Not much is known of the roles of southern elephant seals while they are at sea. However, they are known to be important to their ecosystem as predators of fish, sharks, squid, crabs, and shrimp and as prey for large sharks, killer whales, and leopard seals. Another important role they play in the ecosystem is as a host for many kinds of parasites. Some of the known parasites include tapeworms, acanthocephalans, and the louse Lepidophthirus macrorhini.
Commensal/Parasitic Species:
Knowledge of predation of M. leonina is limited due to their deep ocean habitat. Known predators include large sharks, specifically great white sharks, and killer whales. Leopard seals are also known to prey on pups. In order to avoid predation, southern elephant seals have dark dorsal surfaces with lighter undersides. This allows some camouflage by blending in with the lighter water when viewed from below and the darker water when seen from above.
Known Predators:
Anti-predator Adaptations: cryptic
Male southern elephant seals are the largest pinnipeds, larger even than northern elephant seals, Mirounga angustirostris, their closest relatives. Mirounga leonina males have been documented reaching over six meters long and weighing over 4000 kg. This is in sharp contrast to females, which are rarely over 800 kg or four meters long. In fact, both species of the genus Mirounga are more sexually dimorphic than any other mammal. This dimorphism stretches beyond just size. Males also have a large, inflatable proboscis, which enhances vocalizations used to challenge other males for mating rights. The southern elephant seal proboscis is slightly smaller than the proboscis of northern elephant seals, overhanging the mouth by only about 10 centimeters compared to 30 centimeters in their northern relatives.
Breeding populations vary in size. In the South Georgian population, males average 450 cm in length and weigh 4,000 kg. Females average 280 cm and weigh 900 kg. The seals from Macquarie Island population are somewhat smaller, with males averaging 420 cm in length and 3,000 kg and females averaging 260 cm and 400 kg.
Despite the large difference in size, male and female southern elephant seals do share many physical traits. They have a similar body type. This includes short front flippers used primarily for steering in the water, and very strong, fully webbed, rear flippers that can propel them through the water with remarkable speed and agility. They also have a layer of short, stiff hair covering their bodies. At birth this fur is very dark in color, but lightens after the first molt. New fur after a molt is typically a dark gray/brown with lighter underside and lightens over the course of the year. It is also common for the bodies of both sexes to have scars, usually around the neck, from fighting and mating.
Range mass: 300 to 4000 kg.
Average mass: 2000 (male) 500 (female) kg.
Range length: 260 to 600 cm.
Average length: 500 (male) 300 (female) cm.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger; sexes shaped differently; ornamentation
There is little known about lifespan in southern elephant seals. This is largely due to the lack of substantial information concerning the periods of the year when they are at sea. Average life expectancy in the wild, as seen during the breeding season, is about 23 years. However, about 30% of pups die in their first year. Captive M. leonina have lived to 15 years of age. Not much is known concerning the deaths of these mammals but, in addition to predation, weather and disease may play a large role in limiting their lifespan.
Range lifespan
Status: captivity: 15 (high) years.
Average lifespan
Status: wild: 23 years.
Average lifespan
Sex: male
Status: wild: 20.0 years.
Average lifespan
Sex: female
Status: wild: 18.0 years.
Average lifespan
Status: wild: 23.0 years.
Average lifespan
Status: wild: 20.0 years.
When southern elephant seals are on land, they are typically found along the coast of sub-Antarctic islands on smooth beaches of sand or small rocks. Although they used to breed well into temperate regions, M. leonina are now only found farther south. They are found on land during the breeding season, from August to November, and the molting season, which lasts 3 to 5 weeks in the spring. The rest of the year is spent entirely at sea. During this time they can be found from sub-Antarctic waters to almost as far north as the equator, often venturing thousands of kilometers from their breeding grounds. While males typically forage on the Antarctic continental shelf, females travel farther into open waters. During their time at sea, southern elephant seals can sustain dives for up to two hours, but most dives last only around thirty minutes. Amazingly, they only spend 2 to 3 minutes on the surface between dives. During most trips at sea, they are underwater for 90% of the time, day and night. While most dives are only between 300 and 800 m, dives of over 1500 m have been recorded, nearing depths only surpassed in mammals by sperm whales.
