Sea otter
Sea otter | |
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In Morro Bay, California | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Family: | Mustelidae |
Subfamily: | Lutrinae |
Genus: | Enhydra |
Species: | E. lutris
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Binomial name | |
Enhydra lutris | |
Subspecies | |
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Sea otter range | |
Synonyms | |
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The sea otter (Enhydra lutris) is a marine mammal native to the coasts of the northern and eastern North Pacific Ocean. Adult sea otters typically weigh between 14 and 45 kg (30 and 100 lb), making them the heaviest members of the weasel family, but among[3] the smallest marine mammals. Unlike most marine mammals, the sea otter's primary form of insulation is an exceptionally thick coat of fur, the densest in the animal kingdom. Although it can walk on land, the sea otter is capable of living exclusively in the ocean.
The sea otter inhabits nearshore environments, where it dives to the sea floor to
Sea otters, whose numbers were once estimated at 150,000–300,000, were hunted extensively for their fur between 1741 and 1911, and the world population fell to 1,000–2,000 individuals living in a fraction of their historic range.[5] A subsequent international ban on hunting, sea otter conservation efforts, and reintroduction programs into previously populated areas have contributed to numbers rebounding, and the species occupies about two-thirds of its former range. The recovery of the sea otter is considered an important success in marine conservation, although populations in the Aleutian Islands and California have recently declined or have plateaued at depressed levels. For these reasons, the sea otter remains classified as an endangered species.
Evolution
The sea otter is the heaviest (the
Taxonomy
Cladogram showing relationships between sea otters and other otters[17][18] |
The first scientific description of the sea otter is contained in the field notes of
Subspecies
Three subspecies of the sea otter are recognized with distinct geographical distributions. Enhydra lutris lutris (
Description
The sea otter is one of the smallest marine mammal species, but it is the heaviest mustelid.[8] Male sea otters usually weigh 22 to 45 kg (49 to 99 lb) and are 1.2 to 1.5 m (3 ft 11 in to 4 ft 11 in) in length, though specimens up to 54 kg (119 lb) have been recorded.[26] Females are smaller, weighing 14 to 33 kg (31 to 73 lb) and measuring 1.0 to 1.4 m (3 ft 3 in to 4 ft 7 in) in length.[27] The average weight for adult sea otters that are in more densely populated areas, at 28.3 kg (62 lb) in males and 21.1 kg (47 lb) in females, was considerably lighter than the average weight of otters in more sparse populations, at 39.3 kg (87 lb) in males and 25.2 kg (56 lb) in females[5] Presumably less populous otters are more able to monopolize food sources,[5] For its size, the male otter's baculum is very large, massive and bent upwards, measuring 150 mm (5+7⁄8 in) in length and 15 mm (9⁄16 in) at the base.[28]
Unlike most other marine mammals, the sea otter has no
The sea otter displays numerous adaptations to its marine environment. The nostrils and small ears can close.[35] The hind feet, which provide most of its propulsion in swimming, are long, broadly flattened, and fully webbed.[36] The fifth digit on each hind foot is longest, facilitating swimming while on its back, but making walking difficult.[37] The tail is fairly short, thick, slightly flattened, and muscular. The front paws are short with retractable claws, with tough pads on the palms that enable gripping slippery prey.[38] The bones show osteosclerosis, increasing their density to reduce buoyancy.[39]
The sea otter presents an insight into the evolutionary process of the mammalian invasion of the aquatic environment, which has occurred numerous times over the course of mammalian evolution.[40] Having only returned to the sea about 3 million years ago,[41] sea otters represent a snapshot at the earliest point of the transition from fur to blubber. In sea otters, fur is still advantageous, given their small nature and division of lifetime between the aquatic and terrestrial environments.[42] However, as sea otters evolve and adapt to spending more and more of their lifetimes in the sea, the convergent evolution of blubber suggests that the reliance on fur for insulation would be replaced by a dependency on blubber. This is particularly true due to the diving nature of the sea otter; as dives become lengthier and deeper, the air layer's ability to retain heat or buoyancy decreases,[31] while blubber remains efficient at both of those functions.[42] Blubber can also additionally serve as an energy source for deep dives,[43] which would most likely prove advantageous over fur in the evolutionary future of sea otters.
