RESPONSE OF NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS) TO PLAYBACK OF CALLS RECORDED FROM SURFACE ACTIVE GROUPS IN BOTH THE NORTH AND SOUTH ATLANTIC

The surface active group (SAG) is the most obvious social interaction of the North Atlantic right whale ( Eubalaena glacialis ). SAGs are typically composed of an adult female with two or more males engaged in social behavior near the surface. Distinct calls, believed to be produced by the female, are associated with these groups. Calls recorded from three North Atlantic right whale SAGs and three South Atlantic right whale ( Eubalaena australis ) SAGs were played back to North Atlantic right whales to determine if these sounds are sufficient to attract males to the groups. Playbacks of gunshot sounds produced by North Atlantic right whales were used as a control stimulus. Thirty-six trials were carried out from 1999 to 2001 in the Bay of Fundy, Canada. Whales approached 27 of 31 SAG playbacks and 0 of 5 gunshot playbacks. Where sex was determined ( n = 28), all approaches to North Atlantic SAG recordings were by males. Individuals ( n = 22) of all age and sex classes approached South Atlantic SAG playbacks. These trials indicate that SAG calls from both populations are sufficient to attract right whales to SAGs and that males and females respond differently to stimuli from the North Atlantic. The difference in response to North and South Atlantic SAG stimuli was unexpected. Novelty, species differences in calls, and different seasonal or behavioral context for the recorded stimuli may be responsible for the differences in response.     

INTER-OBSERVER COUNT DISCREPANCIES IN A SHORE-BASED CENSUS OF GRAY WHALES (ESCHRICHTIUS ROBUSTUS)

Estimations of gray whale abundance have generally assumed that shore-based observers record all whales migrating through the viewing area during periods uncompromised by visibility. We tested the repeatability of data collected at the standard gray whale census site at Granite Canyon Marine Laboratory in central California by using pairs of observers maintaining independent sighting records. Proximal shore sites were occupied 6 d (60 h) in January 1986 where one team counted 845 whales in 427 groups while the other team counted 990 whales in 477 groups. A comparison of the records showed that the first team missed 290 whales seen by the second team, and the second team missed 204 whales seen by the first team. The total number of whales in the viewing area was calculated for each team by the Petersen estimate, using mutually sighted whale groups as "recaptures". On average, observers recorded only 79% of the whales. More whales (68%) were missed when entire groups of whales were not seen rather than when groups were undercounted (32%). Visibility did not appear to affect observed rates of missed whales. Whales migrating at intermediate distances from the shore were less often missed than were those > 6 km or < 1 km offshore. This count discrepancy test confirms that an uncorrected calculation of population size for gray whales based on sighting records from solitary observers will be underestimated.     

NORTH ATLANTIC RIGHT WHALE DISTRIBUTION IN RELATION TO SEA-SURFACE TEMPERATURE IN THE SOUTHEASTERN UNITED STATES CALVING GROUNDS

Standardized aerial surveys were used to document the winter (December–March) distribution of North Atlantic right whales in their calving area off the coasts of Georgia and northeastern Florida (1991–1998). Survey data were collected within four survey zones in and adjacent to federally designated critical habitat. These data, including whale‐sighting locations and sampling effort, were used to describe right whale distribution in relation to sea‐surface temperature (SST) from satellite‐derived images. Locations where whales were sighted (n= 609) had an overall mean SST of 14.3°C ± 2.1° (range 8°–22°C). Data from two survey zones having sufficient data (including the “early warning system” (EWS) zone and the Florida nearshore) were pooled by season and stratified by month to investigate changes in monthly ambient SST and fine‐scale distribution patterns of right whales in relation to SST within spatially explicit search areas. Using Monte Carlo techniques, SSTs and latitudes (means and standard deviations) of locations where whales were sighted were compared to a sampling distribution of each variable derived from daily‐search areas. Overall, results support a nonrandom distribution of right whales in relation to SST: during resident months (January and February), whales exhibited low variability in observed SST and a suggested southward shift in whale distribution toward warmer SSTs in the EWS zone; while in the relatively warmer and southernmost survey zone (Florida nearshore), right whales were concentrated in the northern, cooler portion. Our results support that warm Gulf Stream waters, generally found south and east of delineated critical habitat, represent a thermal limit for right whales and play an important role in their distribution within the calving grounds. These results affirm the inclusion of SST in a multivariate predictive model for right whale distribution in their southeastern habitat.     

