View source for Salmon run

{{Short description|Annual migration of salmon}}
{{about|the annual salmon migration|the video game|Salmon Run (video game)|the mode in the Splatoon series|Splatoon 2|and|Splatoon 3}}
{{Use British English|date=August 2021}}
{{Use dmy dates|date=August 2021}}
[[File:Grizzly Bear Fishing Brooks Falls (flipped).jpg|thumb|right|A [[grizzly bear]] ambushing a jumping salmon during an annual salmon run]]
<!-- [[File:Moricetown Canyon Rapids.jpg|thumb|300px|right|{{center|Fishermen capture running salmon with nets<br />before [[Tag and release|tagging and releasing]] them}}]] -->

A '''salmon run''' is an annual [[fish migration]] event where many [[salmonid]] species, which are typically hatched in [[fresh water]] and live most of their adult life downstream in the [[ocean]], swim back against the stream to the upper reaches of rivers to [[spawn (biology)|spawn]] on the [[gravel]] beds of small [[stream|creek]]s. After spawning, most [[Atlantic salmon]]<ref>[https://www.nature.scot/plants-and-animals/fish/freshwater-fish/atlantic-salmon Atlantic salmon], ''[[Scottish Natural Heritage]]''. Retrieved 25 January 2018.</ref> and all species of [[Oncorhynchus|Pacific salmon]]{{citation needed|date=December 2024}} die, and the salmon [[biological life cycle|life cycle]] starts over again with the new generation of [[hatchling]]s.

[[Salmon]] are [[anadromous]], spending their [[juvenile fish|juvenile]] life in rivers or lakes, and then migrating out to sea where they spend [[adult]] lives and gain most of their [[body mass]]. When they reach [[sexual maturity]], the adults return to the upstream rivers to [[reproduce]]. Usually they [[natal homing|return with uncanny precision]] to the natal river where they were born, and even to the very spawning ground of their birth. It is thought that, when they are in the ocean, they use [[magnetoreception]] to locate the general position of their natal river, and once close to the river, that they use their sense of smell to [[homing (biology)|home in on]] the river entrance and even their natal spawning ground.

[[Trout]], which are sister species of salmon, also perform similar migrations, although they mostly move [[potamodromous]]ly between creeks and large freshwater lakes, except for some coastal/[[estuary]] subspecies such as [[steelhead]] and [[sea trout]] that migrate seasonally between [[seawater|salty]]/[[brackish]] and fresh water just like salmon do. There are also landlocked populations of some salmon species that have adapted to spend their entire life in freshwater like trout.

In [[Pacific Northwest|Northwest America]], salmons are [[keystone species]], which means the ecological impact they have on other wildlife is greater than would be expected in relation to their [[biomass (ecology)|biomass]]. Most salmon species migrate during the [[autumn]] (September through November),<ref>{{cite web |title=Questions and Answers About Salmon |url=https://www.usgs.gov/centers/western-fisheries-research-center/questions-and-answers-about-salmon |website=Western Fisheries Research Center |publisher=U.S. Geological Survey}}</ref> which coincides with the pre-[[winter]] activities of many [[hibernating]] animals. The annual salmon run can be a major feeding event for [[predator]]s such as [[grizzly bears]] and [[bald eagle]]s, as well as an important [[fishing season|window period]] for [[recreational fishing|sport fishermen]].

The post-spawning death of salmon also has important ecological consequences, because the significant [[nutrient]]s in their [[carrion|carcass]]es, rich in [[nitrogen]], [[sulfur]], [[carbon]] and [[phosphorus]], are transferred from the ocean and released to inland [[aquatic ecosystem]]s, [[terrestrial animal]]s (such as bears) and the [[wetland]]s and [[riparian]] [[woodland]]s adjacent to the rivers. This has knock-on effects not only for the next generation of salmon, but to every wildlife species living in the riparian zones the salmon reach.<ref name=Helfield2006 /> The nutrients can also be washed downstream into [[estuaries]] where they accumulate and provide significant support for [[invertebrate]]s and estuarine-breeding [[waterbird]]s.

==Background==
{{multiple image
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| image1            = Ocean stage and spawning pink salmon.gif
| alt1              = 
| caption1          = Adult ocean phase and spawning phase pink salmon (male)
| image2            = Atlantic salmon redd.jpg
| alt2              = 
| caption2          = ''Sac fry'' remain in the gravel habitat of their redd (nest) until their yolk sac, or "lunch box", is depleted.
| image3            = Atlantic salmon parr emerging from streambed.jpg
| alt3              = 
| caption3          = After depleting their yolk sac nutrients, the young salmon emerge from the gravel habitat as ''parr'' to feed.
}}

{{See also|Juvenile fish#Juvenile salmon}}

Most salmon are [[anadromous]], a term which comes from the Greek ''anadromos'', meaning "running upward".<ref name=Webster/> Anadromous fish grow up mostly in the saltwater in oceans. When they have matured they [[Fish migration|migrate]] or "run up" freshwater rivers to [[Spawn (biology)|spawn]] in what is called the salmon run.<ref name=Moyle188>[[#Moyle|Moyle]], p. 188</ref>

Anadromous salmon are Northern Hemisphere fish that spend their ocean phase in either the [[Atlantic Ocean]] or the [[Pacific Ocean]]. They do not thrive in warm water. There is only one species of salmon found in the Atlantic, commonly called the [[Atlantic salmon]]. These salmon run up rivers on both sides of the ocean. Seven different species of salmon inhabit the Pacific (see table), and these are collectively referred to as [[Pacific salmon]]. Five of these species run up rivers on both sides of the Pacific, but two species are found only on the Asian side.<ref name=FAQ /> In the early 19th century, Chinook salmon were successfully established in the Southern Hemisphere, far from their native range, in New Zealand rivers. Attempts to establish anadromous salmon elsewhere have not succeeded.<ref name=Walrond/>