Range depth: 1500+ to surface m.
Average depth: 300-800 m.
Habitat Regions: polar ; terrestrial ; saltwater or marine
Aquatic Biomes: pelagic ; coastal
Other Habitat Features: intertidal or littoral
Mirounga leonina (southern elephant seals) are found along the coast of Antarctica and on sub-Antarctic islands when breeding or molting. However, before human exploitation they were more common farther north. The largest present population occurs on the island of South Georgia, in the South Atlantic Ocean. Southern elephant seals are also common on Macquarie Island, Heard Island, Kerguelen Island, and the Peninsula Valdez in Argentina. When at sea, M. leonina often journey thousands of miles from their breeding grounds. Despite the occasional sighting, not much is known about their range outside of the breeding season.
Biogeographic Regions: antarctica (Native ); oceanic islands (Native ); indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native )
Southern elephant seals feed exclusively when they are at sea. For this reason not much is known about what they eat. The main known sources of food are squid, crabs, shrimp, fish, and sharks. This prey is obtained both near the surface and also during very deep dives. They have been known to eat bottom dwelling fish.
Animal Foods: fish; mollusks; aquatic crustaceans
Primary Diet: carnivore (Eats non-insect arthropods, Molluscivore )
In the past, southern elephant seals were hunted for their blubber that was boiled down into oil. A typical male could produce about 350 liters of oil. Some aboriginal people also hunted them for food and skins. This activity has ceased and killing is now controlled by the Convention for the Conservation of Antarctic Seals. The only use of M. leonina to man today is for purely scientific purposes.
Positive Impacts: body parts are source of valuable material; research and education
Southern elephant seals may occasionally compete with some fisheries, but this is unlikely. Southern elephant seals live in remote regions where they have few interactions with humans.
When at sea, Mirounga leonina rarely encounter each other and thus have no need for communication. The only time communication is used is during breeding. Males use their large proboscis as a sound chamber for amplifying their bellows. These sounds are made to establish territories and challenge males for established harems. Upright posturing often accompanies these vocalizations and males are known to visually assess their competitor before fighting. Lesser males will also exhibit a flattened posture without inflating their proboscis when near another male’s harem to demonstrate that they are not threats.
A threat vocalization is a low-pitched harsh vocalization. While the seal is doing this it will raise its head and forequarters off of the ground, supporting itself without fore flippers. A lunge from an animal is a rapid movement of the head towards an opponent or invader. This is done with an open mouth. A high rear is the raising of the front half of the body then delivering blows to another animal with the neck or chin. A bite may also be used, mainly from a low rear or a high rear position.
Females are known to communicate with newborn pups through vocalizations. Females and pups recognize each other through these vocal cues and through their individual smells.
Communication Channels: visual ; acoustic ; chemical
Perception Channels: visual ; tactile ; acoustic ; chemical
During the Quaternary, it is thought that elephant seals spread north across the equator from the southern oceans. However, as ice ages ended, northern populations were separated from southern populations. The rewarming of the ocean waters forced both populations in different directions. Now these two populations have been separated by 8,000 km for thousands of years, forming two different species; M. angustirostris, northern elephant seals, and southern elephant seals, M. leonina.
Although once hunted by humans, southern elephant seals were never near extinction like northern elephant seals. This is largely because most of the breeding grounds of M. leonina were out of reach of hunting boats. Hunting did have some impact, but numbers have recovered since hunting has ceased. Some populations are are experiencing declines. This may be normal population fluctuations, however.
US Federal List: no special status
CITES: appendix ii
IUCN Red List of Threatened Species: least concern
Male southern elephant seals arrive at breeding grounds several weeks before females and, through vocalizations, body positions, and occasional fighting, claim territories on the beach. The best and largest territories go to the largest and strongest males. These “alpha” males become the head of a harem when the females arrive, often mating with up to 60 females in their harem. If harems exceed this size, additional “beta” males may be present, each claiming as many females as they can. Females become a part of a harem simply through their position on the beach and may move from one harem to another incidentally.
