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Atlantic salmon

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Atlantic salmon
Breeding male, caught in Germany
Illustration
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Salmoniformes
Family: Salmonidae
Genus: Salmo
Species:
S. salar
Binomial name
Salmo salar
Distribution of Atlantic salmon

The Atlantic salmon (Salmo salar) is a species of ray-finned fish in the family Salmonidae. It is the third largest of the Salmonidae, behind Siberian taimen and Pacific Chinook salmon, growing up to 1 m (3.3 ft) in length. Atlantic salmon are found in the northern Atlantic Ocean and in rivers that flow into it. Most populations are anadromous, hatching in streams and rivers but moving out to sea as they grow where they mature, after which the adults seasonally move upstream again to spawn.[2]

When the mature fish re-enter rivers to spawn, they change in colour and appearance. Some populations of this fish only migrate to large lakes, and are "landlocked", spending their entire lives in freshwater. Such populations are found throughout the range of the species. Unlike Pacific species of salmon, S. salar is iteroparous, which means it can survive spawning and return to sea to repeat the process again in another year with 5–10% returning to the sea to spawn again. Such individuals can grow to extremely large sizes, although they are rare. The different life stages of the fish are known by several different names in English; alevin, fry, parr and smolt.

Atlantic salmon meat is a particularly nutritious food and is considered one of the more refined types of fish meat in many cultures. As such it features in numerous popular traditional cuisines and can fetch a higher price than some other fish. It has thus long been the target of recreational and commercial fishing, and this, as well as habitat destruction, has impacted the population in some areas. As a result, the species is the subject of conservation efforts in several countries, which appear to have been somewhat successful since the 2000s. Techniques to farm this species using aquacultural methods have also been developed, and at present it is farmed in great numbers in many countries, with Norway producing over 50% of the farmed world supply.[3] Although this is now a viable alternative to wild-caught fish, farming methods have attracted criticism from environmentalists.[4]

Nomenclature

[edit]

The Atlantic salmon was given its scientific binomial name by Swedish zoologist and taxonomist Carl Linnaeus in 1758. The name, Salmo salar, derives from the Latin salmo, meaning salmon, and salar, meaning leaper, according to M. Barton,[5] but more likely meaning "resident of salt water" [citation needed]. Lewis and Short's Latin Dictionary (Clarendon Press, Oxford, 1879) translates salar as a kind of trout from its use in the Idylls of the poet Ausonius (4th century CE). Later, the differently coloured smolts were found to be the same species.[citation needed]

Other names used for the Atlantic salmon are: bay salmon, black salmon, caplin-scull salmon, fiddler, sebago salmon, silver salmon, outside salmon and winnish. At different points in their maturation and life cycle, they are known as parr, smolt, grilse, grilt, kelt, slink, and spring salmon. Atlantic salmon that do not journey to sea are known as landlocked salmon.[6][citation needed]

Description

[edit]
In Québec, showing the kype characteristic of breeding males

Atlantic salmon are the largest species in the genus Salmo. After two years at sea, the fish average 71 to 76 cm (28 to 30 in) in length and 3.6 to 5.4 kg (7.9 to 11.9 lb) in weight.[7] Specimens that spend four or more winters feeding at sea can be much larger. An Atlantic salmon netted in 1960 in Scotland, in the estuary of the river Hope, weighed 49.44 kg (109.0 lb), the heaviest recorded in all available literature. Another netted in 1925 in Norway measured 160.65 cm (63.25 in) in length, the longest Atlantic salmon on record.[8]

Juveniles of various sizes, in Scotland

The colour of young Atlantic salmon does not resemble the adult stage. While they live in fresh water, they have blue and red spots. At maturity, they take on a silver-blue sheen. The easiest way of identifying them as an adult is by the black spots predominantly above the lateral line, though the caudal fin is usually unspotted. When they reproduce, males take on a slight green or red colour. The salmon has a fusiform body, and well-developed teeth. All fins, except the adipose fin, are bordered with black.[citation needed]

Distribution and habitat

[edit]
Ocean migration of Atlantic salmon[9]
Ocean phase, in Newfoundland
At Atlantic Sea-Park, in Norway

The natural breeding grounds of Atlantic salmon are rivers in Europe and the northeastern coast of North America. In Europe, Atlantic salmon are still found as far south as Spain, and as far northeast as the Pechora River in Russia.[10] Atlantic salmon have spread north to colonise Svalbard (78°N) from 2002–2006 onward, due to the warming of the climate there.[11] Because of sport-fishing, some of the species' southern populations in northern Spain are growing smaller.[12] The species distribution is easily influenced by changes in freshwater habitat and climate. Atlantic salmon are a cold-water fish species and are particularly sensitive to changes in water temperature.[13]

The Housatonic River, and its Naugatuck River tributary, hosted the southernmost Atlantic salmon spawning runs in the United States.[14][15] However, there is a 1609 account by Henry Hudson that Atlantic salmon once ran up the Hudson River.[16] In addition, fish scale evidence dating to 10,000 years BP places Atlantic salmon in a coastal New Jersey pond.[17]

Two publications from 1988 and 1996 questioned the notion that Atlantic salmon were prehistorically plentiful in New England, when the climate was warmer as it is now. This argument was primarily based on a paucity of bone data in archaeological sites relative to other fish species, and the assertion that historical claims of abundance may have been exaggerated.[18][19] This argument was later challenged in another paper which claimed that lack of archaeological bone fragments could be explained by salmon bones being rare at sites that still have large salmon runs and that salmonid bones in general are poorly recovered relative to other fish species.[20][21]

Atlantic salmon populations were significantly reduced in the United States following European settlement. The fur trade, timber harvesting, dams and mills and agriculture degraded freshwater habitats and lowered the carrying capacity of most North American streams. Beaver populations were trapped to near-extinction by 1800, and log drives and clear-cutting further exacerbated stream erosion and habitat loss. As timber and fur gave way to agriculture, freshwater Atlantic salmon habitat was further compromised. According to historian D.W. Dunfield (1985) "over half of the historical Atlantic salmon runs had been lost in North America by 1850". As early as 1798, a bill for the preservation of Atlantic Salmon was introduced in Canadian Parliament, to protect populations in Lake Ontario.[22] In the Gulf Region of Nova Scotia it was reported that 31 of the 33 Atlantic salmon streams were blocked off by lumber dams, leading to the extirpation of early-run fish in many watersheds. The inshore Atlantic salmon fishery became a major export of the New World, with major fishing operations establishing along the shores of major river systems. The southernmost populations were the first to disappear.[citation needed]

Young salmon spend one to four years in their natal river. When they are large enough (c. 15 centimetres (5.9 in)), they smoltify, changing camouflage from stream-adapted with large, gray spots to sea-adapted with shiny sides. They also undergo some endocrinological changes to adapt to osmotic differences between fresh water and seawater habitat. When smoltification is complete, the parr (young fish) now begin to swim with the current instead of against it. With this behavioural change, the fish are now referred to as smolt. When the smolt reach the sea, they follow sea surface currents and feed on plankton or fry from other fish species such as herring. During their time at sea, they can sense the change in the Earth magnetic field through iron in their lateral line.[citation needed]

When they have had a year of good growth, they will move to the sea surface currents that transport them back to their natal river. It is a major misconception that salmon swim thousands of kilometres at sea; instead they surf through sea surface currents.[citation needed] It is possible they find their natal river by smell, although this is not confirmed;[23] only 5% of Atlantic salmon go up the wrong river.[citation needed] The range of an individual Atlantic salmon can thus be the river where they are born and the sea surface currents that are connected to that river in a circular path.[citation needed]

Wild salmon continued to disappear from many rivers during the twentieth century due to overfishing and habitat change.[23]

Ecology

[edit]

Diet

[edit]

Young salmon begin a feeding response within a few days. After the yolk sac is absorbed by the body, they begin to hunt. Juveniles start with tiny invertebrates, but as they mature, they may occasionally eat small fish. During this time, they hunt both in the substrate and in the current. Some have been known to eat salmon eggs. Plankton such as euphausiids are important food for pre-grilse but amphipods and decapods are also consumed.[24] The most commonly eaten foods include caddisflies, blackflies, mayflies, stoneflies,[23] and chironomids, as well as terrestrial insects.[24]

As adults, the salmon prefer capelin as their meal of choice. Capelin are elongated silvery fish that grow up to 20–25 centimetres (8–10 in) long.[25] Other fish consumed include herring, alewives, smelts, scomberids, sand lance, and small cod.[24]

Behaviour

[edit]

Fry and parr have been said[by whom?] to be territorial, but evidence[example needed] showing them to guard territories is inconclusive. While they may occasionally[when?] be aggressive towards each other, the social hierarchy is still unclear. Many[quantify] have been found[by whom?] to school, especially when leaving the estuary.

Adult Atlantic salmon are considered[by whom?] much more aggressive than other salmon, and are more likely to attack other fish than others.[23]

Life stages

[edit]
Life cycle of Atlantic salmon

Most Atlantic salmon follow an anadromous migration pattern,[2] in that they undergo their greatest feeding and growth in saltwater; however, adults return to spawn in native freshwater streams where the eggs hatch and juveniles grow through several distinct stages.[citation needed]

Atlantic salmon do not require saltwater. Numerous examples of fully freshwater (i.e., "landlocked") populations of the species exist throughout the Northern Hemisphere,[2] including a now extinct population in Lake Ontario, which has been shown in recent studies to have spent its entire life cycle in the watershed of the lake.[26] In North America, the landlocked strains are frequently known as ouananiche.[citation needed]

Freshwater phase

[edit]

The freshwater phases of Atlantic salmon vary between two and eight years, according to river location.[27] While the young in southern rivers, such as those to the English Channel, are only one year old when they leave, those further north, such as in Scottish rivers, can be over four years old, and in Ungava Bay, northern Quebec, smolts as old as eight years have been encountered.[27]

The first phase is the alevin stage, when the fish stay in the breeding ground and use the remaining nutrients in their yolk sacs. During this developmental stage, their young gills develop and they become active hunters. Next is the fry stage, where the fish grow and subsequently leave the breeding ground in search of food. During this time, they move to areas with higher prey concentration. The final freshwater stage is when they develop into parr, in which they prepare for the trek to the Atlantic Ocean.[citation needed]

During these times, the Atlantic salmon are very susceptible to predation. Nearly 40% are eaten by trout alone. Other predators include other fish and birds.[citation needed] Egg and juvenile survival is dependent on habitat quality as Atlantic salmon are sensitive to ecological change.[citation needed]

Saltwater phases

[edit]
Atlantic salmon are among the largest salmon species

When parr develop into smolt, they begin the trip to the ocean, which predominantly happens between March and June. Migration allows acclimation to the changing salinity. Once ready, young smolt leave, preferring an ebb tide.[citation needed]

Having left their natal streams, they experience a period of rapid growth during the one to four years they live in the ocean. Typically, Atlantic salmon migrate from their home streams to an area on the continental plate off West Greenland. During this time, they face predation from humans, seals, Greenland sharks, skate, cod, and halibut. Some dolphins have been noticed playing with dead salmon, but it is still unclear whether they consume them.[citation needed]

Once large enough, Atlantic salmon change into the grilse phase, when they become ready to return to the same freshwater tributary they departed from as smolts. After returning to their natal streams, the salmon will cease eating altogether prior to spawning. Although largely unknown, odor – the exact chemical signature of that stream – may play an important role in how salmon return to the area where they hatched. Once heavier than about 250 g, the fish no longer become prey for birds and many fish, although seals do prey upon them. Grey and common seals commonly eat Atlantic salmon. Survivability to this stage has been estimated at between 14 and 53%.[23]

Breeding

[edit]
Jumping at the Falls of Shin, Scotland

Atlantic salmon breed in the rivers of Western Europe from northern Portugal north to Norway, Iceland, and Greenland, and the east coast of North America from Connecticut in the United States north to northern Labrador and Arctic Canada.[citation needed]

Carcass half-eaten by animals, in Scotland.