{| class="wikitable"
|+ Species of anadromous salmon
|-
! rowspan="2" style="vertical-align: middle;" | Oceans
! rowspan="2" style="vertical-align: middle;" | Coasts
! rowspan="2" style="vertical-align: middle;" | Species<ref name=FAQ/>
! colspan="3" | Maximum
! rowspan="2" style="vertical-align: middle;" | Comment
|-
! length
! weight
! life span
|-
! North Atlantic
| style="text-align: center;" | Both sides
| [[Atlantic salmon]]<ref name=FBAtlantic/>
| style="text-align: right" | 150&nbsp;cm
| style="text-align: right" | 46.8&nbsp;kg
| style="text-align: right" | 13&nbsp;years
| 
|-
! rowspan="8" | North Pacific
| rowspan="5" style="text-align: center; vertical-align: middle;" | Both sides
| style="border-bottom-style: none;" | [[Chinook salmon]]<ref name=FBChinook/>
| style="text-align: right; border-bottom-style: none;" | 150&nbsp;cm
| style="text-align: right; border-bottom-style: none;" | 61.4&nbsp;kg
| style="text-align: right; border-bottom-style: none;" | 9 years
| style="border-bottom-style: none;" | Also established in New Zealand
|-
| style="border-style: none solid;" | [[Chum salmon]]<ref name=FBChum/>
| style="text-align: right; border-style: none solid;" | 100&nbsp;cm
| style="text-align: right; border-style: none solid;" | 15.9&nbsp;kg
| style="text-align: right; border-style: none solid;" | 7 years
| style="border-style: none solid;" |
|-
| style="border-style: none solid;" | [[Coho salmon]]<ref name=FBCoho/>
| style="text-align: right; border-style: none solid;" | 108&nbsp;cm
| style="text-align: right; border-style: none solid;" | 15.2&nbsp;kg
| style="text-align: right; border-style: none solid;" | 5 years
| style="border-style: none solid;" |
|-
| style="border-style: none solid;" | [[Pink salmon]]<ref name=FBPink/>
| style="text-align: right; border-style: none solid;" | 76&nbsp;cm
| style="text-align: right; border-style: none solid;" | 6.8&nbsp;kg
| style="text-align: right; border-style: none solid;" | 3 years
| style="border-style: none solid;" |
|-
| style="border-style: none solid;" | [[Sockeye salmon]]<ref name=FBSockeye/>
| style="text-align: right; border-style: none solid;" | 84&nbsp;cm
| style="text-align: right; border-style: none solid;" | 7.7&nbsp;kg
| style="text-align: right; border-style: none solid;" | 8 years
| style="border-style: none solid;" |
|-
| rowspan="2" style="text-align: center; vertical-align: middle;" |Asian side
| style="border-bottom-style: none;" | [[Masu salmon]]<ref name=FBMasu/>
| style="text-align: right; border-bottom-style: none;" | 79&nbsp;cm
| style="text-align: right; border-bottom-style: none;" | 10.0&nbsp;kg
| style="text-align: right; border-bottom-style: none;" |
| style="text-align: right; border-bottom-style: none;" |
|-
| style="border-top-style: none;" | [[Biwa trout|Biwa salmon]]<ref name=FBBiwa/>
| style="text-align: right; border-top-style: none;" | 44&nbsp;cm  
| style="text-align: right; border-top-style: none;" | 1.3&nbsp;kg
| style="text-align: right; border-top-style: none;" | 
| style="text-align: right; border-top-style: none;" |
|}

The [[Parr (fish)|life cycle]] of an anadromous salmon begins and, if it survives the full course of its natural life, usually ends in a gravel bed in the upper reaches of a stream or river. These are the salmon [[spawning]] grounds where salmon [[Fish egg|eggs]] are deposited, for safety, in the gravel. The salmon spawning grounds are also the salmon nurseries, providing a more protected environment than the ocean usually offers. After 2 to 6 months the eggs hatch into tiny larvae called ''sac fry'' or ''alevin''. The alevin have a sac containing the remainder of the yolk, and they stay hidden in the gravel while they feed on the yolk. When the yolk has gone they must find food for themselves, so they leave the protection of the gravel and start feeding on [[plankton]]. At this point the baby salmon are called ''fry''. At the end of the summer the fry develop into juvenile fish called ''parr''. Parr feed on small [[invertebrate]]s and are camouflaged with a pattern of spots and vertical bars. They remain in this stage for up to three years.<ref name=Bley1988/><ref name=Lindberg2011/>

As they approach the time when they are ready to migrate out to the sea the parr lose their camouflage bars and undergo a process of physiological changes which allows them to survive the shift from freshwater to saltwater. At this point salmon are called ''smolt''. Smolt spend time in the brackish waters of the river estuary while their body chemistry adjusts their [[osmoregulation]] to cope with the higher salt levels they will encounter in the ocean.<ref name=SalmonTrust/> Smolt also grow the silvery scales which visually confuse ocean predators. When they have matured sufficiently in late spring, and are about 15 to 20 centimetres long, the smolt swim out of the rivers and into the sea. There they spend their first year as a ''post-smolt''. Post-smolt form [[Shoaling and schooling|schools]] with other post-smolt, and set off to find deep-sea feeding grounds. They then spend up to four more years as adult ocean salmon while their full swimming ability and reproductive capacity develop.<ref name=Bley1988 /><ref name=Lindberg2011 /><ref name=SalmonTrust />

Then, in one of the animal kingdom's more extreme [[Animal migration|migrations]], the salmon return from the saltwater ocean back to a freshwater river to spawn afresh.<ref name=Crossin2009/>

<!-- {{Image gallery
| title=
| lines=4
| width=160
| height=140
| align=left
| Salmo salar eggs.jpg||Salmon eggs. The growing larvae can be seen through the transparent egg envelope. The black spots are the eyes.
| Salmonlarvakils.jpg||Salmon egg hatching into an ''alevin''. In a few days, the alevin will absorb the yolk sac and become a salmon fry
| SalmonoidsBergeau.jpg||The juvenile salmon, ''parr'', grow up in the relatively protected natal river
| Coho.jpg|| The parr lose their camouflage bars and become ''smolt'' as they become ready for the transition to the ocean
| Salmo salar GLERL 1.jpg||Salmon enter the ocean as ''post-smolt'' and mature into adult salmon. They gain most of their weight in the ocean. 
}} -->
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==Return from the ocean==
<!-- [[File:Running salmon.JPG|thumb|right|{{center|[[Sockeye salmon]] run up a river}}]] -->
[[File:Salmon jumping.jpg|thumb|left|{{center|Salmon jumping a fall}}]]
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 | image1    = Jumping Salmon.jpg
 | alt1      = 
 | caption1  = [[Steelhead]] jumping a rapid
 | image2    = Salmon leaping at Buchanty Spout on River Almond - geograph.org.uk - 1347586.jpg
 | alt2      = 
 | caption2  = Syncronised leaping
}} -->

After several years wandering huge distances in the ocean, most surviving salmon return to the same natal rivers where they were spawned. Then most of them swim up the rivers until they reach the very spawning ground that was their original birthplace.<ref name=Moyle190>[[#Moyle|Moyle]], p. 190</ref>

There are various theories about how this happens. One theory is that there are geomagnetic and chemical cues which the salmon use to guide them back to their birthplace. The fish may be sensitive to the Earth's magnetic field, which could allow the fish to orient itself in the ocean, so it can navigate back to the [[estuary]] of its natal stream.<ref name=Lohmann2008/>

