Dangerous dining: surface foraging of North Atlantic right whales increases risk of vessel collisions

North Atlantic right whales are critically endangered and, despite international protection from whaling, significant numbers die from collisions with ships. Large groups of right whales migrate to the coastal waters of New England during the late winter and early spring to feed in an area with large numbers of vessels. North Atlantic right whales have the largest per capita record of vessel strikes of any large whale population in the world. Right whale feeding behaviour in Cape Cod Bay (CCB) probably contributes to risk of collisions with ships. In this study, feeding right whales tagged with archival suction cup tags spent the majority of their time just below the water's surface where they cannot be seen but are shallow enough to be vulnerable to ship strike. Habitat surveys show that large patches of right whale prey are common in the upper 5 m of the water column in CCB during spring. These results indicate that the typical spring-time foraging ecology of right whales may contribute to their high level of mortality from vessel collisions. The results of this study suggest that remote acoustic detection of prey aggregations may be a useful supplement to the management and conservation of right whales.     

RATES AND POTENTIAL CAUSES OF MORTALITY IN NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS)

North Atlantic right whale mortality rates range from 2% to 17% over the first four years of life. Sources of mortality in this population include ship collisions, entanglements, and natural causes. A combined analysis of stranding data, entanglement records, and photographic information indicates that approximately one third of all right whale mortality is caused by human activities. Anthropogenic sources of right whale mortality may be a significant factor inhibiting growth in the North Atlantic population.     

State–space mark–recapture estimates reveal a recent decline in abundance of North Atlantic right whales

North Atlantic right whales (Eubalaena glacialis Müller 1776) present an interesting problem for abundance and trend estimation in marine wildlife conservation. They are long lived, individually identifiable, highly mobile, and one of the rarest of cetaceans. Individuals are annually resighted at different rates, primarily due to varying stay durations among several principal habitats within a large geographic range. To date, characterizations of abundance have been produced that use simple accounting procedures with differing assumptions about mortality. To better characterize changing abundance of North Atlantic right whales between 1990 and 2015, we adapted a state–space formulation with Jolly‐Seber assumptions about population entry (birth and immigration) to individual resighting histories and fit it using empirical Bayes methodology. This hierarchical model included accommodation for the effect of the substantial individual capture heterogeneity. Estimates from this approach were only slightly higher than published accounting procedures, except for the most recent years (when recapture rates had declined substantially). North Atlantic right whales' abundance increased at about 2.8% per annum from median point estimates of 270 individuals in 1990 to 483 in 2010, and then declined to 2015, when the final estimate was 458 individuals (95% credible intervals 444–471). The probability that the population's trajectory post‐2010 was a decline was estimated at 99.99%. Of special concern was the finding that reduced survival rates of adult females relative to adult males have produced diverging abundance trends between sexes. Despite constraints in recent years, both biological (whales' distribution changing) and logistical (fewer resources available to collect individual photo‐identifications), it is still possible to detect this relatively recent, small change in the population's trajectory. This is thanks to the massive dataset of individual North Atlantic right whale identifications accrued over the past three decades. Photo‐identification data provide biological information that allows more informed inference on the status of this species.     

Simulation of the entanglement of a North Atlantic right whale (Eubalaena glacialis) with fixed fishing gear

Population estimates of the critically endangered North Atlantic right whale (Eubalaena glacialis) put the number of individuals at 458 with the actual number likely being lower due to a recent unusual mortality event. Entanglement with fixed fishing gear is the most significant cause of mortality of North Atlantic right whales. There remains little documentation of how North Atlantic right whales become enwrapped during an encounter with fixed fishing gear. In order to gain a better understanding of how entanglements might occur, an interactive simulator was developed that allows the user to swim a virtual whale model using a standard game controller through a gear field in an attempt to re‐create an entanglement. The morphologically accurate right whale model produces realistic swimming motions and is capable of pectoral fin motions in response to user input. Using the simulator, gear entanglements involving the pectoral flippers including ropes wrapping around the body and entanglements involving the tailstock were re‐created. Entanglements involving the pectoral flippers with body wraps were more easily generated than entanglements involving the tailstock only. The simulator should aid scientists, fisheries experts, fishing gear designers, and bycatch reduction scientists in understanding entanglement dynamics and testing potential new gear configurations.     

