sequestration through the sinking of their carcasses after natural death. We provide the first temporal dynamics of this carbon pump from 1890 to 2100, considering both the effects of exploitation and climate change on whale populations. We reveal that at their pre-exploitation abundance, the five species of southern whales could sequester 4.0 × 105 tonnes of carbon per year (tC yr-1). This estimate dropped to 0.6 × 105 tC yr-1 by 1972 following commercial whaling. However, with the projected restoration of whale populations under a RCP8.5 climate scenario, the sequestration would reach 1.7 × 105 tC yr-1 by 2100, while without climate change, recovered whale populations could sequester nearly twice as much (3.2 × 105 tC yr-1) by 2100. This highlights the persistence of whaling damages on whale populations and associated services as well as the predicted harmful impacts of climate change on whale ecosystem services.

Adaptation of sperm whales to open-boat whalers: rapid social learning on a large scale?

Animals can mitigate human threats, but how do they do this, and how fast can they adapt? Hunting sperm whales was a major nineteenth century industry. Analysis of data from digitized logbooks of American whalers in the North Pacific found that the rate at which whalers succeeded in harpooning ('striking') sighted whales fell by about 58% over the first few years of exploitation in a region. This decline cannot be explained by the earliest whalers being more competent, as their strike rates outside the North Pacific, where whaling had a longer history, were not elevated. The initial killing of particularly vulnerable individuals would not have produced the observed rapid decline in strike rate. It appears that whales swiftly learned effective defensive behaviour. Sperm whales live in kin-based social units. Our models show that social learning, in which naive social units, when confronted by whalers, learned defensive measures from grouped social units with experience, could lead to the documented rapid decline in strike rate. This rapid, large-scale adoption of new behaviour enlarges our concept of the spatio-temporal dynamics of non-human culture.

Subcutaneous, abdominal, and thoracic encapsulated fat necrosis in bowhead whales Balaena mysticetus from Alaska, USA.

We describe a case series of encapsulated fat necrosis with subcutaneous, abdominal, and thoracic locations in 7 subsistence-harvested bowhead whales Balaena mysticetus. Masses had a variably-dense fibrous capsule surrounding necrotic adipocytes and calcium salts (saponification). One animal also had prior concussive injury, pleural fibrosis, and hepatic lipoma; the other animals had no significant findings. The described condition is uncommon in bowhead whales, with 7/575 (1.2%) observed from 1996 to 2015. The exact mechanisms of development of encapsulated fat necrosis in bowhead whales remain to be determined. Encapsulated fat necrosis has been reported in other baleen whales, humans, and cows. It is usually an incidental finding during post-mortem examination that needs to be differentiated from neoplastic and inflammatory lesions, as the latter may have public health implications. Assessment of further cases in bowhead whales and other baleen whales is warranted to better understand their pathogenesis.

Whaling, Bullfighting, and the Conditional Value of Tradition.

The paper develops an account of the value of tradition that completes that of Samuel Scheffler and employs it to discuss whaling and bullfighting. The discussion, however, is applicable to many other practices the paper describes, and its relevance extends also beyond animal ethics. Some of the arguments discussed here for maintaining these traditions appeal to their positive aspects, such as their contribution to social or environmental harmony; other arguments focus on the impermissibility of one group criticizing another group's practices when its own are vulnerable to comparable criticism. Reflecting on the first kind of argument, the paper responds, building on the work of G. A. Cohen and T. M. Scanlon, that the value of tradition, if any, must be conditional. Reflecting on the second, however, the paper disagrees with Cohen and Scanlon on the impermissibility of casting the first stone.

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.

Whales, science, and scientific whaling in the International Court of Justice.

