Assessing the lethality of ship strikes on whales using simple biophysical models

Studies of ship strikes on whales often focus on large vessels (>20 m), with attention to their speeds and the resulting risk of lethality. Smaller coastal vessels also co-occur with whales, resulting in collisions that merit study. To cast light on injuries caused by vessels of all sizes, we used knowledge of right whale anatomy and Newtonian mechanics to construct simple models that predict the mechanical stresses experienced by whales during collisions. By comparing our predictions with published models and with data from ship strikes on various whale species, we developed a model for lethal injury as a function of several vessel and whale properties, finding that collisions that create stresses in excess of 0.241 MPa were likely to cause lethal injuries to large whales. Furthermore, this model has revealed that (1) vessels of all sizes can yield stresses higher than this critical level, and (2) large vessels produce stresses much larger than this even when travelling at reduced speeds (i.e., 10 knots). The model is fast enough to power an interactive GUI-based tool (in R) and flexible enough to simulate strikes by vessels of different masses and speeds upon whales of different species, sizes, and physical conditions.     

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.     

VESSEL COLLISIONS WITH WHALES: THE PROBABILITY OF LETHAL INJURY BASED ON VESSEL SPEED

Historical records demonstrate that the most numerous, per capita, ocean-going-vessel strikes recorded among large-whale species accrue to the North Atlantic right whale ( Eubalaena glacialis ). As vessel speed restrictions are being considered to reduce the likelihood and severity of vessel collisions with right whales, we present an analysis of the published historical records of vessels striking large whales. We examine the influence of vessel speed in contributing to either a lethal injury (defined as killed or severely injured) or a nonlethal injury (defined as minor or no apparent injury) to a large whale when struck. A logistic regression model fitted to the observations, and consistent with a bootstrap model, demonstrates that the greatest rate of change in the probability of a lethal injury ( P _lethal) to a large whale occurs between vessel speeds of 8.6 and 15 knots where P _lethal increases from 0.21 to 0.79. The probability of a lethal injury drops below 0.5 at 11.8 knots. Above 15 knots, P _lethal asymptotically approaches 1. The uncertainties in the logistic regression estimates are relatively large at relatively low speeds ( e.g. , at 8 knots the probability is 0.17 with a 95% CI of 0.03–0.6). The results we provide can be used to assess the utility of vessel speed limits that are being considered to reduce the lethality of vessels striking the critically endangered North Atlantic right whale and other large whales that are frequent victims of vessel strikes.     

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.     

Do ship strikes threaten the recovery of endangered eastern North Pacific blue whales?

Blue whales were targeted in the North Pacific from 1905–1971 and are listed as endangered by the IUCN. Despite decades without whaling, abundance estimates for eastern North Pacific (ENP) blue whales (Balaenoptera musculus) suggest little evidence for a recent increase. One possible reason is fatal strikes by large ships, which have affected populations of other cetaceans and resulted in mitigation. We used a population dynamics model to assess the trends and status of ENP blue whales, and the effects of ship strikes. We estimate the population likely never dropped below 460 individuals, and is at 97% of carrying capacity (95% interval 62%–99%). These results suggest density dependence, not ship strikes, is the key reason for the observed lack of increase. We also estimate future strikes will likely have a minimal impact; for example, an 11‐fold increase in vessels would lead to a 50% chance the long‐term population would be considered depleted. Although we estimate ship strike mitigation would have minimal impacts on population trends and status, current levels of ship strikes are likely above legal limits set by the U.S. The recovery of ENP blue whales from whaling demonstrates the ability of blue whale populations to rebuild under careful management.     