The sea otter propels itself underwater by moving the rear end of its body, including its tail and hind feet, up and down,
Long, highly sensitive
An adult's 32
Behavior
The sea otter is diurnal. It has a period of foraging and eating in the morning, starting about an hour before sunrise, then rests or sleeps in mid-day.[60] Foraging resumes for a few hours in the afternoon and subsides before sunset, and a third foraging period may occur around midnight.[60] Females with pups appear to be more inclined to feed at night.[60] Observations of the amount of time a sea otter must spend each day foraging range from 24 to 60%, apparently depending on the availability of food in the area.[61]
Sea otters spend much of their time grooming, which consists of cleaning the fur, untangling knots, removing loose fur, rubbing the fur to squeeze out water and introduce air, and blowing air into the fur. To casual observers, it appears as if the animals are scratching, but they are not known to have
Foraging
The sea otter hunts in short dives, often to the
Under each foreleg, the sea otter has a loose pouch of skin that extends across the chest. In this pouch (preferentially the left one), the animal stores collected food to bring to the surface. This pouch also holds a rock, unique to the otter, that is used to break open shellfish and clams.[64] At the surface, the sea otter eats while floating on its back, using its forepaws to tear food apart and bring it to its mouth. It can chew and swallow small mussels with their shells, whereas large mussel shells may be twisted apart.[65] It uses its lower incisor teeth to access the meat in shellfish.[66] To eat large sea urchins, which are mostly covered with spines, the sea otter bites through the underside where the spines are shortest, and licks the soft contents out of the urchin's shell.[65]
The sea otter's use of rocks when hunting and feeding makes it one of the few mammal species to use tools.[67] To open hard shells, it may pound its prey with both paws against a rock on its chest. To pry an abalone off its rock, it hammers the abalone shell using a large stone, with observed rates of 45 blows in 15 seconds.[27] Releasing an abalone, which can cling to rock with a force equal to 4,000 times its own body weight, requires multiple dives.[27]
Social structure
Although each adult and independent juvenile forages alone, sea otters tend to rest together in single-sex groups called rafts. A raft typically contains 10 to 100 animals, with male rafts being larger than female ones.[69] The largest raft ever seen contained over 2000 sea otters. To keep from drifting out to sea when resting and eating, sea otters may wrap themselves in kelp.[70]
A male sea otter is most likely to mate if he maintains a breeding territory in an area that is also favored by females.[71] As autumn is the peak breeding season in most areas, males typically defend their territory only from spring to autumn.[71] During this time, males patrol the boundaries of their territories to exclude other males,[71] although actual fighting is rare.[69] Adult females move freely between male territories, where they outnumber adult males by an average of five to one.[71] Males that do not have territories tend to congregate in large, male-only groups,[71] and swim through female areas when searching for a mate.[72]
The species exhibits a variety of vocal behaviors. The cry of a pup is often compared to that of a
Reproduction and life cycle
Sea otters are polygynous: males have multiple female partners, typically those that inhabit their territory. If no territory is established, they seek out females in estrus. When a male sea otter finds a receptive female, the two engage in playful and sometimes aggressive behavior. They bond for the duration of estrus, or 3 days. The male holds the female's head or nose with his jaws during copulation. Visible scars are often present on females from this behavior.[6][76]
Births occur year-round, with peaks between May and June in northern populations and between January and March in southern populations.