Hexavalent chromium is cytotoxic and genotoxic to the North Atlantic right whale (Eubalaena glacialis) lung and testes fibroblasts

Although hexavalent chromium is a known genotoxic agent in human and terrestrial mammals and is present in seawater and air, its effects on marine mammals including the endangered North Atlantic right whale are unknown and untested. The present study investigated the cytotoxic and genotoxic effects of hexavalent chromium in primary cultured North Atlantic right whale lung and testes fibroblasts and levels of total chromium in skin biopsies from North Atlantic right whales. Cytotoxicity was measured by clonogenic survival assay. Genotoxicity was measured as production of chromosome aberrations. Tissue chromium levels were determined from skin biopsies of healthy free-ranging whales in the Bay of Fundy using inductively coupled plasma optical emission spectroscopy. Hexavalent chromium-induced concentration-dependent increases in right whale lung and testes fibroblast cytotoxicity with the testes more sensitive to the cytotoxic effects. It also induced concentration-dependent increases in chromosomal aberrations in both cell types with no significant difference in sensitivity. Skin biopsy data indicate that North Atlantic right whales are exposed to chromium and accumulate a range of 4.9-10 microg Cr/g tissue with a mean of 7.1 microg/g. Hexavalent chromium is cytotoxic and genotoxic to North Atlantic right whale cells. The whales have tissue chromium levels that are concerning. These data support a hypothesis that chromium may be a concern for the health of the North Atlantic right whales. Considering these data with chromium chemistry, whale physiology and atmospheric chromium levels further suggest that inhalation may be an important exposure route.     

Stable isotope analysis and fin whale subpopulation structure in the eastern North Atlantic

The North Atlantic fin whale was subject to heavy exploitation in the past and, despite partial recovery, it is still considered endangered. Recent research has questioned its currently accepted subpopulation structure and migratory movements, challenging management and conservation efforts. Here we contribute to this discussion by presenting results of stable isotope analysis of fin whale bones and krill samples collected from fin whale stomachs from two exploited areas, west Iceland and northwest Spain, and comparing these results with North Atlantic isoscapes. In krill, δ¹⁵N values were highly variable and no significant differences in δ¹⁵N and δ¹³C between areas emerged. δ¹⁵N and δ¹³C values in bone were higher than in krill, due to trophic enrichment, and were not significantly different between areas. Both krill and bone results were slightly inconsistent with the local isoscapes, maybe due to seasonal variations in local productivity and in krill diet and, in the case of bone, to its capability to integrate long‐term isotopic values, derived from food consumed in distant areas. Conversely, δ¹⁸O values, which reflect the basal signal of sea water, were consistent with isoscapes and significantly higher in whales from northwestern Spain, possibly indicating migration to breeding grounds located at lower latitudes.     

Natal origin of Namibian grey whale implies new distance record for in-water migration

We use genomics to identify the natal origin of a grey whale found in the South Atlantic, at least 20 000 km from the species core range (halfway around the world). The data indicate an origin in the North Pacific, possibly from the endangered western North Pacific population, thought to include only approximately 200 individuals. This contributes to our understanding of Atlantic sightings of this species known primarily from the North Pacific, and could have conservation implications if grey whales have the potential for essentially global dispersion. More broadly, documenting and understanding rare extreme migration events have potential implications for the understanding of how a species may be able to respond to global change.     