I provide a brief review of the origins of the International Convention on the Regulation of Whaling and the failure to successfully regulate whaling that led to the commercial moratorium in 1986. I then describe the Japanese Whale Research Programs Under Special Permit in the Antarctica (JARPA I, JARPA II) and the origins of the case Whaling in the Antarctic (Australia v. Japan: New Zealand Intervening) in the International Court of Justice. I explain that the International Court of Justice chose to conduct an objective review of JARPA II, the standard that it used for the review, and the pathway that it took to adjudicate the case without providing a definition of science to be used in international law. I conclude with a brief discussion of the implications of the Judgment for the International Convention on the Regulation of Whaling, and the International Whaling Commission in particular, for other international treaties, and for the interaction of science and law more generally.

Cetacean citations and the covenant of iron.

By the early decades of the nineteenth century, with surveys established as the weapon of choice for the fiscal military state, their instrumentation provided a focal point for radical attacks on political establishments. This paper considers a notorious dispute over mastery of iron in the instrumentation of magnetic surveying that took place in the 1830s between an Admiralty committee and the Reverend William Scoresby, a whaler-turned-clergyman. Scoresby staked his claim by drawing on the labour law of the whaleboats, a culture peculiarly preoccupied with the properties of bone and blubber, ink and skin, parchment and iron, where magnetism was forged in the 'combinations', as Scoresby put it, of such specific materials. The enterprises of his most avid reader, peer and fellow labour rights activist, Herman Melville, showcase the salience of Scoresby's struggle with Admiralty authority. The eminent Australian scholar Greg Dening's approach to ethnohistory proves the appropriate instrument with which to analyse such an encounter between traditions, negotiated through material forms. In the fraught exchange between whaler and maritime state, the combination laws that helped prompt the threat of revolution in early nineteenth-century Britain were translated into Scoresby's iron. Extant material and archival collections in Greenwich and Whitby offer traces of a battle between ways of knowing this protean metal: 'not down in any map; true places never are'.

Multiple processes drive genetic structure of humpback whale (Megaptera novaeangliae) populations across spatial scales.

Elucidating patterns of population structure for species with complex life histories, and disentangling the processes driving such patterns, remains a significant analytical challenge. Humpback whale (Megaptera novaeangliae) populations display complex genetic structures that have not been fully resolved at all spatial scales. We generated a data set of nuclear markers for 3575 samples spanning the seven breeding stocks and substocks found in the South Atlantic and western and northern Indian Oceans. For the total sample, and males and females separately, we assessed genetic diversity, tested for genetic differentiation between putative populations and isolation by distance, estimated the number of genetic clusters without a priori population information and estimated rates of gene flow using maximum-likelihood and Bayesian approaches. At the ocean basin scale, structure is governed by geographical distance (IBD P < 0.05) and female fidelity to breeding areas, in line with current understanding of the drivers of broadscale population structure. Consistent with previous studies, the Arabian Sea breeding stock was highly genetically differentiated (FST 0.034-0.161; P < 0.01 for all comparisons). However, the breeding stock boundary between west South Africa and east Africa was more porous than expected based on genetic differentiation, cluster and geneflow analyses. Instances of male fidelity to breeding areas and relatively high rates of dispersal for females were also observed between the three substocks in the western Indian Ocean. The relationships between demographic units and current management boundaries may have ramifications for assessments of the status and continued protections of populations still in recovery from commercial whaling.

Testing the feasibility of a hypothetical whaling-conservation permit market in Norway.

A cap-and-trade system for managing whale harvests represents a potentially useful approach to resolve the current gridlock in international whale management. The establishment of whale permit markets, open to both whalers and conservationists, could reveal the strength of conservation demand, about which little is known. This lack of knowledge makes it difficult to predict the outcome of a hypothetical whale permit market. We developed a bioeconomic model to evaluate the influence of economic uncertainty about demand for whale conservation or harvest. We used simulations over a wide range of parameterizations of whaling and conservation demands to examine the potential ecological consequences of the establishment of a whale permit market in Norwegian waters under bounded (but substantial) economic uncertainty. Uncertainty variables were slope of whaling and conservation demand, participation level of conservationists and their willingness to pay for whale conservation, and functional forms of demand, including linear, quadratic, and log-linear forms. A whale-conservation market had the potential to yield a wide range of conservation and harvest outcomes, the most likely outcomes were those in which conservationists bought all whale permits.