In addition to their mating duties, alpha males are responsible for keeping unwanted males away from the harems. This is done through the same vocalizations and aggressive body postures that were used originally to claim their harem. Males must remain on their territory to defend it and, therefore, go for periods of months without eating. This, and the stress of aggressive encounters with other males and the energy expense of mating with multiple females, can take a significant toll on male physical condition. Only males in the best physical condition at the beginning of breeding season will successfully defend their territory and breed with multiple females. Subordinate males attempt to copulate with females on the edges of territories or in the surf as they leave the beach.
Females that were pregnant from the previous year’s mating give birth to one pup shortly after arriving on land. A period of lactation follows the birth. Then, several days before the pups are weaned from their mother’s milk, females enter estrus and mate with the alpha male or a successful beta male. Shortly following mating, males return to sea. Females return to the sea immediately after the pups are weaned.
Mating System: polygynous
Once a year, from August to November, southern elephant seals return to land to breed. Amazingly, most return to the very same breeding grounds on which they were born. Five to seven days after pregnant females arrive on the beaches, they give birth to one pup. Occasionally twin pups are born but one typically dies soon afterwards. The mothers then nurse their young for about 23 days. Females may nurse longer if their energy reserves allow them to do so. During their time on the breeding grounds females eat little or not at all. Towards the end of this time, females enter estrus and mate with a male. Shortly after mating, females wean their young. At this point, they abandon their young and return to the ocean. Pups then forage on their own for several weeks before venturing out to sea in small groups. Female southern elephant seals typically reach sexual maturity by the age of 3 and participate in the annual breeding cycle by age 6. Males reach sexual maturity by age 5 or 6, but rarely are developed enough to compete for mates until they reach 10 to 12 years of age. The gestation period of female M. leonina is about eight months. There is a period of several weeks during late October when all mature females mate. In order to maintain the yearly birthing cycle with an eight-month gestation period, there is delayed implantation of the fertilized egg for about three months. After the three-month delay, the egg implants and begins to develop to become mature enough for birth during the next breeding season.
Breeding interval: Southern elephant seals mate once each year.
Breeding season: Breeding season lasts from August to November.
Range number of offspring: 1 to 2.
Average number of offspring: 1.
Range gestation period: 9 to 7 months.
Range weaning age: 20 to 35 days.
Average weaning age: 23 days.
Range time to independence: 20 to 25 days.
Average time to independence: 23 days.
Range age at sexual or reproductive maturity (female): 2 to 4 years.
Range age at sexual or reproductive maturity (male): 5 to 6 years.
Average age at sexual or reproductive maturity (male): 6 years.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization ; viviparous ; delayed implantation
Average birth mass: 42500 g.
Average number of offspring: 1.
Female southern elephant seals are the sole caregivers for their young from the moment of conception until weaning, a period that lasts around one year. After delayed implantation, which follows mating, the nine-month gestation period of the pregnancy begins. During this time, the pup develops inside the mother as she is diving and feeding in sub-Antarctic waters. Shortly after coming to land, females give birth to their pups, typically weighing between 25 and 50 kg at birth. Following birth, mothers bond vocally and through smell with their pup. For the next 20 to 25 days (sometimes as long as 35 days) mothers are responsible for providing milk and protecting pups. Mothers are typically less than one-meter from their pups during the stage of suckling, regardless of tide, the position in the harem, or the time in the breeding season. A pup might get separated from its mother due to male harassment and herding of females. This can result in an abandoned pup. Once a pup is separated from its mother the results are fatal. Alien suckling (nursing between unrelated cows and pups) isn't tolerated in this species. If an orphan pup attempts to steal milk from a sleeping or resting cow, it usually is bitten and will succumb to starvation or the effects of the bites. The most dire threat to young pups is adult males who crush pups as they travel and fight on beach territories. During lactation, mothers do not return to the water to feed and instead live on fat reserves built up during the previous foraging season. At weaning pups weigh from 120 to 130 kg, a weight gain of as much as 105 kg in a few weeks!
Immediately following weaning, female southern elephant seals return to sea, leaving their pups alone on the beach. Eventually the pups begin to get hungry and find their way to the ocean, learning to feed and swim on their own. After weaning, there is no interaction between parents and pups. Approximately 30% of these pups will not live through their first year.
