Temporal range: Quaternary–Recent
|At the Georgia Aquarium|
|Size compared to an average human|
The beluga whale (//) (Delphinapterus leucas) is an Arctic and sub-Arctic cetacean. It is one of two members of the family Monodontidae, along with the narwhal, and the only member of the genus Delphinapterus. It is also known as the white whale, as it is the only marine mammal of this colour; the sea canary, due to its high-pitched calls; and the melonhead, though that more commonly refers to the melon-headed whale, which is an oceanic dolphin.
The beluga is adapted to life in the Arctic, so has anatomical and physiological characteristics that differentiate it from other cetaceans. Amongst these are its all-white colour and the absence of a dorsal fin. It possesses a distinctive protuberance at the front of its head which houses an echolocation organ called the melon, which in this species is large and deformable. The beluga's body size is between that of a dolphin's and a true whale's, with males growing up to 5.5 m (18 ft) long and weighing up to 1,600 kg (3,530 lb). This whale has a stocky body. A large percentage of its weight is blubber (subcutaneous fat), as is true of many cetaceans. Its sense of hearing is highly developed and its echolocation allows it to move about and find breathing holes under sheet ice.
Belugas are gregarious and form groups of up to 10 animals on average, although during the summer, they can gather in the hundreds or even thousands in estuaries and shallow coastal areas. They are slow swimmers, but can dive to 700 m (2,300 ft) below the surface. They are opportunistic feeders and their diets vary according to their locations and the season. The majority of belugas live in the Arctic Ocean and the seas and coasts around North America, Russia and Greenland; their worldwide population is thought to number around 150,000. They are migratory and the majority of groups spend the winter around the Arctic ice cap; when the sea ice melts in summer, they move to warmer river estuaries and coastal areas. Some populations are sedentary and do not migrate over great distances during the year.
The native peoples of North America and Russia have hunted belugas for many centuries. They were also hunted by non-natives during the 19th century and part of the 20th century. Hunting of belugas is not controlled by the International Whaling Commission, and each country has developed its own regulations in different years. Currently some Inuit in Canada and Greenland, Alaska Native groups, and Russians are allowed to hunt belugas to consume and sell; aboriginal whaling is excluded from the International Whaling Commission 1986 moratorium on hunting. The numbers have dropped substantially in Russia and Greenland, but not in Alaska and Canada. Other threats include natural predators (polar bears and killer whales), contamination of rivers (as with Polychlorinated biphenyl (PCBs) which bioaccumulate up the food chain), and infectious diseases. The beluga was placed on the International Union for Conservation of Nature's Red List in 2008 as being "near threatened"; the subpopulation from the Cook Inlet in Alaska, however, is considered critically endangered and is under the protection of the United States' Endangered Species Act. Of seven Canadian beluga populations, those inhabiting eastern Hudson Bay, Ungava Bay, and the St. Lawrence River are listed as endangered.
Belugas are one of the most commonly kept cetaceans in captivity and are housed in aquariums, dolphinariums, and wildlife parks in North America, Europe, and Asia. They are popular with the public due to their colour and expression.
- 1 Taxonomy
- 2 Evolution
- 3 Description
- 4 Behaviour
- 5 Distribution
- 6 Population
- 7 Threats
- 8 Relationship with humans
- 9 Conservation status
- 10 Cultural references
- 11 See also
- 12 References
- 13 Further reading
- 14 External links
The beluga was first described in 1776 by Peter Simon Pallas. It is a member of the Monodontidae family, which is in turn part of the parvorder Odontoceti (toothed whales). The Irrawaddy dolphin was once placed in the same family; recent genetic evidence suggests these dolphins belong to the family Delphinidae. The narwhal is the only other species within the Monodontidae besides the beluga. A skull has been discovered with intermediate characteristics supporting the hypothesis that hybridization is possible between these two families.
The name of the genus, Delphinapterus, means "dolphin without fin" (from the Greek δελφίν (delphin), dolphin and απτερος (apteros), without fin) and the species name leucas means "white" (from the Greek λευκας (leukas), white). The Red List of Threatened Species gives both beluga and white whale as common names, though the former is now more popular. The English name comes from the Russian белуха (belukha), which derives from the word белый (bélyj), meaning "white". The name beluga in Russian refers to an unrelated species, a fish, beluga sturgeon.
The whale is also colloquially known as the sea canary on account of its high-pitched squeaks, squeals, clucks, and whistles. A Japanese researcher says he taught a beluga to "talk" by using these sounds to identify three different objects, offering hope that humans may one day be able to communicate effectively with sea mammals. A similar observation has been made by Canadian researchers, where a beluga which died in 2007 "talked" when he was still a subadult. Another example is NOC, a beluga whale that could mimic the rhythm and tone of human language. Beluga whales in the wild have been reported to imitate human voices.
Mitochondrial DNA studies have shown modern cetaceans last shared a common ancestor between 30 and 34 million years ago. The family Monodontidae separated relatively early from the other odontoceti; it split from the Delphinoidea between 11 and 15 million years ago, and from the Phocoenidae, its closest relatives in evolutionary terms, more recently still. In 2017 the genome of a beluga whale was sequenced, comprising 2.327 Gbp of assembled genomic sequence that encoded 29,581 predicted genes. The authors estimated that the genome-wide sequence similarity between beluga whales and killer whales to be 97.87% ± 2.4 × 10−7% (mean ± standard deviation).
The beluga's earliest known ancestor is the prehistoric Denebola brachycephala from the late Miocene period (9–10 million years ago). A single fossil from the Baja California Peninsula indicates the family once inhabited warmer waters. The fossil record also indicates, in comparatively recent times, the beluga's range varied with that of the polar ice packs expanding during ice ages and contracting when the ice retreated. Counter-evidence to this theory comes from the finding in 1849 of fossilised beluga bones in Vermont in the United States, 240 km (150 mi) from the Atlantic Ocean. The bones were discovered during construction of the first railroad between Rutland and Burlington in Vermont, when workers unearthed the bones of a mysterious animal in Charlotte. Buried nearly 10 ft (3.0 m) below the surface in a thick blue clay, these bones were unlike those of any animal previously discovered in Vermont. Experts identified the bones as those of a beluga. Because Charlotte is over 150 mi (240 km) from the nearest ocean, early naturalists were at a loss to explain the presence of the bones of a marine mammal buried beneath the fields of rural Vermont. The remains were found to be preserved in the sediments of the Champlain Sea, an extension of the Atlantic Ocean within the continent resulting from the rise in sea level at the end of the ice ages some 12,000 years ago. Today, the Charlotte whale is the official Vermont State Fossil (making Vermont the only state whose official fossil is that of a still extant animal).
Its body is round, particularly when well fed, and tapers less smoothly to the head than the tail. The sudden tapering to the base of its neck gives it the appearance of shoulders, unique among cetaceans. The tailfin grows and becomes increasingly and ornately curved as the animal ages. The flippers are broad and short—making them almost square-shaped.
Preliminary investigations suggested a beluga's life expectancy was rarely more than 30 years. The method used to calculate the age of a beluga is based on counting the layers of dentin and dental cement in a specimen's teeth, which were originally thought to be deposited once or twice a year. The layers can be readily identified as one layer consists of opaque dense material and the other is transparent and less dense. It is therefore possible to estimate the age of the individual by extrapolating the number of layers identified and the estimated frequency with which the deposits are laid down. A 2006 study using radiocarbon dating of the dentine layers showed the deposit of this material occurs with a lesser frequency (once per year) than was previously thought. The study therefore estimated belugas can live for 70 or 80 years.
The species presents a moderate degree of sexual dimorphism, as the males are 25% longer than the females and are sturdier. Adult male belugas can range from 3.5 to 5.5 m (11 to 18 ft), while the females measure 3 to 4.1 m (9.8 to 13.5 ft). Males weigh between 1,100 and 1,600 kg (2,430 and 3,530 lb), occasionally up to 1,900 kg (4,190 lb) while females weigh between 700 and 1,200 kg (1,540 and 2,650 lb). They rank as mid-sized species among toothed whales.
Individuals of both sexes reach their maximum size by the time they are 10 years old. The beluga's body shape is stocky and fusiform (cone-shaped with the point facing backwards), and they frequently have folds of fat, particularly along the ventral surface. Between 40% and 50% of their body weight is fat, which is a higher proportion than for cetaceans that do not inhabit the Arctic, where fat only represents 30% of body weight. The fat forms a layer that covers all of the body except the head, and it can be up to 15 cm (5.9 in) thick. It acts as insulation in waters with temperatures between 0 and 18 °C, as well as being an important reserve during periods without food.
The adult beluga is rarely mistaken for any other species, because it is completely white or whitish-grey in colour. Calves are usually born grey, and by the time they are a month old, have turned dark grey or blue grey. They then start to progressively lose their pigmentation until they attain their distinctive white colouration, at the age of seven years in females and 9 in males. The white colouration of the skin is an adaptation to life in the Arctic that allows belugas to camouflage themselves in the polar ice caps as protection against their main predators, polar bears and killer whales. Unlike other cetaceans, the belugas seasonally shed their skin. During the winter, the epidermis thickens and the skin can become yellowish, mainly on the back and fins. When they migrate to the estuaries during the summer, they rub themselves on the gravel of the riverbeds to remove the cutaneous covering.
Head and neck
Like most toothed whales, it has a compartment found at the centre of the forehead that contains an organ used for echolocation called a melon, which contains fatty tissue. The shape of the beluga's head is unlike that of any other cetacean, as the melon is extremely bulbous, lobed, and visible as a large frontal prominence. Another distinctive characteristic it possesses is the melon is malleable; its shape is changed during the emission of sounds. The beluga is able to change the shape of its head by blowing air around its sinuses to focus the emitted sounds. This organ contains fatty acids, mainly isovaleric acid (60.1%) and long-chain branched acids (16.9%), a very different composition from its body fat, and which could play a role in its echolocation system.
Unlike many dolphins and whales, the seven vertebrae in the neck are not fused together, allowing the animal to turn its head laterally without needing to rotate its body. This gives the head a lateral manoeuvrability that allows an improved field of view and movement and helps in catching prey and evading predators in deep water. The rostrum has about eight to 10 small, blunt, and slightly curved teeth on each side of the jaw and a total of 36 to 40 teeth. Belugas do not use their teeth to chew, but for catching hold of their prey; they then tear them up and swallow them nearly whole.
Belugas only have a single spiracle, which is located on the top of the head behind the melon, and has a muscular covering, allowing it to be completely sealed. Under normal conditions, the spiracle is closed and an animal must contract the muscular covering to open the spiracle. A beluga's thyroid gland is larger than that of terrestrial mammals – weighing three times more than that of a horse – which helps it to maintain a greater metabolism during the summer when it lives in river estuaries. It is the marine cetacean that most frequently develops hyperplastic and neoplastic lesions of the thyroid.
The fins retain the bony vestiges of the beluga's mammalian ancestors, and are firmly bound together by connective tissue. The fins are small in relation to the size of the body, rounded and oar-shaped, and slightly curled at the tips. These versatile extremities are mainly used as a rudder to control direction, to work in synchrony with the tailfin and for agile movement in shallow waters up to 3 m (9.8 ft) deep. The fins also contain a mechanism for regulating body temperature, as the arteries feeding the fin's muscles are surrounded by veins that dilate or contract to gain or lose heat. The tailfin is flat with two oar-like lobes, it does not have any bones, and is made up of hard, dense, fibrous connective tissue. The tailfin has a distinctive curvature along the lower edge. The longitudinal muscles of the back provide the ascending and descending movement of the tailfin, which has a similar thermoregulation mechanism to the pectoral fins.
Belugas have a dorsal ridge, rather than a dorsal fin. The absence of the dorsal fin is reflected in the genus name of the species—apterus the Greek word for "wingless". The evolutionary preference for a dorsal ridge rather than a fin is believed to be an adaptation to under-ice conditions, or possibly as a way of preserving heat. The crest is hard and, along with the head, can be used to open holes in ice up to 8 cm (3.1 in) thick.