Potential vessel collisions with Southern Hemisphere humpback whales wintering off Pacific Panama

Vessel collision is a threat to many whale species, and the risk has increased with expanding maritime traffic. This compromises international conservation efforts and requires urgent attention from the world's maritime industry. Humpback whales (Megaptera novaeangliae) are at the top of the death toll, and although Central America is a wintering area for populations from both the Northern and Southern Hemispheres, existing efforts to reduce ship‐whale collisions are meager. Herein, we evaluated the potential collisions between vessels and humpback whales wintering off Pacific Panama by following the movements of 15 whales tagged with satellite transmitters and comparing these data with tracks plotted using AIS real‐time latitude‐longitude points from nearly 1,000 commercial vessels. Movements of whales (adults and calves) in the Gulf of Panama coincide with major commercial maritime routes. AIS vessel data analyzed for individual whale satellite tracks showed that 53% (8 whales) of whales had 98 encounters within 200 m with 81 different vessels in just 11 d. We suggest implementing a 65 nmi Traffic Separation Scheme and a 10 kn maximum speed for vessel routing into the Gulf of Panama during the wintering season. In so doing, the area for potential whale‐vessel collisions could be reduced by 93%.     

Cetacean mitochondrial DNA control region: sequences of all extant baleen whales and two sperm whale species

The sequence of the mitochondrial control region was determined in all 10 extant species commonly assigned to the suborder Mysticeti (baleen or whalebone whales) and to two odontocete (toothed whale) species (the sperm and the pygmy sperm whale). In the mysticetes, both the length and the sequence of the control region were very similar, with differences occurring primarily in the first approximately 160 bp of the 5' end of the L-strand of the region. There were marked differences between the mysticete and sperm whale sequences and also between the two sperm whales. The control region, less its variable portion, was used in a comparison including the 10 mysticete sequences plus the same region of an Antarctic minke whale specimen and the two sperm whales. The difference between the minke whales from the North Atlantic and the Antarctic was greater than that between any acknowledged species belonging to the same genus (Balaenoptera). The difference was similar to that between the families Balaenopteridae (rorquals) and Eschrichtiidae (gray whales). The findings suggest that the Antarctic minke whale should have a full species status, B. bonaerensis. Parsimony analysis separated the bowhead and the right whale (family Balaenidae) from all remaining mysticetes, including the pygmy right whale. The pygmy right whale is usually included in family Balaenidae. The analysis revealed a close relationship between the gray whale (family Eschrichtiidae) sequence and those of the rorquals (family Balaenopteridae). The gray whale was included in a clade together with the sei, Bryde's, fin, blue, and humpback whales. This clade was separated from the two minke whale types, which branched together.      

Vessel collision injuries on live humpback whales,Megaptera novaeangliae,in the southern Gulf of Maine

North Atlantic humpback whales (Megaptera novaeangliae) in the Gulf of Maine overlap with both recreational and commercial vessel activity. Vessel strikes are one source of anthropogenic impact that has the potential to inhibit the recovery of this protected species. There are currently no regulations or guidelines specifically devised to reduce the likelihood of collisions for vessels transiting in the vicinity of humpback whales, except for vessels actively engaged in whale watching. To understand interactions between vessels and humpback whales better, we analyzed injuries on 624 individuals photographed in the southern Gulf of Maine from 2004 to 2013. Multiple reviewers evaluated 210,733 photos for five categories of injury consistent with a vessel strike. In total, 14.7% (n = 92) of individuals photographed showed injuries consistent with one or more vessel strikes. These results likely underestimate vessel collision rates and impacts because multiple events, events resulting in mortality, and those that involved only blunt force trauma could not be detected. Nevertheless, our results indicate that vessel strikes are underreported and that healing is dependent on the severity and location of the injury. We recommend that a management strategy be developed for all classes of vessels transiting in the vicinity of whales.     