BLUE WHALE, BALAENOPTERA MUSCULUS, VOCALIZATIONS FROM THE WATERS OFF CENTRAL CALIFORNIA

Low-frequency calls produced by blue whales, Balaenoptera musculus , were recorded in the northeastern Pacific Ocean off central California. Two blue whales were sighted during a vessel-based marine mammal survey, and when sonobuoys were subsequently deployed, blue whale calls were recorded. A third recording was obtained during the survey from a blue whale that was not seen. Recordings with 15, 25, and 55 min of calls were obtained from these individuals. The three recordings all contain two-part, low-frequency calls with slight interindividual variation. The calls consist of an amplitude modulated (AM) signal with a mean center frequency of 16.5 Hz, followed by a downsweep whose mean center frequency sweeps from 18.2 Hz to 16.6 Hz. The recordings are compared with blue whale recordings from the Pacific and Atlantic Oceans. The geographic variability suggests that blue whale calls may be used as an acoustic indicator of stock identity.     

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.     

RIGHT WHALES (EUBALAENA GLACIALIS) ON JEFFREYS LEDGE: A HABITAT OF UNRECOGNIZED IMPORTANCE?

North Atlantic right whales ( Eubalaena glacialis ) are known to spend the majority of the year between the Great South Channel southeast of Cape Cod, and the Nova Scotian shelf. We examined sightings of right whales on and around Jeffreys Ledge, a 54-km-long glacial deposit off the coast of northern Massachusetts, New Hampshire, and Maine. Sightings on Jeffreys Ledge were extracted from three data sets: (1) a systematic survey of the entire northeastern continental shelf between 1979 and 1982, (2) whale-watch and research-cruise sighting data from 1984 to 1997, and (3) a collaborative database of sightings collected by organizations conducting right whale research and all other available sources. Each database supported two seasonal sighting peaks. During summer (especially July and August) sightings were primarily of mother-calf pairs. Several cow-calf pairs were seen over several days to weeks. Several females were resighted in more than one year, but only when calves were present. During October, November, and December, sightings included all age classes, surface-feeding behavior was frequently observed, and some animals were resighted over several weeks. Given the relatively reduced sighting effort during fall, this number of sightings is surprising. During the 20 yr of observations, 52 of 374 photo-identified North Atlantic right whales (13.9%) were seen at least once on Jeffreys Ledge. We suggest that Jeffreys Ledge may be a more important right whale habitat than previously believed, and that it may play an important role in annual movements and distribution of this population.     

KILLER WHALE PREDATION ON SPERM WHALES: OBSERVATIONS AND IMPLICATIONS

In October 1997 we observed a herd of approximately 35 killer whales ( Orcinus orca ) attack a pod of nine sperm whales ( Physeter macrocephalus ) 130 km off the coast of central California. During the four hours we watched, adult female killer whales, including some with calves, attacked in waves of four to five animals in what was apparently a "wound and withdraw" strategy. Adult male killer whales stood by until the very end when one charged in and quickly killed a seriously wounded sperm whale that had been separated from the group. The sperm whales appeared largely helpless: their main defensive behavior was the formation of a rosette ("marguerite"-heads together, tails out). When the killer whales were successful in pulling an individual out of the rosette, one or two sperm whales exposed themselves to increased attack by leaving the rosette, flanking the isolated individual, and leading it back into the formation. Despite these efforts, one sperm whale was killed and eaten and the rest were seriously, perhaps mortally, wounded. We also present details of two other encounters between sperm whales and killer whales that we observed. Although sperm whales, because of various behavioral and morphological adaptations, were previously thought to be immune to predation, our observations clearly establish their vulnerability to killer whales. We suggest that killer whale predation has potentially been an important, and underrated, selective factor in the evolution of sperm whale ecology, influencing perhaps the development of their complex social behavior and at-sea distribution patterns.     

Acoustic detection and satellite-tracking leads to discovery of rare concentration of endangered North Pacific right whales

The North Pacific right whale, Eubalaena japonica, is one of the most endangered species of whale in the world. On 10 August 2004, two right whales were located in the Bering Sea using headings to right whale calls provided by directional sonobuoys. A satellite-monitored radio tag attached to one of these whales functioned for 40 days. Over the 40-day period, this whale moved throughout a large part of the southeast Bering Sea shelf, including areas of the outer-shelf where right whales have not been seen in decades. In September, multiple right whales were acoustically located and subsequently sighted by another survey vessel approaching a near-real-time position from the tag. An analysis of photographs confirmed at least 17 individual whales (not including the tagged whales). Genetic analysis of biopsy samples identified 17 individuals: 10 males and 7 females. The discovery of seven females was significant, as only one female had been identified in the past. Genetics also confirmed the presence of at least two calves. Although the future of this population is highly uncertain, the discovery of additional females and calves gives some hope that this most critically endangered of all whale populations may still possess the capacity to recover.     