The beluga has a very specialized sense of hearing and its auditory cortex is highly developed. It can hear sounds within the range of 1.2 to 120 kHz, with the greatest sensitivity between 10 and 75 kHz, where the average hearing range for humans is 0.02 to 20 kHz. The majority of sounds are most probably received by the lower jaw and transmitted towards the middle ear. In the toothed whales, the lower jawbone is broad with a cavity at its base, which projects towards the place where it joins the cranium. A fatty deposit inside this small cavity connects to the middle ear. Toothed whales also possess a small external auditory hole a few centimetres behind their eyes; each hole communicates with an external auditory conduit and an eardrum. It is not known if these organs are functional or simply vestigial.
Belugas are able to see within and outside of water, but their vision is relatively poor when compared to dolphins. Their eyes are especially adapted to seeing under water, although when they come into contact with the air, the crystalline lens and the cornea adjust to overcome the associated myopia (the range of vision under water is short). A beluga's retina has cones and rods, which also suggests they can see in low light. The presence of cone cells indicates they can see colours, although this suggestion has not been confirmed. Glands located in the medial corner of their eyes secrete an oily, gelatinous substance that lubricates the eye and helps flush out foreign bodies. This substance forms a film that protects the cornea and the conjunctiva from pathogenic organisms.
Studies on captive animals show they seek frequent physical contact with other belugas. Areas in the mouth have been found that could act as chemoreceptors for different tastes, and they can detect the presence of blood in water, which causes them to react immediately by displaying typical alarm behaviour. Like the other toothed whales, their brains lack olfactory bulbs and olfactory nerves, which suggests they do not have a sense of smell.
Social structure and play
These cetaceans are highly sociable and they regularly form small groups, or pods, that may contain between two and 25 individuals, with an average of 10 members. Pods tend to be unstable, meaning individuals tend to move from pod to pod. Radio tracking has even shown belugas can start out in one pod and within a few days be hundreds of miles away from that pod. These pods contain animals of both sexes, and are led by a dominant male. Many hundreds and even thousands of individuals can be present when the pods join together in river estuaries during the summer. This can represent a significant proportion of the total population and is when they are most vulnerable to being hunted.
They are cooperative animals and frequently hunt in coordinated groups. The animals in a pod are very sociable and often chase each other as if they are playing or fighting, and they often rub against each other.
In captivity, they can be seen to be constantly playing, vocalizing, and swimming around each other. They show a great deal of curiosity towards humans and frequently approach the windows in the tanks to observe them. Belugas may also playfully spit at humans or other whales. It is not unusual for an aquarium handler to be drenched by one of his charges. Some researchers believe spitting originated with blowing sand away from crustaceans at the sea bottom.
Belugas also show a great degree of curiosity towards humans in the wild, and frequently swim alongside boats. They also play with objects they find in the water; in the wild, they do this with wood, plants, dead fish, and bubbles they have created. During the breeding season, adults have been observed carrying objects such as plants, nets, and even the skeleton of a dead reindeer on their heads and backs. Captive females have also been observed displaying this behaviour, carrying items such as floats and buoys, after they have lost a calf; experts consider this interaction with the objects could be acting as a substitute behaviour.
Swimming and diving
Belugas are slower swimmers than the other toothed whales, such as the killer whale and the common bottlenose dolphin, because they are less hydrodynamic and have limited movement of their tailfins, which produce the greatest thrust. They frequently swim at between 3 and 9 km/h (1.9 and 5.6 mph), although they are able to maintain a speed of 22 km/h for up to 15 min. Unlike most cetaceans, they are capable of swimming backwards. Belugas swim on the surface between 5% and 10% of the time, while for the rest of the time they swim at a depth sufficient to cover their bodies. They do not jump out of the water like dolphins or killer whales.
These animals usually only dive to depths to 20 m (66 ft), although they are capable of diving to greater depths. Individual captive animals have been recorded at depths between 400 and 647 m below sea level, while animals in the wild have been recorded as diving to a depth of more than 700 m, with the greatest recorded depth being 872 m. A dive normally lasts 3 to 5 min, but can last up to 18 min. In the shallower water of the estuaries, a diving session may last around two minutes; the sequence consists of five or six rapid, shallow dives followed by a deeper dive lasting up to one minute. The average number of dives per day varies between 31 and 51.
All cetaceans, including belugas, have physiological adaptations designed to conserve oxygen while they are under water. During a dive, these animals will reduce their heart rate from 100 beats a minute to between 12 and 20. Blood flow is diverted away from certain tissues and organs and towards the brain, heart and lungs, which require a constant oxygen supply. The amount of oxygen dissolved in the blood is 5.5%, which is greater than that found in land-based mammals and is similar to that of Weddell seals (a diving marine mammal). One study found a female beluga had 16.5 l of oxygen dissolved in her blood. Lastly, the beluga's muscles contain high levels of the protein myoglobin, which stores oxygen in muscle. Myoglobin concentrations are several times greater than for terrestrial mammals, which help prevent oxygen deficiency during dives.
Belugas play an important role in the structure and function of marine resources in the Arctic Ocean, as they are the most abundant toothed whales in the region. They are opportunistic feeders; their feeding habits depend on their locations and the season. For example, when they are in the Beaufort Sea, they mainly eat Arctic cod (Boreogadus saida) and the stomachs of belugas caught near Greenland were found to contain rose fish (Sebastes marinus), Greenland halibut (Reinhardtius hippoglossoides), and northern shrimp (Pandalus borealis), while in Alaska their staple diet is Pacific salmon (Oncorhynchus kisutch). In general, the diets of these cetaceans consist mainly of fish; apart from those previously mentioned, other fish they feed on include capelin (Mallotus villosus), smelt, sole, flounder, herring, sculpin, and other types of salmon. They also consume a great quantity of invertebrates, apart from shrimp, such as squid, crabs, clams, octopus, sea snails, bristle worms, and other deep-sea species. Animals in captivity eat 2.5% to 3.0% of their body weight per day, which equates to 18.2 to 27.2 kg.
Foraging on the seabed typically takes place at depths between 20 and 40 m, although they can dive to depths of 700 m in search of food. Their flexible necks provide a wide range of movement while they are searching for food on the ocean floor. Some animals have been observed to suck up water and then forcefully expel it to uncover their prey hidden in the silt on the seabed. As their teeth are neither large nor sharp, belugas must use suction to bring their prey into their mouths; it also means their prey has to be consumed whole, which in turn means it cannot be too large or the belugas run the risk of it getting stuck in their throats. They also join together into coordinated groups of five or more to feed on shoals of fish by steering the fish into shallow water, where the belugas then attack them. For example, in the estuary of the Amur River, where they mainly feed on salmon, groups of six or eight individuals join together to surround a shoal of fish and prevent their escape. Individuals then take turns feeding on the fish.
Estimations of the age of sexual maturity for beluga whales vary considerably; the majority of authors estimate males reach sexual maturity when they are between four and seven years old, and females reach maturity when they are between four and nine years old. The average age at which females first give birth is 8.5 years and fertility begins to decrease when they are 25, eventually undergoing menopause, and ceasing reproductive potential with no births recorded for females older than 41.
Female belugas typically give birth to one calf every three years. Most mating occurs usually February through May, but some mating occurs at other times of year. The beluga may have delayed implantation. Gestation has been estimated to last 12.0 to 14.5 months, but information derived from captive females suggests a longer gestation period up to 475 days (15.8 months).
Calves are born over a protracted period that varies by location. In the Canadian Arctic, calves are born between March and September, while in Hudson Bay, the peak calving period is in late June, and in Cumberland Sound, most calves are born from late July to early August. Births usually take place in bays or estuaries where the water is warm with a temperature of 10 to 15 °C. Newborns are about 1.5 m (4.9 ft) long, weigh about 80 kg (180 lb), and are grey in colour. They are able to swim alongside their mothers immediately after birth. The newborn calves nurse under water and initiate lactation a few hours after birth; thereafter, they feed at intervals around an hour. Studies of captive females have indicated their milk composition varies between individuals and with the stage of lactation; it has an average content of 28% fat, 11% protein, 60.3% water, and less than 1% residual solids. The milk contains about 92 cal per ounce.
The calves remain dependent on their mothers for nursing for the first year, when their teeth appear. After this, they start to supplement their diets with shrimp and small fish. The majority of the calves continue nursing until they are 20 months old, although occasionally lactation can continue for more than two years, and lactational anoestrus may not occur. Alloparenting (care by females different from the mother) has been observed in captive belugas, including spontaneous and long-term milk production. This suggests this behaviour, which is also seen in other mammals, may be present in belugas in the wild.
Communication and echolocation
Belugas use sounds and echolocation for movement, communication, to find breathing holes in the ice, and to hunt in dark or turbid waters. They produce a rapid sequence of clicks that pass through the melon, which acts as an acoustic lens to focus the sounds into a beam that is projected forward through the surrounding water. These sounds spread through the water at a speed of nearly 1.6 km per second, some four times faster than the speed of sound in air. The sound waves reflect from objects and return as echoes that are heard and interpreted by the animal. This enables them to determine the distance, speed, size, shape and the object's internal structure within the beam of sound. They use this ability when moving around thick Arctic ice sheets, to find areas of unfrozen water for breathing, or air pockets trapped under the ice.
Some evidence indicates that belugas are highly sensitive to noise produced by humans. In one study, the maximum frequencies produced by an individual located in San Diego Bay, California, were between 40 and 60 kHz. The same individual produced sounds with a maximum frequency of 100 to 120 kHz when transferred to Kaneohe Bay in Hawaii. The difference in frequencies is thought to be a response to the difference in environmental noise in the two areas.
These animals communicate using sounds of high frequency; their calls can sound like bird songs, so belugas were nicknamed "canaries of the sea". Like the other toothed whales, belugas do not possess vocal cords and the sounds are probably produced by the movement of air between the nasal sacks, which are located near to the blowhole.
Belugas are among the most vocal cetaceans. They use their vocalisations for echolocation, during mating and for communication. They possess a large repertoire, emitting up to 11 different sounds, such as cackles, whistles, trills and squawks. They make sounds by grinding their teeth or splashing, but they rarely use body language to make visual displays with their pectoral fins or tailfins, nor do they perform somersaults or jumps in the way other species do, such as dolphins.
The beluga inhabits a discontinuous circumpolar distribution in Arctic and sub-Arctic waters. During the summer, they can mainly be found in the deep waters ranging from 76°N to 80°N, particularly along the coasts of Alaska, northern Canada, western Greenland, and northern Russia. The southernmost extent of their range includes isolated populations in the St. Lawrence River in the Atlantic, and the Amur River delta, the Shantar Islands, and the waters surrounding Sakhalin Island in the Sea of Okhotsk.
Belugas have a seasonal migratory pattern. Migration patterns are passed from parents to offspring. Some travel as far as 6,000 kilometers per year. When the summer sites become blocked with ice during the autumn, they move to spend the winter in the open sea alongside the pack ice or in areas covered with ice, surviving by using polynyas to surface and breathe. In summer after the sheet ice has melted, they move to coastal areas with shallower water (1–3 m deep), although sometimes they migrate towards deeper waters (>800 m). In the summer, they occupy estuaries and the waters of the continental shelf, and on occasion, they even swim up the rivers. A number of incidents have been reported where groups or individuals have been found hundreds or even thousands of kilometres from the ocean. One such example comes from 9 June 2006, when a young beluga carcass was found in the Tanana River near Fairbanks in central Alaska, nearly 1,700 kilometers (1,100 mi) from the nearest ocean habitat. Belugas sometimes follow migrating fish, leading Alaska state biologist Tom Seaton to speculate it had followed migrating salmon up the river at some point in the previous autumn. The rivers they most often travel up include: the Northern Dvina, the Mezen, the Pechora, the Ob and the Yenisei in Asia; the Yukon and the Kuskokwim in Alaska, and the Saint Lawrence in Canada. Spending time in a river has been shown to stimulate an animal's metabolism and facilitates the seasonal renewal of the epidermal layer. In addition, the rivers represent a safe haven for newborn calves where they will not be preyed upon by killer whales. Calves often return to the same estuary as their mother in the summer, meeting her sometimes even after becoming fully mature.