Parental Investment: no parental involvement; precocial ; female parental care ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female)
The southern elephant seal (Mirounga leonina) is one of two species of elephant seals. It is the largest member of the clade Pinnipedia and the order Carnivora, as well as the largest extant marine mammal that is not a cetacean. It gets its name from its massive size and the large proboscis of the adult male, which is used to produce very loud roars, especially during the breeding season. A bull southern elephant seal is about 40% heavier than a male northern elephant seal (Mirounga angustirostris), twice as heavy as a male walrus (Odobenus rosmarus),[6][7] and 6–7 times heavier than the largest living mostly-terrestrial carnivoran, the polar bear (Ursus maritimus) and the Kodiak bear (Ursus arctos middendorffi),[8][9].
The southern elephant seal was one of the many species originally described by Swedish zoologist Carl Linnaeus in the landmark 1758 10th edition of his Systema Naturae, where it was given the binomial name of Phoca leonina.[3] John Edward Gray established the genus Mirounga in 1827.[10]
In the nineteenth century the species was often called "bottle-nosed seal".[11]
The southern elephant seal is distinguished from the northern elephant seal (which does not overlap in range with this species) by its greater body mass and a shorter proboscis. The southern males also appear taller when fighting, due to their tendency to bend their backs more strongly than the northern species. This species may also exhibit the greatest sexual dimorphism of any mammal in terms of mass ratio, with males typically five to six times heavier than females.[12] On average female southern elephant seals weigh 350 to 900 kg (770 to 1,980 lb) and measure 2.6 to 3 m (8.5 to 9.8 ft) long, whereas bulls can range from 1,500 to 3,700 kg (3,300 to 8,200 lb) and grow to 4.2 to 5.8 m (14 to 19 ft) in length.[13][14] For comparison, among the northern elephant seal and the sperm whale (Physeter macrocephalus)—two other large marine mammals that are highly sexual dimorphic by size—males typically outweigh females by a factor of three to four.[15]
Southern elephant seal size also varies regionally. Studies have indicated elephant seals from South Georgia are around 30% heavier and 10% longer on average than those from Macquarie Island.[12] The record-sized bull, shot in Possession Bay, South Georgia, on 28 February 1913, measured 6.85 m (22.5 ft) long and was estimated to weigh 5,000 kg (11,000 lb), although it was only partially weighed piecemeal.[9][16] The maximum size of a female is 1,000 kg (2,200 lb) and 3.7 m (12 ft).[9][17]
A southern elephant seal's eyes are large, round, and black. The width of the eyes, and a high concentration of low-light pigments, suggest sight plays an important role in the capture of prey. Like all seals, elephant seals have hind limbs whose ends form the tail and tail fin. Each of the "feet" can deploy five long, webbed fingers. This agile dual palm is used to propel water. The pectoral fins are used little while swimming. While their hind limbs are unfit for locomotion on land, elephant seals use their fins as support to propel their bodies. They are able to propel themselves quickly (as fast as 8 km/h (5.0 mph)) in this way for short-distance travel, to return to water, to catch up with a female, or to chase an intruder.
Pups are born with fur and are completely black. Their coats are unsuited to water, but protect infants by insulating them from the cold air. The first moulting accompanies weaning. After moulting, the coats may turn grey and brown, depending on the thickness and moisture of hair. Among older males, the skin takes the form of a thick leather which is often scarred.
Like other seals, the vascular system of elephant seals is adapted to the cold; a mixture of small veins surround arteries, capturing heat from them. This structure is present in extremities such as the hind legs.
The world population was estimated at 650,000 animals in the mid-1990s,[1] and was estimated in 2005 at between 664,000 and 740,000 animals.[18] Studies have shown the existence of three geographic subpopulations, one in each of the three oceans.
Tracking studies have indicated the routes traveled by elephant seals, demonstrating their main feeding area is at the edge of the Antarctic continent. While elephant seals may come ashore in Antarctica occasionally to rest or to mate, they gather to breed in subantarctic locations.
The largest subpopulation is in the South Atlantic, with more than 400,000 individuals, including about 113,000 breeding females on South Georgia;[19] the other breeding colonies of the Atlantic subpopulation are located on the Falkland Islands and Valdes Peninsula in Argentina (the only continental breeding population).
The second subpopulation, in the south Indian Ocean, consists of up to 200,000 individuals, three-quarters of which breed in the Kerguelen Islands and the rest in the Crozet Islands, Marion and Prince Edward Islands, and Heard Island. Some individuals also breed on Amsterdam Island.