DNA profile of a sixteenth century western North Atlantic right whale (Eubalaena glacialis)

Low levels of genetic variability identified within the North Atlantic right whale (Eubalaena glacialis), when compared to the Southern right whale (E. australis) and other large whales, have been suggested to result from population reductions due to whaling. Previous genetic analysis of 218 whale bones from sixteenth century Basque whaling sites in the western North Atlantic revealed only a single right whale bone. We determined the genotypes of 27 microsatellite loci using DNA isolated from this bone. All alleles from the historic specimen occur in the extant western North Atlantic population and both the probability of identity of the specimen and the number of heterozygous loci are similar to that in the extant population. Assessments of how genetically different the historical population might have been suggest genetic characteristics have not changed substantially over four centuries of whaling.     

Mechanics of the right whale mandible: Full scale testing and finite element analysis

In an effort to better understand the mechanics of ship-whale collision and to reduce the associated mortality of the critically endangered North Atlantic right whale, a comprehensive biomechanical study has been conducted by the Woods Hole Oceanographic Institution and the University of New Hampshire. The goal of the study is to develop a numerical modeling tool to predict the forces and stresses during impact and thereby the resulting mortality risk to whales from ship strikes.Based on post-mortem examinations, jaw fracture was chosen as a fatal endpoint for the whales hit by a vessel. In this paper we investigate the overall mechanical behavior of a right whale mandible under transverse loading and develop a finite element analysis model of the bone. The equivalent elastic modulus of the cortical component of right whale mandible is found by comparing full-scale bending tests with the results of numerical modeling. The finite element model of the mandible can be used in conjunction with a vessel-whale collision event model to predict bone fracture for various ship strike scenarios.     

Right whale occurrence in the coastal waters of Virginia,U.S.A.: Endangered species presence in a rapidly developing energy market

The endangered North Atlantic right whale is susceptible to increased risks associated with the developing energy market off the Virginia coast of the U.S.A., including increased noise exposure from shipping and energy exploration and extraction, and shipping activity. Primarily viewed as a migratory corridor between northern feeding and mating areas and the southern calving area, Virginia waters have not been intensely monitored for right whales. We conducted a 1 yr, continuous, passive acoustic survey from the Virginia coast to the continental shelf to better describe the seasonal occurrence and spatial distribution of right whales in this area. We found that whales were acoustically detected in every month of the year, with increased seasonal occurrences in fall and late winter/early spring, and not just during limited periods of the year. As industry activities continue to increase in the area, these new right whale occurrence data have implications for siting and permitting of offshore energy activities.     

ESTIMATION OF PREY DENSITIES REQUIRED BY WESTERN NORTH ATLANTIC RIGHT WHALES

The apparent lack of significant recovery of western North Atlantic right whale populations may be related to the availability of prey in exploitable densities. We have attempted to estimate the concentration of zooplankton required by right whales to obtain a net energetic benefit over the long term, using estimated values for body weight, metabolic rate, assimilation efficiency, time spent feeding, mouth size and swimming speed. The estimated range of required energy densities is 7.57 to 2,394 kcal m⁻³ (3.17 × 10⁴ to 1.00 × 10⁷ joule m⁻³). These values are from one to more than three orders of magnitude greater than the densest concentration sampled in the vicinity of right whale aggregations in the Great South Channel. Right whales must seek out and exploit extremely dense patches of prey organisms in order to feed efficiently. The presence of such dense patches of zooplankton is likely to be a primary characteristic of spring, summer and fall right whale habitats.     

Buoyant balaenids: the ups and downs of buoyancy in right whales

A variety of marine mammal species have been shown to conserve energy by using negative buoyancy to power prolonged descent glides during dives. A new non-invasive tag attached to North Atlantic right whales recorded swim stroke from changes in pitch angle derived from a three-axis accelerometer. These results show that right whales are positively buoyant near the surface, a finding that has significant implications for both energetics and management. Some of the most powerful fluke strokes observed in tagged right whales occur as they counteract this buoyancy as they start a dive. By contrast, right whales use positive buoyancy to power glides during ascent. Right whales appear to use their positive buoyancy for more efficient swimming and diving. However, this buoyancy may pose added risks of vessel collision. Such collisions are the primary source of anthropogenic mortality for North Atlantic right whales, whose population is critically endangered and declining. Buoyancy may impede diving responses to oncoming vessels and right whales may have a reduced ability to manoeuvre during free ascents. These risk factors can inform efforts to avoid collisions.     