The migration season is relatively predictable, as it is basically determined by the amount of daylight and not by other variable physical or biological factors, such as the condition of the sea ice. Vagrants may travel further south to areas such as Irish and Scottish waters, islands of Orkney and Hebrides, and to Japanese waters. There had been several vagrant individuals demonstrated seasonal residencies at Volcano Bay, and a unique whale were used to return annually to areas adjacent to Shibetsu in Nemuro Strait in the 2000s. On rarer occasions, individuals of vagrancy can reach the Korean Peninsula. A few other individuals have been confirmed to return to the coasts of Hokkaido, and one particular individual became a resident in brackish waters of Lake Notoro since in 2014.
Some populations are not migratory and certain resident groups will stay in well-defined areas, for example in Cook Inlet, the estuary of the Saint Lawrence River and Cumberland Sound. The population in Cook Inlet stays in the waters furthest inside the inlet during the summer and until the end of autumn, then during the winter, they disperse to the deeper water in the centre of the inlet, but without completely leaving it.
In April, the animals that spend the winter in the centre and southwest of the Bering Sea move to the north coast of Alaska and the east coast of Russia. The populations living in the Ungava Bay and the eastern and western sides of Hudson Bay overwinter together beneath the sea ice in Hudson Strait. Whales in James Bay spend winter months within the basin could be a distinct group from these in Hudson Bay. The populations of the White Sea, the Kara Sea and the Laptev Sea overwinter in the Barents Sea. In the spring, the groups separate and migrate to their respective summer sites.
Belugas exploit a varied range of habitats; they are most commonly seen in shallow waters close to the coast, but they have also been reported to live for extended periods in deeper water, where they feed and give birth to their young.
In coastal areas, they can be found in coves, fjords, canals, bays, and shallow waters in the Arctic Ocean that are continuously lit by sunlight. They are also often seen during the summer in river estuaries, where they feed, socialize, and give birth to young. These waters usually have a temperature between 8 and 10 °C. The mudflats of Cook Inlet in Alaska are a popular location for these animals to spend the first few months of summer. In the eastern Beaufort Sea, female belugas with their young and immature males prefer the open waters close to land; the adult males live in waters covered by ice near to the Canadian Arctic Archipelago, while the younger males and females with slightly older young can be found nearer to the ice shelf. Generally, the use of different habitats in summer reflects differences in feeding habits, risk from predators, and reproductive factors for each of the subpopulations.
The estimate of population sizes is complicated because the boundaries for some of these groups overlap geographically or seasonally. The IUCN estimated the world beluga population in 2008 to be well in excess of 150,000.
The native populations of the Arctic in Alaska, Canada, Greenland, and Russia hunt belugas, consuming part, and selling part. Belugas have been easy prey for hunters due to their predictable migration patterns and the high population density in estuaries and surrounding coastal areas during the summer.
The number of animals killed is about 1,000 per year, with 300 to 400 killed in Alaska, another 300 to 400 in Canada, and fewer in Greenland and Russia (see table below. and its sources). Questions remain whether the number of whales killed is sustainable.
Hunters in Hudson's Bay rarely eat the meat. They give a little to dogs, and leave the rest for wild animals. Other areas may dry the meat for later consumption by humans. In Greenland the skin (Muktuk) is sold commercially to fish factories, and in Canada to other communities. An average of one or two vertebrae and one or two teeth per beluga are carved and sold. One estimate of the annual gross value received from Beluga hunts in Hudson Bay in 2013 was CA$600,000 for 190 belugas, or CA$3,000 per beluga. However the net income, after subtracting costs in time and equipment, was a loss of CA$60 per person. Hunts receive subsidies, but they continue as a tradition, rather than for the money, and the economic analysis noted that whale watching may be an alternate revenue source. Of the gross income, CA$550,000 was for skin and meat, to replace beef, pork and chickens which would otherwise be bought, CA$50,000 was received for carved vertebrae and teeth.
Russia now harvests 5 to 30 per year for meat and captures an additional 20 to 30 per year for live export to Chinese aquaria. However in 2018, 100 were illegally captured for live export.
Previous levels of commercial whaling have put the species in danger of extinction in areas such as Cook Inlet, Ungava Bay, the St. Lawrence River and western Greenland. Continued hunting by the native peoples may mean some populations will continue to decline. Northern Canadian sites are the focus of discussions between local communities and the Canadian government, with the objective of permitting sustainable hunting that does not put the species at risk of extinction.
Commercial whaling by European, American and Russian whalers during the 18th and 19th centuries decreased beluga populations in the Arctic. The animals were hunted for their meat and blubber, while the Europeans used the oil from the melon as a lubricant for clocks, machinery, and lighting in lighthouses. Mineral oil replaced whale oil in the 1860s, but the hunting of these animals continued unabated. In 1863, the cured skin could be used to make horse harnesses, machine belts for saw mills, and shoelaces. These manufactured items ensured the hunting of belugas continued for the rest of the 19th century and the beginning of the 20th century. The cured skin is the only cetacean skin that is sufficiently thick to be used as leather.
Russia had large hunts, peaking in the 1930s at 4,000 per year and 1960s at 7,000 per year, for a total of 86,000 from 1915-2014. Canada hunted a total of 54,000 from 1731-1970. Between 1868 and 1911, Scottish and American whalers killed more than 20,000 belugas in Lancaster Sound and Davis Strait.
During the 1920s, fishermen in the Saint Lawrence River estuary considered belugas to be a threat to the fishing industry, as they eat large quantities of cod, salmon, tuna, and other fish caught by the local fishermen. The presence of belugas in the estuary was, therefore, considered to be undesirable; in 1928, the Government of Quebec offered a reward of 15 dollars for each dead beluga. The Quebec Department of Fisheries launched a study into the influence of these cetaceans on local fish populations in 1938. The unrestricted killing of belugas continued into the 1950s, when the supposed voracity of the belugas was found to be overestimated and did not adversely affect fish populations. L'Isle-aux-Coudres is the setting for the classic 1963 National Film Board of Canada documentary Pour la suite du monde, which depicts a one-off resurrection of the beluga hunt; one animal is caught live, and transported by truck to an aquarium in the big city. The method of capture is akin to dolphin drive hunting.
Beluga catches by location
|Beaufort Sea, Mackenzie, Paulatuk, Ulukhaktok, Canada||Nunavut, Canada||Nunavik, Quebec, Canada||Western Arctic Meat, Russia||Eastern Arctic Meat, Russia||Sea of Okhotsk, Meat, Russia||Total Live Export from Russia||Year||Canada Total||Greenland||USSR+ Russia Total||USA (Alaska)||World Total, Incomplete||Lost at Sea as % of Caught|
|1960–1969 1970–99 2000–2012 2013–15 2014||Arviat||1996–2002
|1954–99||1954–1985 cites Russian papers||NMFS cites Russian paper||Western Okhotsk||Sources||Total of columns at left, incomplete||1954–2016||Total of columns at left, incomplete||1954–84 1987–90 Cook Inlet 1990–2011 2012–2015 +Cook Inlet||Total of other columns||Greenland source 1954–1999, Beaufort source 2000–2012|
During the winter, belugas commonly become trapped in the ice without being able to escape to open water, which may be several kilometres away. Polar bears take particular advantage of these situations and are able to locate the belugas using their sense of smell. The bears swipe at the belugas and drag them onto the ice to eat them. They are able to capture large individuals in this way; in one documented incident, a bear weighing between 150 and 180 kg was able to capture an animal that weighed 935 kg.
Killer whales are able to capture both young and adult belugas. They live in all the seas of the world and share the same habitat as belugas in the sub-Arctic region. Attacks on belugas by killer whales have been reported in the waters of Greenland, Russia, Canada, and Alaska. A number of killings have been recorded in Cook Inlet, and experts are concerned the predation by killer whales will impede the recovery of this subpopulation, which has already been badly depleted by hunting. The killer whales arrive at the beginning of August, but the belugas are occasionally able to hear their presence and evade them. The groups near to or under the sea ice have a degree of protection, as the killer whale's large dorsal fin, up to 2 m in length, impedes their movement under the ice and does not allow them to get sufficiently close to the breathing holes in the ice.
The beluga is considered an excellent sentinel species (indicator of environment health and changes), because it is long-lived, at the top of the food web, bears large amounts of fat and blubber, relatively well-studied for a cetacean, and still somewhat common.
Human pollution can be a threat to belugas' health when they congregate in river estuaries. Chemical substances such as DDT and heavy metals such as lead, mercury and cadmium have been found in individuals of the Saint Lawrence River population. Local beluga carcasses contain so many contaminants, they are treated as toxic waste. Levels of polychlorinated biphenyls between 240 and 800 ppm have been found in belugas' brains, liver and muscles, with the highest levels found in males. These levels are significantly greater than those found in Arctic populations. These substances have a proven adverse effect on these cetaceans, as they cause cancers, reproductive diseases, and the deterioration of the immune system, making individuals more susceptible to pneumonias, ulcers, cysts, tumours, and bacterial infections. Although the populations that inhabit the river estuaries run the greatest risk of contamination, high levels of zinc, cadmium, mercury, and selenium have also been found in the muscles, livers, and kidneys of animals that live in the open sea.
From a sample of 129 beluga adults from the Saint Lawrence River examined between 1983 and 1999, a total of 27% had suffered cancer. This is a higher percentage than that documented for other populations of this species and is much higher than for other cetaceans and for the majority of terrestrial mammals; in fact, the rate is only comparable to the levels found in humans and some domesticated animals. For example, the rate of intestinal cancer in the sample is much higher than for humans. This condition is thought to be directly related to environmental contamination, in this case by polycyclic aromatic hydrocarbons, and coincides with the high incidence of this disease in humans residing in the area. The prevalence of tumours suggests the contaminants identified in the animals that inhabit the estuary are having a direct carcinogenic effect or they are at least causing an immunological deterioration that is reducing the inhabitants' resistance to the disease.
Indirect human disturbance may also be a threat. While some populations tolerate small boats, most actively try to avoid ships. Whale-watching has become a booming activity in the St. Lawrence and Churchill River areas, and acoustic contamination from this activity appears to have an effect on belugas. For example, a correlation appears to exist between the passage of belugas across the mouth of the Saguenay River, which has decreased by 60%, and the increase in the use of recreational motorboats in the area. A dramatic decrease has also been recorded in the number of calls between animals (decreasing from 3.4 to 10.5 calls/min to 0 or <1) after exposure to the noise produced by ships, the effect being most persistent and pronounced with larger ships such as ferries than with smaller boats. Belugas can detect the presence of large ships (for example icebreakers) up to 50 km away, and they move rapidly in the opposite direction or perpendicular to the ship following the edge of the sea ice for distances of up to 80 km to avoid them. The presence of shipping produces avoidance behaviour, causing deeper dives for feeding, the break-up of groups, and asynchrony in dives.
As with any animal population, a number of pathogens cause death and disease in belugas, including viruses, bacteria, protozoans, and fungi, which mainly cause skin, intestinal, and respiratory infections.
Papillomaviruses have been found in the stomachs of belugas in the Saint Lawrence River. Animals in this location have also been recorded as suffering infections caused by herpesviruses and in certain cases to be suffering from encephalitis caused by the protozoan Sarcocystis. Cases have been recorded of ciliate protozoa colonising the spiracle of certain individuals, but they are not thought to be pathogens or are not very harmful.:26, 303, 359
The bacterium Erysipelothrix rhusiopathiae, which probably comes from eating infected fish, poses a threat to belugas kept in captivity, causing anorexia and dermal plaques and lesions that can lead to sepsis.:26, 303, 359 This condition can cause death if it is not diagnosed and treated in time with antibiotics such as ciprofloxacin.:316–7
A study of infections caused by parasitic worms in a number of individuals of both sexes found the presence of larvae from a species from the genus Contracaecum in their stomachs and intestines, Anisakis simplex in their stomachs, Pharurus pallasii in their ear canals, Hadwenius seymouri in their intestines, and Leucasiella arctica in their rectums.
