Predicting the effects of whale population recovery on Northeast Pacific food webs and fisheries: an ecosystem modelling approach

The recovery of whale populations from historical depletion may have the potential to noticeably affect Northeast Pacific ecosystems and fisheries. Surplus production models based on whaling catch records were used to reconstruct the historical abundances of five large whale species in the waters surrounding Haida Gwaii, British Columbia. The results suggest that the local abundances of all five species were vastly higher before the onset of modern whaling. A comparison of ecosystem models representing the states of the local marine food web before and after full whale recovery indicates that abundant whales could consume large proportions of the annual production of their principal prey, ranging up to 87% for Pacific herring (Clupea pallasii) and 72% for piscivorous rockfish (Sebastes spp.). Dynamic modelling of the food web effects of whale recovery, including simulations of simultaneous top‐down and bottom‐up forcing and a Monte Carlo sensitivity analysis, revealed noticeable (～6–12%) top‐down effects on Pacific herring biomass owing to increased predation by humpback and fin whales. However, these effects cannot explain the magnitude of recent declines in local herring biomass. The dynamic modelling results also suggest that top‐down effects of whale recovery could result in reduced biomasses of large rockfish as a result of predation by sperm whales, as well as potential cascading effects on many demersal fish groups. These findings have numerous practical implications for ecosystem‐based fisheries management and whale conservation strategies in Northeast Pacific waters.     

Australian sperm whales from different whaling stocks belong to the same population

1. Understanding the factors driving population structure in marine mammals is needed to evaluate the impacts of previous exploitation, current anthropogenic threats, conservation status, and success of population recovery efforts.
2. Sperm whales are characterized by a worldwide distribution, low genetic diversity, complex patterns of social and genetic structure that differ significantly within and between ocean basins, and a long history of being commercially whaled. In Australia, sperm whales from the (International Whaling Commission assigned) southern hemisphere ‘Division 5’ stock were very heavily exploited by whaling.
3. The present study assessed the potential effects of whaling on the genetic diversity of sperm whales in Australia and the population genetic structure of these whales within a global context. A combination of historical and contemporary sperm whale samples (n = 157) were analysed across six regions, from south-eastern Australia (‘Division 6’ stock in the Pacific Ocean) to south-western Australia (‘Division 5’ stock in the Indian Ocean).
4. Sperm whales sampled from the ‘Division 5’ and ‘Division 6’ stocks belong to the same population based on nuclear and mitochondrial DNA (mtDNA) analyses. Four novel sperm whale mtDNA haplotypes were identified in animals from Australian waters. Levels of genetic diversity were low in Australian sperm whales but were similar to those previously reported for populations in the Indian and Pacific Oceans.
5. Given the genetic distinctiveness of sperm whales in Australian waters from other regions in the Pacific and Indian Oceans, and the lack of recovery in population numbers, further scientific studies are needed to increase our understanding of population dynamics and the effectiveness of threat management strategies in this species.

     

Ecosystem modelling to quantify the impact of historical whaling on Southern Hemisphere baleen whales

Many baleen whales were commercially harvested during the 20th century almost to extinction. Reliable assessments of how this mass depletion impacted whale populations, and projections of their recovery, are crucial but there are uncertainties regarding the status of Southern Hemisphere whale populations. We developed a Southern Hemisphere spatial “Model of Intermediate Complexity for Ecosystem Assessments” (MICE) for phytoplankton, krill (Euphausia superba) and five baleen whale species, to estimate whale population trajectories from 1890 to present. To forward project to 2100, we couple the predator–prey model to a global climate model. We used the most up to date catch records, fitted to survey data and accounted for key uncertainties. We predict Antarctic blue (Balaenoptera musculus intermedia), fin (Balaenoptera physalus) and southern right (Eubalaena australis) whales will be at less than half their pre‐exploitation numbers (K) even given 100 years of future protection from whaling, because of slow growth rates. Some species have benefited greatly from cessation of harvesting, particularly humpbacks (Megaptera novaeangliae), currently at 32% of K, with full recovery predicted by 2050. We highlight spatial differences in the recovery of whale species between oceanic areas, with current estimates of Atlantic/Indian area blue (1,890, <1% of K) and fin (16,950, <4% of K) whales suggesting slower recovery from harvesting, whilst Pacific southern right numbers are <7% of K (2,680). Antarctic minke (Balaenoptera bonaerensis) population trajectories track future expected increases in primary productivity. Population estimates and plausible future predicted trajectories for Southern Hemisphere baleen whales are key requirements for management and conservation.     

