Beluga whale

Beluga whale; Temporal range:Quaternary–Recent PreꞒ Ꞓ O S D C P T J K Pg N ↓
	AtCity of Arts and Sciences,Spain
	Size compared to an average human
Conservation status
	; Least Concern(IUCN 3.1)
	CITESAppendix II(CITES)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Infraorder:	Cetacea
Family:	Monodontidae
Genus:	Delphinapterus; Lacépède,1804
Species:	D. leucas
Binomial name
	Delphinapterus leucas; (Pallas,1776)
	Beluga range

Thebeluga whale(/bɪˈluːɡə/;^[4]Delphinapterus leucas) is anArcticand sub-Arcticcetacean.It is one of two members of the familyMonodontidae,along with thenarwhal,and the only member of thegenusDelphinapterus.It is also known as thewhite whale,as it is the only cetacean to regularly occur with this colour; thesea canary,due to its high-pitched calls; and themelonhead,though that more commonly refers to themelon-headed whale,which is anoceanic dolphin.

The beluga is adapted to life in the Arctic, with anatomical and physiological characteristics that differentiate it from other cetaceans. Amongst these are its all-white colour and the absence of adorsal fin,which allows it to swim under ice with ease.^[5]It possesses a distinctive protuberance at the front of its head which houses anecholocation organcalled themelon,which in this species is large and deformable. The beluga's body size is between that of a dolphin and a true whale, 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. Like many cetaceans, a large percentage of its weight isblubber(subcutaneous fat). Its sense of hearing is highly developed and itsecholocationallows it to move about and find breathing holes under sheet ice.

Belugas aregregariousand form groups of 10 animals on average, although during the summer, they can gather in the hundreds or even thousands inestuariesand shallow coastal areas. They are slow swimmers, but can dive to 700 m (2,300 ft) below the surface. They areopportunistic feedersand their diets vary according to their locations and the season. The majority of belugas live in theArctic Oceanand the seas and coasts around North America, Russia, and Greenland; their worldwide population is thought to number around 200,000.^[6]^[7]They aremigratoryand the majority of groups spend the winter around theArctic ice cap;when thesea icemelts 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 theInternational Whaling Commission,and each country has developed its own regulations in different years. Currently, someInuitin Canada and Greenland,Alaska Nativegroups and Russians are allowed to hunt belugas for consumption as well as for sale, asaboriginal whalingis excluded from theInternational Whaling Commission1986 moratorium on hunting. The numbers have dropped substantially in Russia and Greenland, but not in Alaska and Canada. Other threats include natural predators (polar bearsandkiller whales), contamination of rivers (as withpolychlorinated biphenyl(PCBs) whichbioaccumulateup the food chain),climate changeandinfectious diseases.The beluga was placed on theInternational Union for Conservation of Nature'sRed Listin 2008 as being "near threatened"; the subpopulation from theCook Inletin Alaska is consideredcritically endangeredand is under the protection of the United States'Endangered Species Act.Of all seven extant Canadian beluga populations, those inhabiting easternHudson 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,dolphinariumsand wildlife parks in North America, Europe and Asia. They are consideredcharismaticbecause of their docile demeanour and characteristic smile, communicative nature,^[8]and supple, graceful movement.^[9]

Taxonomy

The beluga was first described in 1776 byPeter Simon Pallas.^[1]It is a member of thefamily Monodontidae,which is in turn part of theparvorder Odontoceti(toothed whales).^[1]TheIrrawaddy dolphinwas once placed in the same family, though recent genetic evidence suggests these dolphins belong to the familyDelphinidae.^[10]^[11]Thenarwhalis the only other species within the Monodontidae.^[12]A skull has been discovered with intermediate characteristics supporting the hypothesis thathybridisationis possible between these two species.^[13]

The name of thegenus,Delphinapterus,means "dolphin without fin" (from theGreekδελφίν (delphin), dolphin and απτερος (apteros), without fin) and the species nameleucasmeans "white" (from the Greek λευκας (leukas), white).^[14]TheRed List of Threatened Speciesgives both beluga and white whale as common names, though the former is now more popular. The English name comes from theRussianбелу́га, which derives from the word белый (bélyj), meaning "white".^[14]Nowadays the word белу́га in Russian refers to thebeluga sturgeon,while the whale is called almost similarly - белу́ха ( "belúha" ).^{[citation needed]}

The whale is also colloquially known as the "sea canary" on account of its high-pitched squeaks, squeals, clucks, and whistles. A Japanese researcher claimed that 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.^[15]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 isNOC,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.^[16]

Evolution

Skeleton ofD. leucas

Mitochondrial DNAstudies have shown modern cetaceans last shared a common ancestor between 25 and 34 million years ago^[17]^[18]The superfamily Delphinoidea (which contains monodontids, dolphins and porpoises) split from other toothed whales, odontoceti, between 11 and 15 million years ago. Monodontids then split from dolphins (Delphinidae) and later fromporpoises(Phocoenidae), their closest relatives in evolutionary terms.^[17]In 2017 the genome of a beluga whale was sequenced, comprising 2.327 Gbp of assembled genomic sequence that encoded 29,581 predicted genes.^[19]The authors estimated that the genome-wide sequence similarity between beluga whales andkiller whalesis 97.87%.^{[citation needed]}

The beluga's earliest known distinctive ancestors include the prehistoricDenebola brachycephalafrom the late Miocene epoch (9–10 million years ago),^[20]^[21]andBohaskaia monodontoides,from the early Pliocene (3–5 million years ago).^[22]Fossil evidence from Baja California^[23]and Virginia indicate the family once inhabited warmer waters.^[22]A fossil of the monodontidCasatia thermophila,from five million years ago, provides the strongest evidence that monodontids once inhabited warmer waters, as the fossil was found alongside fossils of tropical species such as bull and tiger sharks.^[24]

The fossil record also indicates that, in comparatively recent times, the beluga's range varied with that of the polar ice packs expanding duringice agesand contracting when the ice retreated.^[25]Counter-evidence to this theory comes from the finding in 1849 of fossilised beluga bones inVermontin the United States, 240 km (150 mi) from the Atlantic Ocean. The bones were discovered during construction of the first railroad betweenRutlandandBurlingtonin Vermont, when workers unearthed the bones of a mysterious animal inCharlotte.Buried nearly 10 ft (3.0 m) below the surface in a thick blueclay,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 amarine mammalburied beneath the fields of rural Vermont.^{[citation needed]}

The remains were found to be preserved in the sediments of theChamplain 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.^[26]Today, theCharlotte whaleis the official Vermont State Fossil (making Vermont the only state whose official fossil is that of a still extant animal).^[27]

Description

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 tail-fin grows and becomes increasingly and ornately curved as the animal ages. The flippers are broad and short—making them almost square-shaped.