Global distribution of fin whales Balaenoptera physalus in the post‐whaling era (1980–2012)

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CATCHES OF HUMPBACK AND OTHER WHALES FROM SHORE STATIONS AT MOSS LANDING AND TRINIDAD, CALIFORNIA, 1919–1926

Logbook data from California shore whaling stations at Moss Landing (1919–1922 and 1924) and Trinidad (1920 and 1922–1926) are analyzed. The logs for the two stations record the taking of 2,111 whales, including 1,871 humpbacks, 177 fin whales, 26 sei whales, 3 blue whales, 12 sperm whales, 7 gray whales, 1 right whale, 1 Baird's beaked whale, and 13 whales of unspecified type (probably humpbacks). Most whales were taken from spring to autumn, but catches were made in all months of some years. The sex ratios of humpback, fin, and sei whales (the three species with sufficient sample sizes to test) did not differ from parity. Primary prey, determined from stomach contents, included sardines and euphausiids for both humpback and fin whales, and 'plankton' (probably euphausiids) for sei whales. The prevalence of pregnancy was 0.46 among mature female humpbacks and 0.43 among mature female fin whales, although these values are reported with caution. Information on length distribution for all species is summarized. Analysis of the catch data for this and other areas supports the current view that humpback whales along the west coast of the continental United States comprise a single feeding stock and also suggests that the present population is well below pre-exploitation levels.     

North Atlantic right whales (Eubalaena glacialis) ignore ships but respond to alerting stimuli

North Atlantic right whales were extensively hunted during the whaling era and have not recovered. One of the primary factors inhibiting their recovery is anthropogenic mortality caused by ship strikes. To assess risk factors involved in ship strikes, we used a multi-sensor acoustic recording tag to measure the responses of whales to passing ships and experimentally tested their responses to controlled sound exposures, which included recordings of ship noise, the social sounds of conspecifics and a signal designed to alert the whales. The whales reacted strongly to the alert signal, they reacted mildly to the social sounds of conspecifics, but they showed no such responses to the sounds of approaching vessels as well as actual vessels. Whales responded to the alert by swimming strongly to the surface, a response likely to increase rather than decrease the risk of collision.     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

Evidence that ship noise increases stress in right whales

Baleen whales (Mysticeti) communicate using low-frequency acoustic signals. These long-wavelength sounds can be detected over hundreds of kilometres, potentially allowing contact over large distances. Low-frequency noise from large ships (20-200 Hz) overlaps acoustic signals used by baleen whales, and increased levels of underwater noise have been documented in areas with high shipping traffic. Reported responses of whales to increased noise include: habitat displacement, behavioural changes and alterations in the intensity, frequency and intervals of calls. However, it has been unclear whether exposure to noise results in physiological responses that may lead to significant consequences for individuals or populations. Here, we show that reduced ship traffic in the Bay of Fundy, Canada, following the events of 11 September 2001, resulted in a 6 dB decrease in underwater noise with a significant reduction below 150 Hz. This noise reduction was associated with decreased baseline levels of stress-related faecal hormone metabolites (glucocorticoids) in North Atlantic right whales (Eubalaena glacialis). This is the first evidence that exposure to low-frequency ship noise may be associated with chronic stress in whales, and has implications for all baleen whales in heavy ship traffic areas, and for recovery of this endangered right whale population.     

Status of the world's baleen whales

No global synthesis of the status of baleen whales has been published since the 2008 IUCN Red List assessments. Many populations remain at low numbers from historical commercial whaling, which had ceased for all but a few by 1989. Fishing gear entanglement and ship strikes are the most severe current threats. The acute and long‐term effects of anthropogenic noise and the cumulative effects of multiple stressors are of concern but poorly understood. The looming consequences of climate change and ocean acidification remain difficult to characterize. North Atlantic and North Pacific right whales are among the species listed as Endangered. Southern right, bowhead, and gray whales have been assessed as Least Concern but some subpopulations of these species ‐ western North Pacific gray whales, Chile‐Peru right whales, and Svalbard/Barents Sea and Sea of Okhotsk bowhead whales ‐ remain at low levels and are either Endangered or Critically Endangered. Eastern North Pacific blue whales have reportedly recovered, but Antarctic blue whales remain at about 1% of pre‐exploitation levels. Small isolated subspecies or subpopulations, such as northern Indian Ocean blue whales, Arabian Sea humpback whales, and Mediterranean Sea fin whales are threatened while most subpopulations of sei, Bryde's, and Omura's whales are inadequately monitored and difficult to assess.     