The species constructs a nest or "redd" in the gravel bed of a stream. The female creates a powerful downdraught of water with her tail near the gravel to excavate a depression. After she and a male fish have eggs and milt (sperm), respectively, upstream of the depression, the female again uses her tail, this time to shift gravel to cover the eggs and milt which have lodged in the depression.[citation needed]

Unlike the various Pacific salmon species which die after spawning (semelparous), the Atlantic salmon is iteroparous, which means the fish may recondition themselves and return to the sea to repeat the migration and spawning pattern several times, although most spawn only once or twice.[2][28] Migration and spawning exact an enormous physiological toll on individuals, such that repeat spawners are the exception rather than the norm.[28] Atlantic salmon show high diversity in age of maturity and may mature as parr, one- to five-sea-winter fish, and in rare instances, at older sea ages. This variety of ages can occur in the same population, constituting a 'bet hedging' strategy against variation in stream flows. So in a drought year, some fish of a given age will not return to spawn, allowing that generation other, wetter years in which to spawn.[27]

Hybridization

[edit]

When in shared breeding habitats, Atlantic salmon will hybridize with brown trout (Salmo trutta).[29][30][31] Hybrids between Atlantic salmon and brown trout were detected in two of four watersheds studied in northern Spain. The proportions of hybrids in samples of salmon ranged from 0 to 7-7% but these proportions were not significantly homogeneous among locations, resulting in a mean hybridization rate of 2-3%. This is the highest rate of natural hybridization so far reported and is significantly greater than rates observed elsewhere in Europe.[32]

Beaver impact

[edit]

The decline in anadromous salmonid species over the last two to three centuries is correlated with the decline in the North American beaver and European beaver, although some fish and game departments[example needed] continue to advocate removal of beaver dams as potential barriers to spawning runs. Migration of adult Atlantic salmon may be limited by beaver dams during periods of low stream flows, but the presence of juvenile salmon upstream of the dams suggests they are penetrated by parr.[33] Downstream migration of Atlantic salmon smolts was similarly unaffected by beaver dams, even in periods of low flows.[33]

In a 2003 study, Atlantic salmon and sea-run brown trout spawning in the Numedalslågen River and 51 of its tributaries in southeastern Norway was unhindered by beavers.[34] In a restored, third-order stream in northern Nova Scotia, beaver dams generally posed no barrier to Atlantic salmon migration except in the smallest upstream reaches in years of low flow where pools were not deep enough to enable the fish to leap the dam or without a column of water over-topping the dam for the fish to swim up.[35]

The importance of winter habitat to salmonids afforded by beaver ponds may be especially important in streams of northerly latitudes without deep pools where ice cover makes contact with the bottom of shallow streams.[33] In addition, the up to eight-year-long residence time of juveniles in freshwater may make beaver-created permanent summer pools a crucial success factor for Atlantic salmon populations. In fact, two-year-old Atlantic salmon parr in beaver ponds in eastern Canada showed faster summer growth in length and mass and were in better condition than parr upstream or downstream from the pond.[36]

Relationship to humans

[edit]

Atlantic salmon is a popular fish for human consumption[2] and is commonly sold fresh, canned, or frozen.[citation needed]

Seine fishing for salmon – Wenzel Hollar, 1607–1677

Wood and stone weirs along streams and ponds were used for millennia to harvest salmon in the rivers of New England.[37] European fishermen gillnetted for Atlantic salmon in rivers using hand-made nets for many centuries[38] and gillnetting was also used in early colonial America.[39]

In its natal streams, Atlantic salmon are considered prized recreational fish, pursued by fly anglers during its annual runs. At one time, the species supported an important commercial fishery, but having become endangered throughout its range globally, wild-caught Atlantic salmon are now virtually absent from the market.[40] Instead, nearly all are from aquaculture farms, predominantly in Norway, Chile, Canada, the UK, Ireland, Faroe Islands, Russia and Tasmania in Australia.[28]

Aquaculture

[edit]
Atlantic salmon marine cages in the Faroe Islands
Capture (blue) and aquaculture (green) production of Atlantic salmon (Salmo salar) in million tonnes from 1975 to 2022, as reported by the FAO[41]
Sea lice on farmed Atlantic salmon

Adult male and female fish are anaesthetised; their eggs and sperm are "stripped" after the fish are cleaned and cloth dried. Sperm and eggs are mixed, washed, and placed into freshwater. Adults recover in flowing, clean, well-aerated water.[42] Some researchers have even studied cryopreservation of their eggs.[43]

Fry are generally reared in large freshwater tanks for 12 to 20 months. Once the fish have reached the smolt phase, they are taken out to sea, where they are held for up to two years. During this time, the fish grow and mature in large cages off the coasts of Canada, the US, or parts of Europe.[28]

There are many different commercially available cage designs built to operate in a wide variety of aquatic conditions. High-density polyethylene (HDPE) cages are widely used, with HDPE pipes forming a floating collar ring onto which the fish net pen is secured and suspended in the water below.[44]

Advancements in cage technologies have allowed for reduction in fish escapes, improvement in growing conditions, and maximization of aquaculture production volume per unit area of growing space.[44]

Controversy

[edit]

Farmed Atlantic salmon are known to occasionally escape from cages and enter the habitat of wild populations. Interbreeding between escaped farm fish and wild fish decreases genetic diversity and introduces "the potential to genetically alter native populations, reduce local adaptation and negatively affect population viability and character".[45] A study in 2000 demonstrated that the genes of farmed Atlantic salmon intrude wild populations mainly through wild males breeding with farmed females, though farmed specimens showed reduced capacity for breeding success overall compared to their wild counterparts.[46] Further study in 2018 discovered extensive cross-breeding of wild and farmed Atlantic salmon in the Northwest Atlantic, showing that 27.1% of fish in 17 out of 18 rivers examined are artificially stocked or hybrids. Farming of Atlantic salmon in open cages at sea has also been linked, at least in part, to a decline in wild stocks attributed to the passing of parasites from farmed to wild individuals.[47]

On the west coast of the United States and Canada, aquaculturists are generally under scrutiny to ensure that non-native Atlantic salmon cannot escape from their open-net pens, however occasional incidents of escape have been documented.[48] During one incident in 2017, for example, up to 300,000 potentially invasive Atlantic salmon escaped a farm among the San Juan Islands in Puget Sound, Washington.[49] Washington went on in 2019 to implement a gradual phase out of salmon farming to be completed by 2025.[50]

Despite being the source of considerable controversy,[51] the likelihood of escaped Atlantic salmon establishing an invasive presence in the Pacific Northwest is considered minimal, largely because a number of 20th century efforts aimed at deliberately introducing them to the region were ultimately unsuccessful.[52] From 1905 until 1935, for example, in excess of 8.6 million Atlantic salmon of various life stages (predominantly advanced fry) were intentionally introduced to more than 60 individual British Columbia lakes and streams. Historical records indicate, in a few instances, mature sea-run Atlantic salmon were captured in the Cowichan River; however, a self-sustaining population never materialized. Similarly unsuccessful results were realized after deliberate attempts at introduction by Washington as late as the 1980s.[53] Consequently, environmental assessments by the US National Marine Fisheries Service (NMFS), the Washington Department of Fish and Wildlife and the BC Environmental Assessment Office have concluded the potential risk of Atlantic salmon colonization in the Pacific Northwest is low.[54]

Future prospects

[edit]

A study of Næve et al. (2022) estimated the impact of 50 years of genetic selection and tried to predict the impact it could have until 2050. In order to do this, a common garden experiment was used to model and simulate past and future effects for 11 generations of genetic selection of increased growth rate in Atlantic salmon. To model the contribution that breeding has made in the industry from generation 0 (harvested in 1975– 1978) to generation 11 (harvested in 2017 – 2019), and to simulate growth until 2050 (generation 24), the Norwegian salmon aquaculture production between 2016 and 2019 was used as a base case. The simulation of the expected growth until 2050 (generation 24) gave five different scenarios : Historical (H1), Forecast 1 (F1), Forecast 2 (F2), Forecast 3 (F3) and Forecast 4 (F4). Changes in thermal growth coefficient (TGC) per generation were used in the model to simulate the differences in the five scenarios. The genetic data, H1, and the most conservative forecast scenario, F1, simulate what can be expected in 2050 if the trend from generation 0 through 11 is maintained. The following forecast scenarios assume a greater increase in genetic growth with a larger increase in the TGC in the generations to come. In the next two generations, more advanced selection methods such as marker assisted selection (from generation 10) and genomic selection (from generation 11) were implemented. This resulted in increased gain in selection for growth and simulated F2 and F3. The most progressive scenario, F4, aimed at exploring the effect in the industry when the full genetic potential is utilized. This assumes a further development of advanced techniques in the years to come. The authors of the article found that the daily yield of the biomass increased with increasing generations in the historic and forecast scenarios. Further, the production time in seawater to reach the harvest weight of 5100 g is expected to be reduced by 53% in 2050. When production time can be reduced, this will also reduce e.g. time at risk of diseases. In the most progressive scenario, mortality in seawater was expected to be reduced by up to 50%. Further, the authors found that production per license can increase by up to 121%. Additionally, 77% of the new volume needed to achieve five million tonnes in 2050, may be provided by genomic selection. However, one should keep in mind that this article was published by the firm Aquagen, and can possibly be biased and too optimistic.[55]

Conservation

[edit]
A man fishing for Atlantic salmon in the Pabos River of Quebec as recreation.
A fishmonger in Lysekil, Sweden shows a Norwegian salmon.