Longevity

Preliminary investigations suggested a beluga'slife expectancywas rarely more than 30 years.^[28]The method used to calculate the age of a beluga is based on counting the layers ofdentineanddental cementin 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.^[29]A 2006 study usingradiocarbon datingof the dentin 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.^[30]However, recent studies suggest that it is unclear as to whether belugas receive a different number of layers per year depending on the age of the animal (for example young belugas may only receive an additional one layer per year), or simply just one layer per year or every other year.^[31]^{[citation needed]}

Size

The species presents a moderate degree ofsexual dimorphism,as the males are 25% longer than the females and are sturdier.^[32]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).^[33]Males weigh between 1,100 and 1,600 kg (2,430 and 3,530 lb), and occasionally up to 1,900 kg (4,190 lb) while females weigh between 700 and 1,200 kg (1,540 and 2,650 lb).^[34]^[35]They rank as mid-sized species among toothed whales.^[36]

Individuals of both sexes reach their maximum size by the time they are 10 years old.^[37]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.^[38]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.^[39]^[40]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.^[41]

Colour

The adult beluga is rarely mistaken for any other species, because it is completely white or whitish-grey in colour.^[42]Calves are usually born grey,^[33]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 nine in males.^[42]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 bearsandkiller whales.^[43]Unlike other cetaceans, the belugas seasonallyshedtheir skin.^[44]During the winter, theepidermisthickens 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.^[44]

Head and neck

Like mosttoothed whales,the beluga has a compartment found at the centre of the forehead that contains an organ used for echolocation called amelon,which contains fatty tissue.^[45]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.^[45]Another distinctive characteristic it possesses is the melon is malleable; its shape is changed during the emission of sounds.^[12]The beluga is able to change the shape of its head by blowing air around itssinusesto focus the emitted sounds.^[46]^[47]This organ containsfatty acids,mainlyisovaleric 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.^[48]

Unlike many dolphins and whales, the sevenvertebraein the neck are not fused together, allowing the animal to turn its head laterally without needing to rotate its body.^[49]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.^[43]Therostrumhas about eight to ten small, blunt and slightly curved teeth on each side of the jaw and a total of 36 to 40 teeth.^[50]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.^[51]

Belugas only have a singlespiracle,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.^[52]A beluga'sthyroidgland is larger than that of terrestrialmammals—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.^[53]It is the marine cetacean that most frequently developshyperplasticandneoplasticlesions of the thyroid.^[54]

Fins

The fins retain the bony vestiges of the beluga's mammalian ancestors, and are firmly bound together byconnective tissue.^[38]The fins are small in relation to the size of the body, rounded and oar-shaped and slightly curled at the tips.^[14]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.^[37]The fins also contain a mechanism forregulating body temperature,as thearteriesfeeding the fin's muscles are surrounded byveinsthat dilate or contract to gain or lose heat.^[38]^[55]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.^[38]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.^[38]

Belugas have a dorsal ridge, rather than adorsal fin.^[33]The absence of the dorsal fin is reflected in the genus name of the species—apterustheGreekword 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.^[12]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.^[56]

Senses

The beluga has a very specialised sense of hearing and itsauditory cortexis highly developed. It can hear sounds within the range of 1.2 to 120kHz,with the greatest sensitivity between 10 and 75 kHz,^[57]where the average hearing range for humans is 0.02 to 20 kHz.^[58]The majority of sounds are most probably received by the lowerjawand transmitted towards themiddle ear.In the toothed whales, the lower jawbone is broad with a cavity at its base, which projects towards the place where it joins thecranium.A fatty deposit inside this small cavity connects to the middle ear.^[59]Toothed whales also possess a small external auditory hole a few centimetres behind their eyes; each hole communicates with an external auditory conduit and aneardrum.It is not known if these organs are functional or simplyvestigial.^[59]

Belugas are able to see within and outside of water, but their vision is relatively poor when compared to dolphins.^[60]Their eyes are especially adapted to seeing under water, although when they come into contact with the air, the crystallinelensand thecorneaadjust to overcome the associatedmyopia(the range of vision under water is short).^[60]A beluga'sretinahasconesandrods,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.^[60]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 theconjunctivafrom pathogenic organisms.^[60]

Studies on captive animals show they seek frequent physical contact with other belugas.^[43]Areas in the mouth have been found that could act aschemoreceptorsfor different tastes, and they can detect the presence of blood in water, which causes them to react immediately by displaying typical alarm behaviour.^[43]Like the other toothed whales, their brains lackolfactory bulbsand olfactory nerves, which suggests they do not have a sense of smell.^[45]

Behaviour

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.^[61]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.^[62]Beluga whale pods can be grouped into three categories, nurseries (which consist of mother and calves), bachelors (which consist of all males) and mixed groups. Mixed groups contain animals of both sexes.^[63]^[50]Many hundreds and even thousands of individuals can be present when the pods join 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.^[64]

They are cooperative animals and frequently hunt in coordinated groups.^[65]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.^[66]Often individuals will surface and dive together in a synchronized manner, in a behavior known as milling.

In captivity, they can be seen to be constantly playing, vocalising and swimming around each other.^[67]In one case, one whale blew bubbles, while the other one popped them. There have also been reports of beluga whales copying and imitating one another, similar to a game of Simon-says. There have also been reports of them displaying physical affection, via mouth to mouth contact. They also show a great deal of curiosity towards humans and frequently approach the windows in the tanks to observe them.^[68]

Belugas also show a great degree of curiosity towards humans in the wild, and frequently swim alongside boats.^[69]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.^[39]During the breeding season, adults have been observed carrying objects such as plants, nets, and even the skeleton of a deadreindeeron their heads and backs.^[67]Captive females have also been observed displaying this behavior, carrying items such as floats and buoys, after they have lost a calf. Experts consider this interaction with the objects to be asubstitute behavior.^[70]

In captivity, mothering behavior among belugas depends on the individual. Some mothers are extremely attentive while other mothers are so blasé, that they have actually lost their calves. In aquaria, there have been cases where dominant females have stolen calves from mothers, particularly if they have lost a calf or if they are pregnant. After giving birth, dominant females will return the calf back to their mother. Additionally, male calves will temporarily leave their mothers to interact with an adult male who can serve as a role model for the calf, before they return to their mothers. Male calves are also frequently seen interacting with each other.