Historical trends in the incidence of strandings of sperm whales (Physeter macrocephalus) on North Sea coasts: An association with positive temperature anomalies

Information on the migration patterns of sperm whales (Physeter macrocephalus) in the North Atlantic is preserved in historical strandings records, particularly for the North Sea, where sperm whale strandings have been documented since the 16th century, reflecting general public interest in large whales ashore. Most strandings in this area occur during or following the southward migration from the feeding grounds, when some animals enter the North Sea (in which they are thought to have difficulty navigating) instead of following their usual route through deep water to the west of the British Isles. There was much speculation about the causes of the high incidence of strandings on North Sea coasts in the 1990s, among which a recently published analysis of long-term trends in strandings indicated an effect of sunspot cycle length. We show that long-term interannual variation in the incidence of sperm whale strandings on North Sea coasts is related to positive temperature anomalies: the incidence of strandings was higher in warmer periods. The effect of temperature anomalies explains between 8 and 9% of variation in the strandings series. Inclusion of sunspot cycle length as an additional predictor did not significantly improve this model. It is suggested that this link with positive temperature anomalies may reflect changes in the distribution of the sperm whales’ main squid prey.     

Habitat and prey selection of common minke,sei, and Bryde’s whales in mesoscale during summer in the subarctic and transition regions of the western North Pacific

This study represents the first quantitative analysis of the characteristics of the distribution areas and stomach contents of common minke whale Balaenoptera acutorostrata, sei whale B. borealis, and Bryde’s whale B. edeni in relation to oceanographic and prey environments in mid summer in the western North Pacific. Common minke whales were distributed within subarctic regions and the northernmost region of the transitional domain, coinciding with the main habitat of their preferred prey, Pacific saury Cololabis saira. Sei whales were mainly found in the northernmost part of the transition zone and showed prey preference for Japanese anchovy Engraulis japonica, which was significantly more abundant in the main distribution area of the whale than in its adjacent areas. “Hot spots” of Bryde’s whales were found in several regions of the transition zone between the subarctic boundary and the Kuroshio front. This whale species preferred Japanese anchovy as prey, for which the distribution density was significantly higher in the main distribution area of the whale than in the adjacent areas. These results indicate that the summer distributions of Pacific saury and Japanese anchovy greatly influence the distributions of these whale species, suggesting that the whales’ habitat selection is closely related to their prey selection.     

Historical and contemporary habitat use of sperm whales around the Galápagos Archipelago: Implications for conservation

1. Sperm whales have occupied the waters off the Galápagos Islands, Ecuador, for at least the past 200 years. During the 19^th century, they were the target of intensive whaling that severely depleted the population. In recent times, after commercial whaling ended, sperm whales in the region remain vulnerable to multiple threats, especially potential entanglement in fishing gear, which may hinder their ability to recover from the whaling era.
2. As a highly mobile, long-lived species, long-term analysis of the habitat use of sperm whales is necessary to establish effective conservation and management strategies. Here, contemporary (1985–2014) and historical (1830–1850) sperm whale habitat use off the Galápagos Islands was analysed and contrasted to the extent of the Galápagos Marine Reserve (GMR). Contemporary habitat use and its variability over time were modelled as a function of geographic, oceanographic, and topographic variables using generalized additive models.
3. The fine-scale habitat (<50 km) used by sperm whales was associated with topographic (i.e. depth and slope) and oceanographic characteristics (i.e. relative sea surface temperature and standard deviation of sea surface temperature), but these preferences varied over time.
4. While historical and contemporary data indicate that sperm whale habitat primarily occurred within the boundaries of the GMR, in recent years, whales were found up to 30.1% of the time outside the GMR, potentially overlapping with commercial fisheries operating in the area.
5. The dynamic nature of the relationship of this nomadic species with its habitat highlights the need of large-scale conservation efforts across the Eastern Tropical Pacific region, including the wide-scale enforcement of regulations requiring the use of Automatic Identification System in fishing vessels, the promotion of on-board fisheries observer programmes, the development of adaptive management strategies, and international collaboration to identify and mitigate threats.