Birth usually takes place in the water and typically produces a single pup weighing 1.4 to 2.3 kilograms (3 lb 1 oz to 5 lb 1 oz).[79] Twins occur in 2% of births; however, usually only one pup survives.[6] At birth, the eyes are open, ten teeth are visible, and the pup has a thick coat of baby fur.[80] Mothers have been observed to lick and fluff a newborn for hours; after grooming, the pup's fur retains so much air, the pup floats like a cork and cannot dive.[81] The fluffy baby fur is replaced by adult fur after about 13 weeks.[19]
Females perform all tasks of feeding and raising offspring, and have occasionally been observed caring for orphaned pups.[87] Much has been written about the level of devotion of sea otter mothers for their pups – a mother gives her infant almost constant attention, cradling it on her chest away from the cold water and attentively grooming its fur.[88] When foraging, she leaves her pup floating on the water, sometimes wrapped in kelp to keep it from floating away;[89] if the pup is not sleeping, it cries loudly until she returns.[90] Mothers have been known to carry their pups for days after the pups' deaths.[82]
Females become sexually mature at around three or four years of age and males at around five; however, males often do not successfully breed until a few years later.
Population and distribution
Sea otters live in coastal waters 15 to 23 metres (49 to 75 ft) deep,
The sea otter population is thought to have once been 150,000 to 300,000,
In about two-thirds of its former range, the species is at varying levels of recovery, with high population densities in some areas and threatened populations in others. Sea otters currently have stable populations in parts of the Russian east coast, Alaska, British Columbia, Washington, and California, with reports of recolonizations in Mexico and Japan.[102] Population estimates made between 2004 and 2007 give a worldwide total of approximately 107,000 sea otters.[19][103][104][105][106]
Japan
Adele Ogden wrote in The California Sea Otter Trade that western sea otter were hunted "from Yezo northeastward past the Kuril Group and Kamchatka to the Aleutian Chain".[101] "Yezo" refers to the island province of Hokkaido, in northern Japan, where the country's only confirmed population of western sea otter resides.[1] Sightings have been documented in the waters of Cape Nosappu, Erimo, Hamanaka and Nemuro, among other locations in the region. [107]
Russia
Currently, the most stable and secure part of the western sea otter's range is along the
British Columbia
Along the North American coast south of Alaska, the sea otter's range is discontinuous. A remnant population survived off Vancouver Island into the 20th century, but it died out despite the 1911 international protection treaty, with the last sea otter taken near
United States
Alaska
Alaska is the central area of the sea otter's range. In 1973, the population in Alaska was estimated at between 100,000 and 125,000 animals.[113] By 2006, though, the Alaska population had fallen to an estimated 73,000 animals.[104] A massive decline in sea otter populations in the Aleutian Islands accounts for most of the change; the cause of this decline is not known, although orca predation is suspected.[114] The sea otter population in Prince William Sound was also hit hard by the Exxon Valdez oil spill, which killed thousands of sea otters in 1989.[63]
Washington
In 1969 and 1970, 59 sea otters were translocated from
In Washington, sea otters are found almost exclusively on the outer coasts. They can swim as close as six feet off shore along the Olympic coast. Reported sightings of sea otters in the San Juan Islands and Puget Sound almost always turn out to be North American river otters, which are commonly seen along the seashore. However, biologists have confirmed isolated sightings of sea otters in these areas since the mid-1990s.[19]
Oregon
The last native sea otter in Oregon was probably shot and killed in 1906. In 1970 and 1971, a total of 95 sea otters were transplanted from
California
The historic population of California sea otters was estimated at 16,000 before the fur trade decimated the population, leading to their assumed extinction. Today's population of California sea otters are the descendants of a single colony of about 50 sea otters located near
Sea otters were once numerous in