Swimming and diving

Belugas are slower swimmers than the other toothed whales, such as the killer whale and thecommon bottlenose dolphin,because they are less hydrodynamic and have limited movement of their tail-fins, which produce the greatest thrust.^[71]They frequently swim at speeds 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.^[50]Unlike most cetaceans, they are capable of swimming backwards.^[37]^[72]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 enough to cover their bodies.^[37]They do not jump out of the water like dolphins or killer whales.^[14]

These animals usually only dive to depths to 20 m (66 ft),^[73]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,^[74]while animals in the wild have been recorded as diving to a depth of more than 700 m, with the greatest recorded depth being over 900 m.^[75]A dive normally lasts 3 to 5 minutes, but can last up to over 20 minutes.^[76]^[50]^[75]^[77]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.^[37]The average number of dives per day varies between 31 and 51.^[75]

All cetaceans, including belugas, have physiological adaptations designed to conserve oxygen while they are under water.^[78]During a dive, these animals will reduce their heart rate from 100 beats a minute to between 12 and 20.^[78]Blood flow is diverted away from certain tissues and organs and towards thebrain,heartandlungs,which require a constant oxygen supply.^[78]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 ofWeddell seals(a diving marine mammal). One study found a female beluga had 16.5 L of oxygen dissolved in her blood.^[79]Lastly, the beluga's muscles contain high levels of the proteinmyoglobin,which stores oxygen in muscle. Myoglobin concentrations in belugas are several times greater than for terrestrial mammals, which help prevent oxygen deficiency during dives.^[80]

Beluga whales often accompanybowhead whales,for curiosity and to securepolynyafeasibility to breathe as bowheads are capable of breaking through ice from underwater by headbutting.^[81]

Diet

Belugas play an important role in the structure and function of marine resources in theArctic Ocean,as they are the most abundant toothed whales in the region.^[82]They are opportunistic feeders; their feeding habits depend on their locations and the season.^[32]For example, when they are in theBeaufort Sea,they mainly eatArctic cod(Boreogadus saida) and the stomachs of belugas caught near Greenland were found to containrose fish(Sebastes marinus),Greenland halibut(Reinhardtius hippoglossoides) andnorthern shrimp(Pandalus borealis),^[83]while in Alaska their staple diet isCoho salmon(Oncorhynchus kisutch).^[84]In general, the diets of these cetaceans consist mainly of fish; apart from those previously mentioned, other fish they feed on includecapelin(Mallotus villosus),smelt,sole,flounder,herring,sculpinand other types ofsalmon.^[85]They also consume a great quantity of invertebrates, such asshrimp,squid,crabs,clams,octopus,sea snails,bristle wormsand other deep-sea species.^[85]^[86]Belugas feed mainly in winter as their blubber is thickest in later winter and early spring, and thinnest in the fall. Inuit observation has led scientists to believe that belugas do not hunt during migration, at least in Hudson Bay.^[87]

The diet of Alaskan belugas is quite diverse and varies depending on season and migratory behavior. Belugas in the Beaufort Sea mainly feed on staghorn and shorthorn sculpin,walleye pollock,Arctic cod,saffron codandPacific sand lance.Shrimp are the most common invertebrate eaten, with octopus,amphipodsandechiuridsbeing other sources of invertebrate prey. The most common prey species for belugas in the Eastern Chukchi Sea appears to be shrimp, echiurid worms, cephalopods andpolychaetes.The largest prey item consumed by beluga whales in the Eastern Chukchi Sea seems to be saffron cod. Beluga whales in the Eastern Bering Sea feed on a variety of fish species including saffron cod,rainbow smelt,walleye pollock,Pacific salmon,Pacific herringand several species of flounder and sculpin. The primary invertebrate consumed is shrimp. The primary prey item in regard to fish species for belugas in Bristol Bay appears to be the five species of salmon, with sockeye being the most prevalent. Smelt is also another common fish family eaten by belugas in this region. Shrimp is the most prevalent invertebrate prey item. The most common prey items for belugas in Cook Inlet appear to be salmon, cod and smelt.^[88]

Animals in captivity eat 2.5% to 3.0% of their body weight per day, which equates to 18.2 to 27.2 kg (40 to 60 lb).^[89]Like their wild counterparts, captive belugas were found to eat less in the fall.^[90]

Foraging on the seabed typically takes place at depths between 20 and 40 m (66 and 131 ft),^[91]although they can dive to depths of 700 m (2,300 ft) in search of food.^[75]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.^[65]As theirteethare 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.^[92]They also join into coordinated groups of five or more to feed onshoalsof fish by steering the fish into shallow water, where the belugas then attack them.^[65]For example, in the estuary of theAmur River,where they mainly feed on salmon, groups of six or eight individuals join to surround a shoal of fish and prevent their escape. Individuals then take turns feeding on the fish.^[56]

Reproduction

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 nine and fifteen years old, and females reach maturity between eight and fourteen years old.^[93]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,^[94]^[95]and ceasing reproductive potential with no births recorded for females older than 41.^[93]There is a slight difference on the sexual maturation period between males and females. The male beluga whales take seven to nine years to become sexually mature, while the females take four to seven years.^[96]

Female belugas typically give birth to one calf every three years.^[33]Most mating occurs from February to May, but some occurs at other times of year.^[12]The beluga may havedelayed implantation.^[12]Gestation has been estimated to last 12.0 to 14.5 months,^[33]but information derived from captive females suggests a period up to 475 days (15.8 months).^[97]During the mating season, the testes of belugas double in weight. Testosterone levels increase, but seems to be independent of copulation. Copulation typically takes place between 3 and 4 AM.

Calves are born over a protracted period that varies by location. In the Canadian Arctic, calves are born between March and September, while inHudson Bay,the peak calving period is in late June, and inCumberland Sound,most calves are born from late July to early August.^[98]Births usually take place in bays or estuaries where the water is between 10 and 15 °C.^[61]Newborns are about 1.5 m (4 ft 11 in) long, weigh about 80 kg (180 lb), and are grey in colour.^[50]They are able to swim alongside their mothers immediately after birth.^[99]The newborn calves nurse under water and initiatelactationa few hours after birth; thereafter, they feed at intervals around an hour.^[65]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.^[100]The milk contains about 92calperounce.^[101]

The calves remain dependent on their mothers for nursing for the first year, when their teeth appear.^[61]After this, they start to supplement their diets with shrimp and small fish.^[45]The majority of the calves continue nursing until they are 20 months old, although occasionally lactation can continue for more than two years,^[50]and lactationalanoestrusmay 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.^[102]

Hybrids have been documented between the beluga and the narwhal (specifically offspring conceived by a beluga father and a narwhal mother), as one, perhaps even as many as three, such hybrids were killed and harvested during a sustenance hunt. Whether or not these hybrids could breed remains unknown. The unusual dentition seen in the single remaining skull indicates the hybrid hunted on the seabed, much as walruses do, indicating feeding habits different from those of either parent species.^[103]^[104]