The third subpopulation of about 75,000 seals is found in the subantarctic islands of the Pacific Ocean south of Tasmania and New Zealand, mainly Macquarie Island.
Colonies once existed in Tasmania, Saint Helena, and the Juan Fernández Islands off the coast of Chile. Some individuals at the time of moulting have been found in South Africa, Australia or Uruguay. Lost animals have also been reported from time to time on the shores of Mauritius, with two reports from the Río Guayas estuary area in Ecuador[18] and a beach in Lima, Peru.[20] Reality of the creature so called Manatee of Helena had been pointed out as possible misidentification of elephant seals historically present on Saint Helena.[21]
After the end of large-scale seal hunting in the 19th century, the southern elephant seal recovered to a sizable population in the 1950s; since then, an unexplained decline in the subpopulations of the Indian Ocean and Pacific Ocean has occurred. The population now seems to be stable; the reasons for the fluctuation are unknown. Suggested explanations include a phenomenon of depression following a rapid demographic rebound that depletes vital resources, a change in climate, competition with other species whose numbers also varied, or even an adverse influence of scientific monitoring techniques.[22]
Elephant seals are among the seals that can stay on land for the longest periods of time, as they can stay dry for several consecutive weeks each year. Males arrive in the colonies earlier than the females and fight for control of harems when they arrive.[23] Large body size confers advantages in fighting, and the agonistic relationships of the bulls gives rise to a dominance hierarchy, with access to harems and activity within harems being determined by rank.[24] The dominant bulls (“harem masters”) establish harems of several dozen females. The least successful males have no harems, but they may try to copulate with a harem male's females when the male is not looking. The majority of primiparous females and a significant proportion of multiparous females mate at sea with roaming males away from harems.[25]
An elephant seal must stay in his territory to defend it, which could mean months without eating, having to live on his blubber storage. Two fighting males use their weight and canine teeth against each other. The outcome is rarely fatal, and the defeated bull will flee; however, bulls can suffer severe tears and cuts. Some males can stay ashore for more than three months without food. Males commonly vocalize with a coughing roar that serves in both individual recognition and size assessment.[24] Conflicts between high-ranking males are more often resolved with posturing and vocalizing than with physical contact.[24]
Generally, pups are born rather quickly in the breeding season.[26] After being born, a newborn will bark or yap and its mother will respond with a high-pitched moan.[27] The newborn begins to suckle immediately. Lactation lasts an average of 23 days. Throughout this period, the female fasts. Newborns weigh about 40 kg (88 lb) at birth, and reach 120 to 130 kg (260 to 290 lb) by the time they are weaned. The mother loses significant weight during this time. Young weaned seals gather in nurseries until they lose their birth coats. They enter the water to practice swimming, generally starting their apprenticeship in estuaries or ponds. In summer, the elephant seals come ashore to moult. This sometimes happens directly after reproduction.
Satellite tracking revealed the seals spend very little time on the surface, usually a few minutes for breathing. They dive repeatedly, each time for more than 20 minutes, to hunt their prey—squid and fish—at depths of 400 to 1,000 m (1,300 to 3,300 ft). They are the deepest diving air-breathing non-cetaceans and have been recorded at a maximum of 2,388 m (7,835 ft) in depth.[28][29]
As for the duration, depth and sequence of dives, the southern elephant seal is the best performing seal. In many regards, they exceed even most cetaceans. These capabilities result from nonstandard physiological adaptations, common to marine mammals, but particularly developed in elephant seals. The coping strategy is based on increased oxygen storage and reduced oxygen consumption.