In the late 1980s, the USFWS relocated about 140 southern sea otters to San Nicolas Island in southern California, in the hope of establishing a reserve population should the mainland be struck by an oil spill. To the surprise of biologists, the majority of the San Nicolas sea otters swam back to the mainland.[128] Another group of twenty swam 74 miles (119 km) north to San Miguel Island, where they were captured and removed.[129] By 2005, only 30 sea otters remained at San Nicolas,[130] although they were slowly increasing as they thrived on the abundant prey around the island.[128] The plan that authorized the translocation program had predicted the carrying capacity would be reached within five to 10 years.[131] The spring 2016 count at San Nicolas Island was 104 sea otters, continuing a 5-year positive trend of over 12% per year.[132] Sea otters were observed twice in Southern California in 2011, once near Laguna Beach and once at Zuniga Point Jetty, near San Diego. These are the first documented sightings of otters this far south in 30 years.[133]
When the USFWS implemented the translocation program, it also attempted, in 1986, to implement "zonal management" of the Californian population. To manage the competition between sea otters and fisheries, it declared an "otter-free zone" stretching from
Although the southern sea otter's range has continuously expanded from the remnant population of about 50 individuals in
For southern sea otters to be considered for removal from threatened species listing, the
Mexico
Historian Adele Ogden described sea otters are particularly abundant in "Lower California", now the
Ecology
Diet
High energetic requirements of sea otter metabolism require them to consume at least 20% of their body weight a day.[31] Surface swimming and foraging are major factors in their high energy expenditure due to drag on the surface of the water when swimming and the thermal heat loss from the body during deep dives when foraging.[149][31] Sea otter muscles are specially adapted to generate heat without physical activity.[150]
Sea otters consume over 100 prey species.
In a few northern areas, fish are also eaten. In studies performed at
The individuals within a particular area often differ in their foraging methods and prey types, and tend to follow the same patterns as their mothers.[159] The diet of local populations also changes over time, as sea otters can significantly deplete populations of highly preferred prey such as large sea urchins, and prey availability is also affected by other factors such as fishing by humans.[19] Sea otters can thoroughly remove abalone from an area except for specimens in deep rock crevices,[160] however, they never completely wipe out a prey species from an area.[161] A 2007 Californian study demonstrated, in areas where food was relatively scarce, a wider variety of prey was consumed. Surprisingly, though, the diets of individuals were more specialized in these areas than in areas where food was plentiful.[128]
As a keystone species
Sea otters are a classic example of a
Reintroduction of sea otters to British Columbia has led to a dramatic improvement in the health of coastal ecosystems,
Sea otters affect rocky ecosystems that are dominated by mussel beds by removing mussels from rocks. This allows space for competing species and increases species diversity.[165]
Predators
Leading mammalian predators of this species include
Relationship with humans
Fur trade
Sea otters have the thickest fur of any mammal, which makes them a common target for many hunters. Archaeological evidence indicates that for thousands of years,
In the early 18th century, Russians began to hunt sea otters in the Kuril Islands
Russian fur-hunting expeditions soon depleted the sea otter populations in the Commander Islands, and by 1745, they began to move on to the
Other nations joined in the hunt in the south. Along the coasts of what is now Mexico and
Russian hunting expanded to the south, initiated by American ship captains, who subcontracted Russian supervisors and Aleut hunters[177] in what are now Washington, Oregon, and California. Between 1803 and 1846, 72 American ships were involved in the otter hunt in California, harvesting an estimated 40,000 skins and tails, compared to only 13 ships of the Russian-American Company, which reported 5,696 otter skins taken between 1806 and 1846.[178] In 1812, the Russians founded an agricultural settlement at what is now Fort Ross in northern California, as their southern headquarters.[176]
Eventually, sea otter populations became so depleted, commercial hunting was no longer viable. It had stopped in the Aleutian Islands, by 1808, as a conservation measure imposed by the Russian-American Company. Further restrictions were ordered by the company in 1834.
Recovery and conservation
During the 20th century, sea otter numbers rebounded in about two-thirds of their historic range, a recovery considered one of the greatest successes in marine conservation.
