BEHAVIORAL RESPONSE OF FOUR SPECIES OF BALAENOPTERID WHALES TO BIOPSY SAMPLING

Behavioral responses to biopsy sampling of four species of northwestern Atlantic balaenopterid whales summering in the estuary and Gulf of St. Lawrence, Quebec, from 1990 to 1995 were studied to determine if this technique was an important disturbance to the whales. A total of 447 biopsy samples were taken using a small punch-type biopsy tip fired from a crossbow. Biopsies were successfully taken from 91.2% of the whales approached. Whales displayed no reaction to 45.2% of the successful biopsy attempts. Whales that responded to biopsy sampling typically resumed their normal behavior immediately or within a few minutes. Most humpback whales displayed a hard tail flick, and the majority of fin and blue whales submerged following biopsy sampling. Significantly different frequencies and intensities of responses were found between whale species. Minke and humpback whales were found to be more sensitive to biopsy sampling than fin and blue whales. Response frequencies were similar between females and males for all species, with the exception of fin whales where females had a higher response frequency than males. Biopsy sample length, i. e., penetration depth, did not explain variations in response intensity but may influence response frequency to biopsy sampling. Group size, geographical region, and number of biopsies taken per whale were not factors that explained variation in behavioral responses. The biopsy technique was found to be an efficient method for obtaining high-quality whale skin and blubber samples with limited behavioral disturbance to balaenopterid whales.     

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.     

Cetaceans in British waters

Most information on the distribution, movements and ecology of cetaceans in the N.E. Atlantic have come from whale catches mainly in the early part of this century, and from strandings records collected by the British Museum (Nat. Hist.). With the formation of the Cetacean Group in 1973, a scheme for recording live cetaceans at sea was started. This paper summarizes the results of about two thousand sightings involving nearly 25,000 individual animals between the years 1958– 1978 (but mainly from the last 10 years), and relates them to existing information collected from other sources. Difficulties of identification and potential sources of bias are discussed. Most large cetaceans are present in British waters as part of a latitudinal feeding migration whereas smaller species may be present in the N.E. Atlantic throughout the year with movements being mainly of an offshore-inshore nature. Some species are clearly very rare probably as a result of over-exploitation in the last century and early part of this century. These include the Right whale, Blue whale and probably Humpback whale. Other species are rarely recorded because their usual range is some distance from British waters. These include narwhal and White whale (from Arctic waters), Pygmy sperm whale, smaller beaked whales and Euphrosyne dolphin (from warm temperate to tropical waters). The Harbour porpoise is by far the most common and widespread species in British waters, occurring mainly in inshore waters, although it has apparently declined in certain regions (e.g. Southern North Sea, English Channel, Irish Sea) in recent years probably as a result of pollution, disturbance and/or over-exploitation of food resources. Bottle-nosed and Risso's dolphins are also widely distributed close to the coast, although the latter is restricted to the west and south coasts and the former is associated particularly with some large estuaries. Common dolphins are relatively abundant and widespread, and are more pelagic than the previous three species. White-sided dolphins have a mainly pelagic distribution centred on the Northern North Sea whilst the White-sided dolphin has a wider distribution which includes all the western seaboard. Of larger cetaceans, the Killer whale is relatively common particularly on the west coasts and the Pilot whale is locally and seasonally abundant at the north and south ends of Britain and Ireland where they probably represent distinct populations. The Bottlenose whale, Minke, Fin and Sei whales are confined to the west and north coasts, all but the Minke whale having a primarily pelagic distribution. Sperm whales although increasingly commonly stranded on British coasts, are rarely sighted in inshore waters. The west coast of Britain and Ireland are the most important regions for cetaceans whereas the Southern North Sea has the smallest number although in previous decades numbers were probably higher. Most cetacean species occur mainly in the summer months, particularly August and September, although some species, e.g. White-sided Dolphin, Pilot whale and Minke whale show peaks later in the year. A number of species show secondary spring peaks, e.g. Bottle-nosed and Common dolphins, Risso's dolphins, and Pilot whales. Present evidence suggests that only the large whales exhibit definite latitudinal migrations, all other species being resident at high latitudes although they may show offshore-inshore or possibly small latitudinal movements. Many of the movements indicated from the present analysis can be linked to the seasonal changes in food availability and to the timing and geographical location of breeding, and these are described in detail. Many concentrations of a particular cetacean species occur regularly in the same area year after year and these may often be related to spawning concentrations of a particular fish species. Variations in herd size are noted between species and within species at different times of the year. These are related to aggregations associated with feeding, breeding, and long-distance movements winch will vary according to the biology and ecology of different cetacean species.     