Communication and echolocation

Vocalisations ofDelphinapterus leucaspublished byNOAA

Belugas use sounds andecholocationfor movement, communication, to find breathing holes in the ice, and to hunt in dark or turbid waters.^[46]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.^[101]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.^[46]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.^[61]

Some evidence indicates that belugas are highly sensitive to noise produced by humans. In one study, the maximum frequencies produced by an individual located inSan 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 toKaneohe BayinHawaii.The difference in frequencies is thought to be a response to the difference in environmental noise in the two areas.^[105]In special circumstances, beluga whale sounds have been reported to resemble human speech.^[106]^[107]

These animals communicate using sounds of high frequency; their calls can sound like bird songs, so belugas were nicknamed "canaries of the sea".^[108]Like the other toothed whales, belugas do not possessvocal cordsand the sounds are probably produced by the movement of air between the nasal sacks, which are located near to the blowhole.^[46]

Belugas are among the most vocal cetaceans.^[109]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.^[46]They make sounds by grinding their teeth or splashing, but they rarely use body language.^[46]

There is debate as to whether cetacean vocalizations can constitute a language. A study conducted in 2015 determined that European beluga signals share physical features comparable to vowels. These sounds were found to be stable throughout time, but varied among different geographical locations. The further away the populations were from each other, the more varied the sounds were in relation to one another.^[110]

Distribution

The beluga inhabits a discontinuous circumpolar distribution in Arctic and sub-Arctic waters.^[111]During the summer, they can mainly be found in deep waters ranging from76°Nto80°N,particularly along the coasts ofAlaska,northern Canada, westernGreenlandand northern Russia.^[111]The southernmost extent of their range includes isolated populations in theSt. Lawrence Riverin the Atlantic,^[112]and theAmur Riverdelta, theShantar Islandsand the waters surroundingSakhalin Islandin theSea of Okhotsk.^[113]

Migration

Belugas have a seasonalmigratorypattern.^[114]Migration patterns are passed from parents to offspring. Some travel as far as 6,000 kilometres (3,700 mi) per year.^[115]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 usingpolynyasto surface and breathe.^[116]Insummerafter the sheet ice has melted, they move to coastal areas with shallower water (1–3 m [3 ft 3 in – 9 ft 10 in] deep), although sometimes they migrate towards deeper waters (deeper than 800 m [2,600 ft]).^[114]In the summer, they occupyestuariesand the waters of thecontinental shelf,and, on occasion, they even swim up the rivers.^[114]A number of incidents have been reported where groups or individuals have been found hundreds or even thousands of kilometres from the ocean.^[117]^[118]One such example comes from June 9, 2006, when a young beluga carcass was found in theTanana RivernearFairbanksin central Alaska, nearly 1,700 km (1,100 mi) from the nearest ocean habitat. Belugas sometimes follow migrating fish, leading Alaska state biologist Tom Seaton to speculate it had followed migratingsalmonup the river at some point in the previous autumn.^[119]The rivers they most often travel up include: theNorthern Dvina,theMezen,thePechora,theOband theYeniseiin Asia; theYukonand theKuskokwimin Alaska, and the Saint Lawrence in Canada.^[111]Spending time in a river has been shown to stimulate an animal's metabolism and facilitates the seasonal renewal of the epidermal layer.^[53]In addition, the rivers represent a safe haven for newborn calves where they will not be preyed upon by killer whales.^[12]Calves often return to the same estuary as their mother in the summer, meeting her sometimes even after becoming fully mature.^[120]However, not all beluga whale populations summer in estuaries. Belugas from the Beaufort Sea stock were found to summer along the Eastern Beaufort Sea shelf,Amundsen Gulfand slope regions north and west ofBanks Island,in addition to core areas in the Mackenzie River Estuary. Male belugas have been observed summering in deeper waters alongViscount Melville Sound,in depths of up to 600 metres (2,000 ft). The bulk of Eastern Chukchi Sea belugas summer over Barrow canyon.^[121]

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.^[122]Vagrants may travel further south to areas such asIrish^[123]andScottishwaters,^[124]the islands ofOrkney^[125]andHebrides,^[126]and toJapanesewaters.^[127]There had been several vagrant individuals^[128]that have demonstrated seasonal residencies atVolcano Bay,^[129]^[130]^[131]and a unique whale were used to return annually to areas adjacent toShibetsuinNemuro Straitin the 2000s.^[132]On rarer occasions, individuals of vagrancy can reach theKorean Peninsula.^[133]A few other individuals have been confirmed to return to the coasts of Hokkaido, and one particular individual became a resident inbrackish watersofLake Notorosince in 2014.^[134]^[135]

Some populations are not migratory and certain resident groups will stay in well-defined areas, such as inCook Inlet,the estuary of the Saint Lawrence River andCumberland Sound.^[136]The population in Cook Inlet stays in the waters furthest inside the inlet during the summer until the end of autumn. Then during the winter, they disperse to the deeper water in the center of the inlet, but without completely leaving it.^[137]^[138]

In April, the animals that spend the winter in the center and southwest of theBering Seamove to the north coast of Alaska and the east coast of Russia.^[136]The populations living in theUngava Bayand the eastern and western sides ofHudson Bayoverwinter together beneath the sea ice inHudson Strait.Whales inJames Baythat spend winter months within the basin, could be a distinct group from those in Hudson Bay.^[139]The populations of theWhite Sea,theKara Seaand theLaptev Seaoverwinter in the Barents Sea.^[136]In the spring, the groups separate and migrate to their respective summer sites.^[136]

Habitat

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.^[136]

In coastal areas, they can be found incoves,fjords,canals,baysand shallow waters in the Arctic Ocean that are continuously lit by sunlight.^[39]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.^[39]Themudflatsof Cook Inlet in Alaska are a popular location for these animals to spend the first few months of summer.^[140]In the eastern Beaufort Sea, female belugas with their young and immature males prefer the open waters close to land, while the adult males live in waters covered by ice near theCanadian Arctic Archipelago.The younger males and females with slightly older young can be found nearer to the ice shelf.^[141]Generally, the use of different habitats in summer reflects differences in feeding habits, risk from predators and reproductive factors for each of the subpopulations.^[32]

Population

There are currently 22 stocks of beluga whales recognized:^[142]