The most significant threat to sea otters is oil spills,[67] to which they are particularly vulnerable, since they rely on their fur to keep warm. When their fur is soaked with oil, it loses its ability to retain air, and the animals can quickly die from hypothermia.[67] The liver, kidneys, and lungs of sea otters also become damaged after they inhale oil or ingest it when grooming.[67] The Exxon Valdez oil spill of 24 March 1989 killed thousands of sea otters in Prince William Sound, and as of 2006, the lingering oil in the area continues to affect the population.[184] Describing the public sympathy for sea otters that developed from media coverage of the event, a U.S. Fish and Wildlife Service spokesperson wrote:
As a playful, photogenic, innocent bystander, the sea otter epitomized the role of victim ... cute and frolicsome sea otters suddenly in distress, oiled, frightened, and dying, in a losing battle with the oil.[19]
The small geographic ranges of the sea otter populations in California, Washington, and British Columbia mean a single major spill could be catastrophic for that state or province.[19][57][63] Prevention of oil spills and preparation to rescue otters if one happens is a major focus for conservation efforts. Increasing the size and range of sea otter populations would also reduce the risk of an oil spill wiping out a population.[19] However, because of the species' reputation for depleting shellfish resources, advocates for commercial, recreational, and subsistence shellfish harvesting have often opposed allowing the sea otter's range to increase, and there have even been instances of fishermen and others illegally killing them.[185]
In the Aleutian Islands, a massive and unexpected disappearance of sea otters has occurred in recent decades. In the 1980s, the area was home to an estimated 55,000 to 100,000 sea otters, but the population fell to around 6,000 animals by 2000.[186] The most widely accepted, but still controversial, hypothesis is that killer whales have been eating the otters. The pattern of disappearances is consistent with a rise in predation, but there has been no direct evidence of orcas preying on sea otters to any significant extent.[114]
Another area of concern is California, where recovery began to fluctuate or decline in the late 1990s.
Sea otter habitat is preserved through several
Economic impact
Some of the sea otter's preferred prey species, particularly abalone, clams, and crabs, are also food sources for humans. In some areas, massive declines in shellfish harvests have been blamed on the sea otter, and intense public debate has taken place over how to manage the competition between sea otters and humans for seafood.[194]
The debate is complicated because sea otters have sometimes been held responsible for declines of shellfish stocks that were more likely caused by
Many facets of the interaction between sea otters and the human economy are not as immediately felt. Sea otters have been credited with contributing to the
Roles in human cultures
Left: |
For many maritime indigenous cultures throughout the North Pacific, especially the
Some Ainu folk-tales portray the sea-otter as an occasional messenger between humans and the creator.[202] The sea otter is a recurring figure in Ainu folklore. A major Ainu epic, the Kutune Shirka, tells the tale of wars and struggles over a golden sea-otter. Versions of a widespread Aleut legend tell of lovers or despairing women who plunge into the sea and become otters.[203] These stories have been associated with the many human-like behavioral features of the sea otter, including apparent playfulness, strong mother-pup bonds and tool use, yielding to ready anthropomorphism.[204] The beginning of commercial exploitation had a great impact on the human, as well as animal, populations. The Ainu and Aleuts have been displaced or their numbers are dwindling, while the coastal tribes of North America, where the otter is in any case greatly depleted, no longer rely as intimately on sea mammals for survival.[205]
Since the mid-1970s, the beauty and charisma of the species have gained wide appreciation, and the sea otter has become an icon of environmental conservation.[187] The round, expressive face and soft, furry body of the sea otter are depicted in a wide variety of souvenirs, postcards, clothing, and stuffed toys.[206]
Aquariums and zoos
Sea otters can do well in captivity, and are featured in over 40 public aquariums and zoos.[207] The Seattle Aquarium became the first institution to raise sea otters from conception to adulthood with the birth of Tichuk in 1979, followed by three more pups in the early 1980s.[208] In 2007, a YouTube video of two sea otters holding paws drew 1.5 million viewers in two weeks, and had over 22 million views as of July 2022[update].[209] Filmed five years previously at the Vancouver Aquarium, it was YouTube's most popular animal video at the time, although it has since been surpassed. The lighter-colored otter in the video is Nyac, a survivor of the 1989 Exxon Valdez oil spill.[210] Nyac died in September 2008, at the age of 20.[211] Milo, the darker one, died of lymphoma in January 2012.[212]
Current conservation
Sea otters, being a known keystone species, need a humanitarian effort to be protected from endangerment through "unregulated human exploitation".[213] This species has increasingly been impacted by the large oil spills and environmental degradation caused by overfishing and entanglement in fishing gear.[214] Current efforts have been made in legislation: the international Fur Seal Treaty, The Endangered Species Act, IUCN/The World Conservation Union, Convention on international Trade in Endangered Species of Wild Fauna and Flora, and the Marine Mammal Protection Act of 1972. Other conservation efforts are done through reintroduction and zoological parks.