The IUCN rates this as a common species with a conservation status of "least concern", however it has been 25 years since the IUCN has released this status.[56][57] A more recent regional assessment revealed that the European population of this species is vulnerable, and this might be the same or a similar status globally. Location-specific assessments have shown population declines across parts of the Atlantic Salmon's natural range, with populations along the coast of Maine and the Inner Bay of Fundy now listed as "endangered" under the Endangered Species Act, and the Canadian Species at Risk Act, respectively.[58][56]

Human activities have impacted salmon populations across parts of its range. The major threats are from overfishing and habitat change.[22] Salmon decline in Lake Ontario goes back to the 18th–19th centuries, due to logging and soil erosion, as well as dam and mill construction. By 1896, the species was declared extirpated from the lake.[26][59]

In the 1950s, salmon from rivers in the United States and Canada, as well as from Europe, were discovered to gather in the sea around Greenland and the Faroe Islands. A commercial fishing industry was established, taking salmon using drift nets. After an initial series of record annual catches, the numbers crashed; between 1979 and 1990, catches fell from four million to 700,000.[60]

In captivity

Beginning around 1990, the rates of Atlantic salmon mortality at sea more than doubled in the western Atlantic. Rivers of the coast of Maine, southern New Brunswick and much of mainland Nova Scotia saw runs drop precipitously, and even disappear. An international effort to study the increased mortality rate was organized by the North Atlantic Salmon Conservation Organization.[6] In 2000 the numbers of Atlantic salmon dropped to very low levels in Newfoundland, Canada.[61] In 2007 at least one sport fishing organization from Iceland and Scandinavia blamed less fish caught by recreational anglers on overfishing at sea, and thus created the North Atlantic Salmon Fund to buy commercial quotas in the Atlantic from commercial fishermen in an effort to preserve wild Salmo salar stocks.[60]

Possibly because of improvements in ocean feeding grounds, returns in 2008 were very positive. On the Penobscot River in Maine, returns were about 940 in 2007, and by mid-July 2008, the return was 1,938. Similar stories were reported in rivers from Newfoundland to Quebec. In 2011, more than 3,100 salmon returned to the Penobscot, the most since 1986, and nearly 200 ascended the Narraguagus River, up from the low two digits just a decade before.[6][62]

Recreational fishing of stocked, landlocked Atlantic salmon is now authorized in much of the US and Canada where it occurs in large numbers, but this is subject to regulations in many states or provinces which are designed to maintain the continuity of the species. Strict catch limits, catch and release practices and forced fly fishing are examples of those regulations.[63][64] However, catch and release angling can be an additional stressor on Atlantic salmon populations, especially when its impacts are combined with the existing pressures of climate change, overfishing, and predation.[65][63]

Restoration efforts

[edit]
At Lake Ontario

Around the North Atlantic, efforts to restore salmon to their native habitats are underway, with slow progress. Habitat restoration and protection are key to this process, but issues of excessive harvest and competition with farmed and escaped salmon are also primary considerations. In the Great Lakes, Atlantic salmon have been reintroduced, but the percentage of salmon reproducing naturally is very low. Most areas are re-stocked annually.[22] Since the extirpation of Atlantic salmon from Lake Ontario in the late 19th century, the state of New York has stocked its adjoining rivers and tributaries, and in many cases does not allow active fishing.[2][66][22]

Jumping up a waterfall at Oak Orchard Creek, Lake Ontario
Jumping up a fish ladder in Norway

The province of Ontario started the Atlantic Salmon Restoration Program[67] in 2006, which is one of the largest freshwater conservation programs in North America. It has since stocked Lake Ontario and surrounding tributaries with upwards of 6,000,000 young Atlantic salmon, with efforts growing each year.[68] In New England, many efforts are underway to restore salmon to the region by knocking down obsolete dams and updating others with fish ladders and other techniques that have proven effective in the West with Pacific salmon. There is some success thus far, with populations growing in the Penobscot and Connecticut Rivers. Lake Champlain now has Atlantic salmon. The Atlantic Salmon Federation is involved in restoration efforts along the eastern United States and Canada, where their projects are focused on removing barriers to fish passage and eradicating invasive species.[69]

Recent documented successes in the reintroduction of Atlantic salmon include the following:

  • In October 2007, salmon were video-recorded running in Toronto's Humber River by the Old Mill.[59]
  • A migrating salmon was observed in Ontario's Credit River in November 2007.[59]
  • As of 2013, there has been some success in establishing Atlantic salmon in Fish Creek, a tributary of Oneida Lake in central New York.[70]
  • In November 2015, salmon nests were observed in Connecticut in the Farmington River, a tributary of the Connecticut River where Atlantic salmon had not been observed spawning since "probably the Revolutionary War".[71] However, both state and federal experts indicated that this find likely represented a dwindling wave of returning stocked fish from massive salmon restoration efforts that had concluded years earlier in 2012. Significant doubt was cast on fish returning to spawn in meaningful numbers after 2017, when the last generation of stocked salmon would return.[72]

NASCO

[edit]

The North Atlantic Salmon Conservation Organization is an international council made up of Canada, the European Union, Iceland, Norway, the Russian Federation, the United Kingdom, and the United States, with its headquarters in Edinburgh. It was established in 1983 to help protect Atlantic salmon stocks, through the cooperation between nations. They work to restore habitat and promote conservation of the salmon.[73] In December 2021, NASCO published an updated interactive map of their Rivers Database, showing the stock status of wild Atlantic salmon populations across the species range.

Legislation

[edit]

England and Wales

[edit]

Edward I instituted a penalty for collecting salmon during certain times of the year. His son Edward II continued, regulating the construction of weirs. Enforcement was overseen by those appointed by the justices of the peace. Because of confusing laws and the appointed conservators having little power, most laws were barely enforced.[citation needed]

Based on this, a royal commission was appointed in 1860 to thoroughly investigate the Atlantic salmon and the laws governing the species, resulting in the 1861 Salmon Fisheries Act. The act placed enforcement of the laws under the Home Office's control, but it was later transferred to the Board of Trade, and then later to the Board of Agriculture and Fisheries. Another act passed in 1865 imposed charges to fish and catch limits. It also caused the formation of local boards having jurisdiction over a certain river. The next significant act, passed in 1907, allowed the board to charge 'duties' to catch other freshwater fish, including trout.[citation needed]

Despite legislation, board effects decreased until, in 1948, the River Boards Act gave authority of all freshwater fish and the prevention of pollution to one board per river. In total, it created 32 boards. In 1974, the 32 boards, which by then were integrated into regional river authorities, were reduced to 10 regional water authorities (RWAs). Although only the Northumbrian Water Authority, Welsh National Water Development Authority, Northwest Water Authority and Southwest Water Authority had significant salmon populations, all ten also regulated and conserved trout and freshwater eel fisheries

The Salmon and Freshwater Fisheries Act was passed in 1975. Among other things, it regulated fishing licences, seasons, and size limits, and banned obstructing the salmon's migratory paths.[23]

Scotland

[edit]

Salmon was greatly valued in medieval Scotland, and various fishing methods, including the use of weirs, cruives, and nets, were used to catch the fish. Fishing for salmon was heavily regulated in order to conserve the resource.[74] In 1318, King Robert I enacted legislation setting a minimum size for cruives, "so that no fry of fish are impeded from ascending and descending..." Laws on catching fish upon royal lands were frequently updated, demonstrating their importance.[74] Because the fish were held in such high regard, poachers were severely punished; a person twice convicted of poaching salmon on a royal estate could be sentenced to death.[75] The export of salmon was economically important in Aberdeen; beginning in the 15th century, the fish could be preserved through salting and barreling, allowing them to be exported abroad, including as far away as the Baltic. The volume of the early Scottish salmon trade is impossible to determine, since surviving custom records date only from the 1420 onward, and since Aberdeen burgesses enjoyed an exemption on salmon customs until the 1530s.[76]

During the 15th century, many laws were passed; many regulated fishing times, and worked to ensure smolts could safely pass downstream. James III even closed a meal mill because of its history of killing fish attracted to the wheel.[citation needed]

More recent legislation has established commissioners who manage districts. Furthermore, the Salmon and Freshwater Fisheries Act in 1951 required the Secretary of State be given data about the catches of salmon and trout to help establish catch limits.[23][42]

United States

[edit]

Commercial and recreational fishing of wild, anadromous Atlantic salmon is prohibited in the United States.[7] Several populations of Atlantic salmon are in serious decline, and are listed as endangered under the Endangered Species Act (ESA). Currently, runs of 11 rivers in Maine are on the list – Kennebec, Androscoggin, Penobscot, Sheepscot, Ducktrap, Cove Brook, Pleasant, Narraguagus, Machias, East Machias and Dennys. The Penobscot River is the "anchor river" for Atlantic salmon populations in the US. Returning fish in 2008 were around 2,000, more than double the 2007 return of 940.[citation needed]

Section 9 of the ESA makes it illegal to take an endangered species of fish or wildlife. The definition of "take" is to "harass, harm, pursue, hunt, shoot, wound, kill, trap, capture, or collect, or to attempt to engage in any such conduct".[77]

Canada

[edit]

The federal government has prime responsibility for protecting the Atlantic salmon, but over the last generation, effort has continued to shift management as much as possible to provincial authorities through memoranda of understanding, for example. A new Atlantic salmon policy is in the works, and in the past three years,[when?] the government has attempted to pass a new version of the century-old Fisheries Act through Parliament.[citation needed]

Federal legislation regarding at-risk populations is weak.[citation needed] Inner Bay of Fundy Atlantic salmon runs were declared endangered in 2000. A recovery and action plan is in place.[78]

Nongovernmental organizations, such as the Atlantic Salmon Federation,[79] constantly demand for improvements in management, and for initiatives to be considered. For example, the ASF and the Nova Scotia Salmon Association desire the use of technology for mitigation of acid rain-affected rivers such as used in Norway is in 54 Nova Scotia rivers and managed to raise the funds to get a project in place in one river.[citation needed]

In Quebec, the daily catch limit for Atlantic salmon is dependent on the individual river. Some rivers are strictly catch and release with a limit of 3 released fish. Each catch must be declared. Some rivers allow you to keep between 1 or 2 grilse (30 cm to 63 cm), while some of the more prolific rivers (mainly on the north coast) will allow you to keep 1 salmon over 63 cm. The annual catch limit is 4 Atlantic salmon of small size and only 1 of those may be bigger than 63 cm.[citation needed]

In Lake Ontario, the historic populations of Atlantic salmon became extinct, and cross-national efforts have been under way to reintroduce the species, with some areas already having restocked naturally reproducing populations.[80][81]

See also

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Notes

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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The Atlantic salmon (Salmo salar Linnaeus, 1758) is an anadromous ray-finned fish in the family Salmonidae, characterized by its silvery body, small scales, and dorsal fin, with adults typically measuring 70–100 cm in length and weighing 3–12 kg, though exceptional individuals exceed 1.5 m and 30 kg.[1][2] Native to rivers draining into the North Atlantic Ocean, its range spans from the Arctic Circle southward to Portugal and Connecticut in the west, and from the Baltic to western Russia in the east, where populations exhibit high fidelity to natal spawning sites.[3][2] Exhibiting a complex life cycle, Atlantic salmon hatch as alevins in gravel nests (redds) within freshwater streams, develop into parr that feed on aquatic insects for 1–5 years, then smoltify and migrate seaward to grow rapidly on marine prey like herring and capelin, attaining maturity after 1–3 years at sea before returning hundreds or thousands of kilometers upstream to spawn, often ceasing feeding and relying on stored energy.[4][5] This migration, guided by olfaction, geomagnetic cues, and celestial navigation, underscores their ecological role as nutrient vectors between marine and freshwater ecosystems, supporting biodiversity in rivers and coastal food webs.[5] Economically vital, Atlantic salmon underpin global aquaculture production exceeding 2 million tonnes annually, dominated by farming in Norway, Chile, and Scotland, which supplies high-value markets while wild catches have plummeted due to overfishing, habitat degradation, and marine mortality factors, prompting a 2023 IUCN reassessment to Near Threatened globally amid regional extirpations and ongoing restoration efforts.[6][7][8]