Salmon have a strong sense of smell. Speculation about whether odours provide homing cues go back to the 19th century.<ref name=Trevanius/> In 1951, Hasler hypothesised that, once in vicinity of the [[estuary]] or entrance to its birth river, salmon may use chemical cues which they can smell, and which are unique to their natal stream, as a mechanism to [[Homing (biology)|home onto]] the entrance of the stream.<ref name=Hasler1951/> In 1978, Hasler and his students found that the way salmon locate their home rivers with such precision was because they could recognise its characteristic smell. They further demonstrated that the smell of their river becomes imprinted in salmon when they transform into smolts, just before they migrate out to sea.<ref name=Moyle190 /><ref name=Hasler1978/><ref name=Dittman/> Homecoming salmon can also recognise characteristic smells in tributary streams as they move up the main river. They may also be sensitive to characteristic [[pheromone]]s given off by juvenile [[conspecific]]s. There is evidence that they can "discriminate between two populations of their own species".<ref name=Moyle190 /><ref name=Groot1986/>

The recognition that each river and tributary has its own characteristic smell, and the role this plays as a navigation aid, led to a widespread search for a mechanism or mechanisms that might allow salmon to navigate over long distances in the open ocean. In 1977, Leggett identified, as mechanisms worth investigating, the use of the sun for navigation, and orientation to various possible gradients, such as temperature, salinity or chemicals gradients, or geomagnetic or geoelectric fields.<ref name=Leggett/><ref name=Moyle191>[[#Moyle|Moyle]], p. 191</ref>

There is little evidence salmon use clues from the sun for navigation. Migrating salmon have been observed maintaining direction at nighttime and when it is cloudy. Likewise, electronically tagged salmon were observed to maintain direction even when swimming in water much too deep for sunlight to be of use.<ref name=Ogura/>

In 1973, it was shown that Atlantic salmon have conditioned cardiac responses to electric fields with strengths similar to those found in oceans. "This sensitivity might allow a migrating fish to align itself upstream or downstream in an ocean current in the absence of fixed references."<ref name=Rommel/> In 1988, researchers found iron, in the form of [[Single domain (magnetic)|single domain]] [[magnetite]], resides in the skulls of sockeye salmon. The quantities present are sufficient for [[magnetoception]].<ref name=Quinn1988/>

Tagging studies have shown a small number of fish do not find their natal rivers, but travel instead up other, usually nearby streams or rivers.<ref name=Quinn1991/><ref name=Tallman/> It is important some salmon stray from their home areas; otherwise new habitats could not be colonized. In 1984, Quinn hypothesized there is a dynamic equilibrium, controlled by genes, between homing and straying.<ref name=Quinn1984/> If the spawning grounds have a uniform high quality, then natural selection should favour the descendants that home accurately. If the spawning grounds have a variable quality, then natural selection should favour a mixture of the descendants that stray and the descendants that home accurately.<ref name=Lohmann2008 /><ref name=Quinn1984 />

[[File:The kype of a spawning male salmon.jpg|thumb|upright|right|The [[kype]] of a spawning male salmon]]

Prior to the run up the river, the salmon undergo profound physiological changes. [[Undulatory locomotion#Muscle architecture|Fish swim]] by contracting longitudinal red muscle and obliquely oriented white muscles. Red muscles are used for sustained activity, such as ocean migrations. White muscles are used for bursts of activity, such as bursts of speed or jumping.<ref name=Kapoor/> As the salmon comes to end of its ocean migration and enters the estuary of its natal river, its energy metabolism is faced with two major challenges: it must supply energy suitable for swimming the river rapids, and it must supply the sperm and eggs required for the reproductive events ahead. The water in the estuary receives the freshwater discharge from the natal river. Relative to ocean water, this has a high chemical load from [[surface runoff]]. Researchers in 2009 found evidence that, as the salmon encounter the resulting drop in salinity and increase in olfactory stimulation, two key metabolic changes are triggered: there is a switch from using red muscles for swimming to using white muscles, and there is an increase in the sperm and egg load. "Pheromones at the spawning grounds [trigger] a second shift to further enhance reproductive loading."<ref name=Miller/>

The salmon also undergo radical morphological changes as they prepare for the spawning event ahead. All salmon lose the silvery blue they had as ocean fish, and their colour darkens, sometimes with a radical change in hue. Salmon are [[Sexual dimorphism|sexually dimorphic]], and the male salmon develop canine-like teeth and their jaws develop a pronounced curve or hook (''[[kype]]''). Some species of male salmon grow large humps.<ref name=DFWa/>

==Obstacles==
{{multiple image
| align             = right
| caption_align     = center
| direction         = horizontal
| image1            = Fish ladder at Tawe Barrage - geograph.org.uk - 1607929.jpg
| width1            = 205
| alt1              = 
| caption1          = A fish ladder makes it easier for salmon to negotiate a [[weir]].
| image2            = RockyReachDam-FishLadder.jpg
| width2            = 115
| alt2              = 
| caption2          = An extended bypass
}}

Salmon start the run in peak condition, the culmination of years of development in the ocean. They need high swimming and leaping abilities to battle the rapids and other obstacles the river may present, and they need a full sexual development to ensure a successful spawn at the end of the run. All their energy goes into the physical rigours of the journey and the dramatic morphological transformations they must still complete before they are ready for the spawning events ahead.

The run up the river can be exhausting, sometimes requiring the salmon to battle hundreds of miles upstream against strong currents and rapids. They cease feeding during the run.<ref name=Moyle188 /> Chinook and sockeye salmon from central Idaho must travel {{convert|900|mi|km}} and climb nearly {{convert|7000|ft|m}} before they are ready to spawn. Salmon deaths that occur on the upriver journey are referred to as ''en route mortality''.<ref name=Jeffries2011/>

Salmon negotiate [[waterfall]]s and [[rapids]] by leaping or jumping. They have been recorded making vertical jumps as high as 3.65 metres (12&nbsp;ft).<ref name=Beach1984/> The height that can be achieved by a salmon depends on the position of the [[standing wave]] or [[hydraulic jump]] at the base of the fall, as well as how deep the water is.<ref name=Beach1984 />

[[Fish ladder]]s, or fishways, are specially designed to help salmon and other fish to bypass dams and other man-made obstructions, and continue on to their spawning grounds further upriver.<ref name=MichiganDNR/> Data suggest that navigation locks have a potential to be operated as vertical slot fishways to provide increased access for a range of biota, including poor swimmers.<ref>{{Cite journal |last1=Silva |first1=Sergio |last2=Lowry |first2=Maran |last3=Macaya-Solis |first3=Consuelo |last4=Byatt |first4=Barry |last5=Lucas |first5=Martyn C. |date=2017 |title=Can navigation locks be used to help migratory fishes with poor swimming performance pass tidal barrages? A test with lampreys |journal=Ecological Engineering |volume=102 |pages=291–302 |doi=10.1016/j.ecoleng.2017.02.027 |doi-access=free |bibcode=2017EcEng.102..291S }}</ref>{{clarify|date=March 2018}}