Hepatic lipomas and myelolipomas in subsistence-harvested bowhead whales Balaena mysticetus, Alaska (USA): a case review 1980-2016.

We describe a case series of benign hepatic fatty tumors in 10 subsistence-harvested bowhead whales. Microscopic features included lipomatous and myelolipomatous masses. Extensive atrophy and/or destruction of hepatic parenchyma was not observed. No other significant disease was present except in an animal with unrelated chronic pleuritis. Based on our longitudinal case series (1980-2016) which identified 1-2 hepatic lipomas and myelolipomas in landed whales annually at Barrow, Alaska (USA), since 2012, hepatic lipomas and myelolipomas are occasionally seen in hunter-harvested bowhead whales. A conservative estimate for the percentage of bowhead whales with hepatic fatty tumors in landed whales in Barrow from 2012 to 2016 was 6% (7/111). The pathogenesis and exact cell origin of these benign fatty tumors in bowhead whales is undetermined. Assessment of further cases is warranted to better define the tissue distribution and pathogenesis of these tumors in bowhead whale liver.

"The singular state of the ice". Das kartografische Wissen des Walfängers William Scoresby.

"The singular state of the ice". The Cartographic Knowledge of the Whaler William Scoresby. The English whaler William Scoresby, Jr. (1790-1857) made use of his annual voyages to the Greenland Sea for distinguished scientific work, detailed records and the production of amazing maps. Due to his intensive contacts to scientists as Robert Jameson and politicians as Joseph Banks and John Barrow his research achieved a great deal of attention and set a benchmark for at least half a century. Scoresby combined the adventurous world of Arctic fishery with academic sciences. He attained the northernmost point anybody reached in his time, he extended the cartographic knowledge and forced the conquest and utilisation of the oceans for commercial fishing. But his biography enquires also about who got an opportunity for research and for what. Especially it demonstrates the strong impact practical knowledge of a whaler could have on geographic research and Arctic cartography.

A spatially explicit estimate of the prewhaling abundance of the endangered North Atlantic right whale.

The North Atlantic right whale (NARW) (Eubalaena glacialis) is one of the world's most threatened whales. It came close to extinction after nearly a millennium of exploitation and currently persists as a population of only approximately 500 individuals. Setting appropriate conservation targets for this species requires an understanding of its historical population size, as a baseline for measuring levels of depletion and progress toward recovery. This is made difficult by the scarcity of records over this species' long whaling history. We sought to estimate the preexploitation population size of the North Atlantic right whale and understand how this species was distributed across its range. We used a spatially explicit data set on historical catches of North Pacific right whales (NPRWs) (Eubalaena japonica) to model the relationship between right whale relative density and the environment during the summer feeding season. Assuming the 2 right whale species select similar environments, we projected this model to the North Atlantic to predict how the relative abundance of NARWs varied across their range. We calibrated these relative abundances with estimates of the NPRW total prewhaling population size to obtain high and low estimates for the overall NARW population size prior to exploitation. The model predicted 9,075-21,328 right whales in the North Atlantic. The current NARW population is thus <6% of the historical North Atlantic carrying capacity and has enormous potential for recovery. According to the model, in June-September NARWs concentrated in 2 main feeding areas: east of the Grand Banks of Newfoundland and in the Norwegian Sea. These 2 areas may become important in the future as feeding grounds and may already be used more regularly by this endangered species than is thought.

The International Legal Framework for Marine Mammal Conservation in the Mediterranean Sea.