In the ocean, the seals apparently live alone. Most females dive in pelagic zones for foraging, while males dive in both pelagic and benthic zones.[30] Individuals will return annually to the same hunting areas. Due to the inaccessibility of their deep-water foraging areas, no comprehensive information has been obtained about their dietary preferences, although some observation of hunting behavior and prey selection has occurred.[31]
While hunting in the dark depths, elephant seals seem to locate their prey, at least in part, using vision; the bioluminescence of some prey animals can facilitate their capture. Elephant seals do not have a developed system of echolocation in the manner of cetaceans, but their vibrissae (facial whiskers), which are sensitive to vibrations, are assumed to play a role in search of food. When at the subantarctic or Antarctic coasts, the seals forage largely on deep-sea cephalopod species such as Psychroteuthis glacialis, Alluroteuthis antarcticus, Histioteuthis eltaninae, Onykia ingens, Gonatus antarcticus. Martialia hyadesi[32][33] and other molluscs, various fish species, including lanternfish (i.e. Electrona spp. and Gymnoscopelus spp.), nothothens (i.e. Genera Lepidonotothen, Pleuragramma, Trematomus, Pagothenia,), Channichthyidsae spp., Bathylagidae spp.,[34] krill (mostly Euphausia spp.) and other crustaceans, and even algae.
Weaned pups and juveniles may fall prey to orcas.[35] Cases where weaned pups have been attacked and killed by leopard seals (Hydrurga leptonyx) and New Zealand sea lions (Phocarctos hookeri), exclusively small pups in the latter case, have been recorded. Great white sharks (Carcharodon carcharias) have hunted elephant seals near Campbell Island, while bite marks from a southern sleeper shark (Somniosus antarcticus) have been found on surviving elephant seals in the Macquarie Islands.[36][37]
After their near extinction due to hunting in the 19th century, the total population was estimated at between 664,000 and 740,000 animals in 2005,[18] but as of 2002, two of the three major populations were declining.[38] The reasons for this are unclear, but are thought to be related to the distribution and declining levels of the seals' primary food sources.[38] Most of their important breeding sites are now protected by international treaty, as UNESCO World Heritage Sites, or by national legislation.
Climate change likely has a wide-ranging and diverse impact on marine species such as southern elephant seals. As top predators in the Southern Ocean, southern elephant seals inhabit one of the most sensitive and vulnerable regions to rapid climate change. Global efforts such as the Southern elephant seals as oceanographic samplers[39] and the Marine Mammals Exploring the Oceans Pole to Pole projects[40] have led to the collection of a large suite of long-term coupled behavioural-oceanographic data. As part of these projects, satellite-relay data loggers were attached to southern elephant seals to collect physical and biological data. This simultaneous collection of behavioural and environmental data, spanning more than ten years, has enabled researchers to study the impact of climate change on southern elephant seals and the Antarctic ecosystem. This data provides information on how the Southern Ocean is changing in relation to climate change as well as how these species respond to changes.[41][42]
These projects focused on southern elephant seals because they are deep divers as well as a major predator in the Southern Ocean in terms of their population size and food consumption. As predators of the upper levels of the food-web, their foraging success and population dynamics impart valuable information about productivity at different trophic levels.[43] The well-being of southern elephant seals therefore mirrors that of the whole Antarctic ecosystem.
Southern elephant seals from different colonies frequent specific oceanographic regions to forage. They feed most successfully in areas with specific hydrographic properties, e.g. the upwelling regions of Circumpolar Deep Water within the Antarctic Circumpolar Current.[44] In the Southern Ocean, southern elephant seals associate more frequently with southerly, higher‐latitude fronts and frontal zones. However, the foraging success in association with these regions varies strongly according to year, season and sex. Some of the seasonal and interannual variations in foraging success can be linked to climatic changes such as positional shifts in fronts and variability associated with frontal positions.[45] Generally, southern elephant seals appear to be resilient against the apparent variability in the location and productivity of frontal systems.
Distant, William Lucas (September 1899). "The "Sea-Elephant" (Macrorhinus elephantinus)". The Zoologist. 4th series, vol 3 (699): 385/7. The southern elephant seal (Mirounga leonina) is one of two species of elephant seals. It is the largest member of the clade Pinnipedia and the order Carnivora, as well as the largest extant marine mammal that is not a cetacean. It gets its name from its massive size and the large proboscis of the adult male, which is used to produce very loud roars, especially during the breeding season. A bull southern elephant seal is about 40% heavier than a male northern elephant seal (Mirounga angustirostris), twice as heavy as a male walrus (Odobenus rosmarus), and 6–7 times heavier than the largest living mostly-terrestrial carnivoran, the polar bear (Ursus maritimus) and the Kodiak bear (Ursus arctos middendorffi),.
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