Population bottlenecks
A principal target of the fur trade in the eighteenth through early twentieth centuries, sea otter populations were decimated up and down the coast from Alaska to Baja California.[215] For several decades before their rediscovery, it was presumed that sea otters were completely extinct south of the Pacific Northwest. However, in 1938, a raft of about fifty individuals was discovered under a bridge by the Bixby River between Santa Cruz and Big Sur, California.[215] This discovery was soon followed by the taking into captivity of numerous members of the populations in order to begin captive breeding for population restoration. However, with a population size of fifty and the low genetic diversity amongst the population post-fur trade but pre-discovery that was so low, this produced an evolutionary bottleneck. An evolutionary bottleneck is a phenomenon that occurs when population size is greatly diminished from prior numbers due to factors like overhunting, disease, or environmental changes such as decreased carrying capacity and natural disasters. This, in turn, leads to low genomic diversity within the population, which can have adverse effects due to interbreeding and an increase in genetic drift.
Major bottlenecks in history
Sea otter populations, decimated in the eighteenth through early twentieth centuries by the fur trade, experienced a significant population bottleneck in genetic diversity. Following their divergence from their most common ancestor five million years ago, sea otters have developed traits dependent on polygenic selection, or the evolution of numerous traits to create hallmark features like thick and oily fur and large bones, compared to their freshwater sister species.[216] Sea otters require these traits to survive the cold waters of the northern Pacific Ocean, in which they spend their entire lives despite occasionally coming out of the water as pups. Sea otters have the thickest fur of any animal (~1,000,000 hairs per square inch), as they do not have a blubber layer, while their oil glands help matt down their fur and keep it from holding air.[217] Thick bones also prove crucial in increasing buoyancy, as sea otters spend long hours floating atop the ocean. In a study, southern and northern Sea Otter populations were compared against the African clawless otter, and it was determined that aquatic traits like loss of smell and hair thickness independently evolved, evidencing a complex genome of polygenic traits resulting in complex systems.[216] This study was only able to take place after sequencing of Sea Otter nuclear genomes and through phylogeny to find a close ancestor with which to compare genomes.
Previously, it was suspected that sea otters came from the same evolutionary branch as earless seals, such as harbor and monk seals. Sea Otters have experienced numerous population bottlenecks throughout their history, with significant numbers being wiped out 9,000-10,000 generations ago and 300-700 generations ago, long before the fur trade.[216] These previous genetic bottlenecks are responsible for already low genetic diversity amongst species members, making the secondary bottleneck caused by the fur trade more significant. These primary bottlenecks were most likely caused by disease, a common cause for genetic bottlenecks. Estimates place these bottlenecks at leaving around ten to forty animals for about eight to forty-four years.[216] This led to genetic drift, as the populations of northern and southern sea otters were cut off from one another by thousands of miles, leading to significant genomic differences. However, the modern population bottleneck caused by the fur trade of the eighteenth and early twentieth centuries presents the most significant concern to scientists and conservationists attempting to recover population numbers and genetic diversity.