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.     

Determination of steroid hormones in whale blow: It is possible

The feasibility of using analysis of hormone content of whale blow samples to assess reproductive function is addressed. A suitable collection method and analytical technique using liquid chromatography-mass spectrometry (LC-MS) has been developed. Blow samples were collected opportunistically from free-ranging humpback whales ( Megaptera novaeangliae ) ( n = 35) and North Atlantic right whales ( Eubalaena glacialis ) ( n = 18) using a 13-m carbon fiber pole with a collection device. Samples were analyzed for the presence of testosterone and progesterone using a 55% isocratic gradient with LC-MS. Testosterone was detected in four humpback whale samples and eight northern right whale samples. Progesterone was detected in seven humpback whale samples and eight northern right whale samples. This is the first documented use of lung mucosa to determine the presence of reproductive hormones in any free-swimming cetacean and may provide a novel non-invasive technique to quantify the hormonal condition of free-swimming animals that spend brief periods of time at the water's surface.     

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.     

ESTIMATES OF THE NUMBERS OF HUMPBACK WHALES OBSERVED MIGRATING PAST CAPE VIDAL, SOUTH AFRICA, 1988–1991

A recovery-monitoring program of the African east coast humpback whale population was carried out through shore-based visual surveys from Cape Vidal, northern Natal. Surveys of the northward migration were undertaken each winter from 1988 to 1991, and a survey of the southward migration was undertaken in 1990. Independent observer surveys were undertaken during June 1990 and during the entire 1991 survey. Hourly densities of groups sighted each day were adjusted for groups missed by observers with distance from the shore and under different sighting conditions. Densities were multiplied by 24 h and the mean group size of the survey year to give resulting daily densities of individuals, which were summed to provide totals of whales sighted during each year's survey. The best estimate of population size was 1,711 (made during the northward migration of 1990), although this is likely to be biased downwards by a proportion of the population passing outside of observers'view. Bootstrapping of the 1991 daily data resulted in CVs between 11.4% and 12.2%. The numbers sighted show the population to have undergone considerable recovery since protection in October 1963.     

An assessment of sperm whale occurrence and social structure off São Miguel Island,Azores using fluke and dorsal identification photographs

Female and immature sperm whales form stable social units in tropical to subtropical waters. One such area is the Azores archipelago, where details of their year-round occurrence and social organization are not well known. We used year-round sightings data collected from whale watching vessels to assess sperm whale occurrence and social structure off São Miguel Island, Azores from 2010 to 2017. Individuals were photo-identified by their flukes (n = 393) and dorsal pigmentation marks were examined to explore their use in assisting with reidentifications. Of all cataloged whales, 78.8% were sufficiently distinctive to be reidentified from dorsal pigmentation patterns. Associations between individuals were analyzed to determine social structure and delineate social units. We identified 12 units comprising 2–13 members that had stable, and perhaps preferred associations for periods up to the eight years of the study. Preferences between some pairs of units may exist, but more research is required to better understand dynamics within and between units. This local scale study is an important contributor to our knowledge of geographic variation on a global scale. We recommend making use of all available information from the entire archipelago, to further increase our understanding of sperm social organization in the Azores.     

Repeated call sequences and behavioural context in long-finned pilot whales off Cape Breton,Nova Scotia,Canada

Repeated calls are part of the vocal repertoire of a diverse array of species, often presented in sequences that take time and effort on the part of the signal producer. Rhythmic repeated call sequences make up a significant portion of long-finned pilot whale (Globicephala melas) vocal production, yet the function of these sequences has not been investigated until now. In this study, we explored the relationship between behavioural context and the presence of these vocal sequences using recordings of a population of pilot whales found off Cape Breton, Nova Scotia, Canada. We used a binomial logit-link generalized linear model to look for possible predictors of the presence of repeated call sequences. They were more common in recordings of socializing whales than in those of whales in other behavioural states, and least common in resting whales. These vocal repetitions were also more common with larger group size. These results suggest that sequences function in maintaining contact and cohesion within this social species, possibly also serving in individual or group identification. The context of repeated call sequences indicate that they are not primarily mother–calf interactions, as they are heard just as commonly from groups without young. Future studies of pilot whale repeated call sequences should include individual-level behaviour and detailed acoustic calling context.     