James Bay – 14,500 individuals (belugas remain here all year round)
Western Hudson Bay – 55,000 individuals
Eastern Hudson Bay – 3,400–3,800 individuals
Cumberland Sound – 1,151 individuals
Ungava Bay – 32 individuals (maybe functionally extinct)
St. Lawrence River Estuary – 889 individuals
Eastern Canadian Arctic – 21,400 individuals
Southwest Greenland – Extinct
Eastern Chukchi Sea – 20,700 individuals
Eastern Bering Sea – 7,000–9,200 individuals
Eastern Beaufort Sea – 39,300 individuals
Bristol Bay – 2,000–3,000 individuals
Cook Inlet – 300 individuals
White Sea – 5,600 individuals
Kara Sea/Laptev Sea/Barents Sea – Data Deficient
Ulbansky – 2,300
Anadyr – 3,000
Shelikhov – 2,666
Sakhalin/Amur – 4,000 individuals
Tugurskiy – 1,500 individuals
Udskaya – 2,500 individuals
Svalbard – 549 individuals^[7]

The Yakutat Bay belugas are not considered to be a true stock because they have only been present in these waters since the 1980s, and are believed to be of Cook Inlet origin. It is estimated that less than 20 whales inhabit the bay year-round.^[142]Overall the beluga population is estimated to be 150,000–200,000 animals.

Threats

Hunting

The native populations of the Arctic in Alaska, Canada, Greenland and Russia hunt belugas, for both consumption and profit. 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.^[143]

Present

The number of animals killed is about 1,000 per year, (see table below. and its sources). Beluga whale hunting quotas in Canada and the United States are established using the Potential Biological Removal equation PBR = Nmin * 0.5 * Rmax * FR, to determine what constitutes a sustainable hunt. Nmin represents a conservative estimation of the population size, Rmax, represents the maximum rate of population increase and FR represents the recovery factor.^[142]

Hunters in Hudson's Bay rarely eat the meat. They give a little to dogs, and leave the rest for wild animals.^[144]Other areas may dry the meat for later consumption by humans. In Greenland the skin (muktuk) is sold commercially to fish factories,^[145]and in Canada to other communities.^[144]An average of one or two vertebrae and one or two teeth per beluga are carved and sold.^[144]One estimate of the annual gross value received from Beluga hunts inHudson Bayin 2013 wasCA$600,000for 190 belugas, orCA$3,000per beluga. However, the net income, after subtracting costs in time and equipment, was a loss ofCA$60per 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$550,000was for skin and meat, to replace beef, pork and chickens which would otherwise be bought.CA$50,000was received for carved vertebrae and teeth.^[144]

Russia now harvests 5 to 30 belugas per year for meat and captures an additional 20 to 30 per year for live export to Chineseaquaria.^[146]^[147]However, in 2018, 100 were illegally captured for live export.^[148]^[149]

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 permittingsustainable huntingthat does not put the species at risk of extinction.^[150]

The total amount of landed (defined as belugas successfully hunted and retrieved) belugas averages 275 in regard to the Bering, Chukchi and Beaufort stocks from 1987 to 2006. The average annual landed harvest of belugas in the Beaufort Sea consisted of 39 individuals while the Chukchi harvest averaged 62 individuals. Bristol bay's annual average landed harvest was 17 while the Bering Sea's was 152. Statistical studies have demonstrated that subsistence hunting in Alaska did not significantly impact the population of the Alaskan beluga whale stocks. The number of belugas struck and lost did not seem to profoundly impact Chukchi and Bering Sea belugas.^[151]

Past

Commercial whaling by European, American and Russian whalers during the 18th and 19th centuries decreased beluga populations in the Arctic.^[143]^[152]^[153]The animals were hunted for their meat and blubber, while the Europeans used the oil from the melon as alubricantforclocks,machinery and lighting inlighthouses.^[143]Mineral oil replaced whale oil in the 1860s, but into the early 20th century the cured skin was still used to makehorse harnessesand machine belts forsaw millsandshoelaces.The cured skin is the only cetacean skin that is sufficiently thick to be used asleather,and was used to manufacture some of the firstbulletproof vests.^[143]^[154]^[155]

Russia had large hunts, peaking in the 1930s at 4,000 per year and the 1960s at 7,000 per year, for a total of 86,000 from 1915 to 2014.^[146]^[152]Canada hunted a total of 54,000 from 1731 to 1970.^[156]Between 1868 and 1911,Scottishand American whalers killed more than 20,000 belugas inLancaster SoundandDavis Strait.^[143]

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.^[155]The presence of belugas in the estuary was, therefore, considered to be undesirable; in 1928, theGovernment of Quebecoffered a reward of 15 dollars for each dead beluga.^[157]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.^[155]L'Isle-aux-Coudresis the setting for the classic 1963National Film Board of CanadadocumentaryPour 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 todolphin drive hunting.