Each bottleneck has lowered genomic diversity and thus increased the chance of deleterious genetic drift, in which a population, because of low effective and census size is driven to interbreed, which increases effective size through increased mutations. However, these mutations, like most mutations, are neutral if not decreasing the population's fitness, thus leading to complications in populations. This phenomenon is studied more widely in cheetah populations, where due to overhunting throughout the eighteenth and early twentieth centuries, the size of the population was greatly diminished, leading to interbreeding and the rise of deleterious mutations.[218] However, thanks to conservation agencies worldwide, the cheetah population is slowly returning, along with its genetic diversity, thanks to purposeful interbreeding with several different cheetah populations.[218]
Conservation
The recent population constraints put on the sea otter have led to low genomic diversity among species members, with much evidence of interbreeding.[217] This interbreeding has led to the mutation of deleterious missense mutations, which may make fast-paced population growth difficult for conservation reasons. While longer-term recovery goals bolstering genetic diversity by interbreeding are costly and challenging, they could significantly aid in avoiding the further evolution of deleterious variation, thus aiding sea otter population stabilization. This method has already been utilized in returning cheetah populations to higher numbers and higher genetic diversity, and captive breeding programs through organizations such as the Monterey Bay Aquarium and The Marine Mammal Center make the chances of getting sea otter populations back up to pre-fur trade numbers possible. The population of sea Otters in California has risen to around 3,000 in the wild.[215] While this figure is far below pre-fur trade numbers, it represents a massive improvement in the conservation of the species and a massive increase in genetic diversity. On the other hand, northern sea otters have reached back up to pre-fur trade population numbers, with populations living all along the state's coast from Ketchikan in the south to Attu in the west. Historical populations, however, are estimated to have been between 150,000 and 300,000 individuals living along the northern Pacific rim from Baja California to Hokkaido Island in Japan. Modern conservation techniques have included breeding northern and southern populations of sea otters to increase genetic diversity and prevent both inbreeding and genetic drift. Moreover, the introduction of the Marine Mammal Protection Act in the 1970s made their hunting highly illegal in the United States.
See also
Notes
- ^ Enhydra lutris nereis is included in Appendix I
References
Citations
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Cited works
- Kenyon, Karl W. (1969). The Sea Otter in the Eastern Pacific Ocean. Washington, D.C.: U.S. Bureau of Sport Fisheries and Wildlife. ISBN 978-0-486-21346-0.
- Love, John A. (1992). Sea Otters. Golden, Colorado: Fulcrum Publishing. OCLC 25747993.
- Nickerson, Roy (1989). Sea Otters, a Natural History and Guide. San Francisco, CA: Chronicle Books. OCLC 18414247.
- Silverstein, Alvin; Silverstein, Virginia and Robert (1995). The Sea Otter. Brookfield, Connecticut: The Millbrook Press, Inc. OCLC 30436543.
- Middleton, John (2001). Maritime Activities And Their Perception Today. San Francisco, California: California Academy of Science. )
- VanBlaricom, Glenn R. (2001). Sea Otters. Stillwater, MN: Voyageur Press Inc. OCLC 46393741.
- McLeish, Todd (2018). Return of the Sea Otter: The Story of the Animal That Evaded Extinction on the Pacific Coast. Seattle, WA: Sasquatch Books. ISBN 978-1-63217-137-5.
External links
Enhydra lutris
(Sea otter).
- Enhydra lutris (Linnaeus, 1758) at the Integrated Taxonomic Information System
- Enhydra lutris (Linnaeus, 1758) at the World Register of Marine Species
- De Bestiis Marinis, or, The Beasts of the Sea (1751) (PDF), pp. 68–82, transcribed field notes from 18th-century German zoologist Georg Wilhelm Steller
- Precipice of Survival: The Southern Sea Otter Archived 22 April 2008 at the Wayback Machine (Adobe Flash), a 48-minute program on the southern sea otter's history by the United States Geological Survey