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.     

Abundance and Distribution of Sperm Whales in the Canary Islands: Can Sperm Whales in the Archipelago Sustain the Current Level of Ship-Strike Mortalities?

Sperm whales are present in the Canary Islands year-round, suggesting that the archipelago is an important area for this species in the North Atlantic. However, the area experiences one of the highest reported rates of sperm whale ship-strike in the world. Here we investigate if the number of sperm whales found in the archipelago can sustain the current rate of ship-strike mortality. The results of this study may also have implications for offshore areas where concentrations of sperm whales may coincide with high densities of ship traffic, but where ship-strikes may be undocumented. The absolute abundance of sperm whales in an area of 52933 km², covering the territorial waters of the Canary Islands, was estimated from 2668 km of acoustic line-transect survey using Distance sampling analysis. Data on sperm whale diving and acoustic behaviour, obtained from bio-logging, were used to calculate g(0) = 0.92, this is less than one because of occasional extended periods when whales do not echolocate. This resulted in an absolute abundance estimate of 224 sperm whales (95% log-normal CI 120–418) within the survey area. The recruitment capability of this number of whales, some 2.5 whales per year, is likely to be exceeded by the current ship-strike mortality rate. Furthermore, we found areas of higher whale density within the archipelago, many coincident with those previously described, suggesting that these are important habitats for females and immature animals inhabiting the archipelago. Some of these areas are crossed by active shipping lanes increasing the risk of ship-strikes. Given the philopatry in female sperm whales, replacement of impacted whales might be limited. Therefore, the application of mitigation measures to reduce the ship-strike mortality rate seems essential for the conservation of sperm whales in the Canary Islands.     

Spatial and Temporal Occurrence of Blue Whales off the U.S. West Coast,with Implications for Management

Mortality and injuries caused by ship strikes in U.S. waters are a cause of concern for the endangered population of blue whales (Balaenoptera musculus) occupying the eastern North Pacific. We sought to determine which areas along the U.S. West Coast are most important to blue whales and whether those areas change inter-annually. Argos-monitored satellite tags were attached to 171 blue whales off California during summer/early fall from 1993 to 2008. We analyzed portions of the tracks that occurred within U.S. Exclusive Economic Zone waters and defined the ‘home range’ (HR) and ‘core areas’ (CAU) as the 90% and 50% fixed kernel density distributions, respectively, for each whale. We used the number of overlapping individual HRs and CAUs to identify areas of highest use. Individual HR and CAU sizes varied dramatically, but without significant inter-annual variation despite covering years with El Niño and La Niña conditions. Observed within-year differences in HR size may represent different foraging strategies for individuals. The main areas of HR and CAU overlap among whales were near highly productive, strong upwelling centers that were crossed by commercial shipping lanes. Tagged whales generally departed U.S. Exclusive Economic Zone waters from mid-October to mid-November, with high variability among individuals. One 504-d track allowed HR and CAU comparisons for the same individual across two years, showing similar seasonal timing, and strong site fidelity. Our analysis showed how satellite-tagged blue whales seasonally used waters off the U.S. West Coast, including high-risk areas. We suggest possible modifications to existing shipping lanes to reduce the likelihood of collisions with vessels.     