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{{multiple image
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| image1            = Black Bear Caught A Salmon.jpg
| width1            = 210
| alt1              = 
| caption1          = The black fur of [[American black bear|black bear]]s is easily spotted by salmon in daylight, and the bears fish more successfully using auditory clues at night.
| image2            = Spiritbear.jpg
| width2            = 110
| alt2              = 
| caption2          = White-coated [[Kermode bear]]s have more success fishing in daylight.
}}

Skilled predators, such as [[bear]]s, [[bald eagle]]s and [[Recreational fishing|fishermen]] can await the salmon during the run. Normally solitary animals, [[grizzly bear]]s congregate by streams and rivers when the salmon spawn.<ref name=Helfield2006/><ref name=Hilderbrand/> Predation from [[harbor seals]], [[California sea lions]], and [[Steller sea lions]] can pose a significant threat, even in river ecosystems.<ref>{{citation |url=http://www.nwr.noaa.gov/marine-mammals/seals-and-sea-lions/upload/cr-pinniped-fs.pdf |date=March 2008 |title=Seal & Sea Lion Facts of the Columbia River & Adjacent Nearshore Marine Areas |publisher=[[NOAA]] |access-date=2012-04-16 |archive-url=https://web.archive.org/web/20120723020046/http://www.nwr.noaa.gov/Marine-Mammals/Seals-and-Sea-Lions/upload/CR-Pinniped-FS.pdf |archive-date=2012-07-23 }}</ref><ref>{{citation |url=https://www.npr.org/templates/story/story.php?storyId=90213669 |title=Endangered Seals Eating Endangered Salmon |publisher=[[NPR]] |work=[[Bryant Park Project]] |date=May 6, 2008 }}</ref>

[[American black bear|Black bear]]s also fish the salmon. Black bears usually operate during the day, but when it comes to salmon they tend to fish at night.<ref name=Klinka2009a/> This is partly to avoid competition with the more powerful brown bears, but it is also because they catch more salmon at night.<ref name=Reimchen2009/> During the day, salmon are very evasive and attuned to visual clues, but at night they focus on their spawning activities, generating acoustic clues the bears tune into.<ref name=Klinka2009a /> Black bears may also fish for salmon during the night because their black fur is easily spotted by salmon in the daytime. In 2009, researchers compared the foraging success of black bears with the white-coated [[Kermode bear]], a [[Polymorphism (biology)|morphed]] subspecies of the black bear. They found the Kermode bear had no more success catching salmon at night time, but had greater success than the black bears during the day.<ref name=Klinka2009b/>

[[Otter]]s are also common predators. In 2011, researchers showed that when otters predate salmon, the salmon can "sniff them out". They demonstrated that once otters have eaten salmon, the remaining salmon could detect and avoid the waters where otter faeces was present.<ref name=Roberts/><ref name=PlanetEarth/>

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==Spawning==
{{see also|Juvenile salmon}}
{{multiple image
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| width             = 300
| header            = Salmon redds
| header_align      = center
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| footer            = 
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| background color  = 
| caption_align     = center
| image1            = Becharof Wilderness Salmon.jpg
| alt1              = 
| caption1          = Spawning salmon building redds on a riffle
| image2            = Salmon redds.jpg
| alt2              = 
| caption2          = The white areas on the river bottom are completed redds.
| image3            = Redds.jpg
| alt3              = 
| caption3          = Closeup of redds on a riverbed
}}

The term ''prespawn mortality'' is used to refer to fish that arrive successfully at the spawning grounds, and then die without spawning. Prespawn mortality is surprisingly variable, with one study observing rates between 3% and 90%.<ref name=Jeffries2011 /><ref name=Gilhousen1990/> Factors that contribute to these mortalities include high temperatures,<ref name=Crossin2008/><ref name=Farrell/> high river discharge rates,<ref name=Rand/> and parasites and diseases.<ref name=Gilhousen1990 /><ref name=Jones/> "At present there are no reliable indicators to predict whether an individual arriving at a spawning area will in fact survive to spawn."<ref name=Jeffries2011 />

The eggs of a female salmon are called her [[roe]]. To lay her roe, the female salmon builds a spawning nest, called a ''redd'', in a riffle with gravel as its [[streambed]]. A [[riffle]] is a relatively shallow length of stream where the water is turbulent and flows faster. She builds the redd by using her tail ([[caudal fin]]) to create a low-pressure zone, lifting gravel to be swept downstream, and excavating a shallow depression.  The redd may contain up to 5,000 eggs, each about the size of a pea, covering {{convert|30|sqft|m2}}.<ref name=McGrath/> The eggs usually range from orange to red. One or more males will approach the female in her redd, depositing his sperm, or milt, over her eggs.<ref name="USFWS"/> The female then covers the eggs by disturbing the gravel at the upstream edge of the depression before moving on to make another redd. The female will make as many as seven redds before her supply of eggs is exhausted.<ref name="USFWS" /><ref name=DFWb/>

Male pink salmon and some sockeye salmon develop pronounced humps just before they spawn. These humps may have evolved because they confer species advantages. The humps make it less likely the salmon will spawn in the shallow water at margins of the streambed, which tend to dry out during low water flows or freeze in winter. Further, riffles can contain many salmon spawning simultaneously, as in the image on the right. Predators, such as bears, will be more likely to catch the more visually prominent humped males, with their humps projecting above the surface of the water. This may provide a protective buffer for the females.<ref name=Groot1991 />

Dominant male salmon defend their redds by rushing at and chasing intruders. They butt and bite them with the canine-like teeth they developed for the spawning event. The kypes are used to clamp around the base of the tail ([[caudal peduncle]]) of an opponent.<ref name=Groot1991 />