Taxonomy and Nomenclature

Scientific Classification

The Atlantic salmon (Salmo salar) is classified in the domain Eukarya, kingdom Animalia, phylum Chordata, class Actinopterygii (ray-finned fishes), order Salmoniformes, family Salmonidae, genus Salmo, and species S. salar.[3][9][10] The binomial nomenclature Salmo salar was established by Carl Linnaeus in his Systema Naturae (10th edition) published on October 26, 1758, making it one of the earliest formally described salmonid species.[10][9]
Taxonomic RankNameNotes
KingdomAnimaliaMulticellular, heterotrophic eukaryotes with motility in some life stages.[3]
PhylumChordataCharacterized by a notochord, dorsal nerve cord, and pharyngeal slits at some stage.[3]
ClassActinopterygiiRay-finned fishes with leptoid scales and fin rays supported by lepidotrichia.[11]
OrderSalmoniformesIncludes salmonids and allies, adapted for freshwater and anadromous lifestyles.[11][10]
FamilySalmonidaeSalmon and trout family, featuring adipose fin and migratory behavior in many species.[3]
GenusSalmoEuropean salmonids, distinguished from Pacific Oncorhynchus by karyotype and morphology.[10][9]
SpeciesSalmo salarType species of genus Salmo; North Atlantic native with distinct parr marks and spawning migration.[11][10]
This hierarchy reflects molecular and morphological consensus from phylogenetic studies, confirming S. salar's placement within Salmonidae as a monophyletic group basal to Pacific salmon genera.[9] Subspecies distinctions are not formally recognized, though ecotypes vary by river drainage due to genetic isolation rather than taxonomic rank.[3]

Common Names and Synonyms

The Atlantic salmon (Salmo salar) is primarily known by its common name, Atlantic salmon, which distinguishes it from Pacific salmon species in the genus Oncorhynchus.[12] This name reflects its native distribution in the northern Atlantic Ocean and associated rivers.[13] Synonyms and regional variants in English include lake salmon, freshwater salmon, sebago salmon, and ouananiche, often applied to landlocked populations adapted to freshwater lakes, such as those in Maine's Sebago Lake or Quebec's landlocked strains.[12] Additional historical or localized English names encompass bay salmon, black salmon (referring to spawning adults with darkened coloration), caplin-scull salmon, fiddler, silver salmon (for bright oceanic adults), outside salmon, and winnish.[14] These terms arise from North American fishing traditions and morphological observations, with caplin-scull salmon alluding to feeding on capelin fish.[13] In non-English contexts, equivalents include lax in Icelandic, losos in Belarusian, and Atlanterhavslaks in Danish, underscoring its cultural significance in Nordic and Eastern European fisheries.[15] The term grilse specifically denotes young adult individuals returning to fresh water after one sea winter, though it is sometimes used more broadly as a synonym for smaller salmon.[16]

Physical Characteristics

Morphology and Anatomy

The Atlantic salmon (Salmo salar) possesses a fusiform body shape with an oval cross-section, facilitating efficient locomotion through varied aquatic environments.[17] This streamlined form tapers toward the caudal peduncle, supporting high-speed migration and sustained swimming. Body proportions, including head length and depth, vary across life stages and maturity, with juveniles exhibiting more compact forms relative to adults.[12] The skin is covered in small, cycloid scales embedded in a mucous layer that reduces drag and provides protection against pathogens and parasites. Coloration in oceanic adults features bluish-green dorsal hues transitioning to silvery flanks and white ventral surfaces, accented by sparse black spots confined above the lateral line, aiding camouflage in open water.[17] The mouth is moderately large, extending posteriorly to beneath the eye's rear margin, equipped with vomerine teeth and a narrow, pointed tongue bearing small, well-developed dentition.[12] Fins include a slightly emarginated caudal fin with 19 principal rays, a dorsal fin preceded by 3-4 spines, and an anal fin with 9-12 rays, distinguishing it from Pacific congeners by possessing fewer than 13 anal rays.[12] [2] During spawning, sexual dimorphism intensifies: males develop an elongated lower jaw forming a kype that hooks upward, thickened fins, and increased mucus production, while both sexes may redden in spawning grounds.[18] [17] Internally, the species features a physostomous swim bladder, a thin-walled, gas-filled sac connected to the digestive tract via a pneumatic duct, enabling buoyancy regulation through gas secretion or resorption during depth changes and salinity shifts.[19] Four gill arches support respiration and osmoregulation, with gill rakers filtering food particles and facilitating ion exchange critical for anadromous life history.[20] The digestive system comprises a short esophagus leading to a muscular stomach, pyloric caeca for nutrient absorption enhancement, and a spiral valve intestine adapted for processing both invertebrate and fish prey across freshwater and marine phases.[21] The liver produces bile stored in the gallbladder to emulsify lipids, supporting a diet high in fats during oceanic growth. Kidneys handle osmotic balance, with anterior sections aiding marine hyperosmoregulation by excreting excess salts.[22] These anatomical adaptations underpin the species' physiological plasticity in exploiting divergent habitats.[23]

Size, Growth, and Variations

Adult Atlantic salmon (Salmo salar) typically reach lengths of 70–100 cm and weights of 3–12 kg upon returning to freshwater to spawn after one or two years at sea.[11] [24] After two years in the ocean, averages are 71–76 cm in length and 3.6–5.4 kg in weight.[11] Maximum recorded lengths are 150 cm for males and 120 cm for females, with weights up to 46.8 kg documented.[24] [12] Growth in Atlantic salmon is characterized by slow development in freshwater as parr, followed by accelerated rates during the marine phase as post-smolts.[11] Freshwater growth varies with habitat quality, density, and temperature, typically yielding modest annual increments before smoltification at 1–3 years of age.[11] Marine growth is more rapid due to abundant prey, though recent analyses indicate temporal declines in first-year-at-sea growth rates across populations, potentially linked to environmental changes.[25] Factors influencing overall growth include sea age at maturity, sex, and river origin, with multi-sea-winter individuals exhibiting larger sizes than grilse (one-sea-winter salmon).[11] Variations in size and growth occur across life history forms and environments. Landlocked populations, confined to freshwater lakes, attain smaller average sizes of 40–46 cm and 0.5–0.7 kg, with maxima around 3–5 kg, due to limited nutrient availability compared to anadromous counterparts.[26] Farmed Atlantic salmon, under controlled aquaculture conditions, achieve harvest sizes of 4–6 kg in 18–24 months, often exceeding wild counterparts in growth velocity owing to optimized feed and density management, though wild-reared farmed strains show reduced plasticity.[27] Regional differences persist; for instance, salmon from Newfoundland and Labrador rivers average under 70 cm and 4.5 kg, reflecting localized productivity and migration patterns.[4]

Geographic Distribution and Habitat

Native Range and Historical Distribution

The Atlantic salmon (Salmo salar) is natively distributed across rivers draining into the North Atlantic Ocean and adjacent Arctic seas. Its range spans the eastern Atlantic from the Barents Sea and Arctic waters around Svalbard southward to Iberian rivers in Portugal and Spain, encompassing populations in Norway, Sweden, Finland, Russia, Iceland, the British Isles, France, and the Baltic Sea region.[17][2] In the western Atlantic, native stocks occur from Ungava Bay in northern Quebec and Labrador southward along the eastern seaboard to the Connecticut River in the United States, with historical presence extending inland via the St. Lawrence River to the Great Lakes, including Lake Ontario until the late 19th century.[12][11][28] Historically, the species' distribution reflected post-glacial recolonization following the retreat of the Pleistocene ice sheets approximately 10,000 years ago, with distinct North American and European lineages showing limited gene flow due to oceanic barriers.[29] In North America, pre-colonial populations supported abundant runs in nearly every major river from the Hudson River northward to Ungava Bay, sustaining indigenous fisheries and early European settlements.[30] European stocks similarly occupied large river systems like the Rhine, Elbe, and Loire, though many southern peripheral populations have since declined or vanished due to habitat alteration and overexploitation.[31] Genetic and morphological evidence confirms two primary native subpopulations—North American and European—with Baltic salmon forming a distinct subgroup adapted to brackish conditions; transatlantic migrations occur but do not significantly blur these boundaries.[11] The historical extent underscores the species' anadromous life history tied to cold, oligotrophic freshwater habitats and nutrient-rich marine feeding grounds in the subarctic North Atlantic.[17]

Habitat Preferences and Environmental Requirements

Atlantic salmon (Salmo salar) inhabit cold, oxygen-rich freshwater rivers for spawning and juvenile rearing before migrating to marine environments as smolts. Freshwater habitats consist of clear, fast-flowing streams and rivers with low to moderate gradients, featuring gravel and cobble substrates that provide oxygenated interstitial water for egg incubation and fry emergence.[32][4] Spawning adults select sites with water temperatures between 4°C and 12°C, optimally 7.2°C to 10°C, depths of 17-76 cm, and velocities of 0.25-0.9 m/s to facilitate redd construction and oxygenation.[12][33] Substrate gravel sizes range from 20-100 mm in diameter, with median particle diameters typically 14.5-35 mm to ensure permeability and minimize siltation, which can reduce oxygen diffusion to developing embryos.[34][35] Dissolved oxygen levels must exceed 8 mg/L, as lower concentrations impair egg survival and development.[11] Juvenile parr require similar cool waters, with temperatures ideally 10-16°C and avoiding levels above 20-23°C that stress metabolism or coincide with reduced dissolved oxygen solubility.[11][36] They occupy riffles and pools with high oxygen (>5 mg/L, preferably >8 mg/L) and structured substrates for cover, where water flow supports territorial defense and foraging.[37][11] In the marine phase, post-smolts and adults prefer coastal and pelagic waters of the North Atlantic with temperatures of 4-12°C and full salinity (around 35 ppt), actively selecting higher salinity and lower temperatures while avoiding hypoxic zones below critical thresholds that vary with size and activity.[11][38] These conditions support rapid growth and survival, though warming trends beyond thermal tolerances reduce habitat suitability and increase metabolic stress.[39]