Relationship with humans
Belugas were among the first whale species to be kept in captivity. The first beluga was shown at Barnum's Museum in New York City in 1861. For most of the 20th century, Canada was the predominant source for belugas destined for exhibition. Until the early 1960s, they were taken from the St. Lawrence River estuary and from 1967 from the Churchill River estuary. This continued until 1992, when the practice was banned. Since Canada ceased to be the supplier of these animals, Russia has become the largest provider. Individuals are caught in the Amur River delta and the far eastern seas of the country, and then are either transported domestically to aquaria in Moscow, St. Petersburg, and Sochi, or exported to foreign nations, including China and formerly Canada. Canada's Senate has passed a bill outlawing new captivity for cetaceans, and the Commons are expected to pass it by May 2019.
To provide some enrichment while in captivity, aquaria train belugas to perform behaviours for the public and for medical exams, such as blood draws and ultrasound, provide toys, and allow the public to play recorded or live music.
Between 1960 and 1992, the United States Navy carried out a program that included the study of marine mammals' abilities with echolocation, with the objective of improving the detection of underwater objects. The program started with dolphins, but a large number of belugas were also used from 1975 on. The program included training these mammals to carry equipment and material to divers working under water, the location of lost objects, surveillance of ships and submarines, and underwater monitoring using cameras held in their mouths. A similar program was implemented by the Soviet Navy during the Cold War, in which belugas were also trained for antimining operations in Arctic waters. It is possible this program continues within the Russian Navy, as on 24 April 2019 a tame Beluga whale wearing a Russian equipment harness was found by fishermen near the Norwegian island of Ingøya.
Belugas released from captivity have difficulties adapting to life in the wild, but if not fed by humans they may have a chance to join a group of wild belugas and learn to feed themselves, according to Audun Rikardsen of the University of Tromsø.
In 2018-2019, a sanctuary in Iceland is being established for belugas retired from a marine park in China. A British company, Merlin Entertainments, bought the park in 2012, as part of an Australian chain, and it is one of their largest aquaria. Merlin has a policy against captive cetaceans, so they are sponsoring a 32,000 square meter sea pen as a sanctuary. The 12-year old belugas, caught in Russia and raised in captivity, do not know how to live in the wild. The cost is variously listed as ISK 3,000,000 (USD 24,000) or USD 27,000,000. Merlin was owned until 2015 by Blackstone Group, which also owned SeaWorld until selling its last stake in 2017 to a Chinese company which will use SeaWorld's expertise to expand in China; SeaWorld still keeps belugas in captivity.
Belugas are the only whale species kept in aquaria and marine parks. They are displayed across North America, Europe, and Asia. As of 2006, 58 belugas were in captivity in Canada and the United States, and 42 deaths in US captivity had been reported up to that time. A single specimen cost up to US$100,000. As of January 2018, according to the nonprofit Ceta Base, which tracks belugas and dolphins under human care, there were 81 captive belugas in Canada and the United States, and unknown numbers in the rest of the world. The price had dropped to USD 70,000 (RUB 4.2 million). The beluga's popularity with visitors reflects its attractive colour and its range of facial expressions. The latter is possible because while most cetacean "smiles" are fixed, the extra movement afforded by the beluga's unfused cervical vertebrae allows a greater range of apparent expression.
Most belugas found in aquaria are caught in the wild, as captive-breeding programs have not had much success so far. For example, despite best efforts, as of 2010, only two male whales had been successfully used as stud animals in the Association of Zoos and Aquariums beluga population, Nanuq at SeaWorld San Diego and Naluark at the Shedd Aquarium in Chicago, USA. Nanuq has fathered 10 calves, five of which survived birth. Naluark at Shedd Aquarium has fathered four living offspring. Naluark was relocated to the Mystic Aquarium in the hope that he would breed with two of their females, but he did not, and in 2016 he was moved to SeaWorld Orlando. The first beluga calf born in captivity in Europe was born in L'Oceanogràfic marine park in Valencia, Spain, in November 2006. However, the calf died 25 days later after suffering metabolic complications, infections, and not being able to feed properly. A second calf was born in November, 16th 2016, and was successfully maintained by artificial feeding based on enriched milk.
In 2009 during a free-diving competition in a tank of icy water in Harbin, China, a captive beluga brought a cramp-paralyzed diver from the bottom of the pool up to the surface by holding her foot in its mouth, saving the diver's life.
Whale watching has become an important activity in the recovery of the economies of towns in Hudson Bay near to the Saint Lawrence and Churchill Rivers. The best time to see belugas is during the summer, when they meet in large numbers in the estuaries of the rivers and in their summer habitats. The animals are easily seen due to their high numbers and their curiosity regarding the presence of humans.
However, the boats' presence poses a threat to the animals, as it distracts them from important activities such as feeding, social interaction and reproduction. In addition, the noise produced by the motors has an adverse effect on their auditory function and reduces their ability to detect their prey, communicate, and navigate. To protect these marine animals during whale-watching activities, the US National Oceanic and Atmospheric Administration has published a "Guide for observing marine life". The guide recommends boats carrying the whale watchers keep their distance from the cetaceans and it expressly prohibits chasing, harassing, obstructing, touching, or feeding them.
On 25 September 2018, a Beluga was sighted in the Thames Estuary and near towns along the Kent side of the Thames, being nicknamed 'Benny' by newspapers. The whale, who was noticed by conservationists to be travelling alone, appeared to be separated from the rest of its group, and is thought to be a lost individual. Subsequent sightings were reported on the following day.
Male belugas in captivity can mimic the pattern of human speech, several octaves lower than typical whale calls. It is not the first time a beluga has been known to sound human, and they often shout like children, in the wild. One captive beluga, after overhearing divers using an underwater communication system, caused one of the divers to surface by imitating their order to get out of the water. Subsequent recordings confirmed that the beluga had become skilled at imitating the patterns and frequency of human speech. After several years, this beluga ceased making these sounds.
Prior to 2008, the beluga was listed as "vulnerable" by the International Union for Conservation of Nature (IUCN), a higher level of concern. The IUCN cited the stability of the largest subpopulations and improved census methods that indicate a larger population than previously estimated. In 2008, the beluga was reclassified as "near threatened" by the IUCN due to uncertainty about threats to their numbers and the number of belugas over parts of its range (especially the Russian Arctic), and the expectation that if current conservation efforts cease, especially hunting management, the beluga population is likely to qualify for "threatened" status within five years. In June 2017, its status was reassessed to "least concern".
Subpopulations are subject to differing levels of threat and warrant individual assessment. The nonmigratory Cook Inlet subpopulation is listed as "Critically Endangered" by the IUCN as of 2006 and is listed as Endangered under the Endangered Species Act as of October 2008. This was due to overharvesting of belugas prior to 1998. The population has failed to recover, though the reported harvest has been small. The most recently published estimate as of May 2008 was 302 (CV=0.16) in 2006. In addition, the National Marine Fisheries Service indicated the 2007 aerial survey's point estimate was 375.
Despite beluga whales not being threatened overall subpopulations are being listed as critically endangered and are facing increased mortality from human actions. The IUCN cites land development, oil and gas drilling, shipping, dams and water management, pollution, harvesting for consumption, and climate change as threats to the prolonged survival of beluga whale subpopulations.
Beluga whale populations are currently being harvested at levels which are not sustainable and it is difficult for those harvesting beluga whales to know which subpopulation they are from. Because there is little protection of subpopulations, harvest will need to be managed to ensure subpopulations will survive long into the future to discover the importance of their migratory patterns and habitat use.
Beluga whales, like most other arctic species, are being faced with alteration of their habitat due to climate change and melting arctic ice. It is still not well understood why beluga whales use annual ice, but it is hypothesised to be a place of shelter from killer whale predation or a place they go to feed on schooling fish. Changes in arctic ice density have caused massive death tolls to beluga whales through ice entrapment. It is apparent that they can be faced by an increased risk of entrapment from leads and cracks freezing. Abrupt changes in weather can cause these leads and cracks to freeze ultimately causing the whales to die of suffocation.
The US Congress passed the Marine Mammal Protection Act of 1972 outlawing the persecution and hunting of all marine mammals within US coastal waters. The act has been amended a number of times to permit subsistence hunting by native peoples, temporary capture of restricted numbers for research, education and public display, and to decriminalise the accidental capture of individuals during fishing operations. The act also states that all whales in US territorial waters are under the jurisdiction of the National Marine Fisheries Service, a division of NOAA.
To prevent hunting, belugas are protected under the 1986 International Moratorium on Commercial Whaling; however, hunting of small numbers of belugas is still allowed. Since it is very difficult to know the exact population of belugas because their habitats include inland waters away from the ocean, they easily come in contact with oil and gas development centres. To prevent whales from coming in contact with industrial waste, the Alaskan and Canadian governments are relocating sites where whales and waste come in contact.
The beluga whale is listed on appendix II of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). It is listed on Appendix II as it has an unfavourable conservation status or would benefit significantly from international co-operation organised by tailored agreements. All toothed whales are protected under the CITES that was signed in 1973 to regulate the commercial exploitation of certain species.
The isolated beluga population in the Saint Lawrence River has been legally protected since 1983. In 1988 Canadian Department of Fisheries and Oceans and Environment Canada, a governmental agency that supervises national parks, implemented the Saint Lawrence Action Plan with the aim of reducing industrial contamination by 90% by 1993; as of 1992, the emissions had been reduced by 59%. The population of the St. Lawrence belugas decreased from 10,000 in 1885 to around 1,000 in the 1980 and around 900 in 2012.
White Whale Records was an American record company that operated between 1965 and 1971 in Los Angeles, California, it was the record company of The Turtles. The company's logo was the silhouette of a beluga with the words "White Whale" above it.
The children's singer Raffi released an album called Baby Beluga in 1980. The album starts with the sound of whales communicating, and includes songs representing the ocean and whales playing. The song "Baby Beluga" was composed after Raffi saw a recently born beluga calf in Vancouver Aquarium.
Yamaha's Beluga motorcycle (Riva 80/CV80) which had an 80-cc engine was produced from 1981 until 1987 and sold throughout the world, particularly in Canada, the U.S., the Netherlands, Belgium, Sweden, and Japan.
The fuselage design of the Airbus Beluga, one of the world's biggest cargo planes, is very similar to that of a beluga; it was originally called the Super Transporter, but the nickname Beluga became more popular and was then officially adopted.
The German company SkySails GmbH & Co. KG, a subsidiary of the Beluga Shipping group based in Hamburg, tested a new propulsion system for ships that involved a large wing similar to that used in paragliding and which has demonstrated a reduction in fuel use between 10% and 35%. The programme to prove the efficiency of the system was called Project Beluga, as it involved the ship MS Beluga Skysails. The company's insignia, a beluga's tailfin, was printed on the giant wing, which had a surface area of 160 m2.
- Mead, J.G.; Brownell, R. L. Jr. (2005). "Order Cetacea". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 735. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Lowry, L (2017). "Delphinapterus leucas". IUCN Red List of Threatened Species. Version 2017-3. International Union for Conservation of Nature. Retrieved 21 December 2017.
- Arnold, P. "Irrawaddy Dolphin Orcaella brevirostris". Encyclopedia of Marine Mammals. p. 652.
- Grétarsdóttir, S.; Árnason, Ú. (1992). "Evolution of the common cetacean highly repetitive DNA component and the systematic position of Orcaella brevirostris". Journal of Molecular Evolution. 34 (3): 201–208. Bibcode:1992JMolE..34..201G. doi:10.1007/BF00162969. PMID 1588595.
- O'Corry-Crowe, G. "Beluga Whale Delphinapterus leucas". Encyclopedia of Marine Mammals. pp. 94–99.
- Heide-Jørgensen, Mads P.; Reeves, Randall R. (1993). "Description of an Anomalous Monodontid Skull from West Greenland: A Possible Hybrid?". Marine Mammal Science. 9 (3): 258–68. doi:10.1111/j.1748-7692.1993.tb00454.x.
- Leatherwood, Stephen; Reeves, Randall R. (1983). The Sierra Club Handbook of Whales and Dolphins. Sierra Club Books. p. 320. ISBN 978-0-87156-340-8.
- "Japanese whale whisperer teaches beluga to talk". meeja.com.au. 16 September 2008. Archived from the original on 13 January 2009. Retrieved 16 September 2008.
- Kusma, Stephanie (23 October 2012). "Ein "sprechender" Beluga-Wal" [A "talking" Beluga whale] (in German). Neue Zürcher Zeitung. Archived from the original on 24 October 2012. Retrieved 25 October 2012.