== Deterioration ==
The physical condition of the salmon deteriorates the longer they remain in fresh water. Once the salmon have spawned, most of them deteriorate rapidly (a.k.a. "spawned out") and soon die. Some deteriorating salmon are still alive, but their bodies have already begun the process of rotting,<ref name=":2">{{Cite news|last=|first=|date=|title=Zombie salmon are the true 'living dead' and now is the time to see them, experts say|work=[[The Sacramento Bee]]|url=https://www.sacbee.com/news/nation-world/national/article246743471.html}}</ref> and these deteriorating salmon are sometimes colloquially called "zombie fish".<ref>{{Cite web|title=U.S. Fish & Wildlife Service - Zombie Fish|url=https://www.fws.gov/zombiefish/|access-date=2021-02-16|website=www.fws.gov}}</ref> This is because upstream freshwater bodies (especially creeks) typically do not have sufficient food available for the adult salmon diet, and they have used large amounts of energy swimming upriver, thus exhausting their own internal nutrient reserves.<ref name=":0">{{Cite web|title=Why do salmon change color and die after they spawn?|url=https://www.usgs.gov/faqs/why-do-salmon-change-color-and-die-after-they-spawn|access-date=2021-02-16|website=www.usgs.gov}}</ref><ref name=":1">{{Cite web|last=Alessandra Bergamin|date=2013-11-22|title=Why do Pacific Salmon Die After Spawning? -|url=https://baynature.org/2013/11/21/pacific-salmon-die-spawning/|access-date=2021-02-16|website=Bay Nature|language=en-US}}</ref> Spawning salmon also have programmed [[senescence]], which is "characterized by [[immunosuppression]] and organ deterioration", making them more vulnerable to diseases.<ref name="Jeffries2011" /><ref name="Dickhoff" /><ref name="Finch" /> Most zombie fish die within days of spawning, but some can last up to a couple of weeks.<ref name=":2" /> Once the salmon die in the river, they are either [[scavenge]]d by other animals, or they [[decompose]] and release inorganic nutrients to the [[plankton]] in the river and the [[riparian]] vegetation in the [[floodplain]].<ref name=":0" />

The [[Pacific salmon]] are a classic example of a [[semelparous]] animal, which reproduce only once in their lifetime. Semelparity is sometimes called "big bang" reproduction, since the single reproductive event of semelparous organisms is usually large and fatal to the spawners.<ref name="Ricklefs" /> It is an evolutionary strategy that concentrates all available resources into maximizing reproduction, at the expense of individual organism's life, which is common among [[insect]]s but rare among [[vertebrate]]s.<ref name=":1" /> All six species of Pacific salmons live for many years in the ocean before swimming to the freshwater stream of its birth, spawning, and then dying. Most [[Atlantic]] salmon also die after spawning, but about 5 to 10% (mostly female) return to the ocean where they can recover and spawn again the next season.<ref name="SalmonTrust" />

{{multiple image
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| image1            = Oncorhynchus nerka.jpg
| width1            = 305
| alt1              = 
| caption1          = Spawning male [[sockeye salmon]]
| image2            = Salmo salar eggs.jpg
| width2            = 130
| alt2              = 
| caption2          = The pea-sized eggs are laid in redds.
| image3            = Dead salmon in spawning season.jpg
| width3            = 200
| alt3              = 
| caption3          = All Pacific salmon ''(pictured)'' and most Atlantic salmon die after spawning.
}}
{{Clear}}
<!-- 
 A ''kelt'' is an Atlantic salmon which has survived the spawning process and is now on its way back to the ocean.<ref name=SalmonTrust/>
{{Image gallery
| title=
| lines=6
| width=160
| height=140
| align=left
|Oncorhynchus nerka 2.jpg||Salmon are mostly [[semelparous]] or "big bang" spawners, which means they die shortly after spawning
| Mission Accomplished^ Spent salmon carcases on the Lour Burn. - geograph.org.uk - 298415.jpg||Spent carcases of [[Atlantic salmon]] up a tributary of the [[River Spey]] in Scotland. Most, but not all Atlantic salmon die after spawning.
| A typical salmon kelt from the Spey. - geograph.org.uk - 338457.jpg||A salmon kelt which was making its way back down the River Spey. After spawning the fish is slim, having been relieved of the spawn and its body fat over the previous months.
}} -->

{{Clear}}

==Keystone species==
{{multiple image
| align             = right
| caption_align     = center
| width             = 220
| direction         = vertical
| image1            = Brown bears with salmon carcass.jpg
| alt1              = 
| caption1          = Grizzly bears tend to carry salmon carcasses into adjacent riparian areas.
| image2            = Salmon nitrogen cycling in a stream.gif
| alt2              = 
| caption2          = Salmon subsidy to the [[nitrogen cycle]] in a hypothetical stream system
}}

In the Pacific Northwest and Alaska, salmon is a [[keystone species]], supporting wildlife from birds to bears and otters.<ref name=Willson/>  The bodies of salmon represent a transfer of nutrients from the ocean, rich in nitrogen, sulfur, carbon and phosphorus, to the [[forest ecosystem]].

[[Grizzly bear]]s function as [[ecosystem engineer]]s, capturing salmon and carrying them into adjacent wooded areas. There they deposit nutrient-rich urine and feces and partially eaten carcasses. It has been estimated that bears leave up to half the salmon they harvest on the forest floor,<ref name=Reimchen2001/><ref name=Quinn2009/> in densities that can reach 4,000 kilograms per hectare,<ref name=Reimchen2002/> providing as much as 24% of the total nitrogen available to the riparian woodlands.<ref name=Helfield2006 /> The foliage of [[Picea glauca|spruce trees]] up to {{convert|500|m|ft|abbr=on}} from a stream where grizzlies fish salmon have been found to contain nitrogen originating from fished salmon.<ref name=Helfield2006 />

{{quote box 
| quoted = true 
| width = 25%
| align = left 
| salign = right 
| quote = Salmon continue to surprise us, showing us new ways in which their oceanic migrations eventually permeate entire terrestrial ecosystems. In terms of providing food and nutrients to a whole food web, we like to think of them as North America's answer to the Serengeti's [[wildebeest]].<ref name=ScienceDaily2008/>
| source = 
}}

Wolves normally hunt for deer. A 2008 study suggests that when the salmon run starts, wolves choose to fish for salmon, even if plenty of deer are still available.<ref name=Darimont/> "Selecting benign prey such as salmon makes sense from a safety point of view. While hunting deer, wolves commonly incur serious and often fatal injuries. In addition to safety benefits we determined that salmon also provides enhanced nutrition in terms of fat and energy."<ref name=ScienceDaily2008 />

The upper reaches of the [[Chilkat River]] in Alaska has particularly good spawning grounds. Each year these attract a run of up to half a million [[chum salmon]]. As the salmon run up the river, [[bald eagle]]s arrive in their thousands to feast at the spawning grounds. This results in some of the world's largest congregations of bald eagles. The number of participating eagles is directly correlated with the number of spawning salmon.<ref name=Hansen2008/>

Residual nutrients from salmon can also accumulate downstream in estuaries. A 2010 study suggests the density and diversity of many estuarine breeding birds in the summer "were strongly predicted by salmon [[Biomass (ecology)|biomass]] in the autumn".<ref name=Field/> Anadromous salmon provide nutrients to these "diverse assemblages ... ecologically comparable to the migrating herds of wildebeest in the [[Serengeti]]".<ref name=Reimchen2002 />

