Strong and lasting impacts of past global warming on baleen whales and their prey.

Global warming is affecting the population dynamics and trophic interactions across a wide range of ecosystems and habitats. Translating these real-time effects into their long-term consequences remains a challenge. The rapid and extreme warming period that occurred after the Last Glacial Maximum (LGM) during the Pleistocene-Holocene transition (7-12 thousand years ago) provides an opportunity to gain insights into the long-term responses of natural populations to periods with global warming. The effects of this post-LGM warming period have been assessed in many terrestrial taxa, whereas insights into the impacts of rapid global warming on marine taxa remain limited, especially for megafauna. In order to understand how large-scale climate fluctuations during the post-LGM affected baleen whales and their prey, we conducted an extensive, large-scale analysis of the long-term effects of the post-LGM warming on abundance and inter-ocean connectivity in eight baleen whale and seven prey (fish and invertebrates) species across the Southern and the North Atlantic Ocean; two ocean basins that differ in key oceanographic features. The analysis was based upon 7032 mitochondrial DNA sequences as well as genome-wide DNA sequence variation in 100 individuals. The estimated temporal changes in genetic diversity during the last 30,000 years indicated that most baleen whale populations underwent post-LGM expansions in both ocean basins. The increase in baleen whale abundance during the Holocene was associated with simultaneous changes in their prey and climate. Highly correlated, synchronized and exponential increases in abundance in both baleen whales and their prey in the Southern Ocean were indicative of a dramatic increase in ocean productivity. In contrast, the demographic fluctuations observed in baleen whales and their prey in the North Atlantic Ocean were subtle, varying across taxa and time. Perhaps most important was the observation that the ocean-wide expansions and decreases in abundance that were initiated by the post-LGM global warming, continued for millennia after global temperatures stabilized, reflecting persistent, long-lasting impacts of global warming on marine fauna.

Assessing the recovery of an Antarctic predator from historical exploitation.

The recovery of whale populations from centuries of exploitation will have important management and ecological implications due to greater exposure to anthropogenic activities and increasing prey consumption. Here, a Bayesian population model integrates catch data, estimates of abundance, and information on genetics and biology to assess the recovery of western South Atlantic (WSA) humpback whales (Megaptera novaeangliae). Modelling scenarios evaluated the sensitivity of model outputs resulting from the use of different data, different model assumptions and uncertainty in catch allocation and in accounting for whales killed but not landed. A long period of exploitation drove WSA humpback whales to the brink of extinction. They declined from nearly 27 000 (95% PI = 22 800-33 000) individuals in 1830 to only 450 (95% PI = 200-1400) whales in the mid-1950s. Protection led to a strong recovery and the current population is estimated to be at 93% (95% PI = 73-100%) of its pre-exploitation size. The recovery of WSA humpback whales may result in large removals of their primary prey, the Antarctic krill (Euphausia superba), and has the potential to modify the community structure in their feeding grounds. Continued monitoring is needed to understand how these whales will respond to modern threats and to climate-driven changes to their habitats.

Stretching the truth: length data highlight falsification of Japanese sperm whale catch statistics in the Southern Hemisphere.

Falsification of reports on Japanese catches of sperm whales (Physeter macrocephalus) is known to have occurred at both land whaling stations and in North Pacific factory fleets. Here, we conduct an analysis of pelagic sperm whale catches in the Southern Hemisphere: we compare true Soviet length data from the Yuri Dolgorukiy factory fleet during 1960-1975 to data for the same period reported to the International Whaling Commission (IWC) by Japan. Prior to implementation of the International Observer Scheme (IOS) in 1972, the Soviet fleet killed 5536 females, of which only 153 (2.8%) were at or above the minimum legal length of 11.6 m. During the same period, Japan killed 5799 females and reported that 5686 (98.5%) were of legal size, with 88.5% of the entire length distribution reported as being between 11.6 and 12.0 m. This unrealistic distribution, together with the fact that Japanese fleets were supposedly able to catch 37 times the number of legal-sized females as the Soviet fleet, indicates extensive falsification of catch data by Japan. Further evidence of misreporting is that females >11.5 m dropped to 9.1% of the Japanese catch after 1971, when the IOS made cheating much more difficult. That 99.6% of 10 433 males in the pre-IOS catch were also reported to be of legal size, indicates that illegal catches were not confined to females. We caution that the Japanese sperm whale data in the IWC Catch Database are unreliable and should not be used in population assessments. The ease with which illegal catches were apparently made underscores the past failures of the IWC to effectively regulate whaling.