     

Feeding habits of Baird's beaked whale Berardius bairdii, in the western North Pacific and Sea of Okhotsk off Japan

ABSTRACT:   We examined the stomach contents of 26 Baird's beaked whales caught off the coast of Japan by small-type coastal whalers. The main prey for these whales was rat-tails and hakes in the western North Pacific. Pollock and squids were also important food in the whales collected from the southern Sea of Okhotsk. The prey species found in the stomachs of the whales were almost identical to those caught in bottom-trawl nets at depths greater than about 1000 m in the western North Pacific, which suggests that the Baird's beaked whale forages for prey at depths of about 1000 m or more. Baird's beaked whales in the western North Pacific migrate to waters of 1000–3000 m in depth, where demersal fish are abundant. This implies that Baird's beaked whales migrate to waters where demersal fish, especially rat-tails and hakes, are abundant. Although there is limited information on the feeding habits of ziphiid whales, they are generally thought to prefer squid. The present data suggest that demersal fish are also important prey for ziphiid whales.     

Prey selection of common minke (Balaenoptera acutorostrata) and Bryde's (Balaenoptera edeni) whales in the western North Pacific in 2000 and 2001

A study of common minke and Bryde's whales was conducted in the western North Pacific in the 2000 and 2001 summer seasons to estimate prey selection of cetaceans as this is an important parameter in ecosystem models. Whale sighting and sampling surveys and prey surveys using quantitative echosounder and mid‐water trawl were carried out concurrently in the study. Biomasses of Japanese anchovy, walleye pollock and krill, which were major prey species of common minke and Bryde's whales, were estimated using an echosounder. The results suggested that common minke whale showed prey selection for Japanese anchovy while they seemed to avoid krill in both the offshore and coastal regions and walleye pollock in the continental shelf region. Selection for shoaling pelagic fish was similar to that in the eastern North Atlantic. Bryde's whale showed selection for Japanese anchovy in August 2000 and July 2001, while it showed prey selection for krill in May and June in 2001.     

The Norwegian minke whale DNA register: a data base monitoring commercial harvest and trade of whale products

The application of genetics for the management of natural resources is expanding, and within this field, DNA registers will play an increasing role. The Norwegian minke whale DNA register, established in 1996, was designed primarily as a control system to detect any attempts at illegal trade of products derived from other stocks of minke whale, or other whale species, under cover of the legal Norwegian harvest originating from the Northeast Atlantic. The register contains genetic data for 7644 of 7751 whales landed in the period 1997–2010. Profiles are established from sequencing part of the mtDNA control region, analysis of 10 STRs and a sex‐determining marker. Probabilities of genotypes matching between two randomly selected whales are 6.0^?04 and 3.0^?08 for five and eight of the STR loci, respectively. This permits verification of traded whale products via match to the register. The register has also been used in a number of ad hoc scientific studies resulting through the accumulation of genetic, demographic and biological data. Here, we review the register’s logistics, specifications and evaluate the potential to apply similar registers to control the exploitation of other marine species.     

A historical feeding ground for humpback whales in the eastern South Pacific revisited: the case of northern Patagonia,Chile

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Habitat differentiation between sei (Balaenoptera borealis) and Bryde's whales (B. brydei) in the western North Pacific

Two closely related baleen whale species, sei and Bryde's whales, in the western North Pacific were studied to identify differences in habitat use. Data were obtained from May to August 2004 and 2005. This study examined the relationship between oceanographic features derived from satellite data and the distribution of sei and Bryde's whales using basic statistics. We investigated oceanographic features including sea surface temperature (SST), sea surface chlorophyll a (Chl‐a), sea surface height anomalies (SSHAs), and depth of the habitat. These two whale species used habitats with different SST, Chl‐a, and SSHA ranges. The 0.25 mg m^?3 Chl‐a contour (similar to the definition of the Transition Zone Chlorophyll Front) was a good indicator that separated the habitats of sei and Bryde's whales. Then generalized linear models were used to model the probabilities that the whale species would be present in a habitat and to estimate their habitat distribution throughout the study area as a function of environmental variables. The potential habitats of the two species were clearly divided, and the boundary moved north with seasonal progression. The habitat partitioning results indicated that SST contributed to the patterns of habitat‐use and might reflect differences in prey species between the two whales. This study showed that the habitats of the sei and Bryde's whales were clearly divided and their potential habitat‐use changed seasonally.     