Beluga catches by location

Annual table showing the number of belugas caught in Canada, Russia, Greenland and Alaska each year, from 2016 back to 1954
Beaufort Sea, Mackenzie, Paulatuk, Ulukhaktok, Canada	Nunavut, Canada	Nunavik, Quebec, Canada	Western Arctic, Russia, hunted for meat	Eastern Arctic, Russia, hunted for meat	Sea of Okhotsk, Russia, hunted for meat	All areas of Russia, live export	Year	Canada total	Greenland	USSR+ Russia total	US (Alaska)	World total, incomplete	Lost at sea as % of caught
		157					2016	157	246
83		303					2015	386	156		326	868
136		302			30	23	2014	438	317	53	346	1154	2%
92		207			30	23	2013	299	353	53	367	1072	2%
102		207			30	44	2012	309	245	74	360	988	4%
72		207			30	33	2011	279	179	63	288	809	3%
94		207			30	30	2010	301	222	60	318	901	3%
102		207			30	24	2009	309	286	54	253	902	6%
79		207			30	25	2008	286	330	55	254	925	8%
85		207			30	0	2007	292	145	30	576	1043	2%
126		207			30	20	2006	333	169	50	226	778	3%
108		207			30	31	2005	315	231	61	282	889	2%
142		207			30	25	2004	349	246	55	234	884	8%
125	250	207			30	26	2003	582	510	56	251	1399	9%
89	170	210			30	10	2002	469	510	40	362	1381	3%
96		370			30	22	2001	466	560	52	416	1494	1%
91	116	243			30	10	2000	450	733	40	280	1503	8%
102	207	243			30	23	1999	552	590	53	217	1412	19%
93	137	243			30	23	1998	473	873	53	342	1741	8%
123	376	243			30	23	1997	742	682	53	276	1753	8%
139	203	243			30	23	1996	585	681	53	389	1708	16%
143					30	23	1995	143	960	53	171	1327	11%
149					30	23	1994	149	757	53	285	1244	6%
120					30	23	1993	120	930	53	369	1472	9%
130					30	23	1992	130	1014	53	181	1378	7%
144					30	23	1991	144	747	53	315	1259	24%
106					30	23	1990	106	933	53	335	1427	22%
156			27		30	23	1989	156	816	80	13	1065	15%
139			7		30	23	1988	139	428	60	19	646	19%
174			15		30	23	1987	174	928	68	22	1192	13%
199			192		30	23	1986	199	973	245	0	1417	15%
148			248	150	30		1985	148	887	428	0	1463	17%
156			850	150	30		1984	156	930	1030	170	2286	20%
102			450	150	30		1983	102	888	630	235	1855	20%
146			116	150	30		1982	146	1217	296	335	1994	19%
155			294	150	30		1981	155	1506	474	209	2344	20%
85			368	150	30		1980	85	1346	548	249	2228	23%
171			200	26	30		1979	171	1116	256	138	1681	22%
157			63	26	30		1978	157	1112	119	177	1565	25%
172			1196	26	30		1977	172	1264	1252	247	2935	22%
183			472	26	30		1976	183	1260	528	186	2157	28%
177			169	23	30		1975	177	995	222	185	1579	23%
152			194	23	30		1974	152	1149	247	184	1732	25%
212			288	23	30		1973	212	1451	341	150	2154	23%
134			288		30		1972	134	1168	318	180	1800	21%
94			612		30		1971	94	913	642	250	1899	23%
137			990		30		1970	137	861	1020	200	2218	25%
			302	700	30		1969	0	1364	1032	170	2566	25%
14			30	700	30		1968	14	1490	760	150	2414	26%
40			274	700	30		1967	40	825	1004	225	2094	24%
96			3046	700	30		1966	96	828	3776	225	4925	23%
70			3614	700	30		1965	70	595	4344	225	5234	21%
45			5952	700	30		1964	45	403	6682	225	7355	22%
94			2526	700	30		1963	94	278	3256	225	3853	21%
96			2334	700	30		1962	96	409	3064	225	3794	24%
145			3500	700	30		1961	145	438	4230	300	5113	27%
145			6444	700	30		1960	145	398	7174	375	8092	22%
			1945	700	830		1959		472	3475	450	4397	24%
			2103	700	830		1958		411	3633	450	4494	23%
			796	700	830		1957		770	2326	450	3546	26%
			600	700	830		1956		671	2130	450	3251	25%
			329	700	130		1955		507	1159	450	2116	24%
			776	700	130		1954		767	1606	450	2823	28%
1960–1969^[153]1970–99^[158]2000–2012^[159]2013–15^[160]2014^[161]	Arviat^[162]	1996–2002^[163] 2003–16^[164]	1954–99^[152]	1954–1985 cites Russian papers^[153]	NMFS cites Russian paper^[146]	Western^[152]Okhotsk^[146]	Sources	Total of columns at left, incomplete	1954–2016^[165]	Total of columns at left, incomplete	1954–84^[153]1987–90 Cook Inlet^[166]1990–2011^[167]2012–2015 +Cook Inlet^[168]^[169]	Total of other columns	Greenland source 1954–1999, Beaufort source 2000–2012

Predation

During the winter, belugas commonly become trapped in the ice without being able to escape to open water, which may be several kilometres away.^[170]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.^[34]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 a beluga that weighed 935 kg.^[171]

Killer whaleshunt and eat both young and adult belugas.^[34]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.^[172]^[173]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 sub-population, which has already been badly depleted by hunting.^[172]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.^[39]Beluga whale behavior under killer whale predation makes them vulnerable to hunters. When killer whales are present, large numbers of beluga whales congregate in the shallows for protection, which allows them to be hunted in droves.

Contamination

The beluga is considered an excellentsentinel 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 asDDTand heavy metals such aslead,mercuryandcadmiumhave been found in individuals of the Saint Lawrence River population.^[174]Local beluga carcasses contain so many contaminants, they are treated as toxic waste.^[175]Levels ofpolychlorinated biphenylsbetween 240 and 800ppmhave been found in belugas'brains,liverandmuscles,with the highest levels found in males.^[176]These levels are significantly greater than those found in Arctic populations.^[177]These substances have a proven adverse effect on these cetaceans, as they causecancers,reproductive diseases and the deterioration of theimmune system,making individuals more susceptible topneumonias,ulcers,cysts,tumoursand bacterialinfections.^[177]Although the populations that inhabit the river estuaries run the greatest risk of contamination, high levels ofzinc,cadmium,mercury andseleniumhave also been found in the muscles, livers andkidneysof animals that live in the open sea.^[178]The concentration of mercury in Beaufort Sea belugas tripled from the 1980s to the 1990s, but has decreased in Beaufort belugas as of the 21st century, possibly due to changes in dietary preference. Larger body sized belugas tend to have more mercury than smaller sized belugas, because they spend more time offshore, hunting prey such as cod and shrimp, which have more mercury.^[179]

From a sample of 129 beluga adults from the Saint Lawrence River examined between 1983 and 1999, a total of 27% had suffered cancer.^[180]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.^[180]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 bypolycyclic aromatic hydrocarbons,and coincides with the high incidence of this disease in humans residing in the area.^[180]The prevalence of tumours suggests the contaminants identified in the animals that inhabit the estuary are having a directcarcinogeniceffect or they are at least causing an immunological deterioration that is reducing the inhabitants' resistance to the disease.^[181]

Indirect human disturbance may also be a threat. While some populations tolerate small boats, most actively try to avoid ships.Whale-watchinghas 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.^[182]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 asferriesthan with smaller boats.^[183]Belugas can detect the presence of large ships (for exampleicebreakers) 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.^[184]

Pathogens

As with any animal population, a number of pathogens cause death and disease in belugas, includingviruses,bacteria,protozoansandfungi,which mainly cause skin, intestinal and respiratory infections.^[185]

Papillomaviruses,herpesviruses,andencephalitiscaused by the protozoanSarcocystishave been found in belugas in the Saint Lawrence River. Cases have been recorded ofciliateprotozoa colonising the spiracle of certain individuals, but they are not thought to be pathogens or are not very harmful.^[186]^{: 26, 303, 359}The bacteriumErysipelothrix rhusiopathiae,which probably comes from eating infected fish, poses a threat to belugas kept in captivity, causinganorexiaanddermalplaques andlesionsthat can lead tosepsis.^[186]^{: 26, 303, 359}This condition can cause death if it is not diagnosed and treated in time withantibioticssuch asciprofloxacin.^[187]^[186]^: 316–7A study of infections caused byparasitic wormsin a number of individuals of both sexes found the presence of larvae from a species from the genusContracaecumin theirstomachsandintestines,Anisakissimplexin their stomachs,Pharurus pallasiiin their ear canals,Hadwenius seymouriin their intestines andLeucasiella arcticain their rectums.^[188]