Satellite tracking reveals novel migratory patterns and the importance of seamounts for endangered South Pacific humpback whales.

The humpback whale population of New Caledonia appears to display a novel migratory pattern characterized by multiple directions, long migratory paths and frequent pauses over seamounts and other shallow geographical features. Using satellite-monitored radio tags, we tracked 34 whales for between 5 and 110 days, travelling between 270 and 8540 km on their southward migration from a breeding ground in southern New Caledonia. Mean migration speed was 3.53±2.22 km h(-1), while movements within the breeding ground averaged 2.01±1.63 km h(-1). The tag data demonstrate that seamounts play an important role as offshore habitats for this species. Whales displayed an intensive use of oceanic seamounts both in the breeding season and on migration. Seamounts probably serve multiple and important roles as breeding locations, resting areas, navigational landmarks or even supplemental feeding grounds for this species, which can be viewed as a transient component of the seamount communities. Satellite telemetry suggests that seamounts represent an overlooked cryptic habitat for the species. The frequent use by humpback whales of such remote locations has important implications for conservation and management.

What's the catch? Validity of whaling data for Japanese catches of sperm whales in the North Pacific.

The failure of international efforts to manage commercial whaling was exemplified by revelations of large-scale illegal whale catches by the USSR over a 30 year period following World War II. Falsifications of catch data have also been reported for Japanese coastal whaling, but to date there has been no investigation of the reliability of catch statistics for Japanese pelagic (factory fleet) whaling operations. Here, we use data of known reliability from Soviet whaling industry reports to show that body lengths reported to the International Whaling Commission (IWC) by Japanese factory fleets for female sperm whales caught in the North Pacific are not credible. In 1968/1969, Japanese whaling fleets in the North Pacific killed 1568 females, of which 1525 (97.3%) were reported as being at or above the IWC's minimum length of 11.6 m (legal-sized females, LSFs). By contrast, Soviet fleets operating during this period killed 12 578 females; only 824 (6.6%) were LSFs. Adjusting for effort, catches of LSFs were up to 9.1 times higher for Japan compared with the USSR, and even higher for very large females. Dramatic differences in body length statistics were evident when both nations operated in the same area. Significantly, the frequency of LSFs and very large females in the Japanese catch markedly declined after the IWC's International Observer Scheme in 1972 made illegal whaling more difficult. We conclude that the Japanese length data reflect systematic falsification of catch statistics submitted to the IWC, with serious implications for the reliability of data used in current population assessments. The apparent ease with which catch data were falsified in the past underscores the necessity of transparent and independent inspection procedures in any future commercial whaling.

Straight as an arrow: humpback whales swim constant course tracks during long-distance migration.

Humpback whale seasonal migrations, spanning greater than 6500 km of open ocean, demonstrate remarkable navigational precision despite following spatially and temporally distinct migration routes. Satellite-monitored radio tag-derived humpback whale migration tracks in both the South Atlantic and South Pacific include constant course segments of greater than 200 km, each spanning several days of continuous movement. The whales studied here maintain these directed movements, often with better than 1° precision, despite the effects of variable sea-surface currents. Such remarkable directional precision is difficult to explain by established models of directional orientation, suggesting that alternative compass mechanisms should be explored.

Confirmation of right whales near a nineteenth-century whaling ground east of southern Greenland.

North Atlantic right whales (Eubalaena glacialis) were found in an important nineteenth century whaling area east of southern Greenland, from which they were once thought to have been extirpated. In 2007-2008, a 1-year passive acoustic survey was conducted at five sites in and near the 'Cape Farewell Ground', the former whaling ground. Over 2000 right whale calls were recorded at these sites, primarily during July-November. Most calls were northwest of the historic ground, suggesting a broader range in this region than previously known. Geographical and temporal separation of calls confirms use of this area by multiple animals.

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.

Geographic variation of persistent organic pollutant levels in humpback whale (Megaptera novaeangliae) feeding areas of the North Pacific and North Atlantic.