Hydrodynamic performance of the flippers of large‐bodied cetaceans in relation to locomotor ecology

Cetaceans evolved flippers that are unique in both size and shape probably due to selection pressures associated with foraging and body size. Flippers function as control surfaces for maneuverability and stability. Flippers of cetaceans and engineered hydrofoils are similar with streamlined cross‐sections and wing‐like planforms, which affect lift, drag and hydrodynamic efficiency. Scale models of the flippers from large‐bodied (body length > 6 m) cetaceans (fin whale, killer whale, sperm whale) were constructed from computed tomography (CT) scans of flippers. Flipper planforms were highly tapered for the fin whale, a rounded, paddle‐like design for the killer whale, and a square geometry for the sperm whale. Hydrodynamic properties of the models at varying angles of attack (?40º to 40^o) were determined in a water tunnel with a multi‐axis load cell. The flippers were found to have hydrodynamic characteristics similar to engineered wings. Differences in flipper morphology of large‐bodied cetaceans and their hydrodynamic performance are associated with the requirements of aquatic locomotion involved with ecology of the whales. The flippers of the killer whale provided the greatest maneuverability, whereas the flippers of the fin whale had low drag for lunging and the flippers of the sperm whale provided lift for diving.     

Fin whale (Balaenoptera physalus) target strength measurements

Active acoustic techniques can be used to detect whales. The ability to detect whales from a moving vessel or stationary buoy could reduce conflicts between hazardous human activities and whales, enabling implementation of mitigation procedures. In order to identify acoustic targets correctly as whales, knowledge of whale target strength (TS) is required. Active acoustic detections of fin whales (Balaenoptera physalus) were made in the Norwegian Sea; acoustic data were collected using calibrated omnidirectional sonar, operating at a discrete frequency of 110 kHz. Three fin whales of similar size (estimated between 16 and 18 m total length) had an overall average TS for all insonified body aspects of ?11.4 dB [95% CI ?12.05, ?10.8] at 110 kHz, with a total spread of nearly 14 dB. As expected, the received signals were stronger when the fin whales were insonified at broadside (?5.6 dB). Individual fin whale TS varied by approximately 12 dB, probably due to variation in lung volume with breathing, and to dynamic swimming kinematics. Our TS values are consistent with values reported previously for other large whales. All data together pave the way for development of automated acoustic whale detection protocols that could aid whale conservation.     

Predicted decline of protected whales based on molecular genetic monitoring of Japanese and Korean markets

We present a two-tiered analysis of molecular genetic variation in order to determine the origins of whale' products purchased from retail markets in Japan and the Republic of (South) Korea during 1993-1999. This approach combined phylogenetic analysis of mitochondrial DNA sequences for identification of protected species with a statistical comparison of intraspecific haplotype frequencies for distinguishing regional subpopulations or 'stocks' hunted for scientific research by the Japanese and killed incidentally in coastal fisheries by the Koreans. The phylogenetic identification of 655 products included eight species or subspecies of baleen whales, sperm whales, a pygmy sperm whale, two species of beaked whales, porpoises, killer whales and numerous species of dolphins as well as domestic sheep and horses. Six of the baleen whale species (the fin, sei, common-form and small-form Bryde's, blue or blue/fin hybrid, and humpback) and the sperm whale are protected by international agreements dating back to at least 1989 for all species and 1966 for some species. We compared the haplotype frequencies from the Japanese market sample to those reported from scientific hunting in the western North Pacific stock for products derived from the exploited North Pacific minke whale. The market sample differed significantly from the scientific catch (p < 0.001), showing a greater than expected frequency of haplotypes characteristic of the protected Sea of Japan stock. We used a 'mixed-stock' analysis and maximum-likelihood methods to estimate that 31% (95% confidence interval 19-43%) of the market for this species originated from the Sea of Japan stock. The source of these products was assumed to be undocumented 'incidental takes' from fisheries' by-catch, although we cannot exclude the possibility of illegal hunting or smuggling. The demographic impact of this undocumented exploitation was evaluated using the model of population dynamics adopted by the Scientific Committee of the International Whaling Commission. For the range of exploitation consistent with the market sample, this protected stock was predicted to decline towards extinction over the next few decades. These results confirmed the power of molecular methods in monitoring retail markets and pointed to the inadequacy of the current moratorium for ensuring the recovery of protected species. More importantly, the integration of genetic evidence with a model of population dynamics identified an urgent need for actions to limit undocumented exploitation of a 'protected' stock of whales.