A surplus no more? Variation in krill availability impacts reproductive rates of Antarctic baleen whales

The krill surplus hypothesis of unlimited prey resources available for Antarctic predators due to commercial whaling in the 20th century has remained largely untested since the 1970s. Rapid warming of the Western Antarctic Peninsula (WAP) over the past 50 years has resulted in decreased seasonal ice cover and a reduction of krill. The latter is being exacerbated by a commercial krill fishery in the region. Despite this, humpback whale populations have increased but may be at a threshold for growth based on these human-induced changes. Understanding how climate-mediated variation in prey availability influences humpback whale population dynamics is critical for focused management and conservation actions. Using an 8-year dataset (2013–2020), we show that inter-annual humpback whale pregnancy rates, as determined from skin-blubber biopsy samples (n = 616), are positively correlated with krill availability and fluctuations in ice cover in the previous year. Pregnancy rates showed significant inter-annual variability, between 29% and 86%. Our results indicate that krill availability is in fact limiting and affecting reproductive rates, in contrast to the krill surplus hypothesis. This suggests that this population of humpback whales may be at a threshold for population growth due to prey limitations. As a result, continued warming and increased fishing along the WAP, which continue to reduce krill stocks, will likely impact this humpback whale population and other krill predators in the region. Humpback whales are sentinel species of ecosystem health, and changes in pregnancy rates can provide quantifiable signals of the impact of environmental change at the population level. Our findings must be considered paramount in developing new and more restrictive conservation and management plans for the Antarctic marine ecosystem and minimizing the negative impacts of human activities in the region.     

Insights into the population structure of blue whales in the Eastern North Pacific from recent sightings and photographic identification

Blue whales were widely distributed in the North Pacific prior to the primary period of modern commercial whaling in the early 1900s. Despite concentrations of blue whale catches off British Columbia and in the Gulf of Alaska, there had been few documented sightings in these areas since whaling for blue whales ended in 1965. In contrast, large concentrations of blue whales have been documented off California and Baja California and in the eastern tropical Pacific since the 1970s, but it was not known if these animals were part of the same population that previously ranged into Alaskan waters. We document 15 blue whale sightings off British Columbia and in the Gulf of Alaska made since 1997, and use identification photographs to show that whales in these areas are currently part of the California feeding population. We speculate that this may represent a return to a migration pattern that has existed for earlier periods for eastern North Pacific blue whale population. One possible explanation for a shift in blue whale use is changes in prey driven by changes in oceanographic conditions, including the Pacific Decadal Oscillation (PDO), which coincides with some of the observed shifts in blue whale occurrence.     

Population histories of right whales (Cetacea: Eubalaena) inferred from mitochondrial sequence diversities and divergences of their whale lice (Amphipoda: Cyamus)

Right whales carry large populations of three 'whale lice' (Cyamus ovalis, Cyamus gracilis, Cyamus erraticus) that have no other hosts. We used sequence variation in the mitochondrial COI gene to ask (i) whether cyamid population structures might reveal associations among right whale individuals and subpopulations, (ii) whether the divergences of the three nominally conspecific cyamid species on North Atlantic, North Pacific, and southern right whales (Eubalaena glacialis, Eubalaena japonica, Eubalaena australis) might indicate their times of separation, and (iii) whether the shapes of cyamid gene trees might contain information about changes in the population sizes of right whales. We found high levels of nucleotide diversity but almost no population structure within oceans, indicating large effective population sizes and high rates of transfer between whales and subpopulations. North Atlantic and Southern Ocean populations of all three species are reciprocally monophyletic, and North Pacific C. erraticus is well separated from North Atlantic and southern C. erraticus. Mitochondrial clock calibrations suggest that these divergences occurred around 6 million years ago (Ma), and that the Eubalaena mitochondrial clock is very slow. North Pacific C. ovalis forms a clade inside the southern C. ovalis gene tree, implying that at least one right whale has crossed the equator in the Pacific Ocean within the last 1-2 million years (Myr). Low-frequency polymorphisms are more common than expected under neutrality for populations of constant size, but there is no obvious signal of rapid, interspecifically congruent expansion of the kind that would be expected if North Atlantic or southern right whales had experienced a prolonged population bottleneck within the last 0.5 Myr.     