Introduced and Translocated Populations

Atlantic salmon (Salmo salar) have been introduced to several regions outside their native North Atlantic range, primarily through deliberate stocking efforts and escapes from aquaculture facilities, but most attempts have failed to establish self-sustaining wild populations.[12] In the Pacific Northwest of North America, including British Columbia, Washington, and Oregon, millions of Atlantic salmon were released into streams between 1905 and 1935 in dozens of trials aimed at creating sport fisheries, yet no viable reproducing populations resulted due to factors such as competition with native Pacific salmonids, unsuitable freshwater habitats, and high mortality during ocean migration.[40] Similarly, in Alaska, escaped farmed individuals are widespread in marine waters, but no evidence of natural reproduction or establishment exists, as confirmed by monitoring data showing reliance on continuous farm inputs rather than wild spawning success.[41] In the Great Lakes basin, introductions from 1935 to 1939 led to temporary presence, with a purported self-sustaining group in Lake Superior, though ongoing populations there depend heavily on hatchery supplementation and do not represent fully independent wild stocks.[12] One notable exception involves limited natural reproduction in non-native southern regions. In Patagonia, Argentina, Atlantic salmon introduced via hatchery releases since the late 20th century have shown evidence of spawning and early life stages in rivers draining to the Atlantic, marking the first documented wild reproduction outside the native range in 2015, though population sizes remain small and viability uncertain due to environmental mismatches and predation.[42] Escapes from salmon farms in Chile and other southern aquaculture sites have raised concerns for potential establishment in Pacific-draining rivers, but genetic and ecological barriers, including hybridization risks with native species and disease transmission, have prevented widespread invasion to date.[41] Translocation efforts within the native range have focused on restoring depleted or extirpated populations through hatchery propagation and river-specific stocking. In the United States, federal hatcheries initiated Atlantic salmon rearing in 1864 to counter overfishing and habitat loss, releasing juveniles into rivers like those in Maine and southward to Connecticut, where wild runs had vanished by the early 1800s; these programs continue to support remnant Gulf of Maine distinct population segments, comprising fewer than 1,000 adults annually in rivers such as the Penobscot and Kennebec.[3] In Europe, translocations from healthy donor rivers have bolstered stocks in fragmented habitats, such as the River Rhine, where post-1950s barrier removals and fry releases re-established runs exceeding 100,000 smolts by the 2010s, though genetic monitoring reveals ongoing challenges from introgression with farmed strains.[43] Landlocked forms, translocated to inland lakes within historical limits like Quebec's Lake Témiscouata, maintain isolated populations without anadromous migration, providing refugia but highlighting adaptation limits under warming climates.[3] These interventions prioritize native genetic lineages to preserve diversity, yet success varies with habitat restoration and straying rates.[11]

Ecology and Life Cycle

Life Stages and Development

Atlantic salmon (Salmo salar) undergo a distinct anadromous life cycle characterized by sequential freshwater and marine phases. Spawning occurs in freshwater rivers during late autumn to winter, with females excavating gravel nests (redds) where 5,000–8,000 eggs per female are deposited and fertilized externally by males. Eggs incubate within the redds for 150–250 days, influenced by water temperature (optimal 4–10°C), hatching primarily in spring as alevins.[11][4] Alevins, measuring 20–25 mm and bearing a yolk sac for endogenous nutrition, remain concealed in the gravel interstices for 2–8 weeks until yolk absorption is complete, minimizing predation risk during this vulnerable phase. Emergence follows as fry, which initiate active foraging on drift organisms like zooplankton and aquatic insects, typically at lengths of 25–35 mm; high mortality rates (up to 90%) occur due to predation and environmental stressors. Fry soon transition to the parr stage, developing 8–11 dark vertical parr marks for crypsis against stream substrates, and inhabit riffles or pools with gravel-cobble beds.[4][12] Parr growth in freshwater lasts 1–5 years (median 2–3 years), during which they reach 10–20 cm, feeding diurnally on macroinvertebrates via ambush tactics from cover; density-dependent competition and temperature regulate cohort size and development timing. Smoltification, a hormonally driven metamorphosis, occurs in spring for select parr (size threshold ~12 cm), replacing parr marks with a silvery scales, enhancing osmoregulatory capacity for marine migration; smolts descend rivers, enter estuaries, and proceed to oceanic feeding grounds.[11][44] In the ocean, post-smolts undergo rapid somatic growth, consuming schooling forage fish and euphausiids, attaining maturity after 1–4 years at sea (grilse for one-winter fish; multi-sea-winter for larger adults up to 150 cm and 20+ kg). Homing adults re-enter natal rivers, often leaping obstacles, to spawn; iteroparity is rare, with most semelparous individuals succumbing post-spawning from exhaustion and osmoregulatory failure, though survival rates for kelts can reach 20–40% in some populations.[11][45] [11]

Diet, Feeding, and Trophic Role

Juvenile Atlantic salmon (Salmo salar) in freshwater habitats primarily consume aquatic invertebrates, including larvae of mayflies (Ephemeroptera), stoneflies (Plecoptera), chironomids, caddisflies (Trichoptera), and blackflies (Simuliidae), as well as annelids and mollusks.[36] Fry initially feed on microscopic plankton and detritus shortly after emergence, transitioning to more active foraging as parr, where they ambush drifting prey from streambed cover using visual cues in clear, oxygenated waters.[46] This benthically oriented, opportunistic feeding supports rapid growth, with daily rations often exceeding 5% of body weight during peak periods.[46] Upon seaward migration as smolts, Atlantic salmon undergo a dietary shift to marine prey, initially consuming plankton before larger individuals prey on schooling fish such as herring (Clupea harengus), capelin (Mallotus villosus), and sprat (Sprattus sprattus), alongside crustaceans including krill (Meganyctiphanes norvegica), amphipods, and mesopelagic shrimp.[11][47] Adult salmon exploit over 40 fish species and at least 10 invertebrate taxonomic groups, with diet composition varying by region, season, and size; for instance, in the Northwest Atlantic, herring and shrimp dominate stomach contents during summer feeding migrations.[48] Feeding ceases several months prior to spawning, relying on stored lipids accumulated at sea.[46] In freshwater, juveniles function as secondary consumers, exerting predation pressure on benthic macroinvertebrates and potentially limiting their abundance through density-dependent effects.[49] In marine phases, adults occupy higher trophic positions (typically 3.5–4.0 based on stable isotope analysis), preying on primary and secondary consumers to regulate forage fish stocks and contributing to energy transfer up the pelagic food web.[49] The anadromous lifecycle enables cross-ecosystem nutrient subsidies, as returning adults deposit marine-derived nitrogen and phosphorus via excretion (up to 20–25% of body mass as gametes and waste) and carcasses, boosting algal biomass by 20–200%, invertebrate production, and supporting resident fish and bird populations in nutrient-poor rivers.[50][51] Declines in salmon abundance have cascading effects, reducing these subsidies and altering trophic dynamics in affected watersheds.[50]

Behavior, Migration, and Physiology

Juvenile Atlantic salmon (Salmo salar) parr in freshwater streams display territorial behavior, defending discrete feeding positions through aggressive interactions such as chasing and biting.[52] In controlled tests, parr aggression varies individually, with classifications into high-, medium-, low-, and zero-aggression groups based on mirror image stimulation; high-aggression fish exhibit peak striking durations around the sixth minute of observation, while low-aggression individuals peak earlier with reduced intensity.[53] Aggression decreases in slow water flows below 5 cm/s, prompting parr to hide in substrate rather than defend territories.[54] In the marine environment, adults shift to schooling formations alongside residual territorial tendencies.[55] Migration begins with smolt outmigration from rivers, where post-smolts employ rheotaxis to navigate downstream, reorienting directionally in response to current reversals at velocities exceeding a threshold of 8.9 cm/s (95% CI: 7.7–10.4 cm/s).[56] This current-based orientation facilitates initial marine dispersal, often along surface flows in coastal zones.[56] Adults undertake upstream homing to natal rivers for spawning, guided by olfactory cues from differentially expressed genes responsive to environmental odors, supplemented potentially by geomagnetic and hydrodynamic signals.[57] Physiological adaptations center on smoltification, a developmental transition in spring where freshwater-adapted parr remodel for seawater tolerance, marked by elevated gill Na⁺/K⁺-ATPase activity achieving a seawater-to-freshwater isoform ratio of at least 2.0 for effective ion regulation.[58] This involves isoform shifts from α1a (freshwater-dominant) to α1b (seawater-active), alongside morphological silvering and a 16.6% reduction in condition factor (K) in age-0 smolts, enabling hypo-osmoregulation through chloride cell proliferation for salt extrusion.[58] Behavioral correlates include increased activity and downstream orientation, with the process exhibiting moderate heritability (h² = 0.23–0.26), allowing genetic selection for synchronized timing.[58] Salinity shifts post-smoltification provoke transient stress responses, including cortisol elevation, underscoring the precision required for successful oceanic transition.[59]

Reproduction and Genetic Considerations

Atlantic salmon (Salmo salar) reproduce through anadromous spawning, where sexually mature adults migrate from marine environments back to their natal freshwater rivers in late autumn, typically October to November in the Northern Hemisphere. Females select gravelly substrates in upstream riffles to construct nests known as redds by turning on their sides to excavate depressions approximately 0.5–1 meter in diameter and 20–30 cm deep, displacing stones with their caudal fin. Eggs, numbering from 3,500 to 18,000 per female depending on body size, are released in batches and externally fertilized by one or more males, which compete aggressively using displays and physical contests; the eggs are then covered with gravel for protection and oxygenation. Larger females exhibit higher fecundity and produce larger eggs, with egg diameter positively correlating with egg number within river-specific cohorts, though overall fecundity scales with female length and prior sea wintering experience.[60][61][62] Egg development proceeds over 150–250 days at water temperatures of 4–8°C, with hatching typically occurring in spring (March–May), yielding alevins that remain buried in the gravel absorbing yolk sacs for 4–6 weeks before emerging as fry. Hatching success varies with gravel quality, water flow, and temperature, but pre-fertilization gamete exposure to suboptimal thermal regimes can reduce it by up to 20% through impaired embryo metabolism and developmental abnormalities. While most salmon semelparous in practice due to post-spawning mortality from exhaustion and predation, iteroparity occurs in up to 10–20% of survivors in some populations, identifiable via scale circuli analysis combined with DNA profiling, though repeat spawners contribute disproportionately to future generations owing to their experience and size.[63][64][65] Genetically, Atlantic salmon exhibit fine-scale population structure tied to natal rivers, with distinct local adaptations for migration timing, disease resistance, and smoltification driven by natural selection over thousands of years, as evidenced by genomic scans revealing outlier loci under divergent pressures. This diversity underpins resilience to environmental stochasticity, yet aquaculture-escaped farmed strains, selectively bred for rapid growth and reduced aggression over 10–12 generations, pose risks through hybridization; introgression rates in wild populations near farms reach 20–50% in affected rivers, eroding adaptive genetic variation and reducing wild offspring fitness by 10–30% in hybrid crosses due to maladaptive traits like poorer predator avoidance and ocean survival. Empirical studies confirm farmed genotypes exhibit 15–25% lower lifetime reproductive success in wild conditions, amplifying genetic pollution where escape events exceed 1% of farm biomass annually, as documented in Norwegian and Scottish systems. Conservation strategies thus prioritize stock-specific broodstock and genomic monitoring to mitigate dilution of wild gene pools, with peer-reviewed models estimating that sustained introgression could halve effective population sizes within decades absent intervention.[66][67][68]