Relationship between the distribution of euphausiids and baleen whales in the Antarctic (35°E – 145°W)

Simultaneous whale sighting and hydroacoustic surveys were conducted from a research vessel in the Antarctic to examine the relationship between the distribution of euphausiids and baleen whales. High densities of minke whales and large aggregations of euphausiids were observed along the ice edge over the continental slope in the southeast region of area IV and in the southwest region of area V. The results suggest that the continental slope zone that coincides with the ice edge would be an important minke whale feeding area. Minke whales were rarely sighted in the offshore region even if euphausiids were abundant. Distributions of humpback whales were correlated with high euphausiid density zones, regardless of the bottom topographic features. Several groups of blue whales were sighted in the small area along the ice edge where euphausiids were abundant, but sightings were too few to draw any general conclusion about the relationship between blue whales and euphausiids. Both baleen whales and euphausiids were scarce in the area east of 170°W where sea ice covered the continental shelf and slope zone.     

Temperature regulation of marine mammals

A mathematical model of heat loss from an aquatic animal to the surrounding water is presented. Heat is generated in metabolically active tissues and distributed by circulating blood and by conduction. The time dependent radial temperature profile of the animal is numerically solved from heat transfer equations by a computer. The model is applied to large whales, porpoises, and seals. For the whales, blood circulation to the dermal layer below appendage and body skin surfaces proved to be essential for sufficient heat dissipation. When decreasing the blood flow below a certain value (dependent on sea temperature and whale activity) the large whales would overheat. Blubber thickness was found to be of minor importance in whale thermoregulation, because the blubber coat can be bypassed by blood circulation. On the other hand, it is in general not possible for small porpoises and seals to stay warm in the coldest waters using normal mammalian resting metabolic rates, even if the peripheral circulation is shut off (or artery-vein heat exchangers used). Heat loss can be reduced if the outermost tissue layers are allowed to cool. This is achieved by minimizing convective radial heat flow via the circulation. (For large whales even minute radial blood flow raises the muscle temperatures to the core temperature level.) Seasonal acclimatization of harbour seals is explained by changes in their effective insulation thickness. Differences in whale activity induce changes in the temperature profile mainly within the first few centimeters from the skin surface. These superficial temperatures, if known, could be used to estimate whale metabolic rates. Since they drop close to the sea water temperature within minutes after whale death, the measurements should be done of live whales.     

Description and Analysis of the use of Cold Harpoons in the Norwegian Minke Whale Hunt in the 1981,1982 and 1983 Hunting Seasons

Until 1984, cold harpoons, i.e. harpoons with no detonating device, were used to hunt minke whales in Norway. To investigate the effectiveness of such harpoons and compare them with alternatives, data on kills using cold harpoons were collected as part of a project dealing with alternative killing techniques for whales. Data on 353 whale kills were collected in 1981–83. The criteria used to determine the time of death were cessation of flipper movement, that the mandible relaxed, or that the whale hung immobile from the harpoon line. These criteria do not take into account any movements caused by spinal reflexes. About 17% of the animals died instantaneously (≤10 s). The median survival time was 570 s. Animals died most rapidly if hit in the brain, heart or major blood vessels. If only the lungs were injured, minke whales died less rapidly than terrestrial mammals. For whales that did not die immediately, shooting range, animal size and the angle of the shot all influenced the time to death. The efficiency of cold harpoons could be improved, but their use was no longer considered acceptable, and they were replaced by harpoons with penthrite grenades in 1984.     