Southern Ocean iron fertilization by baleen whales and Antarctic krill

Iron is the limiting micronutrient in the Southern Ocean and experiments have demonstrated that addition of soluble iron to surface waters results in phytoplankton blooms, particularly by large diatoms. Antarctic krill (Euphausia superba) eat diatoms and recycle iron in surface waters when feeding. Baleen whales eat krill, and, historically, defecation by baleen whales could have been a major mechanism for recycling iron, if whale faeces contain significant quantities of iron. We analysed the iron content in 27 samples of faeces from four species of baleen whale. Faecal iron content (145.9 ± 133.7 mg kg^?1) is approximately ten million times that of Antarctic seawater, suggesting that it could act as a fertilizer. Furthermore, we analysed the iron content of seven krill species and of muscle tissue of two species of baleen whales; all samples had high iron levels. Using these figures, together with recent estimates of the range and biomass of krill, we calculate that the Antarctic krill population contains ～24% of the total iron in the surface waters in its range. Thus, krill can act as a long‐term reservoir of iron in Antarctic surface waters, by storing the iron in their body tissue. Pre‐exploitation populations of whales and krill must have stored larger quantities of iron and would have also recycled more iron in surface waters, enhancing overall ocean productivity through a positive feedback loop. Thus, allowing the great whales to recover might actually increase Southern Ocean productivity through enhancing iron levels in the surface layer.     

Effects of whale watching on the activity budgets of humpback whales,Megaptera novaeangliae (Borowski, 1781), on a feeding ground

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Estimation of the abundance of the sei whale <Emphasis Type="Italic">Balaenoptera borealis</Emphasis> in the central and eastern North Pacific in summer using sighting data from 2010 to 2012

The abundance of the sei whale Balaenoptera borealis in the central and eastern North Pacific (north of 40°N, south of the Alaskan coast including both the US and Canadian Exclusive Economic Zones between 170°E and 135°W), from July to August, was estimated by the line transect method using sighting data obtained during the 2010–2012 International Whaling Commission-Pacific Ocean Whale and Ecosystem Research cruises. The probability of detecting whales at a perpendicular distance from the transect was estimated using two different models: hazard rate and half-normal models. Because the difference in Akaike’s information criterion between the two models was small, the Akaike weighted average of the two models was taken, which gave an estimated abundance of 29,632 (coefficient of variation, 0.242; 95% confidence interval, 18,576–47,267). This is the first abundance estimate of sei whales in this region based on systematic sighting survey data, which contributes to an understanding of the current status of this species.     

Environmental factors and megafauna spatio‐temporal co‐occurrence with purse‐seine fisheries

Tropical tuna purse‐seine fisheries spatially co‐occur with various megafauna species, such as whale sharks, dolphins and baleen whales in all oceans of the world. Here, we analyzed a 10‐year (2002–2011) dataset from logbooks of European tropical tuna purse‐seine vessels operating in the tropical Eastern Atlantic and Western Indian Oceans, with the aim of identifying the principle environmental variables under which such co‐occurrence appear. We applied a Delta‐model approach using Generalized Additive Models (GAM) and Boosted Regression Trees (BRT) models, accounting for spatial autocorrelation using a contiguity matrix based on a residuals autocovariate (RAC) approach. The variables that contributed most in the models were chlorophyll‐a concentration in the Atlantic Ocean, as well as depth and monsoon in the Indian Ocean. High co‐occurrence between whale sharks, baleen whales and tuna purse‐seine fisheries were mostly observed in productive areas during particular seasons. In light of the lack of a full coverage scientific observer on board program, the large, long‐term dataset obtained from logbooks of tuna purse‐seine vessels is highly important for identifying seasonal and spatial co‐occurrence between the distribution of fisheries and megafauna, and the underlying environmental variables. This study can help to design conservation management measures for megafauna species within the framework of spatial fishery management strategies.     

New urban habitat for endangered humpback whales: San Francisco Bay

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Sperm whales associated with Gulf Stream features off the north-eastern USA shelf

Marine mammal surveys were conducted during sum Mer 1990 and 1991 in shelf-edge and off-shelf waters between Cape Hatteras and Georges Bank. Sperm whales were the most frequently sighted large whale in both years, constituting 27.9% and 12.7% of total sightings in 1990 and 1991, respectively. Sighting rates of sperm whales that were within a 9.3 km buffer (5 n.m.) of the Gulf Stream north wall were not significantly different (x²= 2.86, P > 0.05) from those within the Gulf Stream during 1990. In 1991, however, significantly more (x²= 51.0, P < 0.05) sightings were associated with shelf-edge areas where warm core rings were located.     