Ecological Interactions

Atlantic salmon (Salmo salar) serve as key connectors in aquatic food webs, facilitating the transfer of marine-derived nutrients to freshwater systems upon spawning death, which elevates lipid content and alters fatty acid profiles across multiple trophic levels in rivers, thereby subsidizing invertebrate and fish communities.[51] This nutrient input, documented in studies of spawning runs, can increase benthic invertebrate biomass by up to 25% in salmon-influenced streams, supporting higher-order consumers.[69] Juvenile salmon in freshwater stages, particularly parr, function as predators on macroinvertebrates such as chironomid larvae and ephemeropterans, exerting size-dependent top-down pressure that influences benthic community structure.[46] Their diet shifts ontogenetically, with larger parr incorporating more fish prey, which can intensify inter- and intraspecific competition for resources.[70] In marine phases, adults occupy a higher trophic position (approximately 4.0–4.5 via stable isotope analysis), preying on clupeids, gadoids, and crustaceans like euphausiids, contributing to pelagic food web dynamics.[49] Predation pressure on Atlantic salmon varies by life stage and habitat. In rivers, alevins and fry suffer high mortality from piscivorous fish, amphibians, and birds including common mergansers (Mergus merganser) and great blue herons (Ardea herodias), with predation rates exceeding 50% in some cohorts.[71] Smolts and post-smolts en route to sea face avian predators like gulls and cormorants, as well as riverine mammals such as otters (Lutra lutra).[72] At sea, adults encounter marine predators including harbor seals (Phoca vitulina), Atlantic cod (Gadus morhua), and mackerel (Scomber scombrus), where tag-return data indicate annual ocean mortality from predation averaging 20–40% for first-sea-winter fish.[73] Competitive interactions primarily occur with congeneric species and other salmonids sharing natal streams or feeding grounds. Brown trout (Salmo trutta) compete aggressively for territorial space and drift-feeding positions, reducing salmon parr growth by 10–20% in mixed populations through resource partitioning failures.[46] In estuarine and coastal zones, overlap with herring (Clupea harengus) and sprat (Sprattus sprattus) leads to exploitative competition for zooplankton, potentially depressing salmon condition factors during early marine residency.[11] Introduced species, such as rainbow smelt (Osmerus mordax) in lacustrine habitats, disrupt size-based predator-prey equilibria, with smelt predation on salmon parr documented in systems like the Great Lakes.[74] These interactions underscore salmon's vulnerability to density-dependent effects, where high predator densities amplify compensatory mortality, while nutrient subsidies from carcasses mitigate bottom-up limitations in oligotrophic rivers.[75] Empirical models from long-term monitoring indicate that balanced predator control can enhance salmon recruitment by 15–30% in predator-saturated systems, highlighting the need for ecosystem-level management.[71]

Threats and Population Dynamics

Natural Predators and Mortality Factors

Atlantic salmon (Salmo salar) experience significant natural mortality across life stages, primarily driven by predation, with rates varying by habitat and size. In freshwater, eggs and alevins are vulnerable to predation by benthic invertebrates, small fish such as sculpins, and opportunistic birds, contributing to early-life mortality exceeding 50% in many rivers.[76] Juvenile parr face threats from piscivorous fish like pike (Esox lucius) and brown trout (Salmo trutta), as well as avian predators including goosanders (Mergus merganser) and red-breasted mergansers (Mergus serrator), which consume substantial numbers of young salmon; studies of merganser diets indicate salmonids comprise up to 20-30% of their intake in salmon rivers.[77][76] Mammalian predators such as otters (Lutra lutra) also target parr and smolts in streams, though their impact is localized.[78] During seaward migration as smolts, predation intensifies due to behavioral changes and exposure in estuaries, where seals (Phoca vitulina and Halichoerus grypus) and piscivorous fish like striped bass (Morone saxatilis) and smallmouth bass (Micropterus dolomieu) account for elevated mortality; acoustic tagging reveals predation rates on smolts can reach 10-20% in bottleneck areas near seal haul-outs.[79][80] Cormorants (Phalacrocorax carbo) and gulls further contribute, with documented consumption of emigrating smolts.[77] Marine survival from smolt to adult is low, often below 5-10% in recent decades, with predation implicated as a key proximate factor alongside environmental stressors like low sea temperatures that may concentrate prey vulnerability.[81][11] In the oceanic phase, adult salmon are preyed upon by a broader array of endothermic predators, including Atlantic cod (Gadus morhua), porbeagle sharks (Lamna nasus), and marine mammals such as harbor seals and killer whales (Orcinus orca), with tag recovery data showing endothermic fish responsible for over half of detected predation events in areas like the Gulf of St. Lawrence.[82] Toothed whales and seals target returning adults near river mouths, exacerbating post-oceanic mortality.[71] Beyond predation, natural mortality includes high post-spawning die-off, where 90-95% of adults perish after reproduction due to exhaustion and associated physiological stress, a semelparous trait in most populations.[44] Environmental factors like extreme freshwater flows or ice scour can destroy redds (nests), causing cohort-wide losses, while density-dependent competition and starvation affect juvenile survival in overcrowded streams.[78] Overall, cumulative natural mortality shapes population dynamics, with freshwater stages experiencing 80-90% loss before smolting and marine phases adding further attrition.[11]

Anthropogenic Impacts

Human activities have profoundly influenced Atlantic salmon populations through direct exploitation and habitat modification. Commercial overfishing contributed to sharp declines, with North Atlantic salmon numbers falling from approximately seven million to five million between 1983 and 1990.[8] Historical records indicate even earlier collapses, such as in certain European rivers where annual catches dropped from 300-350 fish in the 14th century to just three or four by the late 15th century due to intensive harvesting.[83] Since the 1980s, sustained commercial and recreational fisheries have exacerbated returns to many wild stocks, leading to collapses in multiple regions.[84] Hydropower dams and river regulation severely disrupt migration patterns essential for the species' anadromous life cycle. These structures block access to spawning grounds, fragment habitats, and alter downstream flows, sediment transport, and water temperatures, which degrade rearing conditions and increase vulnerability to predators.[85] Salmon delayed at dams expend energy reserves in warmer waters, depleting fat stores needed for spawning, as observed in studies from the Penobscot River system.[86] Fish passage facilities, such as lifts and ladders, often fail to fully mitigate these effects, with downstream smolt mortality elevated due to turbine passage and predation in reservoirs.[87] In regulated rivers, such alterations have reduced available spawning and rearing areas, contributing to persistent population declines.[88] Escapes from aquaculture facilities pose a genetic threat via introgression into wild populations. Farmed Atlantic salmon, selectively bred for traits like rapid growth, exhibit lower fitness in natural environments, yet interbreeding dilutes adaptive wild genotypes, affecting the full life cycle from juveniles to adults.[89] Genetic analyses reveal introgression in up to 67% of monitored populations, with no changes detected in only 33%, positioning escaped farmed fish as the primary anthropogenic genetic risk.[89] Notable escape events, such as nearly 75,000 farmed salmon in Scotland following Storm Amy in 2021, amplify this issue, potentially causing lasting hybridization in fragile stocks.[90] Additionally, escaped fish transmit diseases and compete for resources, further pressuring wild salmon.[91] Pollution from industrial, agricultural, and urban sources degrades water quality and habitat suitability. Historical acidification from acid rain and other contaminants reduced spawning success and juvenile survival, though improvements in some areas have aided partial recovery.[92] Land-use practices, including deforestation and agriculture, increase sedimentation and nutrient loads, smothering redds and altering prey availability.[93] Emerging pollutants like pharmaceuticals disrupt migratory behavior and physiological processes in salmon, as evidenced by altered smolt migration timing in contaminated rivers.[94] Farm effluents contribute localized benthic impacts near net pens, though recovery occurs post-fallow periods.[95] These cumulative effects compound other pressures, hindering population resilience.

Disease and Parasite Dynamics

Atlantic salmon (Salmo salar) are susceptible to a range of viral, bacterial, and parasitic pathogens that influence population dynamics in both wild and farmed contexts, often exacerbating mortality during critical life stages such as smolt migration.[96] Parasitic infections, particularly from sea lice (Lepeophtheirus salmonis), have been linked to significant post-smolt mortality in wild populations, with experimental evidence indicating up to 39% reduction in marine survival due to lice-induced osmoregulatory failure and secondary infections.[97] Bacterial diseases like furunculosis, caused by Aeromonas salmonicida subsp. salmonicida, manifest as systemic infections leading to hemorrhaging and tissue necrosis, historically causing epizootics in dense aquaculture settings and spilling over to wild stocks via waterborne transmission.[98] Viral pathogens, including infectious salmon anaemia virus (ISAV), induce severe anemia and immunosuppression primarily in farmed fish, with virulence factors enabling persistence in carriers and outbreaks documented since the 1980s in Norway and subsequent spread to other regions.[99] Sea lice dynamics are driven by high infestation pressures from salmon farms, where adult female lice release planktonic larvae that infect migrating wild smolts, reducing condition and inducing premature returns or death; correlations between farm lice counts and declines in wild catches have been observed across Norwegian salmon rivers, with lice abundance increasing infection risk by orders of magnitude near aquaculture sites.[100] [101] The introduced monogenean Gyrodactylus salaris exemplifies parasite invasion dynamics, proliferating rapidly on naive salmon hosts in Scandinavian rivers since the 1970s, causing up to 100% mortality in affected populations through direct skin damage and osmoregulatory stress, with over 50 Norwegian rivers impacted as of 2024.[102] Amoebic gill disease (AGD), induced by Neoparamoeba perurans, recurs in marine-farmed salmon, leading to gill hyperplasia and hypoxia; host-parasite interactions involve mucus hypersecretion and immune modulation, with prevalence tied to environmental stressors like temperature rises.[103] Transmission dynamics often involve farm-to-wild spillovers, where high-density aquaculture amplifies pathogen loads; for instance, ISAV survives in seawater for weeks under cool conditions, facilitating horizontal spread and vertical transmission in eggs, contributing to farm-level mortality rates exceeding 90% in outbreaks.[104] Furunculosis exhibits facultative intracellular behavior in salmon macrophages, evading early immune responses and persisting in chronic carriers, with genomic studies revealing re-emergent strains in Atlantic Canadian farms as of 2024 adapted to vaccine pressures.[105] Parasite burdens interact with host physiology, such as smoltification stress reducing tolerance thresholds, leading to context-dependent mortality where co-infections amplify effects; selective pressures from these dynamics have prompted genetic resistance breeding, though wild stocks remain vulnerable to novel introductions.[106] Overall, these interactions underscore causal links between intensified aquaculture and elevated disease risks to wild salmon, with empirical models predicting sustained population declines absent mitigation.[107]