Chemical characterization of the oligosaccharides in beluga (Delphinapterus leucas) and Minke whale (Balaenoptera acutorostrata) milk

Carbohydrates were extracted from the milk of a beluga, Delphinopterus leucas (family Odontoceti), and two Minke whales, Balaenoptera acutorostrata (Family Mysticeti), sampled late in their respective lactation periods. Free oligosaccharides were separated by gel filtration and then neutral oligosaccharides were purified by preparative thin layer chromatography and gel filtration, while acidic oligosaccharides were purified by ion-exchange chromatography, gel filtration and high performance liquid chromatography (HPLC). Their structures were determined by 1H-NMR. In one of the Minke whale milk samples, lactose was a dominant saccharide, with Fuc(alpha1-2)Gal(beta1-4)Glc(2'-fucosyllactose), Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc(lacto-N-neotetraose), GalNAc(alpha1-3)[Fuc(alpha1-2)]Gal(beta1-4)Glc(A-tetrasaccharide), Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (para lacto-N-neohexaose), Neu5Ac(alpha2-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (sialyl lacto-N-neotetraose), Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (LST c) and Neu5Ac(alpha2-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (sialyl para lacto-N-neohexaose) also being found in the milk. The second Minke whale sample contained similar amounts of lactose, 2'-fucosyllactose and A-tetrasaccharide, but no free sialyl oligosaccharides. Sialyl lacto-N-neotetraose and sialyl para lacto-N-neohexaose are novel oligosaccharides which have not been previously reported from any mammalian milk or colostrum. These and other oligosaccharides of Minke whale milk may have biological significance as anti-infection factors, protecting the suckling young against bacteria and viruses. The lactose of Minke whale milk could be a source of energy for them. The beluga whale milk contained trace amounts of Neu5Ac(alpha2-3)Gal(beta1-4)Glc(3'-N-acetylneuraminyllactose), but the question of whether it contained free lactose could not be clarified. Therefore, lactose may not be a source of energy for suckling beluga whales.     

High Velocity Projectiles for Killing Whales. Hunting Trials using 20 mm High Velocity Projectiles for Minke Whales in 1982

Norway was requested by the International Whaling Commission (IWC) to explore the use of high-velocity projectiles to replace cold harpoon as killing device for minke whales (Anon 1980). Tests of suitable high-velocity projectiles for minke whales were therefore initiated in 1982 as part of a wider project with the purpose of studying alternative killing methods to the traditional cold harpoon used in the Norwegian minke whale hunt until 1984 (Øen 1995). The results of the trials have previously been presented in unpublished reports to the IWC (Øen 1982, 1983, 1992).     

Underwater acrobatics by the world's largest predator: 360° rolling manoeuvres by lunge-feeding blue whales

The extreme body size of blue whales requires a high energy intake and therefore demands efficient foraging strategies. As an obligate lunge feeder on aggregations of small zooplankton, blue whales engulf a large volume of prey-laden water in a single, rapid gulp. The efficiency of this feeding mechanism is strongly dependent on the amount of prey that can be captured during each lunge, yet food resources tend to be patchily distributed in both space and time. Here, we measured the three-dimensional kinematics and foraging behaviour of blue whales feeding on krill, using suction-cup attached multi-sensor tags. Our analyses revealed 360° rolling lunge-feeding manoeuvres that reorient the body and position the lower jaws so that a krill patch can be engulfed with the whale''s body inverted. We also recorded these rolling behaviours when whales were in a searching mode in between lunges, suggesting that this behaviour also enables the whale to visually process the prey field and maximize foraging efficiency by surveying for the densest prey aggregations. These results reveal the complex manoeuvrability that is required for large rorqual whales to exploit prey patches and highlight the need to fully understand the three-dimensional interactions between predator and prey in the natural environment.     

Modern whaling in Britain and the north-east Atlantic Ocean

Modern whaling, using an explosive harpoon fired from a steam catcher-boat to kill the fast-swimming rorquals, began from shore whaling stations in northern Norway in the 1860s. It spread to Iceland, the Faeroe Islands and Spitsbergen, before reaching the British Isles in 1903.
Whaling took place from four stations in the Shetland Islands, one in the Outer Hebrides, and two in Ireland, before the First World War. Fin whales were the main species caught but Blue, Humpback, Sei, Right, Sperm and Bottle-nosed whales were also taken. Four stations re-opened in 1920, but from 1923 onwards only two continued to operate and whaling ceased in 1929, though the Hebridean station worked again for two seasons in 1950–1951.
The species composition of catches at the Hebridean and Irish stations was very similar and different from that of the stations in the Shetland Islands where few Blue whales and Right whales were taken. There is evidence that Fin whales were being overfished on the Shetland Islands whaling grounds at an early date, and that Blue whales and Right whales, but not Fin whales, declined in numbers on the Hebridean grounds.
The history of modern whaling in the north-east North Atlantic region as a whole indicates that the stocks of Blue, Humpback and Right whales were not large enough to support continuous whaling on the scale which took place there. The development of whaling since 1945 supports the view that there are separate populations of Fin whales in the region. The numbers of this species have declined on the whaling grounds of the Faeroe Islands and western Norway, and possibly also of north Norway, but not on the Icelandic grounds where there is no evidence of overfishing.     