- Arnason, U.; Gullberg, A. (1996). "Cytochrome b nucleotide sequences and the identification of five primary lineages of extant cetaceans". Molecular Biology and Evolution. 13 (2): 407–17. doi:10.1093/oxfordjournals.molbev.a025599. PMID 8587505.
- Waddell, Victor G.; Milinkovitch, Michel C.; Bérubé, Martine; Stanhope, Michael J. (2000). "Molecular Phylogenetic Examination of the Delphinoidea Trichotomy: Congruent Evidence from Three Nuclear Loci Indicates That Porpoises (Phocoenidae) Share a More Recent Common Ancestry with White Whales (Monodontidae) Than They Do with True Dolphins (Delphinidae)". Molecular Phylogenetics and Evolution. 15 (2): 314–8. doi:10.1006/mpev.1999.0751. PMID 10837160.
- Jones, Steven J.M.; et al. (2017). "The Genome of the Beluga Whale (Delphinapterus leucas)". Genes. 8 (12): 378. doi:10.3390/genes8120378. PMC 5748696. PMID 29232881. Archived from the original on 12 December 2017.
- Barnes, Lawrence G. Fossil odontocetes (Mammalia: Cetacea) from the Almejas Formation, Isla Cedros, Mexico. University of California, Museum of Paleontology. p. 46. ASIN B0006YRTU4.
- "Denebola brachycephala". ZipcodeZoo.com. Archived from the original on 20 August 2012. Retrieved 1 August 2010.
- Barnes, L. G. (1977). "Outline of Eastern North Pacific Fossil Cetacean Assemblages". Systematic Zoology. 25 (4): 321–343. doi:10.2307/2412508. JSTOR 2412508.
- Perrin & Wursig 2009, p. 214.
- "Charlotte, The Vermont Whale – An Electronic Museum". University of Vermont. Archived from the original on 19 December 2010. Retrieved 2 August 2010.
- Wright, Wesley (2009). "Our State Fossil". University of Vermont. Archived from the original on 10 August 2016. Retrieved 23 August 2016.
- Burns J.J.; Seaman G.A. (1983). "Investigations of belukha whales in coastal waters of western and northern Alaska, 1982–1983: marking and tracking of whales in Bristol Bay". Biology and Ecology. US Dept Commer, NOAA, OCSEAP Final Rep, II. 56: 221–357. OSTI 6361651.
- Goren, Arthur D.; Brodie, Paul F.; Spotte, Stephen; Ray, G. Carleton; Kaufman, H. W.; Gwinnett, A. John; Sciubba, James J.; Buck, John D. (1987). "Growth Layer Groups (GLGs) in the Teeth of an Adult Belukha Whale (Delphinapterus leucas) of Known Age: Evidence for Two Annual Layers". Marine Mammal Science. 3 (1): 14–21. doi:10.1111/j.1748-7692.1987.tb00148.x.
- Stewart, R.E.A.; Campana, S.E.; Jones, C.M.; Stewart, B.E. (2006). "Bomb radiocarbon dating calibrates beluga (Delphinapterus leucas) age estimates". Can. J. Zool. 84 (12): 1840–1852. doi:10.1139/Z06-182.[permanent dead link]
- Convention on Migratory Species – CMS. "Delphinapterus leucas (Pallas, 1776". Archived from the original on 21 August 2011. Retrieved 3 August 2010.
- Shirihai, H. & Jarrett, B. (2006). Whales, Dolphins and Other Marine Mammals of the World. Princeton: Princeton Univ. Press. pp. 97–100. ISBN 978-0-691-12757-6.
- Reeves, R.; Stewart, B.; Clapham, P. & Powell, J. (2003). Guide to Marine Mammals of the World. New York: A.A. Knopf. pp. 318–321. ISBN 978-0-375-41141-0.
- Aquatic Species at Risk – Beluga Whale (St. Lawrence Estuary population) Archived 11 December 2011 at the Wayback Machine. Fisheries and Oceans Canada
- Stewart BE & Stewart RE (1989). "Delphinapterus leucas" (PDF). Mammalian Species. 336 (336): 1–8. doi:10.2307/3504210. JSTOR 3504210. Archived from the original (PDF) on 15 June 2013. Retrieved 22 September 2011.
- Ridgway, S. & Harrison, R. (1981). Handbook of Marine Mammals (6 ed.). San Diego: Academic Press Limited. p. 486. ISBN 978-0-12-588506-5.
- "Beluga Whales – Physical Characteristics". Sea World.org. Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Paine, S. (1995). The World of the Arctic Whales. San Francisco: Sierra Club. ISBN 978-0-87156-957-8.
- Sergeant, D. E. & Brodie, P. F. (1969). "Body Size in White Whales, Delphinapterus leucas". Journal Fisheries Research Board of Canada. 26 (10): 2561–2580. doi:10.1139/f69-251.
- United Nations Environment Programme – Conservation of Migratory Species of Wild Animals (2006). Review of small cetaceans: distribution, behaviour, migration and threats: 177 (Marine mammal action plan/regional seas reports and studies). UNEP/CMS. p. 356. ISBN 978-3-937429-02-1.
- "Delphinapterus leucas – Morphology, Physical Description". Encyclopedia of Life. Retrieved 3 August 2010.[permanent dead link]
- Friedman W. R. (June 2006). "Environmental Adaptations of the Beluga Whale (Delphinapterus leucas)" (PDF). Cognitive Science. 143. Archived (PDF) from the original on 30 October 2012.
- St. Aubin, D. J.; Smith, T. G.; Geraci, J. R. (1990). "Seasonal Epidermal Molt in Beluga Whales, Delphinapterus leucas". Canadian Journal of Zoology. 68 (2): 339–367. doi:10.1139/z90-051.[permanent dead link]
- Haley, Delphine (1986). Marine Mammals of Eastern North Pacific and Arctic Waters (2 ed.). Seattle: Pacific Search Press. ISBN 978-0-931397-14-1.[page needed]
- "Beluga Whales – Communication and Echolocation". Sea World.org. Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Turl, C.W. (1990). T.G. Smith; D.J. St. Aubin; J.R. Geraci. (eds.). "Echolocation abilities of the beluga, Delphinapterus leucas: a review and comparison with the bottlenose dolphin, Tursiops truncatus". Can. Bull. Fish. Aquat. Sci. 224: 119–128.
- Litchfield, Carter; Ackman, R. G.; Sipos, J. C.; Eaton, C. A. (1971). "Isovaleroyl triglycerides from the blubber and melon oils of the beluga whale (Delphinapterus leucas)". Lipids. 6 (9): 674–81. doi:10.1007/BF02531529. PMID 5141491.
- Bonner, W.N. (1980). Whales. Poole, England: Abe Books. pp. 17, 23–24. ISBN 978-0-7137-0887-5.
- Nowak, Ronald M. (1991). Walker's Mammals of the World. 2 (5 ed.). Baltimore: The Johns Hopkins University Press. ISBN 978-0-8018-5789-8.[page needed]
- The National Oceanic and Atmospheric Administration's National Marine Fisheries Service – Alaska Regional Office (2 October 2018). "Beluga Whales". Archived from the original on 4 June 2010. Retrieved 3 August 2010.
- Enchanted Learning. "Blowhole". Archived from the original on 5 March 2010. Retrieved 6 August 2010.
- Aubin DJST (1989). "Thyroid function and epidermal growth in beluga whales, Delphinapterus leucas". Diss Abst Int Pt B Sci and Eng. 50 (1).
- Mikaelian, I.; Labelle, P.; Kopal, M.; De Guise, S.; Martineau, D. (2003). "Adenomatous Hyperplasia of the Thyroid Gland in Beluga Whales (Delphinapterus leucas) from the St. Lawrence Estuary and Hudson Bay, Quebec, Canada". Veterinary Pathology. 40 (6): 698–703. doi:10.1354/vp.40-6-698. PMID 14608025.
- Kasting NW, Adderley SA, Safford T, Hewlett KG (1989). "Thermoregulation in beluga (Delphinapterus leucas) and killer (Orcinus orca) whales". Physiological Zoology. 62 (3): 687–701. doi:10.1086/physzool.62.3.30157921. JSTOR 30157921.
- Belkovitch, V. M.; Shekotov, M. N. (1993). The Belukha Whale: Natural Behavior and Bioacoustics (PDF). Woods Hole, MA: Woods Hole Oceanographic Inst. Archived (PDF) from the original on 7 April 2014.
- Fay, R. R. (1988). Hearing in Vertebrates: a Psychophysics Databook. Acoustical Society of America Journal. 86. Winnetka IL: Hill-Fay Associates. p. 621. Bibcode:1989ASAJ...86.2044F. doi:10.1121/1.398550. ISBN 978-0-9618559-0-1.
- Olson, Harry F (1967). Music, Physics and Engineering. Dover Journals. p. 249. ISBN 978-0-486-21769-7.
- Sea World.org. "Beluga Whales – Senses". Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Herman, Louis (1988). Cetacean Behavior: Mechanisms and Functions. Nueva York: John Wiley and Sons. p. 480. ISBN 978-0-89464-272-2.
- Katona, Steven K.; Valerie Rough & David T. Richardson (1993). Field Guide to Whales, Porpoises, and Seals from Cape Cod to Newfoundland (4 ed.). Smithsonian Institution Press. p. 336. ISBN 978-1-56098-333-0.
- Illustrated Encyclopedia of North American Mammals: A Comprehensive Guide to Mammals of North America. MobileReference. 2009. ISBN 978-1-60501-279-7.
- Slijper, E. J. (1979). Whales (2 ed.). Ithaca, Nueva York: Cornell University Press. p. 511. ISBN 978-0-8014-1161-8.
- Jefferson TA, Leatherwood S, Webber MA (1993). FAO Species identification guide. Marine mammals of the world (PDF). UNEP / FAO, Rome. p. 320. Archived from the original (PDF) on 2 March 2013.
- MacDonald, David (1993). "The Encyclopedia of Mammals". The Encyclopedia of Mammals. Nueva York: Facts on File, Inc. p. 895. ISBN 978-0-87196-871-5.
- Sjare, Becky L. & Thomas G. Smith (1986). "The Relationship Between Behavioral Activity and Underwater Vocalizations of the White Whale, Delphinapterus leucas". Canadian Journal of Zoology. 64 (12): 2824–2831. doi:10.1139/z86-406.[permanent dead link]
- Sea World.org. "Beluga Whales – Behavior". Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Georgia Aquarium. "Beluga Whale". Archived from the original on 2 October 2008. Retrieved 12 October 2008.
- Alaska Geographic Society (1979). Alaska Whales and Whaling. Edmonds, Washington: Alaska Northwest Publishing Co. Archived from the original on 6 November 2012. Retrieved 12 September 2012.
- Smith, T. G. & Sleno, G. A. (1986). "Do White Whales, Delphinapterus leucas, Carry Surrogates in Response to Early Loss of Their Young". Canadian Journal of Zoology. 64 (7): 1581–1582. doi:10.1139/z86-237.[permanent dead link]
- Fish, FE (1998). "Comparative kinematics and hydrodynamics of odontocete cetaceans: Morphological and ecological correlates with swimming performance". The Journal of Experimental Biology. 201 (Pt 20): 2867–2877. PMID 9739069.
- "Georgia Aquarium – Beluga Whale". Archived from the original on 2 October 2008. Retrieved 12 October 2008.
- Sea World.org. "Beluga Whales – Adaptations for an Aquatic Environment". Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Ridgway, Sam H.; Carder, Donald A.; Kamolnick, Tricia; Smith, Robert R.; Schlundt, Carolyn E.; Elsberry, Wesley R. (2001). "Hearing and whistling in the deep sea: Depth influences whistle spectra but does not attenuate hearing by white whales (Delphinapterus leucas) (Odontoceti, Cetacea)". The Journal of Experimental Biology. 204 (Pt 22): 3829–41. PMID 11807101.
- Heide-Jørgensen, M. P.; Richard, P. R.; Rosing-Asvid, A. (March 1998). "Dive Patterns of Belugas (Delphinapterus lencas) in Waters near Eastern Devon Island". Arctic. 51 (1): 17–26. doi:10.14430/arctic1041. JSTOR 40511799.