The paper reviews the international treaties that are today applicable for the protection of marine mammals in the Mediterranean Sea. They include instruments applicable at the world or the regional level. Emphasis is put on the International Whaling Commission, which is today affected by a confrontation between the two opposing groups of nonwhaling and whaling parties, the Agreement on the Conservation of Cetaceans of the Black Sea, Mediterranean Sea and Contiguous Atlantic Area (ACCOBAMS) and its implementation through the resolutions adopted by the Meeting of the Parties, as well as the subregional Agreement between France, Italy and Monaco for the establishment of a sanctuary for the protection of marine mammals (Pelagos Sanctuary). While several treaty provisions are in place to ensure the protection of marine mammals in the Mediterranean, specific mandatory provisions are still lacking for certain subjects, such as underwater noise, collisions with ships, by catch in fishing gear and ingestion of plastic litter.

Female, juvenile, and calf sperm whale Physeter macrocephalus (Linnaeus 1758) records from Ireland.

Sperm whales spatially segregate by sex and social behavior as they mature. In the North Atlantic, male whales move to higher latitudes as far as Svalbard at 80° N, while females and young whales typically remain around lower latitudes below 40-45° N. The Azores, Madeira, and the Canary Islands constitute important nursery grounds for female and young sperm whales. Irish waters represent a midpoint for this species' spatial segregation in the Northeast Atlantic, where the species occurs along the submarine canyon systems to the west of the country. Historically, just male whales were thought to be found in this region between 51 and 55° N, but one adult female was caught by commercial whalers in 1910, and a 5.49 m calf was found stranded in 1916. Between 1995 and 2023, 10 female sperm whales have been stranded around the coast of Ireland. Eight of these whales have been stranded since 2013, and there has been at least one stranding per year between 2019 and 2023. Four of these strandings have occurred in Donegal in the northwest of Ireland, indicating the presence of female whales along the continental shelf off this region. Two females were stranded within a day of each other and were found in similar states of decomposition in February 2022, indicating that they may have been part of the same group rather than being lone vagrant individuals. Sperm whale calves and juveniles were also sighted in Irish waters in 2001, 2004, and 2010 in the Rockall Trough, along the Porcupine Bank and Goban Spur, where between 1 and 3 individuals were observed on four occasions while one calf live stranded in 2004. These records indicate a historical presence of female and young sperm whales in this region but that an apparent increase in occurrence has taken place over the past decade.

Collecting whales: processes and biases in Nordic museum collections.

Whales are unique museum objects that have entered collections in many ways and for different reasons. This work studies three Nordic natural history museum collections in Norway and Denmark with more than 2,500 whale specimens in total, and gathers the available biological and collection data on the specimens, which include skeletal elements, foetuses and organs preserved in ethanol or formalin, and a few dry-preserved organs. It finds that influx of specimens, which were mainly locally common species that were hunted, to the collections, mainly happened in the latest 1800s and earliest 1900s, fuelled by research trends, nation building, local whaling, and colonial mechanisms. Norway was a major whaling nation, but the largest hunt for whales in the Southern Ocean in the mid-1900s is not reflected in the Norwegian museum collections, probably because of the commercial focus of the whaling industry and logistical challenges, combined with limited research interest in zoological specimens at that time. The results demonstrate that it is important to understand these processes and the resulting biases for future research, outreach, and conservation.

First documented movement of a humpback whale between the Cape Verde Islands and West Greenland.

The endangered population of humpback whales (Megaptera novaeangliae) breeding and calving off the Cape Verde Islands (CVI) are known to migrate to feeding areas located along the eastern margin of the North Atlantic Ocean (Iceland, and Norway). Here, we report for the first time a confirmed migration of an individual humpback whale from CVI breeding ground to a western North Atlantic feeding ground of West Greenland. This individual humpback, which was photographed and identified off the coast of West Greenland in 2021, was previously documented in CVI 22 years before (1999). An annual subsistence hunt for humpbacks occurs in West Greenland and the resighting at this location with a humpback whale from CVI has strong implications for the conservation efforts of the small CVI population.