Human Interactions and Utilization

Commercial Fisheries and Harvesting

Commercial fisheries for wild Atlantic salmon operate primarily in the North Atlantic, targeting migratory adults and post-smolts in coastal and estuarine waters, though activities are severely restricted due to population declines and conservation imperatives. Major regions include West Greenland, where mixed-stock fisheries intercept salmon from North American and European origins; Russia, particularly the Kola Peninsula rivers; Iceland; and limited areas in Canada such as Quebec and Newfoundland, with many North American operations closed since the 1990s.[8][108] In the United States, commercial harvesting is prohibited under federal law, reflecting the endangered status of Gulf of Maine stocks.[109] Harvesting methods emphasize selective gears to reduce bycatch and comply with quotas, including fixed gillnets anchored to shores, trap nets, and weirs in rivers that allow enumeration and escapement of spawners. In Greenland, commercial fishers deploy up to 20 gillnets per vessel, with driftnets banned since earlier regulations; catches are often for local consumption rather than export.[110] These practices stem from international agreements under the North Atlantic Salmon Conservation Organization (NASCO), which mandates total allowable catches (TACs) based on pre-fishery abundance indices from ICES assessments.[111] In 2023, ICES reported record-low catches across NASCO areas, underscoring ongoing stock vulnerability despite management efforts. Greenland's West Greenland TAC was 27 tonnes, with reported landings of 34.3 tonnes (33 tonnes west, 1.3 tonnes east), equivalent to roughly 7,000-10,000 fish depending on size.[112][113][114] North American commercial harvests remain negligible following Canada's closures, which have supported positive trends in adult returns, such as record highs in Labrador.[108] Historical peaks exceeded 3.5 million fish in 1973, primarily from interceptory fisheries, but overharvesting contributed to multi-decadal declines, prompting quota reductions and moratoria. Current wild catches total under 2,000 tonnes annually, a fraction of farmed production surpassing 2.5 million tonnes, with wild Atlantic salmon comprising less than 1% of market supply.[8][115] This shift underscores aquaculture's dominance while highlighting regulatory successes in curbing exploitation, though non-fishing mortality factors limit recovery.[116]

Aquaculture Production

Aquaculture production of Atlantic salmon (Salmo salar) dominates global supply, with farmed output exceeding wild capture by a factor of over 70 percent as of 2023.[115] Commercial farming originated in Norway during the 1960s, building on earlier 19th-century freshwater stocking efforts in the United Kingdom to enhance wild fisheries.[117] By 1990, worldwide production stood at 230,000 tonnes, expanding to over 2.2 million tonnes by the 2020s through advancements in hatchery techniques, smolt transfer to marine sites, and optimized feeds.[118] This growth reflects a more than 1,000 percent increase since 1990, driven by selective breeding for faster growth and disease resistance.[119] Norway leads production, accounting for more than 50 percent of global output as the world's largest farmed salmon producer, followed by Chile at approximately 27-32 percent, with smaller contributions from Scotland (part of the UK at 5 percent), Canada, and the Faroe Islands.[120] [121] Norwegian farmed Atlantic salmon is widely regarded as one of the world's best-tasting salmons due to strict farming standards, clean cold waters, high-quality feed, and resulting firm texture, rich flavor, and high fat content. It is highly prized globally, particularly in Asia and Europe. However, taste preferences are subjective, with some preferring wild Alaskan salmon for its natural diet and flavor. In 2023, total farmed salmonid production surpassed 2.8 million tonnes, predominantly Atlantic salmon.[115] For January to September 2024, global Atlantic salmon supply reached about 2.02 million tonnes, a 1 percent decline year-over-year due to biological constraints like sea lice and pancreas disease in key regions.[122] Harvest volumes in Norway hit record highs in 2023 at 1.53 million tonnes, while Chile recovered to 0.8 million tonnes post-2019 infectious salmon anemia outbreaks.[119] The predominant method involves open net pens or sea cages deployed in coastal marine waters, where juvenile salmon smolts—reared initially in freshwater hatcheries—are transferred to grow to market size over 12-18 months.[123] These floating enclosures, typically 50-200 meters in circumference, allow natural water exchange for oxygenation and waste dispersal but expose fish to environmental pathogens and predators.[124] Emerging alternatives include land-based recirculating aquaculture systems (RAS), which recycle up to 99 percent of water and enable year-round control of conditions, though higher energy costs limit scalability; RAS currently produce less than 1 percent of global salmon but are expanding in regions like Canada and Norway for post-smolt rearing.[125] [126] Semi-closed containment systems, such as in-sea floating bags, offer partial barriers against parasites while retaining open-ocean benefits, with pilot projects demonstrating viability in Norwegian fjords.[124] Production efficiency has improved via triploidy and genetic selection programs, yielding fish that convert feed to biomass at a 1.1-1.3 ratio, though escapes from net pens—estimated at 0.1-0.5 percent of stocked fish annually—pose risks to wild stocks through interbreeding and disease transmission.[119] Industry leaders like Mowi, originating from Norwegian pioneers, integrate vertical operations from broodstock to processing, exporting to over 70 countries and emphasizing biosecurity to mitigate outbreaks that reduced Chilean output by 40 percent in 2016-2017.[127] Despite regulatory pressures for closed systems in Norway—capping open-pen growth at 6 percent annually through 2025—projected global supply anticipates modest increases to 2.5 million tonnes by 2030, supported by innovation in vaccine delivery and feed alternatives to fishmeal.[126][119]

Nutritional Value and Health Implications

Atlantic salmon provides high-quality protein and essential nutrients, including omega-3 fatty acids, vitamin D, vitamin B12, and selenium. A 100-gram serving of raw farmed Atlantic salmon contains approximately 208 calories, 20 grams of protein, 13 grams of total fat (predominantly unsaturated), and negligible carbohydrates.[128] It supplies about 2-3 grams of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), the key long-chain omega-3 fatty acids, which constitute a significant portion of its lipid profile.[129] Wild Atlantic salmon, being leaner, offers fewer total calories (around 142 per 100 grams) and lower absolute omega-3 content due to reduced fat (5-8 grams per 100 grams), though its omega-3 to omega-6 ratio is more favorable (approximately 0.05 versus 0.7 in farmed).[129][130] Regular consumption of Atlantic salmon supports cardiovascular health through its omega-3 content, which lowers triglycerides, reduces inflammation, and decreases the risk of coronary heart disease and stroke when eaten 1-2 times weekly.[131][132] DHA from salmon contributes to brain health by maintaining neural membrane integrity and potentially reducing cognitive decline risk in older adults.[133][134] Astaxanthin, a carotenoid pigment in salmon, acts as an antioxidant, offering additional benefits for heart and nervous system function.[133] Selenium and vitamin D further aid immune function and bone health, with salmon providing over 50% of the daily recommended vitamin D intake per serving.[135] Farmed Atlantic salmon may contain elevated levels of persistent organic pollutants (POPs) such as polychlorinated biphenyls (PCBs) and dioxins compared to wild counterparts, with farmed fillets averaging up to eight times higher PCB concentrations due to feed sources and higher fat content.[136] However, recent monitoring (post-2010) indicates these levels have declined substantially—often below European Union maximum residue limits—and pose minimal cancer or noncancer risks at recommended intake levels of 150-250 grams weekly.[129][137] Mercury concentrations remain low across both farmed and wild salmon (typically under 0.05 mg/kg), far below thresholds that would contraindicate consumption.[138] Overall, the nutrient density and omega-3 benefits outweigh contaminant risks for moderate eaters, though opting for wild salmon minimizes POP exposure where available.[139][140]

Economic and Cultural Importance

![Atlantic salmon total production million tonnes 1975-2022.svg.png][float-right] The Atlantic salmon industry, dominated by aquaculture, generates substantial economic value globally, with farmed production exceeding 2 million metric tons annually as of recent years. Norway leads as the largest producer, exporting 1.2 million tons of salmon valued at 122.5 billion Norwegian kroner (approximately 11.5 billion USD) in 2023, supporting tens of thousands of direct and indirect jobs in coastal regions.[141] Chile follows as a major exporter, shipping 782,076 tons of salmon and trout in 2024, contributing significantly to national GDP through foreign exchange earnings and employment in southern aquaculture zones.[142] In Canada, the sector adds to regional economies, though imports from Norway highlight competitive dynamics, with exported salmon value rising to 42.8 million CAD in the first half of 2024.[143] Wild capture fisheries, while smaller, provide economic benefits through commercial harvests in rivers like those in North America and Europe, alongside recreational angling that generates millions in spending; for instance, high-impact angling camps in Canada alone accounted for 26 million USD in expenditures in 2010.[144] Culturally, Atlantic salmon holds symbolic importance in North Atlantic societies, representing resilience and natural abundance in folklore and traditions. In indigenous communities of eastern North America, such as those along the Miramichi River, salmon feature in stewardship practices and oral histories, akin to broader salmonid reverence, though less documented than Pacific species.[145] European angling heritage, exemplified by 17th-century texts like Izaak Walton's The Compleat Angler, elevates salmon as a prized quarry, fostering traditions of fly-fishing and river management in Scotland, Ireland, and Norway.[146] These cultural roles extend to cuisine, where smoked or fresh salmon forms staples in Nordic and Celtic diets, reinforcing seasonal festivals and community identities tied to migratory cycles.[145]

Conservation Status and Efforts

Global and Regional Population Status

The wild Atlantic salmon (Salmo salar) population across the North Atlantic has declined substantially since the mid-20th century, with an estimated reduction of approximately 23% over the past three generations, prompting the International Union for Conservation of Nature (IUCN) to reclassify the species from Least Concern to Near Threatened in December 2023.[7] This downgrade reflects ongoing pressures including marine mortality, habitat degradation, and mixed-stock fisheries, with annual returns to rivers now estimated at fewer than 2 million adults, down from peaks exceeding 7 million in the early 1980s.[8] [147] The International Council for the Exploration of the Sea (ICES) reported record-low catches in 2023 and persistently low returns in 2024, advising zero exploitation for many stocks due to their poor status. [148] In North America, wild populations are critically depleted, with the Gulf of Maine Distinct Population Segment listed as endangered under the U.S. Endangered Species Act since 2000, confined primarily to rivers in central and eastern Maine where adult returns have not exceeded 2,000 annually in recent years despite restoration efforts.[11] In Canada, 2023 adult returns totaled about 668,600 fish, exceeding recent averages in some areas but with record-low numbers of small (1-sea-winter) salmon in Quebec and Newfoundland, and several rivers like the Restigouche and Miramichi falling below conservation limits for egg deposition.[149] [150] Overall, North American stocks have shown no recovery from 1990s lows, exacerbated by aquaculture escapes introducing genetic risks.[151] European stocks exhibit similar declines, with wild Atlantic salmon in Great Britain reclassified as endangered by the IUCN in 2023 due to a 30-50% population drop since 2006 and projected further losses of 50-80% by 2025 in some regions.[152] In England, 88% of principal salmon rivers were rated at risk or probably at risk in 2024 assessments, while Scotland recorded its lowest rod catch since 1952 at 32,477 fish in 2023.[153] [154] Continental Europe, including France and Norway, reports mixed but generally downward trends, with ICES noting harvesting closures recommended for over 80% of assessed stocks in 2024 owing to failure to meet conservation thresholds.[155] These regional patterns underscore a basin-wide crisis, with post-smolt marine survival rates dropping to historic lows since the 2010s.[156]