Relationship with humans

Captivity

Belugas were among the first whale species to be kept in captivity. The first beluga was shown atBarnum's MuseuminNew York Cityin 1861.^[189]For most of the 20th century, Canada was the predominant source for belugas destined for exhibition. Throughout the early 1960s, belugas were taken from theSt. Lawrence Riverestuary. In 1967, the Churchill River estuary became the main source from which belugas were captured. This continued until 1992, when the practice was banned.^[156]Since then, Russia has become the largest provider.^[156]Individuals are caught in the Amur River delta and the far eastern seas of the country, and then are either transported domestically to aquaria inMoscow,St. PetersburgandSochi,or exported to foreign nations, including China^[147]and formerly Canada.^[156]Canada has now banned the practice of holding new animals in captivity.^[190]

Beluga aims echolocation beam to choose a spot on an acoustic "touch screen" of hydrophones in the water

To provide some enrichment while in captivity, aquaria train belugas to perform behaviours for the public^[191]and for medical exams, such as blood draws,^[192]ultrasound,^[193]providing toys,^[191]and allowing the public to play recorded or live music.^[194]

Between 1960 and 1992, theUnited States Navycarried out a program that included the study ofmarine 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 onwards.^[195]The program included training them to carry equipment and material to divers working under water, the location of lost objects, surveillance of ships andsubmarines,and underwater monitoring using cameras held in their mouths.^[195]A similar program was implemented by theSoviet Navyduring theCold War,in which belugas were also trained for antiminingoperations in Arctic waters.^[174]It is possible this program continues within theRussian Navy,as on April 24, 2019, atame beluga whale wearing a Russian equipment harnesswas found by fishermen near the Norwegian island ofIngøya.^[196]

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 theUniversity of Tromsø.^[197]

In 2019, a sanctuary in Iceland was established for two belugas, Little White and Little Grey, that retired from a marine park in China. The Sea Life Trust Beluga Whale Sanctuary was created with support fromMerlin EntertainmentsandWhale and Dolphin Conservation (WDC).Merlin bought the park in 2012, as part of an Australian chain, and it is one of their largest aquaria.^[198]Merlin has a policy against captive cetaceans, so they sponsored a 32,000-square-metre 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.^[199]^[200]The cost is variously listed as ISK 3,000,000 (US$24,000) or US$27,000,000.^[199]Merlin was owned until 2015 byBlackstone Group,which also ownedSeaWorld^[201]until selling its last stake in 2017 to a Chinese company which will use SeaWorld's expertise to expand in China;^[202]SeaWorld still keeps belugas in captivity.

Belugas are displayed across North America, Europe and Asia.^[156]As of 2006, 58 belugas were held in captivity in Canada and the United States, and 42 deaths in US captivity had been reported up to that time. A single specimen costs up to US$100,000, although the price has now dropped to US$70,000.^[203]^[204]^[156]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.^[205]^[203]^[206]The beluga's popularity with visitors reflects its attractive colour and its range offacial 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.^[49]

Most belugas found in aquaria are caught in the wild, as captive-breeding programs have not had much success so far.^[207]For example, despite best efforts, as of 2010, only two male whales had been successfully used as stud animals in theAssociation of Zoos and Aquariumsbeluga population, Nanuq atSeaWorld San Diegoand Naluark at theShedd AquariuminChicago,US. Nanuq has fathered 10 calves, five of which survived birth.^[208]Naluark at Shedd Aquarium has fathered four living offspring.^[209]Naluark was relocated to theMystic Aquariumin the hope that he would breed with two of their females,^[210]but he did not, and in 2016 he was moved to SeaWorld Orlando.^[211]The first beluga calf born in captivity in Europe was born inL'Oceanogràficmarine park inValencia,Spain, in November 2006.^[212]However, the calf died 25 days later after sufferingmetaboliccomplications, infections and not being able to feed properly.^[213]A second calf was born on 16 November 2016, and was successfully maintained by artificial feeding based on enriched milk.^[214]

In 2009 during a free-diving competition in a tank of icy water inHarbin,China, a captive beluga brought a cramp-paralysed diver from the bottom of the pool up to the surface by holding her foot in its mouth, saving the diver's life.^[215]^[216]

Films which have publicised issues of beluga welfare includeBorn to Be Free,^[217]Sonic Sea,^[218]andVancouver Aquarium Uncovered.^[219]

Whale watching

Whale watchinghas become an important activity in the recovery of the economies of towns in Quebec and Hudson Bay, near the Saint Lawrence and Churchill Rivers.^[220]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.^[221]The animals are easily seen due to their high numbers and their curiosity regarding the presence of humans.^[221]

However, the boats' presence poses a threat, 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.^[222]To protect these marine animals during whale-watching activities, the USNational Oceanic and Atmospheric Administrationhas 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.^[223]

Some regular migrations do occur into RussianEEZofSea of Japansuch as to Rudnaya Bay, where diving with wild belugas became a less-known but popular attraction.^[224]

On 25 September 2018, a beluga was sighted in theThames Estuaryand near towns along theKentside of the Thames, being nicknamedBennyby newspapers. The whale, who was noticed by conservationists to be traveling 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,^[225]and continued into 2019, when local experts concluded that Benny had left the estuary.^[226]

On 13 May 2021, two beluga whales were sighted in waters aroundPrince Edward Island,Atlantic Canada.One whale entered theCharlottetownHarbour and travelled up the Hillsborough River toMount Stewart, Prince Edward Island.As of 30 May the whale was still sighted in the area.^[227]^[228]

In August 2022, a beluga was found on the river Seine, France.^[229]

Human speech

Male belugas in captivity can mimic the pattern of human speech, several octaves lower than typical whale calls.^[230]One captive male beluga namedNOC,pronounced "No-see", caused the diver to surface by imitating orders to get out of the water. Subsequent recordings confirmed that NOC had become skilled at imitating the patterns and frequency of human speech, and intentionally altered his normal methods of vocalization to achieve these sounds. After several years, he ceased making these sounds.^[231]

Conservation status

Video explaining conservation efforts in theSea of Okhotsk,Russia

Prior to 2008, the beluga was listed as "vulnerable" by theInternational Union for Conservation of Nature(IUCN), a higher level of concern. The IUCN cited the stability of the largest sub-populations 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.^[232]In June 2017, its status was reassessed to "least concern".^[2]