KILLER WHALE ATTACKS ON MINKE WHALES: PREY CAPTURE AND ANTIPREDATOR TACTICS

We describe nine incidents of predation or attempted predation of minke whales ( Balaenoptera acutorostrata ) by mammal-hunting "transient" killer whales ( Orcinus orca ) in coastal waters of British Columbia, Washington, and southeastern Alaska. Pursuits of minke whales were characterized by prolonged chases on a straight heading at velocities of 15–30 km/h. In four of the nine cases the adultsized minke whale gradually outdistanced the killer whales, which abandoned the high-speed pursuit after 0.5–1 h. In one case the minke beached itself and died. Four attacks were successful. In one instance a subadult minke was killed in open water following a chase. In two cases the fleeing minke entered a confined bay and was killed by the killer whales. One adult minke was taken after apparently attempting to seek cover beside a large sailboat. Minke whales made no attempt to physically defend themselves and were killed by repeated ramming or by asphyxiation. Although killer whales are capable of sprinting speeds greater than those of minke whales, it appears that adult minkes can maintain higher sustained speeds and evade capture if sufficient space for an extended escape trajectory is available. Successful predation of minke whales in coastal waters is rare compared to pinnipeds and small cetaceans, the main prey of transient killer whales.     

Behavioral responses of minke whales (Balaenoptera acutorostrata) to experimental fishing gear in a coastal environment

Whale entanglement in fishing gear is a global problem, and underwater ropes associated with this gear are often the cause of injuries that can lead to fatalities. Minke whales (Balaenoptera acutorostrata) are especially at risk because they are relatively small, widely distributed, and often occur in coastal habitats where many types of fishing gear are deployed. It is unknown whether minke whales can detect and avoid ropes associated with fishing gear. To address this question we conducted a series of field experiments to measure behavioral responses of nearshore minke whales to underwater ropes simulating crab and whelk fishing gear. We also investigated correlations between whale behaviors and specific environmental variables. Our methods involved both visual and acoustic monitoring of whale behaviors near experimental ropes and buoys of different colors. A remote sensing system was also used to simultaneously monitor oceanographic conditions, record underwater sounds, and capture underwater video of whales swimming near ropes. Minke whales (N = 42) decreased their swimming velocity and altered their bearing when passing near experimental ropes, especially during trials with white and black ropes. Some minkes (N = 7) also altered their underwater swimming trajectories when passing near ropes, and often appeared to produce low-frequency vocalizations. Collectively this information provides strong evidence that minke whales detect and react behaviorally to the presence of underwater ropes that simulate fishing gear in nearshore areas. We hypothesize that visual and perhaps passive acoustic sensory abilities may be the mechanisms behind these rope avoidance behaviors. We recommend that high contrast ropes (white and black) be used with fishing gear in coastal areas to help minimize entanglements of minke and perhaps other whale species.     

Chemical characterization of the oligosaccharides in Bryde's whale (Balaenoptera edeni) and Sei whale (Balaenoptera borealis lesson) milk

Samples of milk from a Bryde's whale and a Sei whale contained 2.7 g/100 mL and 1.7 g/100 mL of hexose, respectively. Both contained lactose as the dominant saccharide along with small amounts of Neu5Ac(alpha2-3)Gal(beta1-4)Glc (3'-N-acetylneuraminyllactose), Neu5Ac(alpha2-6)Gal(beta1-4)Glc (6'-N-acetylneuraminyllactose) and Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc (LST c). The dominance of lactose in the carbohydrate of these milks is similar to that of Minke whale milk and bottlenose dolphin colostrum, but the oligosaccharide patterns are different from those of these two species, illustrating the heterogeneity of milk oligosaccharides among the Cetacea.     