==Prospects==
In 1997, researchers noted that the future of salmon runs worldwide would depend on many factors, most of which are driven by human actions. Among the key driving factors are (1) harvest of salmon by commercial, recreational, and subsistence fishing, (2) alterations in stream and river channels, including construction of dikes and other riparian corridor modifications, (3) electricity generation, flood control, and irrigation supplied by dams, (4) alteration by humans of freshwater, [[Estuary|estuarine]], and marine environments used by salmon, coupled with aquatic changes due to climate and ocean circulatory regimes, (5) water withdrawals from rivers and reservoirs for agricultural, municipal, or commercial purposes,  (6) changes in climate caused at least in part by human activities, (7) competition from non-native fishes, (8) salmon predation by marine mammals, birds, and other fish species, (9) diseases and parasites, including those from outside the native region, and (10) reduced nutrient replenishment from decomposing salmon.<ref>{{Cite book|last=Stouder, Deanna J.|title=Pacific Salmon & Their Ecosystems: Status and Future Options|date=1997|publisher=Springer US|others=Bisson, Peter A., Naiman, Robbert J., Duke, Marcus G.|isbn=978-1-4615-6375-4|location=Boston, MA|oclc=840286102}}</ref>

In 2009, [[NOAA]] advised that continued runoff into North American rivers of three widely used [[pesticide]]s containing [[neurotoxin]]s would "jeopardize the continued existence" of endangered and threatened Pacific salmon.<ref name="NOAA2009" /><ref name="ENS" /> Global warming could see the end of some salmon runs by the end of the century,{{according to whom|date=August 2023}} such as the Californian runs of Chinook salmon.<ref name="Thompson" /><ref name="ScienceDaily2011" /> A 2010 United Nations report said that increases in [[ocean acidification|acidification of oceans]] would mean that shellfish such as [[Pteropod]]s, an important component of the ocean salmon diet, would be finding it more difficult to build their [[aragonite]] shells.<ref name="UNEP" /> There were concerns{{by whom|date=August 2023}} that this too could endanger future salmon runs.<ref name="TheTelegraph" />

==In popular culture==

In a 1982 video game called ''[[Salmon Run (video game)|Salmon Run]]'', the player takes the role of Sam the Salmon, swimming upriver to mate. Along the way he encounters waterfalls, a bear, fishermen, and seagulls.

In the 2003 [[The Walt Disney Company|Disney]] animated feature film, ''[[Brother Bear]]'', Kenai and Koda reached the salmon run and met a large group of bears led by Tug at the Annual Salmon Run. Featuring the song "[[Brother Bear (soundtrack)#Track Listing|Welcome]]" by [[The Blind Boys of Alabama]] and [[Phil Collins]].{{citation needed|date=August 2023}}

[[Splatoon|Splatoon 2 and 3]] introduce the game mode "[https://splatoonwiki.org/wiki/Salmon_Run Salmon Run]" where players fight off fictional [https://splatoonwiki.org/wiki/Salmonid Salmonids], a race of salmon-like beings. Grizzo Industries, owned by [https://splatoonwiki.org/wiki/Mr._Grizz Mr. Grizz], employs players to hunt for salmonids and their eggs, similar to [[Grizzly bear]]s hunting salmon in nature. The game's lore suggests that the Salmonids participate in a massive salmon migration every 70 years, which coincides with the time the Splatoon series takes place.

==Notable runs==
{{external media
|float=right
|width=300px
|video1={{YouTube|0NcJ_63z-mA|Grizzly Bears Catching Salmon}} – Nature's Great Events: The Great Salmon Run
|video2={{YouTube|hecXupPpE9o|Bald Eagle catches salmon}} – BBC Nature's Great Events - The Great Salmon Run 
|video3={{YouTube|B5ToVRwRUT4|The Great Salmon Run}} – ''[[Nature's Great Events|BBC Nature's Great Events]]''
|video4={{YouTube|uqR2g8darqs|Nimbus Hatchery Fish Ladder}}
|video5={{YouTube|EqmGSexPaEk|Life Cycle of Salmon}}
|video6={{YouTube|5DqjsWsY8-g|Life Cycle of Salmon}} – ''Discovery Channel''
|video7=[https://web.archive.org/web/20111201190215/http://www.atlanticsalmontrust.org/salmon-life-cycle-habitat-threats-and-concerns.html The Salmon's Lifecycle] ''Atlantic Salmon Trust''
|video8={{YouTube|gFCA5G83DfQ|Sockeye Salmon Run 2010}}
|video9={{YouTube|AGRpxyojc58|Spawning salmon constructing a redd}}
|video10={{YouTube|rYw_Da9y4oY|Raising salmon}}
|video11={{YouTube|qV30UZ9aF04|Salmon life cycle song}}
}}
* [[Adams River (British Columbia)|Adams River]] (British Columbia)
* Anan Creek (Alaska)<ref>{{cite web |url=http://www.fs.usda.gov/r10/tongass/recreation |title=Anan Wildlife Observatory Site |publisher=U.S. Forest Service |date=August 2023}}</ref>
* [[Bristol Bay]] (Alaska)
* [[Chilkat River]] (Alaska)
* [[Columbia River]] (British Columbia, United States)
* [[Copper River (Alaska)|Copper River]] (Alaska)
* [[Fraser River]] (British Columbia)
* [[Kenai River]] (Alaska)
* [[River Spey]] (Scotland)
* [[Tana (Norway)|River Tana]] (Norway, Finland)
* [[River Tay]] (Scotland)
* [[River Tweed]] (border of Scotland and England)
* [[River Tyne]] (England)
* [[Snake River]] (United States)
* [[Yukon River]] (Alaska, Yukon, British Columbia)

==See also==
* [[Salmonidae]]
* [[Animal navigation]]
* [[Environmental impact of reservoirs]]
* [[June hogs]]
* [[Natal homing]]
* [[Olfactory navigation]]
* [[Pre-spawn mortality in coho salmon]]
* [[Sardine run]]