- "Delphinapterus leucas: Beluga Whale". Marine Bio. Archived from the original on 10 April 2008. Retrieved 26 August 2008.
- Ridgway, Sam H (1972). Mammals of the Sea. Biology and Medicine. Springfield, Illinois: Charles C. Thomas. p. 812. ISBN 978-0-398-02389-8.
- Ridgway, S. H.; Bowers; Miller; Schultz; Jacobs; Dooley; et al. (1984). "Diving and Blood Oxygen in the White Whale". Canadian Journal of Zoology. 6 (11): 2349–2351. doi:10.1139/z84-344.[permanent dead link]
- Noren, S.R.; Williams, T.M. (2000). "Body size and skeletal muscle myoglobin of cetaceans: Adaptations for maximizing dive duration". Comparative Biochemistry and Physiology A. 126 (2): 181–91. doi:10.1016/S1095-6433(00)00182-3. PMID 10936758.
- Dueck L. (2014). "3.2 Habits – 3.2.1 Diving Characteristics and Sightability Estimates of Eastern Arctic Bowhead Whales Based on Satellite-Linked Telemetry". The Fisheries and Oceans Canada. Archived from the original on 30 January 2015. Retrieved 30 January 2015.
- Loseto LL, Stern GA, Connelly TL, Deibel D, Gemmill B, Prokopowicz A, Fortier L, Ferguson SH (2009). "Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web" (PDF). J Exp Mar Biol Ecol. 374 (1): 12–18. doi:10.1016/j.jembe.2009.03.015.[permanent dead link]
- Heide-Jorgensen, M.P.; Teilmann, J (1994). "Growth, reproduction, age structure and feeding habits of white whales (Delphinapterus leucas) in West Greenland waters". Meddr Gronland, Biosci. 39: 195–212.
- Frost, K.J. & Lowry, L.F. (1981). "Trophic importance of some marine gadids in Northern Alaska and their body-otolith size relationships". Fish. Bull. 79: 187–192.
- Lentifer, J (1988). Selected Marine Mammals of Alaska: Species Accounts with Research and Management Recommendations. Marine Mammals Commission. ASIN B000102908.[page needed]
- Perez, Michael A – NOAA (1990). NOAA Technical Memorandum NMFS F/NWC-186. Review of Marine Mammal Population and Prey Information for Bering Sea Ecosystem Studies. NOAA.
- Sea World.org. "Beluga Whales – Diet and Eating Habits". Archived from the original on 12 July 2010. Retrieved 30 July 2010.
- Katona, Steven K.; Valerie Rough & David T. Richardson. (1983). A Field Guide to the Whales, Porpoises and Seals of the Gulf of Maine and Eastern Canada. New York: Charles Scribner's Sons. p. 255. ISBN 978-0-684-17902-5.
- Lentifer, Jack W. (1988). Selected Marine Mammals of Alaska: Species Accounts with Research and Management Recommendations. Washington, D.C.: Marine Mammals Commission. ASIN B000102908. OCLC 18060784.[page needed]
- Suydam, Robert Scott (2009). "Age, growth, reproduction, and movements of beluga whales (Delphinapterus leucas) from the eastern Chukchi Sea" (PDF). University of Washington. Archived (PDF) from the original on 19 September 2011. Retrieved 4 August 2008.
- Ellis, Samuel; Franks, Daniel W.; Nattrass, Stuart; Currie, Thomas E.; Cant, Michael A.; Giles, Deborah; Balcomb, Kenneth C.; Croft, Darren P. (27 August 2018). "Analyses of ovarian activity reveal repeated evolution of post-reproductive lifespans in toothed whales". Scientific Reports. 8 (1): 12833. Bibcode:2018NatSR...812833E. doi:10.1038/s41598-018-31047-8. ISSN 2045-2322. PMC 6110730. PMID 30150784.
- Robeck, Todd R.; Monfort, Steven L.; Calle, Paul P.; Dunn, J. Lawrence; Jensen, Eric; Boehm, Jeffrey R.; Young, Skip; Clark, Steven T. (2005). "Reproduction, Growth and Development in Captive Beluga (Delphinapterus leucas)". Zoo Biology. 24 (1): 29–49. doi:10.1002/zoo.20037.
- Cosens, S. & Dueck, L. (June 1990). "Spring Sightings of Narwhal and Beluga Calves in Lancaster Sound, N.W.T" (PDF). Arctic. 43 (2): 1–2. doi:10.14430/arctic1602. Archived (PDF) from the original on 28 October 2008.
- Animal Diversity Web – Museum of Zoology, University of Michigan. "Delphinapterus leucas". Archived from the original on 6 June 2011. Retrieved 30 July 2010.
- Sea World.org. "Beluga Whales – Birth and Care of the Young". Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Sea World Education Department (1993). Toothed Whales. San Diego: Sea World Education Department Publication.
- Leung, Elaine S.; Vergara, Valeria; Barrett-Lennard, Lance G. (2010). "Allonursing in captive belugas (Delphinapterus leucas)". Zoo Biology. 29 (5): 633–7. doi:10.1002/zoo.20295. PMID 20127963.
- Au, W. W. L.; Carder, D. A.; Penner, R. H.; Scronce, B. L. (1985). "Demonstration of adaptation in beluga whale echolocation signals". The Journal of the Acoustical Society of America. 77 (2): 726–30. Bibcode:1985ASAJ...77..726A. doi:10.1121/1.392341. PMID 3973242.
- "The Canaries of the Sea, granted a pardon, this time…". ePluribus Media. Archived from the original on 29 November 2009. Retrieved 7 August 2010.
- Bonner, W (1989). Whales of the World. New York: Facts on File Journals. p. 191. ISBN 978-0-8160-5216-5. Archived from the original on 29 June 2012.
- Rice DW (1998). Wartzok D (ed.). Marine mammals of the world: systematics and distribution (4 ed.). Lawrence, KS. USA: Society for Marine Mammalogy, Special Journal. p. 231. ISBN 978-1-891276-03-3.
- Fisheries and Oceans Canada. Beluga Whale Archived 5 August 2017 at the Wayback Machine
- Artyukhin, Yu.B.; Burkanov, V.N. (1999). Sea birds and mammals of the Russian Far East: a Field Guide. Moscow: АSТ Publishing. p. 215. ISBN 978-5237044751. (in Russian)
- Suydam RS, Lowry LF, Frost KJ, O' Corry-Crowe GM, Pikok D JR (2001). "Satellite Tracking of Eastern Chukchi Sea Beluga Whales into the Arctic Ocean". Arctic. 54 (3): 237–243. doi:10.14430/arctic784. Archived from the original on 2 February 2015.
- Andrei, Mihai (6 April 2018). "Beluga whales value culture and family ties". ZME Science. Archived from the original on 15 April 2018. Retrieved 14 April 2018.
- Barber DG, Saczuk E, Richard PR (2001). "Examination of Beluga-Habitat Relationships through the Use of Telemetry and a Geographic Information System". Arctic. 54 (3): 305–316. doi:10.14430/arctic790. Archived from the original on 2 February 2015.
- "Whale Carcass in Alaska River Mystifies Experts". Los Angeles Times. 16 June 2006. Archived from the original on 11 January 2012. Retrieved 3 August 2010.
- Gewalt W (1994). "Wale und Delphine 1 -Delphinapterus leucas – Weißwal oder Beluga". Handbuch der Säugetiere Europas. Meeressäuger. Teil IA (in German). Aula-Verlag, Wies-baden. pp. 185–208. ISBN 978-3-89104-560-2.
- "Beluga's body in river baffles scientists". South Bend Tribune. Archived from the original on 13 December 2013. Retrieved 10 September 2012.
- "Beluga Whale". Alaskan Naure. Archived from the original on 2 June 2013.
- Heide Jørgensen MP & Reeves RR (1996). "Evidence of a decline in beluga, Delphinapterus leucas, abundance off West Greenland". ICES J Mar Sci. 53 (1): 61–72. doi:10.1006/jmsc.1996.0006.
- Irish Whale and Dolphin Group. beluga whale Archived 8 March 2016 at the Wayback Machine. Retrieved 2 June 2014
- BBC News. (21 February 2012) Beluga whale sightings in SNH marine renewables report Archived 2 June 2014 at the Wayback Machine. Retrieved 2 June 2014
- Wildlife Extra. Dolphin and whale watching in the Scotland Archived 2 June 2014 at the Wayback Machine. Retrieved 2 June 2014
- Hebridean Whale and Dolphin Trust. Beluga Whale Archived 20 June 2013 at the Wayback Machine. Retrieved 2 June 2014
- Sato, H.; Ichimura, M. (2011). "The Sighting Record of Beluga (White Whale) Delphinapterus leucas in Shiretoko–Nemuro Strait Water, Eastern Hokkaido, Japan. Sea Life Watch in Shibetsu". Bulletin of the Shiretoko Museum. 32: 45–52.
- Uni, Y (2006). "Whales, Dolphins and Porpoises off Shiretoko" (PDF). Bulletin of the Shiretoko Museum. 27: 37–46. Archived (PDF) from the original on 14 July 2014.
- Volcano-bay Marine-animals Waching Association Archived 23 September 2015 at the Wayback Machine of Muroran Renaissance
- Mukai. T., KK-ELM Archived 20 October 2015 at the Wayback Machine. Retrieved 2 June 2014
- Sasamori K., Muroran Dolphin and Whale Watching Archived 14 July 2014 at the Wayback Machine. Retrieved 2 June 2014
- Imai Y., 野生のシロイルカ/ベルーガ in 日本 Beluga, the White Whale observations in Japanese waters. Archived 8 June 2015 at the Wayback Machine. YouTube. Retrieved 2 June 2014
- 흰고래 Delphinaterus leucas 영명 : Beluga, White whale 일명 : 시로이루카 Archived 4 March 2016 at the Wayback Machine. whalelove.com
- ベルーガ（シロイルカ） Delphinapterus leucas Archived 20 November 2015 at the Wayback Machine. Association for Protection of Marine Communities (1 January 2015)
- Nakayama Y.. 2016. 北極のベルーガ、北海道でひとりぼっち 遊び相手は漁師 Archived 25 March 2017 at the Wayback Machine. The Asahi Shimbun. Retrieved 25 March 2017
- Reyes JC (1991). The conservation of small cetaceans: a review. Report prepared for the Secretariat of the Convention on the Conservation of Migratory Species of Wild Animals. UNEP/CMS Secretariat, Bonn.
- Hobbs RC, Laidre KL, Vos DJ, Mahoney BA, Eagleton M (2005). "Movements and Area Use of Belugas, Delphinapterus leucas, in a Subarctic Alaskan Estuary" (PDF). Arctic. 58 (4): 331–340. doi:10.14430/arctic447. Archived (PDF) from the original on 10 February 2012.
- Rugh DJ, Mahoney BA, Smith BK (2004). "Aerial Surveys of Beluga Whales in Cook Inlet, Alaska, Between June 2001 and June 2002" (PDF). NOAA Technical Memorandum NMFS AFSC. no. 145. p. 20. Retrieved 7 August 2010.[permanent dead link]
- Beluga whales in James Bay: a separate entity from eastern Hudson Bay belugas? Archived 24 October 2016 at the Wayback Machine
- Goetz, Kimberly T.; Rugh, David J.; Read, Andrew J. & Hobbs, Roderick C. (2 October 2018). "Habitat use in a marine ecosystem: beluga whales Delphinapterus leucas in Cook Inlet, Alaska" (PDF). National Marine Mammal Laboratory, Alaska Fisheries Science Center, NMFS, NOAA, Washington, USA. Archived (PDF) from the original on 15 June 2010.
- Loseto, L.L.; Richard, P.; Stern, G.A.; Orr, J.; Ferguson, S.H. (1 December 2006). "Segregation of Beaufort Sea beluga whales during the open-water season". Canadian Journal of Zoology. 84 (12): 1743–1751(9). doi:10.1139/Z06-160.[permanent dead link]
- Ellis, Richard (1991). Men and Whales (1st ed.). The Lyons Press. p. 560. ISBN 978-1-55821-696-9.
- Dold, Catherine (July – August 1993). "The Great White Whales". Wildlife Conservation: 45–53. ISSN 1048-4949.