The decline of Steller sea lions (Eumetopias jubatus) in the North Pacific: insights from indigenous people,ethnohistoric records and archaeological data

A number of hypotheses have been proposed to explain the most recent decline (1977–2012) of Steller sea lions (SSL; Eumetopias jubatus) in the Gulf of Alaska and Aleutian Islands. We examined hypotheses about fisheries competition, environmental change, predation, anthropogenic effects and disease using observations of modern Aleut and archaeological, ethnohistoric and ethnographic data from the western Gulf of Alaska and Aleutian Islands. These data indicate that Steller sea lion numbers have declined and recovered repeatedly over the past 4500 years and were last at critically low numbers during the 1870s–1930s. Steller sea lions appear to have been more abundant during the cool periods – and lower during the warmer periods. Observations by local peoples, explorers, early government surveyors and biologists since the late 1800s suggest that low populations of SSL have been associated with high populations of Gadidae fishes (Pacific cod – Gadus macrocephalus and walleye pollock – Theragra chalcogramma) and are consistent with the ocean climate hypothesis to explain the decline of sea lions. They suggest that removals by people and killer whales (Orcinus orca) did not cause the sea lion declines, but could have compounded the magnitude of the decline as sea lion numbers approached low densities. Archaeological, anthropological and ethnohistorical analyses demonstrate that fluctuations have occurred in the North Pacific over hundreds to thousands of years and provide context for understanding the changes that occur today and the changes that will continue to occur in the future.     

Comparative external morphology of cetacean spermatozoa

SUMMARY: To compare size and morphology of spermatozoa in cetaceans, sperm and epididymis samples were collected from 10 species in four families and spermatozoa were observed with phase-contrast and scanning electron microscopes. According to the average total length of the spermatozoa, the 10 species examined were classified into the following four groups in order of increasing size: (i) Baird's beaked (Ziphiidae) and Bryde's whales (Balaenopteridae); (ii) Dall's and finless porpoises (Phocoenidae); (iii) common, bottlenose, and Pacific white-sided dolphins (Delphinidae); and (iv) killer and short-finned pilot whales, and Risso's dolphin (Delphinidae). Spermatozoa head length of Bryde's whale and the finless porpoise were shorter than those of the other species. Spermatozoa head width was widest in the killer whale and thinnest in the Baird's beaked whale. The lateral aspects of sperm heads from the 10 species were characterized as the 'anterior region of the sperm head is thin and flat while the posterior region is thick.' The dorsal aspects of sperm heads were 'paddle-shaped' in Bryde's whales, 'bowling pin-shaped' in Baird's beaked whales, 'Japanese fan-shaped' in killer whales, an 'elongated ellipsoid shape' in Delphinidae except for killer whales, and 'ellipsoid shaped' in Phocoenidae. Size and morphology of the spermatozoa showed interspecific differences among the 10 species examined, which correspond to cetacean taxonomic classification.     

Patterns of stable carbon and nitrogen isotopes in the baleen of common minke whale Balaenoptera acutorostrata from the western North Pacific

ABSTRACT:   Stable carbon and nitrogen isotope ratios were determined in the baleen plates of 17 common minke whales Balaenoptera acutorostrata from the north-western Pacific Ocean off Japan, as well as prey species (krill Euphausia pacifica , Japanese anchovy Engraulis japonicus and Pacific saury Cololabis saira ) collected in the stomach contents, to investigate the trophic relationship between the minke whales and their prey. A few δ¹⁵N-depleted peaks occurred along the length of baleen plates for 10 males irrespective of stomach content (anchovies and sauries). Similar δ¹⁵N-depleted peaks were also found for one female and two immature individuals. It was likely that these δ¹⁵N-depleted peaks formed in early summer. The stable nitrogen isotope ratio (δ¹⁵N) values in Pacific saury (9.3 ± 1.4‰) did not differ significantly from that in Japanese anchovy (8.8 ± 0.9‰). In contrast, δ¹⁵N in krill (7.2 ± 0.5‰ in July and 8.0 ± 0.2‰ in September) were significantly lower than in the Pacific saury. Thus, these peaks may reflect the dietary change from krill to fishes in the feeding migration of the whales. Growth rate of the baleen plate was estimated to be 129 mm/y, and it appeared that a dietary record of about 1.4 years remained in the baleen plate. For two immature whales, the maximum value of δ¹⁵N occurred at the tip of baleen. This δ¹⁵N enrichment may possibly be useful for discriminating weanlings and older whales.