{{Clear}}

==References==
{{Reflist|30em|refs=
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<ref name=FBAtlantic>{{FishBase| genus=Salmo | species=salar | year=2011 | month=December }}</ref>
<ref name=FBChinook>{{FishBase| genus=Oncorhynchus | species=tshawytscha | year=2011 | month=December }}</ref>
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<ref name=FBSockeye>{{FishBase| genus=Oncorhynchus | species=nerka | year=2011 | month=December }}</ref>
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<ref name=Rand>{{cite journal|vauthors=Rand PS, Hinch SG, Morrison J, Foreman MG, MacNutt MJ, Macdonald JS, Healey MC, Farrell AP, Higgs DA |url=http://people.landfood.ubc.ca/anthony.farrell/pubs/p224-Rand_et_al_2006.pdf |title=Effects of river discharge, temperature, and future climates on energetics and mortality of adult migrating Fraser River sockeye salmon|journal=Transactions of the American Fisheries Society |volume=135 |issue=3 |pages=655–667 |doi=10.1577/T05-023.1|year=2006 |bibcode=2006TrAFS.135..655R }}</ref>
<ref name=Quinn1984>Quinn TP (1984) [https://books.google.com/books?id=xYYWAQAAIAAJ&q=%22Mechanisms+of+Migration+in+Fishes%22 "Mechanisms of Migration in Fishes] Eds: McCleave JD, Arnold GP, Dodson JJ, Neill WH, pp.&nbsp;357–362. Plenum Press. {{ISBN|978-0-306-41676-7}}.</ref>
<ref name=Quinn1988>{{cite journal|vauthors=Walker MM, Quinn TP, Kirschvink JL, Groot C |url=http://authors.library.caltech.edu/31920/1/WALjeb88.pdf|year=1988|title=Production of single-domain magnetite throughout life by sockeye salmon, ''Oncorhynchus nerka''|journal=Journal of Experimental Biology|volume=140|pages=51–63|doi=10.1242/jeb.140.1.51|pmid=3204338}}</ref>
<ref name=Quinn1991>{{cite journal|vauthors=Quinn TP, Nemeth RS, McIsaac DO |year=1991|doi=10.1577/1548-8659(1991)120<0150:HASPOF>2.3.CO;2|title=Homing and straying patterns of fall chinook salmon in the lower Columbia River|journal=Trans Am Fish Soc|volume=120|issue=2|pages=150–156|bibcode=1991TrAFS.120..150Q }}</ref>
<ref name=Quinn2009>{{Cite journal |last1=Quinn |first1=T. |last2=Carlson |first2=S. |last3=Gende |first3=S. |name-list-style=amp |last4=Rich |first4=H. |year=2009 |title=Transportation of Pacific Salmon Carcasses from Streams to Riparian Forests by Bears |journal=Canadian Journal of Zoology |url=http://nature.berkeley.edu/carlsonlab/papers/QuinnEtAl_2009_CJZ.pdf |volume=87 |issue=3 |pages=195–203 |doi=10.1139/Z09-004 |archive-url=https://web.archive.org/web/20120616141524/http://nature.berkeley.edu/carlsonlab/papers/QuinnEtAl_2009_CJZ.pdf |archive-date=2012-06-16 }}</ref>
<ref name=Reimchen2001>{{cite journal|vauthors=Reimchen TE|year=2001|url=http://www.web.uvic.ca/~reimlab/reimchen_ecoforestry.pdf|title=Salmon nutrients, nitrogen isotopes and coastal forests|journal=Ecoforestry|volume=16|page=13|archive-date=6 May 2003|access-date=26 October 2016|archive-url=https://web.archive.org/web/20030506215141/http://web.uvic.ca/~reimlab/reimchen_ecoforestry.pdf}}</ref>
<ref name=Reimchen2002>{{cite journal|vauthors=Reimchen TE, Mathewson DD, Hocking MD, Moran J |year=2002|url=http://web.uvic.ca/~reimlab/n15clayoquot.pdf |title=Isotopic evidence for enrichment of salmon-derived nutrients in vegetation, soil, and insects in riparian zones in coastal British Columbia|journal=American Fisheries Society Symposium|volume=20|pages= 1–12}}</ref>
<ref name=Reimchen2009>{{cite journal|vauthors=Reimchen TE |year=2009|url=http://web.uvic.ca/~reimlab/NOCTURNAL%20FORAGING-MORESBY0001.pdf |title=Nocturnal foraging behaviour of black bears ''Ursus americanus'' on Moresby Island British Columbia|journal=Canadian Field-Naturalist|volume=112|issue=3 |pages=446–450|doi=10.5962/p.358447 }}</ref>
<ref name=Ricklefs>Ricklefs RE and Miller GK (2000) [https://books.google.com/books?id=6TMvdZQiySoC&q=Ecology++Ricklefs++Miller ''Ecology''] W.H. Freeman. {{ISBN|978-0-7167-2829-0}}.</ref>
<ref name=Roberts>{{cite journal|vauthors=Roberts LJ, de Leaniz CG |title=Something smells fishy: predator-naive salmon use diet cues, not kairomones, to recognize a sympatric mammalian predator|journal=Animal Behaviour|volume=82|issue=4|pages=619–625|doi=10.1016/j.anbehav.2011.06.019|year=2011|s2cid=53163932}}</ref>
<ref name=Rommel>{{cite journal|vauthors=Rommel SA, McCleave JD |title=Sensitivity of American Eels ''(Anguilla rostrata)'' and Atlantic Salmon ''(Salmo salar)'' to Weak Electric and Magnetic Fields|journal=Journal of the Fisheries Research Board of Canada|volume=30|issue=5|pages=657–663|doi=10.1139/f73-114|year=1973}}</ref>
<ref name=SalmonTrust>''Atlantic Salmon Trust'' [http://www.atlanticsalmontrust.org/knowledge/salmon-facts.html Salmon Facts] {{webarchive|url=https://web.archive.org/web/20111130201523/http://www.atlanticsalmontrust.org/knowledge/salmon-facts.html |date=2011-11-30 }} Retrieved 15 December 2011.</ref>
<ref name=ScienceDaily2008>''ScienceDaily'' (1 September 2008) [https://www.sciencedaily.com/releases/2008/09/080901205633.htm Wolves Would Rather Eat Salmon].</ref>
<ref name=ScienceDaily2011>''ScienceDaily'' (1 September 2011) [https://www.sciencedaily.com/releases/2011/09/110901155257.htm Warming Streams Could Be the End for Spring-Run Chinook Salmon in California].</ref>
<ref name=Tallman>{{cite journal|vauthors=Tallman RF, Healey MC |year=1994|title=Homing, straying, and gene flow among seasonally separated populations of chum salmon (''Oncorhynchus keta'')|journal=Can J Fish Aquat Sci|volume=51|issue=3|pages=577–588|doi=10.1139/f94-060}}</ref>
<ref name=TheTelegraph>''The Telegraph'' (3 December 2010). [https://web.archive.org/web/20101206075353/http://www.telegraph.co.uk/earth/environment/climatechange/8177990/Cancun-climate-summit-Britains-salmon-at-risk-from-ocean-acidification.html Cancun climate summit: Britain's salmon at risk from ocean acidification].</ref>
<ref name=Thompson>{{cite journal|vauthors=Thompson LC, Escobar MI, Mosser CM, Purkey DR, Yates D, Moyle PB |title=Water Management Adaptations to Prevent Loss of Spring-Run Chinook Salmon in California under Climate Change|journal=Journal of Water Resources Planning and Management|volume=138|issue=5|pages=465–478|doi=10.1061/(ASCE)WR.1943-5452.0000194|year=2012|s2cid=109723886|doi-access=free}}</ref>
<ref name=Trevanius>Trevanius GR (1822) [https://archive.org/details/biologieoderphi08trevgoog ''Biologie oder Philosophic der lebenden Natur fur Naturforscher und Arzte'', vol VI] Rower, Gottingen.</ref>
<ref name=UNEP>''[[UNEP]]'' (2010) [http://www.unep.org/dewa/pdf/Environmental_Consequences_of_Ocean_Acidification.pdf Environmental Consequences of Ocean Acidification: A Threat to Food Security] {{Webarchive|url=https://web.archive.org/web/20110125073557/http://www.unep.org/dewa/pdf/Environmental_Consequences_of_Ocean_Acidification.pdf |date=25 January 2011 }}</ref>
<ref name="USFWS">''Fish and Wildlife Services'' (2011) [http://www.fws.gov/species/species_accounts/bio_salm.html Pacific Salmon, (''Oncorhynchus'' spp.)] ''U.S. Fish and Wildlife Services''. Accessed: 28 December 2011.</ref>
<ref name=Walrond>Walrond C (2010) [http://www.teara.govt.nz/en/trout-and-salmon/3 Trout and salmon – Chinook salmon] ''Te Ara - the Encyclopedia of New Zealand''. Updated 9 September 2010.</ref>
<ref name=Webster>{{Cite Merriam-Webster|Anadromous}}</ref>
<ref name=Willson>{{cite journal|vauthors=Willson MF, Halupka KC|year=1995|url=http://nativefishsociety.org/wp-content/uploads/Willison-salmon-as-keystone-species-CB.pdf|title=Anadromous Fish as Keystone Species in Vertebrate Communities|journal=Conservation Biology|volume=9|issue=3|pages=489–497|doi=10.1046/j.1523-1739.1995.09030489.x|bibcode=1995ConBi...9..489W |archive-url=https://web.archive.org/web/20111128194020/http://nativefishsociety.org/wp-content/uploads/Willison-salmon-as-keystone-species-CB.pdf|archive-date=2011-11-28}}</ref>