Major Threats to Wild Stocks

Interactions between escaped farmed Atlantic salmon and wild populations pose a significant genetic threat through introgression, reducing fitness and adaptability in wild stocks; in Norway, escaped farmed salmon and related infections rank among the largest threats to wild salmon viability.[157] Salmon lice originating from aquaculture facilities inflict high mortality on wild post-smolts during marine migration, with infestation levels exceeding tolerance thresholds in multiple regions; Norwegian assessments identify salmon lice as the primary threat, linked to farming density.[158] Disease transmission from farms, including pathogens like infectious salmon anemia, further exacerbates declines, as evidenced by spillover events correlating with reduced wild survival rates.[159] Habitat degradation from barriers such as dams and weirs impedes upstream migration and access to spawning grounds, contributing to population fragmentation; in North America, altered freshwater habitats from land-use practices have degraded essential rearing areas.[160] Water pollution, including nutrient runoff and contaminants, affects all life stages by reducing oxygen levels and increasing toxicity, with direct exposure linked to elevated mortality in juveniles and adults.[161] Climate change amplifies vulnerabilities across the life cycle, with rising sea and river temperatures compressing migration windows and reducing prey availability; global populations have declined by approximately 23% over three generations partly due to these thermal stresses.[7] Marine survival has plummeted, with post-smolt mortality rates increasing amid warmer oceanic conditions and shifting predator-prey dynamics; a 2022 review of marine stressors ranks climate change as the foremost current and projected threat over the next decade.[39] Exploitation through mixed-stock fisheries and bycatch continues to pressure remnant populations, despite regulatory reductions; in the marine phase, incidental capture in non-salmon fisheries compounds losses.[162] Predation by species such as seals and birds has intensified in some areas due to environmental changes, though natural predation alone does not explain observed declines without anthropogenic amplification.[39]

Restoration and Management Strategies

Restoration strategies for Atlantic salmon emphasize habitat rehabilitation to address barriers to migration and spawning, including dam removals, construction of fish passes, and riparian zone enhancements. In the Gulf of Maine, ongoing projects funded by NOAA, such as those receiving $1.2 million in 2023, target river connectivity and water quality improvements to support endangered populations.[163] These efforts aim to reconnect rivers to oceans, fostering resilient ecosystems that benefit salmon and associated species.[164] Hatchery and stocking programs have played a role in bolstering populations, particularly in regions like New England where they have averted local extinctions, though evidence indicates potential genetic dilution and reduced fitness in wild stocks when not managed carefully.[164] Guidelines from the North Atlantic Salmon Conservation Organization (NASCO) recommend stocking only for extirpated or critically low populations, prioritizing wild fry transplants over mass releases to minimize risks.[165] Peer-reviewed syntheses highlight that successful outcomes often require integration with habitat fixes, as isolated stocking yields limited long-term gains.[166] Management frameworks incorporate strict regulatory measures, such as prohibitions on commercial and recreational harvest in U.S. waters since the 1990s to allow stock recovery.[11] In Europe and North America, NASCO-coordinated catch limits and enforcement beyond 12 nautical miles from coasts prevent overexploitation in mixed-stock fisheries.[167] Monitoring programs track adult returns and juvenile survival, informing adaptive strategies like flow regime adjustments in hydropower-regulated rivers to mimic natural conditions.[168] These combined approaches, grounded in population viability analyses, seek to restore self-sustaining runs while accounting for anthropogenic pressures.[169]

International Cooperation and Agreements

The North Atlantic Salmon Conservation Organization (NASCO) serves as the primary international body for the conservation and management of Atlantic salmon (Salmo salar) stocks, established under the Convention for the Conservation of Salmon in the North Atlantic Ocean, signed on March 2, 1982, and entering into force on October 1, 1983. The convention aims to promote the conservation, restoration, enhancement, and rational management of salmon stocks through international cooperation among states whose nationals fish for salmon or in whose rivers salmon migrate.[170] It prohibits commercial fishing for salmon in international waters of the North Atlantic, limiting such activities to coastal zones generally within 12 nautical miles of baselines, with exceptions allowing Greenland up to 40 nautical miles and the Faroe Islands specific historical fisheries.[167] NASCO's membership includes Canada, the United States, the European Union, the United Kingdom, Norway, the Russian Federation, Iceland, and Denmark (representing Greenland and the Faroe Islands), enabling coordinated efforts across the species' range.[171] The organization has developed a series of binding resolutions, agreements, and guidelines addressing key challenges, including the application of a precautionary approach to fisheries management, protection of salmon habitat, regulation of aquaculture impacts such as escaped farmed fish and sea lice infestations on wild stocks, and minimization of incidental catches in mixed-stock fisheries.[172] For instance, the West Greenland Commission under NASCO facilitates agreements to manage interceptory fisheries, such as a 2022 commitment by stakeholders to improve monitoring and reduce unreported catches in Greenland's salmon fishery.[171] Complementing NASCO, Article 66 of the United Nations Convention on the Law of the Sea (UNCLOS), adopted in 1982, obligates states to cooperate in the conservation and management of anadromous species like Atlantic salmon, with home states bearing primary responsibility and prohibiting fishing on the high seas.[173] NASCO collaborates with scientific bodies, including the International Council for the Exploration of the Sea (ICES), which provides independent stock assessments and advice free from political influence to inform annual regulatory measures. These frameworks emphasize data-driven catch limits and restoration targets, though implementation varies by jurisdiction, reflecting the transboundary nature of salmon migrations and the need for ongoing bilateral and multilateral enforcement to counter overexploitation and environmental pressures.[174]

Regulatory Frameworks

North American Regulations

In the United States, the Gulf of Maine Distinct Population Segment of Atlantic salmon (Salmo salar) was listed as endangered under the Endangered Species Act on November 17, 2000, encompassing all native populations in rivers from the Androscoggin River eastward to the Dennys River.[175] This designation prohibits take, including commercial and recreational harvest, possession, and sale of wild Atlantic salmon, with federal agencies required to ensure actions do not jeopardize recovery.[164] NOAA Fisheries implements the Atlantic Salmon Fishery Management Plan, originally established in 1988 and updated to align with Endangered Species Act recovery plans, which mandates habitat protections such as dam removals or modifications to restore access to over 11,000 kilometers of historical spawning habitat and prohibits retention of incidentally caught salmon in commercial fisheries targeting other species.[176][109] State-level regulations, coordinated through bodies like the U.S. Fish and Wildlife Service, further restrict angling to catch-and-release in designated restoration rivers, with no bag limits permitting retention.[177] In Canada, Fisheries and Oceans Canada (DFO) regulates Atlantic salmon under the Fisheries Act, with commercial fisheries progressively closed across most regions starting in the 1980s—fully prohibited in the inner Bay of Fundy by 1985 and expanded nationwide by the early 2000s—to halt declines attributed to overexploitation and habitat degradation.[178][179] Recreational fisheries emphasize conservation, requiring catch-and-release in threatened populations; for example, in Nova Scotia's Salmon Fishing Area 18, the daily limit is four salmon, all released, during provincially set open seasons typically from June to October.[180] Anglers aged 16 and older must obtain a dedicated salmon license, separate from general fishing permits, with provincial variations such as full retention bans in Prince Edward Island rivers to support recovery of designatable units assessed as endangered or threatened.[181][182] Aquaculture regulations in North America prioritize containment and biosecurity to mitigate risks to wild stocks from escapes and pathogens. In Canada, where Atlantic salmon farming produces over 50% of national aquaculture output primarily in New Brunswick and Nova Scotia bays, DFO enforces site-specific licenses under the Fisheries Act, mandatory disease surveillance reporting, and adherence to Finfish Aquaculture Effluent Regulations limiting waste discharges, alongside voluntary third-party certifications for best practices.[183][184] U.S. operations, though minimal and mostly land-based or in closed systems due to environmental concerns, require FDA approval for veterinary drugs and feeds, with NOAA and EPA oversight to prevent non-native genetic introductions via the National Policy on Genetically Modified Organisms in Aquaculture.[6] Cross-border coordination occurs through memoranda like the 2023 Atlantic Salmon Aquaculture Framework, addressing transfers and monitoring shared coastal waters.[185]

European and Other Regional Laws

The European Union participates in the North Atlantic Salmon Conservation Organization (NASCO) through the Convention for the Conservation of Salmon in the North Atlantic Ocean, which prohibits directed fishing for Atlantic salmon outside coastal states' areas of fisheries jurisdiction, limiting such activities to within 12 nautical miles of the coast, with exceptions extending to 40 nautical miles off Greenland and permitting fishing throughout the Faroe Islands' zone.[167][186] This framework, implemented via Council Decisions such as (EU) 2019/864, prioritizes conservation by restricting high-seas interception of migrating stocks.[187] Under the EU Common Fisheries Policy, Atlantic salmon management devolves to member states for coastal and riverine fisheries, but regional seas like the Baltic impose specific controls, including annual total allowable catches (TACs) set by Council regulations.[188] In the Baltic Sea, recreational fishing for wild salmon has been prohibited since at least 2024, with any incidental captures required to be released immediately, alongside gear restrictions and seasonal prohibitions in subdivisions 22-31 to protect declining stocks.[188] For 2025, EU ministers agreed to further limits, including delayed opening seasons in main basin zones and bans on recreational fishing for reared salmon to minimize bycatch impacts.[189] EU aquaculture regulations, harmonized under environmental and health directives rather than exclusive competence, mandate licensing, disease monitoring, and containment measures for salmon farms to curb escapes, which pose risks of disease transmission, resource competition, and genetic introgression with wild populations.[190][191] Member states enforce structural standards for net pens and rapid response protocols for escape events, with ongoing policy updates in 2025 emphasizing ecosystem protection amid production growth.[192] In Norway, outside the EU but a dominant salmon aquaculture producer, the Aquaculture Act of 2005 regulates all farming operations, requiring permits tied to site-specific environmental assessments and maximum allowable biomass limits.[193] A "traffic light" system zones coastal areas green, yellow, or red based on sea lice prevalence and other risks, triggering production halts or relocations in high-risk red zones to safeguard wild salmon.[194] Wild fisheries face stringent national controls, with 2024 seeing multiple rivers closed to angling and strict quotas elsewhere due to conservation limits set by regional directorates.[195] The United Kingdom, post-Brexit, maintains NASCO commitments prohibiting high-seas salmon fishing and enforces domestic protections via byelaws mandating catch-and-release for all rod-caught salmon on rivers classified as vulnerable, affecting most principal salmon rivers where stocks have declined sharply.[196][197] In England, the Environment Agency has intensified enforcement since 2024, including barrier removals and inspections, while Northern Ireland's 2014 regulations require 100% catch-and-release for salmon angling to bolster spawning escapement.[153][198] Aquaculture sites adhere to similar escape prevention and welfare standards, integrated into the UK's Blue Book of fishing regulations.[199]

References

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