Rorqual whale (Balaenopteridae) surface lunge‐feeding behaviors: Standardized classification,repertoire diversity,and evolutionary analyses

Rorqual whales (Family: Balaenopteridae) are the world's largest predators and sometimes feed near or at the sea surface on small schooling prey. Most rorquals capture prey using a behavioral process known as lunge‐feeding that, when occurring at the surface, often exposes the mouth and head above the water. New technology has recently improved historical misconceptions about the natural variation in rorqual lunge‐feeding behavior yet missing from the literature is a dedicated study of the identification, use, and evolution of these behaviors when used to capture prey at the surface. Here we present results from a long‐term investigation of three rorqual whale species (minke whale, Balaenoptera acutorostrata; fin whale, B. physalus; and blue whale, B. musculus) that helped us develop a standardized classification system of surface lunge‐feeding (SLF) behaviors. We then tested for differences in frequency of these behaviors among the three species and across all rorqual species. Our results: (1) propose a unified classification system of six homologous SLF behaviors used by all living rorqual whale species; (2) demonstrate statistically significant differences in the frequency of each behavior by minke, fin, and blue whales; and (3) provide new information regarding the evolution of lunge‐feeding behaviors among rorqual whales.     

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.     

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.     

Changes in North Atlantic right whale (Eubalaena glacialis) cow–calf association times and use of the calving ground: 1993–2005

Like most mysticetes, North Atlantic right whale cows generally separate from their calves on their feeding grounds within a year. Right whale life history data from 1993 to 2005 were analyzed to determine the duration of cow/calf associations and where the pair separated. A change occurred with the 2001 cows; 71% of those available stayed with their calves into the second year and this behavior remained elevated for several years. Less experienced cows, independent of their age, were more likely to extend their associations. The occurrence of cow/yearling associations was not related to the length of the cow's previous interbirth interval, used as a proxy for cow condition, but the hypothesis that body condition impacts how long cows nurse their young could not be adequately tested. Seventy-seven percent of the observed cow/yearling pairs also returned to the calving ground, a substantial physiological investment given the 1,450 km plus migration and the fact that they fast there, indicating that factors other than nutrition also influenced the cow's behavior. The concurrent increase in juveniles in the shallow waters of the winter calving grounds may afford naive whales greater protection from predators or provide a social benefit that improves their overall fitness.     

Population size and migratory connectivity of humpback whales wintering in Las Perlas Archipelago,Panama

From 2003 to 2009, we surveyed Las Perlas Archipelago off the Pacific coast of Panama 53 times between the months of August and October to estimate abundance of humpback whales and to test for a migratory connection with populations from the southern hemisphere. We identified 295 individuals using photo‐identification of dorsal fins, including 58 calves, and the population estimate for a single season was 100–300 solitary adults plus 25–50 mothers with calves; the estimated population of animals across all seasons using a mark and recapture model was over 1,000. Eight of the 139 fluke identifications were matched to whales in photograph catalogues from the Antarctic Peninsula and a ninth was matched to a whale sighted in Chilean waters; four of these nine individuals have also been sighted in Colombia. We conclude that Panama (Las Perlas Archipelago in particular) is an important calving area for humpback whales in the Southern Hemisphere. These data should provide a foundation for monitoring of population change and to increase awareness in Panama about the need to manage vessel traffic and tourism related to the whales at Las Perlas.     

Phenological changes in North Atlantic right whale habitat use in Massachusetts Bay

The North Atlantic right whale (Eubalaena glacialis) is one of the world's most highly endangered baleen whales, with approximately 400–450 individuals remaining. Massachusetts Bay (MB) and Cape Cod Bay (CCB) together comprise one of seven areas in the Gulf of Maine where right whales seasonally congregate. Here, we report on acoustically detected presence of right whales in MB over a nearly 6 year period, July 2007–April 2013, a time of both rapid ocean warming throughout the Gulf of Maine and apparent changes in right whale migratory dynamics. We applied an automated detection algorithm to assess hourly presence of right whale “up‐calls” in recordings from a 19‐channel acoustic array covering approximately 4,000 km² in MB. Over the survey, up‐calls were detected in 95% of 8 day periods. In each year, as expected, we observed a “peak season” of elevated up‐call detections in late winter and early spring corresponding to the season when right whales congregate to feed in CCB. However, we also saw an increase in right whale occurrence during time periods thought to be part of the “off‐season.” With the exception of 2009–2010, when acoustic presence was unusually low, the mean percent of hours in which up‐calls were detected increased every year, both during the peak season (from 38% in 2008 to 70% in 2012), and during the summer–fall season (from 2% in 2007 to 13% in 2012). Over the entire study, the peak season start date varied between 17 January and 26 February. Changes in right whale phenology in MB likely reflect broadscale changes in habitat use in other areas within the species range. This study demonstrates the value of continuous long‐term survey datasets to detect and quantify shifts in cetacean habitat use as environmental conditions change and the long‐term continued survival of right whales remains uncertain.