Visual and acoustic surveys for North Atlantic right whales,Eubalaena glacialis,in Cape Cod Bay,Massachusetts, 2001–2005: Management implications

North Atlantic right whales, Eubalaena glacialis, remain endangered, primarily due to excessive anthropogenic mortality. Current management protocols in US waters are triggered by identifying the presence of at least one right whale in a management area. We assessed whether acoustic detection of right whale contact calls can work as an alternative to visual aerial surveys for establishing their presence. Aerial survey and acoustic monitoring were conducted in Cape Cod Bay, Massachusetts, in 2001–2005 and used to evaluate and compare right whale detections. Over the 58 d with simultaneous aerial and acoustic coverage, aerial surveys saw whales on approximately two-thirds of the days during which acoustic monitoring heard whales. There was no strong relationship between numbers of whales seen during aerial surveys and numbers of contact calls detected on survey days. Results indicate acoustic monitoring is a more reliable mechanism than aerial survey for detecting right whales. Because simple detection is sufficient to trigger current management protocols, continuous, autonomous acoustic monitoring provides information of immediate management utility more reliably than aerial surveillance. Aerial surveys are still required to provide data for estimating population parameters and for visually assessing the frequency and severity of injuries from shipping and fishing and detecting injured and entangled right whales.     

The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea

1. The ecology and status of fin whales Balaenoptera physalus in the Mediterranean Sea is reviewed. The species’ presence, morphology, distribution, movements, population structure, ecology and behaviour in this semi‐enclosed marine region are summarized, and the review is complemented with original, previously unpublished data. 2. Although the total size of the fin whale population in the Mediterranean is unknown, an estimate for a portion of the western basin, where most of the whales are known to live, was approximately 3500 individuals. High whale densities, comparable to those found in rich oceanic habitats, were found in well‐defined areas of high productivity. Most whales concentrate in the Ligurian‐Corsican‐Provençal Basin, where their presence is particularly noticeable during summer; however, neither their movement patterns throughout the region nor their seasonal cycle are clear. 3. Based on genetic studies, fin whales from the Mediterranean Sea are distinct from North Atlantic conspecifics, and may constitute a resident population, separate from those of the North Atlantic, despite the species’ historical presence in the Strait of Gibraltar. Fin whales are known to calve in the Mediterranean, with births peaking in November but occurring at lower rates throughout the year. They feed primarily on krill Meganyctiphanes norvegica which they capture by diving to depths in excess of 470 m. It is suggested that the extensive vertical migratory behaviour of its main prey may have influenced the social ecology of this population. 4. Known causes of mortality and threats, including collisions with vessels, entanglement in fishing gear, deliberate killing, disturbance, pollution and disease, are listed and discussed in view of the implementation of appropriate conservation measures to ensure the species’ survival in the region.     