Seasonal feeding behavior and high fidelity to feeding areas allow humpback whales (Megaptera novaeangliae) to be used as biological indicators of regional contamination. Biopsy blubber samples from male individuals (n = 67) were collected through SPLASH, a multinational research project, in eight North Pacific feeding grounds. Additional male samples (n = 20) were collected from one North Atlantic feeding ground. Persistent organic pollutants were measured in the samples and used to assess contaminant distribution in the study areas. North Atlantic (Gulf of Maine) whales were more contaminated than North Pacific whales, showing the highest levels of polychlorinated biphenyls (PCBs), polybrominated diphenyl ethers (PBDEs), and chlordanes. The highest dichlorodiphenyltrichloroethane (DDT) levels were detected in whales feeding off southern California, USA. High-latitude regions were characterized by elevated levels of hexachlorocyclohexanes (HCHs) but generally nondetectable concentrations of PBDEs. Age was shown to have a positive relationship with SigmaPCBs, SigmaDDTs, Sigmachlordanes, and total percent lipid. Contaminant levels in humpback whales were comparable to other mysticetes and lower than those found in odontocete cetaceans and pinnipeds. Although these concentrations likely do not represent a significant conservation threat, levels in the Gulf of Maine and southern California may warrant further study.

Acoustic monitoring on a humpback whale (Megaptera novaeangliae) feeding ground shows continual singing into late Spring.

Singing by males is a major feature of the mating system of humpback whales, Megaptera novaeangliae (Borowski). Although a few songs have been opportunistically recorded on the whales' high-latitude feeding grounds, singing in these regions was thought to be only sporadic. We report results from the first continuous acoustic monitoring of a humpback whale feeding ground (off Cape Cod, MA, USA) in spring. Using autonomous sea-floor recording systems, we found singing on a daily basis over the entire 25 day monitoring period, from 14 May to 7 June 2000. For much of the period, song was recorded 24 h per day. These results, combined with evidence for aseasonal conceptions in whaling catch data, suggest that the humpback whale breeding season should no longer be considered as confined to lower-latitude regions in winter. Rather, we suggest breeding extends geographically and temporally onto feeding grounds into at least spring and early summer. Singing at these times represents either low-cost opportunistic advertising by (perhaps relatively few) males to court females that failed to conceive during the winter, and/or possibly an intrasexual display.

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.

Utility of North Atlantic Right Whale Museum Specimens for Assessing Changes in Genetic Diversity.

We examined six historical specimens of the endangered North Atlantic right whale (Eubalaena glacialis) using DNA isolated from documented baleen plates from the late nineteenth and early twentieth centuries. Sequences from the mitochondrial DNA (mtDNA) control region from these samples were compared with those from a near-exhaustive survey (269 of approximately 320 individuals) of the remaining right whales in the western North Atlantic Ocean. Our results suggest that there has been only relatively modest change in maternal lineage diversity over the past century in the North Atlantic right whale population. Any significant reduction in genetic variation in the species most likely occurred prior to the late nineteenth century. One historical specimen was from the last documented female capable of propagating one of the maternal lineages in the population today; no females in the existing population have been found to carry this mtDNA haplotype. Analysis of the only specimens from the eastern North Atlantic right whale population ever to be examined revealed that eastern and western North Atlantic right whales may not have been genetically differentiated populations. Loss of gene diversity experienced by North Atlantic right whales over the last century has been modest, and the six decades of protection have been successful in maintaining much of the maternal lineage diversity that was present in the late nineteenth century.

RESUMEN: Examinamos seis especímenes históricos de la ballena franca septentrional Eubalaena glacialis del Atlántico Norte, usando ADN aislado de placas de las barbas documentadas de finales de siglo diecinueve y principios del siglo veinte. Las secuencias del ADN mitocondrial (mtADN) de la región control de estas muestras comparadas con aquéllas de una muestra casi exhaustiva (269 de aproximadamente 320 individuos) de las ballenas francas remanentes en el océano Atlántico Norte Occidental. Nuestros resultados sugieren que solo ha habido ligeros cambios en la diversidad del linaje materno a lo largo del siglo pasado. Probablemente, cualquier reducción en la variación genética en las especies ocurrió antes del final del siglo diecinueve. Uno de los especímenes históricos fue de la última hembra documentada capaz de propagar uno de los linajes maternos en la población actual. Ninguna de las hembras en las poblaciones existentes ha sido identificada como portadora de este haplotipo de mtADN. El análisis de los especímenes de la población de ballenas del Atlántico Norte Oriental examinados revelan que las ballenas francas del este y del oeste del Atlántico Norte podrían no haber sido poblaciones genéticamente diferentes. La pérdida de diversidad genética experimentada en las poblaciones de ballena franca del Atlántico Norte a lo largo del siglo pasado ha sido modesta y las seis décadas de protección han sido exitosas en el mantenimiento de la mayoría de la diversidad del linaje materno que estaba presente a finales del siglo diecinueve.