- Hoover C, Bailey M, Higdon J, Ferguson SH, Sumalia R (March 2013). "Estimating the Economic Value of Narwhal and Beluga Hunts in Hudson Bay, Nunavut". The Arctic Institute of North America. 66: 1–16.
- Heide-Jørgensen, Mads Peter (January 1994). "Distribution, exploitation and population status of white whales (Delphinapterus leucas) and narwhals (Monodon monoceros) in West Greenland". Meddelelser Om Grønland, Bioscience. 39: 135–149.
- Bettridge, Shannon, Robert L. Brownell Jr., Melissa Andersen Garcia, Rod C. Hobbs, Cheri L. McCarty, Richard D. Methot Jr., Debra L. Palka, Patricia E. Rosel, Kathryn S. Swails, and Barbara L. Taylor (1 March 2016). "Status Review of the Sakhalin Bay-Amur River Beluga Whale (Delphinapterus leucas) under the Marine Mammal Protection Act" (PDF). cites Shpak, Meschersky, Hobbs, Andrews, Glazov, Chelintsev, Kuznetsova, Solovyev, Nazarenko, Michaud, and Mukhametov. 2011. Current status of the Sakhalin-Amur beluga aggregation (The Okhotsk Sea, Russia): sustainability assessment. Report for 2007–2010 stages: Report presented at IUCN Independent Scientific Review Panel, Chicago, 6–7 March 2011 (unpublished). 68p and 5 Appendices. Archived (PDF) from the original on 30 April 2017. Retrieved 7 April 2018. This table uses their "high" estimate, since they say, page 23: "The high take scenario is considered the better of the two because it accounts for times when takes are known to have occurred but are not documented. Additionally, we did not account for beluga whales that are struck and lost because these data are not available, so the high take option may even be an underestimate."
- Master, Farah (20 September 2018). "Tidal wave of Chinese marine parks fuels murky cetacean trade". Reuters. Retrieved 17 November 2018.
- Kustikova, Alisa (8 November 2018). "It's like a detention center!, "Это как СИЗО!"". Новая газета – Novayagazeta.ru (in Russian). Retrieved 23 November 2018.
- "In Primorye, the court arrested killer whales and belugas in the case of the illegal export of mammals abroad, В Приморье суд арестовал косаток и белух по делу о незаконном вывозе млекопитающих за границу". Новая газета – Novayagazeta.ru (in Russian). 23 November 2018. Retrieved 23 November 2018.
- Tyrrell, Martina (2007). "Sentient Beings and Wildlife Resources: Inuit, Beluga Whales and Management Regimes in the Canadian Arctic". Human Ecology. 35 (5): 575–86. doi:10.1007/s10745-006-9105-2.
- Boltunov, Andrei N. and Belikov, Stanislav E. (2002). "Belugas (Delphinapterus leucas) of the Barents, Kara and Laptev seas". NAMMCO Sci. Publ. 4: 149–168. doi:10.7557/3.2842 – via Septentrio Academic Publishing, Arctic University of Norway.CS1 maint: Multiple names: authors list (link)
- Burns, John J. and Glenn A. Seaman (1 November 1986). "INVESTIGATIONS OF BELUKHA WHALES IN COASTAL WATERS OF WESTERN AND NORTHERN ALASKA II. BIOLOGY AND ECOLOGY" (PDF). Outer Continental Shelf Environmental Assessment Program. Alaska Department of Fish and Game. Archived (PDF) from the original on 27 March 2018.
- Dionne, Suzan & Gourbilière, Claire (2007). "St. Lawrence Beluga". Encyclopedia of French Cultural Heritage in North America. Archived from the original on 4 March 2013. Retrieved 1 September 2010.
- "Beluga Whales in Captivity" (PDF). Special Report on Captivity 2006. Canadian Marine Environment Protection Society. 2006. Archived from the original (PDF) on 26 December 2014. Retrieved 26 December 2014.
- Reeves, R. R. & Mitchell, E. (1984). "Catch history and initial population of white whales (Delphinapterus leucas) in the river and gulf of St. Lawrence, eastern Canada". Canadian Field-Naturalist (111): 63–121. Archived from the original on 5 November 2012. Retrieved 7 September 2010.
- HARWOOD, LOIS A., PAMELA NORTON, BILLY DAY and PATRICIA A. HALL (1 March 2002). "The Harvest of Beluga Whales in Canada's Western Arctic: Hunter-Based Monitoring of the Size and Composition of the Catch (has data through 1984)". Arctic. 55: 10–20. Archived from the original on 3 May 2018 – via University of Calgary. Includes struck and lost.
- Fisheries Joint Management Committee (2013). "BEAUFORT SEA BELUGA MANAGEMENT PLAN, 4th amended printing" (PDF). Archived (PDF) from the original on 25 March 2018. Retrieved 7 April 2018. Includes struck and lost.
- Fisheries, NOAA (17 September 2018). "Marine Mammal Stock Assessment Reports (SARs) by Region :: NOAA Fisheries". fisheries.noaa.gov. Archived from the original on 3 May 2018. Retrieved 7 April 2018. includes struck and lost.
- Loseto, Lisa L.; Brewster, Jasmine D.; Ostertag, Sonja K.; Snow, Kathleen; MacPhee, Shannon A.; McNicholl, Darcy G.; Choy, Emily S.; Giraldo, Carolina; Hornby, Claire A. (21 February 2018). "Diet and feeding observations from an unusual beluga harvest in 2014 near Ulukhaktok, Northwest Territories, Canada". Arctic Science. 4 (3): 421–431. doi:10.1139/as-2017-0046. hdl:1807/88141. ISSN 2368-7460.
- SULUK, THOMAS K., and SHERRIE L. BLAKNEY (2008). "Land Claims and Resistance to the Management of Harvester Activities in Nunavut" (PDF). Arctic. 61: 62–70. Archived (PDF) from the original on 21 September 2017 – via University of Calgary.
- Kishigami, Noboru (2005). "Co-Management of Beluga Whales in Nunavik (Arctic Quebec), Canada". Senri Ethnological Studies. 67: 121–144. Archived from the original on 3 May 2018.
- Rogers, Sarah (22 August 2016). "Nunatsiaq News 2016-08-22: NEWS: Nunavik's beluga season closes early". Nunatsiaq News. Archived from the original on 12 July 2018. Retrieved 7 April 2018.
- Heide-Jorgensen, M.P. and E. Garde (1 March 2017). "Catch statistics for belugas in Greenland 1862 to 2016" – via NAMMCO/JCNB Joint Working Group on narwhals and belugas, Copenhagen. Paper is not online. This table uses their "high" estimate (which includes struck and lost), without harvests from ice entrapments (which would die even if not harvested).
- Stanek, Ronald (1 July 1994). "The Subsistence Use of Beluga Whale in Cook Inlet by Alaska Natives, 1993" (PDF). Alaska Dept of Fish and Game. Archived (PDF) from the original on 9 August 2017. Retrieved 7 April 2018.
- "Participation in subsistence fishing by residents-Beluga harvested by residents (downloadable data)". 2011. Archived from the original on 27 March 2018. Retrieved 7 April 2018.
- Muto, M.M., V. T. Helker, R. P. Angliss, B. A. Allen, P. L. Boveng, J.M. Breiwick, M. F. Cameron, P. J. Clapham, S. P. Dahle, M. E. Dahlheim, B. S. Fadely, M.C. Ferguson, L. W. Fritz, R. C. Hobbs, Y. V. Ivashchenko, A. S. Kennedy, J.M. London, S. A. Mizroch, R. R. Ream, E. L. Richmond, K. E. W. Shelden, R. G. Towell, P. R. Wade, J. M. Waite, and A. N. Zerbini (2017). "Alaska Marine Mammal Stock Assessments, 2017 (draft)". Marine Mammal Laboratory, Alaska Fisheries Science Center, NMFS, NOAA. Retrieved 8 April 2018.
- Fisheries, NOAA (31 January 2018). "Draft Marine Mammal Stock Assessment Reports, NOAA Fisheries". fisheries.noaa.gov. Archived from the original on 28 April 2018. Retrieved 8 April 2018.
- Hernández, Clara (17 November 2006). "Un grupo de cazadores matará en Canadá a cerca de 80 belugas que han quedado atrapadas por el hielo" (in Spanish). 20 minutos.es – Internacional.
- Freeman, Milton M. R. "Polar Bear Predation on Beluga in the Canadian Arctic" (PDF). McMaster University, Hamilton, Ontario. Archived (PDF) from the original on 15 April 2010. Retrieved 12 August 2010.
- Shelden KE, Rugh DJ, Mahoney BA, Dahlheim ME (2003). "Killer whale predation on belugas in Cook Inlet, Alaska: Implications for a depleted population" (PDF). Mar. Mammal Sci. 19 (3): 529–544. doi:10.1111/j.1748-7692.2003.tb01319.x. Archived (PDF) from the original on 24 December 2012.
- Lowry LF, Nelson RR, Frost KJ (1987). "Observations of killer whales, Orcinus orca, in western Alaska: Sightings, strandings, and predation on other marine mammals". Ont. Field-Nat. 101 (1): 6–12. Archived from the original on 5 November 2012.
- Beland, Pierre (1996). Beluga: A Farewell to Whales (1 ed.). The Lyons Press. p. 224. ISBN 978-1-55821-398-2.
- "Marine Pollution: Causes and Effects". Bamfield Marine Sciences Centre Public Education Program. Archived from the original on 18 February 2013.
- Metcalfe, C; Metcalfe, T; Ray, S; Paterson, G; Koenig, B (1999). "Polychlorinated biphenyls and organochlorine compounds in brain, liver and muscle of beluga whales (Delphinapterus leucas) from the Arctic and St. Lawrence estuary". Marine Environmental Research. 47 (1): 1–15. doi:10.1016/S0141-1136(98)00107-X.
- Smith, T. G.; D. J. St. Aubin & J. R. Geraci (1990). Advances in Research on the Beluga Whale, Delphinapterus leucas. Ottawa: Department of Fisheries and Oceans. ISBN 978-0-660-13817-6.
- Hansen, Carstenthye; Nielsen, Christianovergaard; Dietz, Rune; Hansen, Martinmunk (1990). "Zinc, cadmium, mercury and selenium in minke whales, belugas and narwhals from West Greenland". Polar Biology. 10 (7): 529–39. doi:10.1007/BF00233702.
- Martineau, D.; Lemberger, K.; Dallaire, A.; Labelle, P.; Lipscomb, T. P.; Michel, P.; Mikaelian, I. (2002). "Cancer in Wildlife, a Case Study: Beluga from the St. Lawrence Estuary, Québec, Canada". Environmental Health Perspectives. 110 (3): 285–92. doi:10.1289/ehp.02110285. PMC 1240769. PMID 11882480.
- Guise, S. D.; Lagace, A.; Beland, P. (1994). "Tumors in St. Lawrence Beluga Whales (Delphinapterus leucas)". Veterinary Pathology. 31 (4): 444–9. doi:10.1177/030098589403100406. PMID 7941233.
- Caron LM, Sergeant DE (1988). "Yearly variation in the frequency of passage of beluga whales (Delphinapterus leucas) at the mouth of the Saguenay River, Quebec, over the past decade". Can Nat. 115 (2): 111–116.
- Lesage, Veronique; Barrette, Cyrille; Kingsley, Michael C. S.; Sjare, Becky (January 1999). "The Effect of Vessel Noise on the Vocal Behavior of Belugas in the St. Lawrence River Estuary, Canada" (PDF). Marine Mammal Science. 15 (1): 65–84. doi:10.1111/j.1748-7692.1999.tb00782.x. Archived (PDF) from the original on 2 February 2012.
- Finley KJ, Miller GW, Davis RA, Greene CR (1990). "Reactions of belugas, Delphinapterus leucas, and narwhals, Monodon monoceros, to ice-breaking ships in the Canadian High Arctic. In "Advances in research on the beluga whale, Delphinapterus leucas"". Can Bull Fish Aquat Sci. 224: 97–117. Archived from the original on 5 November 2012.
- Sea World.org (2002). "Beluga Whales – Longevity and Causes of Death". Archived from the original on 19 June 2010. Retrieved 30 July 2010.