Limited MHC polymorphism in whales

Little is known about disease and genetic variation in aquatic mammalian species such as whales. In this paper human HLA class I and class 11 probes were used to study major histocompatibility complex (MHC) genes from two species of whale: Fin (Balaenoptera physalus) and Sei (B. borealis). Stronger signals were obtained on whale than on equivalent concentrations of mouse DNA. Evidence was obtained for severalDRB-related genes, aDNA gene, oneDQA gene, and multiple class I genes in whales. Interestingly, the whale genes, from the small panel studied, were less polymorphic than those of humans or mice. The aquatic environment of this mammalian species may be a unique factor in shaping its immune response through the MHC.     

In vitro maturation and ultrastructural observation of cryopreserved minke whale (Balaenoptera acutorostrata) follicular oocytes

Minke whale (Balaenoptera acutorostrata) follicular oocytes were cryopreserved by a slow-step freezing procedure using ethylene glycol. The morphologically viable proportion of postthawed minke whale follicular oocytes was 39.7%. The maturity of the animals (immature and mature whales) or the presence or absence of cumulus cells (CC) did not affect the proportion of morphologically viable oocytes. Postthawed oocytes were examined for nuclear status after in vitro maturation. The presence of CC (29.1%) significantly enhanced (P < 0.05) the proportion of oocytes at metaphase I/anaphase I/telophase I stages compared to results with the absence of CC (13.5%). A total of 4 of 194 postthawed oocytes matured to the second metaphase stage after culture for 5.5 days with or without CC. The cryopreserved immature oocytes obtained from immature and mature whales were processed to examine the ultrastructure by transmission electron microscopy. Varying ultrastructural damage to the cytoplasm was observed as a result of the cryopreservation procedures. These results show that 20-30% of cryopreserved minke whale follicular oocytes can resume meiosis in vitro, but damage induced by the freezing and thawing procedures was observed.     

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.     

Distribution and abundance of sei whales off the west coast of the Falkland Islands

Little information exists on the current status of Southern Hemisphere sei whales (Balaenoptera borealis). We assessed their distribution and abundance along the west coast of the Falkland Islands (southwest Atlantic) during February and March 2018, using line transect and nonsystematic surveys. Abundance estimates were generated for a single survey stratum using design- and model-based approaches. Sightings of sei whales and unidentified baleen whales (most, if not all, likely to be sei whales) occurred from the coast to the 100 m depth isobath that marked the offshore boundary of the stratum. The modeled distribution predicted highest whale densities in King George Bay and in the waters between Weddell Island and the Passage Islands. Sei whale abundance was estimated as 716 animals (CV = 0.22; 95% CI [448, 1,144]; density = 0.20 whales/km²) using the design-based approach, and 707 animals (CV = 0.11; 95% CI [566, 877]; density = 0.20 whales/km²) using the model-based approach. For sei whales and unidentified baleen whales combined, the equivalent estimates were 916 animals (CV = 0.19; 95% CI [606, 1,384]; density = 0.26 whales/km²) and 895 animals (CV = 0.074; 95% CI [777, 1,032]; density = 0.25 whales/km²). The data indicate that the Falkland Islands inner shelf region may support globally important seasonal feeding aggregations of sei whales, and potentially qualify as a Key Biodiversity Area.     

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.      

Gray whales hear and respond to signals from a 21–25 kHz active sonar

Shore-based theodolite tracking of eastern gray whale (Eschrichtius robustus) movements was conducted to test for potential whale responses to a high-frequency sonar system. Southbound migrating whales were observed from two California shore observation stations as the whales swam past the source vessel that was moored in their migration path. The sonar transducer was deployed from the vessel during all observations, broadcasting 21–25 kHz sweeps for half of each day, the other half remaining silent. The order of control and experimental periods was randomized. No readily apparent response to sonar transmissions was observed in the field or in the visual data. Statistical analysis of tracking data indicates that, compared to control data, gray whales deflected inshore at ranges of 1–2 km from the vessel during sonar transmissions at a received sound pressure level of approximately 148 dB re 1 μPa² (134 dB re 1 μPa²s). These data suggest that the functional hearing sensitivity of gray whales extends to at least 21 kHz.