There are about 21 sub-populations of beluga whales and it is estimated that 200,000 individuals still exist, which are listed as Least Concern on the IUCN Red List.^[233]However, the nonmigratory Cook Inlet sub-population off the Gulf of Alaska is a separate sub-population that is listed as "critically endangered" by the IUCN as of 2006^[2]and as "endangered" under theEndangered Species Actas of October 2008.^[234]^[235]^[236]^[2]This was primarily due to unregulatedoverharvestingof beluga whales prior to 1998. The population has remained relatively consistent, though the reported harvest has been small. As of 2016, the estimated abundance of the endangered Cook Inlet population was 293 individuals.^[237]The most recent estimate in 2018 by NOAA Fisheries suggested that the population declined to 279 individuals.^[237]

Despite beluga whales not being threatened overall, sub-populations are being listed as critically endangered and are facing increased mortality from human actions. For example, even though commercial hunting is now banned due to the Marine Mammal Protection Act, beluga whales are still being hunted to preserve the livelihood of native Alaskan communities.^[238]The IUCN and NOAA Fisheries cite habitat degradation, oil and gas drilling, underwater noise, harvesting for consumption and climate change as threats to the prolonged survival of beluga whale sub-populations.^[238]

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 sub-population they are from.^[239]Because there is little protection of sub-populations, harvest will need to be managed to ensure sub-populations 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.^[239]Changes in sea-ice has resulted in changes in the area used by Chukchi belugas, since belugas spent less time in close proximity to the ice edge in comparison to previous years. Additionally, Chukchi Sea belugas spent a prolonged amount of time in Barrow Canyon on the Beaufort Sea side in October. Chukchi sea belugas also appear to be spending more time in deeper water presently, as opposed to the 1990s. Belugas also seemed to be taking longer and deeper dives. A hypothesis as to why this might be the case is an up-welling of rich Atlantic water in the Beaufort Sea may result in concentrated prey items like Arctic cod. The fall migration of Chukchi belugas is later, although summer and fall habitat selection has not changed. Fall migration of Chukchi belugas appears to be correlated with Beaufort Sea freeze up.^[240]

It is hypothesized that beluga whales utilize ice as protection from killer whale predation or for feeding on schools of fish.^[241]Killer whales can penetrate further into the Arctic and remain in arctic waters for a longer period of time due to reductions in sea ice. For example, residents in Kotzebue, have reported that killer whales have been sighted more frequently in Kotzebue Sound.

As annual ice cover declines, humans may gain access and disrupt beluga whale habitats.^[241]For example, the number of vessels in the Arctic for gas and oil exploration, fishing, and commercial shipping has already increased and a continuous trend may lead to higher risks of injuries and deaths for beluga whales.^[241]

In addition, it is possible that beluga whales may face by an increased risk of entrapment from leads and cracks freezing, due to the erratic nature of climate change. Abrupt changes in weather can cause these leads and cracks to freeze ultimately causing the whales to die of suffocation.^[239]An increase in urbanization will likely lead to higher concentrations of toxic pollutants in the blubber of beluga whales since they are at the top of the food chain and are affected by bio-accumulation.^[241]Loss of sea ice and a change in ocean temperatures may also affect the distribution and composition of prey or affect their competition.^[241]There is also some evidence that climate change can affect males and females differently. Since 1983, belugas have been increasing scarce in Kotzebue sound. However, in 2007, several hundred whales were spotted in the sound, with over 90% of the whales being male. However, more research needs to be conducted to understand how climate change affects beluga whale sex aggregation.^[242]

Legal protection

The US Congress passed theMarine 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.^[243]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.^[243]

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 withoil and gasdevelopment 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^[244]of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). It is listed on appendix II^[244]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 theCITESthat was signed in 1973 to regulate the international import and export of certain species.^[245]

The isolated beluga population in the Saint Lawrence River has been legally protected since 1983.^[246]In 1988Canadian Department of Fisheries and OceansandEnvironment Canada,a governmental agency that supervises national parks, implemented the Saint Lawrence Action Plan^[247]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.^[248]

Conservation research in managed care facilities

As of 2015, there were 33 individuals housed in managed care facilities in North America.^[249]These facilities are members of the Association of Zoos and Aquariums, aiming to understand the complex reproductive physiology of this species to improve their conservation. With the extreme difficulty of studying beluga whales in the wild and the lack of ability to collect biological samples or perform examinations on individuals, managed care facilities play a critical role.^[250]

Managed care facilities in North America have been able to work cooperatively to build upon the research of beluga whale reproduction and have made remarkable advances. Using operant conditioning, these facilities have trained beluga whales for voluntary biological sampling and examinations. Blood,^[251]urine,^[252]and blow samples^[253]have all been collected for longitudinal hormone monitoring studies.

In addition, beluga whales have undergone semen collection,^[249]body temperature data collection,^[251]reproductive tract examinations via transabdominal ultrasound, and endoscopic exams.^[254]With new technology, the reproductive characteristics of both the female and male beluga whale have been accurately described and has benefited captive breeding programs globally.

As more research is done, the management of beluga whales in managed care facilities can be greatly improved and may even help develop other cetacean breeding and contraceptive programs, such as that of the bottlenose dolphin.^[249]Through fetal health and gestation monitoring, facilities can be more equipped to deal with pregnant animals as well.^[251]While training has been done to collect beluga whale semen, only few facilities have been able to successfully do so as both saltwater and urine contamination need to be avoided.^[255]Improvement of this process will help increase the success of captive breeding programs.

Cultural references

Pour la suite du monde,is a Canadian documentary film released in 1963 about traditional beluga hunting carried out by the inhabitants ofL'Isle-aux-Coudreson the Saint Lawrence River.^[256]

The children's singerRaffireleased an album calledBaby Belugain 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 inVancouver Aquarium.^[257]

The fuselage design of theAirbus 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.^[258]The company paints the 2019Beluga XLversion to emphasize the plane's similarity to the Beluga whale.^[259]

In the 2016Disney/Pixaranimated filmFinding Dory,the sequel toFinding Nemo(2003), the character Bailey is a beluga whale and its echolocation abilities are a significant part of the plot.^[260]^[261]

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External links

US National Marine Fisheries Service beluga whale page
Cook Inlet Beluga Population Info
ARkivePhotos
Animal Diversity Web
Video of belugas blowing bubble rings and performing other tricks at a Japanese aquariumonYouTube
Convention on Migratory Species page on the Beluga / White whale
Animals in National Geographic – Beluga WhaleDelphinapterus leucas
Lifestyles of Beluga Whales National Geographic,video
"Beluga whale".Encyclopedia of Life.Archived fromthe originalon 22 September 2009.Retrieved12 September2012.
Information about beluga in animaldiversity
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Information about belugas in marinebio.org
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