- Dierauf, L. & Gulland, F. (2001). CRC Handbook of Marine Mammal Medicine:Health, Disease, and Rehabilitation. CRC Press. pp. 26, 303, 359. ISBN 978-0-8493-0839-0.
- Calle, Paul P.; Kenny, David E.; Cook, Robert A. (1993). "Successful treatment of suspected erysipelas septicemia in a beluga whale (Delphinapterus leucas)". Zoo Biology. 12 (5): 483–90. doi:10.1002/zoo.1430120510.
- Wazura, K. W.; Strong, J. T.; Glenn, C. L.; Bush, Albert O. (1986). "Helminths of the Beluga Whale (Delphinapterus leucas) from the Mackenzie River Delta, Northwest Territories". Journal of Wildlife Diseases. 22 (3): 440–2. doi:10.7589/0090-3558-22.3.440. PMID 3735598.
- "The Whales, New York Tribune, August 9, 1861". New York Tribune. 9 August 1861. Archived from the original on 7 June 2003. Retrieved 5 December 2011.
- Tasker, John Paul (24 October 2018). "Senate passes bill that would ban whale, dolphin captivity in Canada". CBC. Retrieved 21 November 2018.
- "Beluga Whales". Mystic Aquarium. Archived from the original on 13 October 2013. Retrieved 21 November 2014.
- "Species of the Month, Juno". Mystic Aquarium. Archived from the original on 19 January 2013. Retrieved 21 November 2014.
- "Beluga Whales Training". GeorgiaAquarium. 4 March 2008. Archived from the original on 16 March 2016. Retrieved 21 November 2014.
- "Mariachi Band Serenades Beluga Whale". HuffPost. 3 August 2011. Archived from the original on 29 November 2014. Retrieved 21 November 2014.
- "Reddet russisk «militærhval» på norgesbesøk". Dagbladet (in Norwegian). 27 April 2019. Retrieved 29 April 2019.
- Jon Henley (2 May 2019). "As experts fear for its survival, Tufjord residents told not to feed beluga so that it can learn to forage". The Guardian.
- "Merlin Entertainments plc - 2018 Interim Results – Company Announcement - FT.com". Financial Times. 2 August 2018. Retrieved 21 November 2018.
- Ástvaldsson, Jóhann Páll (28 June 2018). "Beluga Whales Train for Sanctuary in Iceland". Iceland Review. Retrieved 21 November 2018.
- Wong, Li Za (10 July 2018). "World's First Open Water Sanctuary For Beluga Whales To Open Spring 2019 In Iceland". Star2.com. Retrieved 21 November 2018.
- Mountain, Michael (15 June 2017). "Merlin Entertainment Seeks to Move Belugas to Sanctuary". The Whale Sanctuary Project. Retrieved 21 November 2018.
- "Les parcs à thème controversés SeaWorld arrivent bientôt en Asie". Sciences et Avenir (in French). 27 March 2017. Retrieved 23 November 2018.
- "Ceta Base – Captive Cetacean Database – Cetacean Inventory • U.S. & Canada". www.cetabase.org. 31 January 2018. Retrieved 22 November 2018.
- "Belugas "floated away" to China. Белухи "уплыли" в Китай. За что задержан директор ФГБНУ "ТИНРО-Центр" Лев Бочаров?". Porto Franko, Vladivostok. Retrieved 23 November 2018. Article says Belugas cost USD 70,000 or 60,000, while Orcas cost USD 7 million, while another article says Orcas cost USD 1 million (RUB 67 million): https://www.novayagazeta.ru/articles/2018/09/14/77814-orfey-2005-g-r-1-sht-gratsiya-2009-g-r-1-sht
- "Arrests made in Russia following illegal whale trafficking scandal". WDC, Whale and Dolphin Conservation. 21 March 2017. Retrieved 23 November 2018.
- "Beluga (Delphinapterus leucas)Facts – Distribution – In the Zoo". WAZA : World Association of Zoos and Aquariums. Archived from the original on 10 February 2012. Retrieved 5 December 2011.
- "Nanuq". Cetacousin. Retrieved 9 September 2012.[permanent dead link]
- "Beluga Whale Calf Born at Shedd Aquarium". Association of Zoos and Aquariums. Archived from the original on 14 September 2012. Retrieved 9 September 2012.
- "Male Beluga Whale Arrives at Mystic Aquarium". Mystic Aquarium. Archived from the original on 20 January 2012. Retrieved 9 September 2012.
- Pedicini, Sandra. "SeaWorld switches beluga whales". OrlandoSentinel.com. Retrieved 22 November 2018.
- Inmaculada Sanz (3 November 2006). "Nace la primera cría de ballena beluga en cautividad en Europa" (in Spanish). Noticias 20minutos. Archived from the original on 7 February 2011.
- Noticias 20minutos Valencia (27 November 2006). "Muere la ballena beluga que nació hace 25 días en el Oceanográfico de Valencia" (in Spanish). Archived from the original on 1 July 2013.
- eldiario.es (16 November 2016). "Kylu, la beluga que nació en el Oceanogràfic de Valencia un día de superluna" (in Spanish). Archived from the original on 17 January 2017.
- Lepisto, Christine (2 August 2009). "Beluga Whale Saves Diver". Archived from the original on 30 March 2010. Retrieved 31 August 2010.
- "Wal Rettet Ertrinkende Taucherin!". Archived from the original on 2 August 2009. Retrieved 27 September 2009.
- "Born to Be Free". 18 July 2017. Archived from the original on 14 April 2017. Retrieved 12 October 2017.
- "Sonic Sea – Wild & Scenic Film Festival". Wild & Scenic Film Festival. Archived from the original on 12 October 2017. Retrieved 12 October 2017.
- Proctor, Jason. "Vancouver Aquarium sues filmmaker over critical documentary". CBC News. Archived from the original on 24 March 2017. Retrieved 12 October 2017.
- UCLA Department of Chemistry and Biochemistry. "Beluga Whale Watching". Archived from the original on 26 May 2010. Retrieved 11 February 2010.
- Blane, JM.; Jackson, R (1994). "The impact of ecotourism boats on the St. Lawrence beluga whales". Environmental Conservation. 21 (3): 267–269. doi:10.1017/S0376892900033282. Archived from the original on 5 November 2012.
- National Oceanic and Atmospheric Administration (January 2004). "Marine Wildlife Viewing Guidelines" (PDF). Archived (PDF) from the original on 17 June 2010. Retrieved 6 August 2010.
- Shpatak A. (2012). "Japan Sea. Rudnaya Bay. Polar White Whale". The 35PHOTO.ru. Archived from the original on 19 January 2015. Retrieved 19 January 2015.
- Berg Olsen, Martine (26 September 2018). "Benny the beluga whale spotted swimming in River Thames again". Metro.
- "Study: Male beluga whale mimics human speech". 23 October 2012. Archived from the original on 14 July 2014.
- "The Story of One Whale Who Tried to Bridge the Linguistic Divide Between Animals and Humans". Smithsonian Magazine. June 2014.
- Jefferson TA, Karkzmarski L, Laidre K, O'Corry-Crowe G, Reeves R, Rojas-Bracho L, Secchi E, Slooten E, Smith BD, Wang JY, Zhou K (2012). "Delphinapterus leucas". IUCN Red List of Threatened Species. 2012: e.T6335A17690692. doi:10.2305/IUCN.UK.2012.RLTS.T6335A17690692.en.
- Rosen, Yereth (17 October 2008). "Beluga whales in Alaska listed as endangered". Reuters. Archived from the original on 20 October 2008. Retrieved 17 October 2008.
- "Endangered and Threatened Species; Endangered Status for the Cook Inlet Beluga Whale" (PDF). National Oceanic and Atmospheric Administration. 22 October 2008. Archived (PDF) from the original on 12 May 2009. Retrieved 26 August 2009.
- Herbert, H. Josef (17 October 2008). "Government declares beluga whale endangered". Associated Press. Archived from the original on 28 October 2008. Retrieved 17 October 2008.
- Laidre, K. L.; Stirling, I.; Lowry, L.; Wiig, Ø; Heide-Jørgensen, M. P.; Ferguson, S. (2008). "Quantifying the sensitivity of arctic marine mammals to climate-induced habitat change". Ecological Applications. 18 (2): S97–S125. doi:10.1890./06-0546.1. PMID 18494365.
- National Oceanic and Atmospheric Administration (January 2004). "MMPA – The Marine Mammal Protection Act of 1972 as Amended (2007)" (PDF). Archived (PDF) from the original on 1 June 2010. Retrieved 6 August 2010.
- "Appendix II Archived 11 June 2011 at the Wayback Machine" of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). As amended by the Conference of the Parties in 1985, 1988, 1991, 1994, 1997, 1999, 2002, 2005 and 2008. Effective: 5 March 2009.
- "Official CITES website". Convention on International Trade in Endangered Species of Wild Fauna and Flora. Archived from the original on 9 December 2009. Retrieved 20 December 2009.
- Shih, Xenia. "In-depth: Belugas – Beluga Whales Under Threat". Jean-Michael Cousteau – Ocean Adventures. Archived from the original on 20 June 2017.
- St. Lawrence Action Plan – Official Page. "St. Lawrence Action Plan For a Sustainable Development". Archived from the original on 25 January 2011. Retrieved 6 August 2010.
- "Beluga Whale (St. Lawrence Estuary population)". Government of Canada. Archived from the original on 21 August 2017. Retrieved 22 August 2017.
- The Internet Movie Database. "Pour la suite du monde". Archived from the original on 4 January 2010. Retrieved 7 August 2010.
- Callahan, Mike; Owens, Mike; Edwards, David & Eyries, Patrice (2005). "White Whale Album Discography". Archived from the original on 25 November 2010.
- "Baby beluga". National Institutes of Health. Archived from the original on 27 May 2010. Retrieved 12 February 2010.
- Motor Scooter Guide (2010). "Yamaha Beluga (Riva 80/CV80)". Archived from the original on 3 November 2010. Retrieved 8 August 2010.
- "Project 1710 Mackerel Beluga class". Federation of American Scientists. Archived from the original on 3 April 2009. Retrieved 11 February 2010.
- "Airbus beluga service". airbus.com. Archived from the original on 23 December 2010.
- Skysails. "Skysails turn wind into profit". Archived from the original on 10 October 2008. Retrieved 12 February 2010.
- Schillaci, Sophie (9 August 2013). "D23: Disney Sets Voice Casts for 'Finding Dory,' 'Inside Out' and 'Good Dinosaur'". The Hollywood Reporter. Archived from the original on 5 March 2016. Retrieved 9 August 2013.
- Lord, Nancy (2004). Beluga days: tracking a white whale's truths. Counterpoint. ISBN 978-1-58243-151-2.
- Outridge P. M.; Hobson K. A.; McNeely R.; Dyke A. (2002). "A Comparison of Modern and Preindustrial Levels of Mercury in the Teeth of Beluga in the Mackenzie Delta, Northwest Territories, and Walrus at Igloolik, Nunavut, Canada". Arctic. 55 (2): 123–132. doi:10.14430/arctic696.
- Perrin, William F.; Wursig, Bernd; Thewissen, J.G.M. 'Hans', eds. (26 February 2009). "Encyclopedia of Marine Mammals". Encyclopedia of Marine Mammals. Academic Press. ISBN 978-0-08-091993-5.
|Wikimedia Commons has media related to Delphinapterus leucas.|
|Wikispecies has information related to Delphinapterus leucas|
- US National Marine Fisheries Service beluga whale page
- Cook Inlet Beluga Population Info
- Vancouver Aquarium Beluga Webcam
- ARkive Photos
- Animal Diversity Web
- on YouTube
- Convention on Migratory Species page on the Beluga / White whale
- Animals in National Geographic – Beluga Whale Delphinapterus leucas
- Lifestyles of Beluga Whales National Geographic, video
- "Beluga whale". Encyclopedia of Life. Archived from the original on 22 September 2009. Retrieved 12 September 2012.
- Information about beluga in animaldiversity
- Video showing the birth of a beluga calf in Vancouver, video
- Information about belugas in marinebio.org
- Voices in the Sea – Sounds of the Beluga Whale