Regional and temporal variation in diet and provisioning rates suggest weather limits prey availability for an endangered raptor

Understanding variation in food requirements of wild animals is of central importance in population ecology and conservation, as it helps to identify where and when food may be limiting. Studies on diet variation or prey provisioning rates may give useful insights when direct information on prey availability is lacking. We assess spatial and temporal variation in the diet of an endangered predator, the Black Harrier Circus maurus. This raptor is endemic to southern Africa and specializes on small mammals but also feeds on birds and reptiles as alternative prey. Using data on 1679 prey identified in 953 pellets collected in inland and coastal regions from 2006 to 2015, we show that diet composition changed little throughout the breeding season in the coastal region, whereas there was a marked seasonal decline in the occurrence of small mammal prey in the inland region, with a concomitant increase in alternative prey. The proportion of small mammals in the diet declined with increasing maximum temperature, the latter being highest at the inland region late in the breeding season. Using camera recordings at nests in 2014, we further analysed daily patterns of prey provisioning to nestlings. A marked reduction in small mammal provisioning rates occurred during the middle of the day in the hotter inland region but not in the cooler coastal region. Reduced availability of the primary prey, small mammals, in hotter conditions, through a reduction in activity or overall abundance, could explain these patterns. Finally, we show a positive relationship between winter rainfall and interannual differences in the proportion of small mammals in the diet of Black Harriers breeding in the coastal region, suggesting relationships between diet and prey abundance that are mediated through rainfall. We discuss the need to consider spatial variation in food availability in conservation strategies.     

Estimated field metabolic rates and prey requirements of resident killer whales

Killer whales are large animals that often feed in groups and thus have the potential to deplete prey populations. Determining predator energy requirements is essential to assessing whether prey availability is sufficient. This is important because one risk factor facing the endangered Southern Resident killer whale distinct population segment is limited prey availability. Body mass, field metabolic rate (FMR), and daily prey energy requirements (DPERs) were estimated for each individual in the population. FMRs were calculated from body mass, assuming they range from five to six times Kleiber‐predicted basal metabolic rates. FMRs of adults were also calculated from resident killer whale activity budgets and the metabolic cost of swimming at speeds associated with daily activities. These two methods yielded similar results. Total FMRs varied by age and sex, which is partly due to the long developmental period and sexual dimorphism in killer whales. FMRs for males (465–4,434 kg) ranged from 35,048 to 228,216 kcal/d while FMRs for females (465–3,338 kg) ranged from 35,048 to 184,444 kcal/d. DPERs were calculated from FMRs assuming a standard digestive efficiency. Corresponding DPERs ranged from 41,376 to 269,458 kcal/d and 41,376 to 217,775 kcal/d, respectively.     

Protecting persistent dynamic oceanographic features: transboundary conservation efforts are needed for the critically endangered Balearic shearwater

The protection of key areas for biodiversity at sea is not as widespread as on land and research investment is necessary to identify biodiversity hotspots in the open ocean. Spatially explicit conservation measures such as the creation of representative networks of marine protected areas (MPAs) is a critical step towards the conservation and management of marine ecosystems, as well as to improve public awareness. Conservation efforts in ecologically rich and threatened ecosystems are specially needed. This is particularly urgent for the Mediterranean marine biodiversity, which includes highly mobile marine vertebrates. Here, we studied the at sea distribution of one of the most endangered Mediterranean seabird, the critically endangered Balearic shearwater Puffinus mauretanicus. Present knowledge, from vessel-based surveys, suggests that this species has a coastal distribution over the productive Iberian shelf in relation to the distribution of their main prey, small pelagic fish. We used miniaturised satellite transmitters to determine the key marine areas of the southern population of Balearic shearwaters breeding on Eivissa and spot the spatial connections between breeding and key marine areas. Our tracking study indicates that Balearic shearwaters do not only forage along the Iberian continental shelf but also in more distant marine areas along the North African coast, in particular W of Algeria, but also NE coast of Morocco. Birds recurrently visit these shelf areas at the end of the breeding season. Species distribution modelling identified chlorophyll a as the most important environmental variable in defining those oceanographic features characterizing their key habitats in the western Mediterranean. We identified persistent oceanographic features across time series available in the study area and discuss our results within the current conservation scenario in relation to the ecology of the species.