}}

==Cited sources==
*{{cite book|ref=Moyle|vauthors=Moyle PB, Cech JJ |year=2004|url=https://books.google.com/books?id=sZYWAQAAIAAJ|title=Fishes, An Introduction to Ichthyology|edition=5th|publisher= Benjamin Cummings|isbn=978-0-13-100847-2}}

==Further reading==
{{Refbegin|2}}
* {{FishBase| genus=Oncorhynchus | species=mykiss | year=2011 | month=December }}
* ''USDA Forest Service'', [http://www.fs.fed.us/r6/fishing/regional/fishresources/salmon.html Salmon/Steelhead] Pacific Northwest Fisheries Program. Retrieved 30 December 2011.
* Knapp G, Roheim CA and Anderson JL (2007) [http://www.traffic.org/species-reports/traffic_species_fish25.pdf The Great Salmon Run: Competition Between Wild and Farmed Salmon] ''World Wildlife Fund''.
* Mozaffari, Ahmad and Alireza Fathi (2013) [https://arxiv.org/abs/1312.4078 "A natural-inspired optimization machine based on the annual migration of salmons in nature"] {{arXiv|1312.4078}}.
* Quinn, Thomas P. (2005) [https://books.google.com/books?id=potjIVogROcC ''The Behavior and Ecology of Pacific Salmon and Trout''] UBC Press. {{ISBN|978-0-7748-1128-6}}.

;Magnetoception and natal homing
* {{cite journal|vauthors=Bandoh H, Kida I, Ueda H |title=Olfactory Responses to Natal Stream Water in Sockeye Salmon by BOLD fMRI|journal=PLOS ONE|volume=6|issue=1|article-number=e16051|doi=10.1371/journal.pone.0016051|pmid=21264223|pmc=3022028|year=2011|bibcode=2011PLoSO...616051B|doi-access=free}}
* Bracis, Chloe (2010) [https://web.archive.org/web/20120426082749/http://wiki.cbr.washington.edu/qerm/sites/qerm/images/2/25/BracisThesis.pdf A model of the ocean migration of Pacific salmon] ''University of Washington''.
* {{cite journal | vauthors = Johnsen S, Lohmann KJ | year = 2005 | title = The physics and neurobiology of magnetoreception | url = http://www.sinnesphysiologie.de/download/magrev05.pdf | journal = Nature Reviews Neuroscience | volume = 6 | issue = 9 | pages = 703–712 | doi = 10.1038/nrn1745 | pmid = 16100517 | s2cid = 13996233 | archive-url = https://web.archive.org/web/20070630043005/http://www.sinnesphysiologie.de/download/magrev05.pdf | archive-date = 2007-06-30 }}
* {{cite journal |vauthors=Johnsen S, Lohmann KJ | year = 2008 | title = Magnetoreception in animals | journal = Physics Today | volume = 61 | issue = 3 | pages = 29–35 | doi = 10.1063/1.2897947 | bibcode = 2008PhT....61c..29J | doi-access =  }}
* {{cite journal |vauthors=Lohmann KJ, Lohmann CM, Endres CS | year = 2008 | title = The sensory ecology of ocean navigation | journal = J Exp Biol | volume = 211 | issue = 11| pages = 1719–1728 | doi=10.1242/jeb.015792| pmid = 18490387 | doi-access = free }}
* {{cite journal | vauthors = Mann S, Sparks NH, Walker MM, Kirschvink JL | year = 1988 | title = Ultrastructure, morphology and organization of biogenic magnetite from sockeye salmon, ''Oncorhynchus nerka'': implications for magnetoreception | url = http://jeb.biologists.org/content/140/1/35.full.pdf | journal = J Exp Biol | volume = 140 | pages = 35–49 | doi = 10.1242/jeb.140.1.35 | pmid = 3204335 | doi-access = free }}
* Metcalfe J, Arnold G and McDowall R (2008) [https://books.google.com/books?id=mhDs4X3KW2MC&pg=PA175 "Migration"] pp.&nbsp;175–199. In: John D. Reynolds, ''Handbook of fish biology and fisheries'', Volume 1, John Wiley & Sons. {{ISBN|978-0-632-05412-1}}.
* Moore A, Privitera L and Riley WD (2013) [https://books.google.com/books?hl=en&lr=&id=ohwbAAAAQBAJ&oi=fnd&pg=PP1 "The behaviour and physiology of migrating Atlantic salmon"] In: H Ueda and K Tsukamoto (eds),''Physiology and Ecology of Fish Migration'', CRC Press, pp.&nbsp;28–55. {{ISBN|9781466595132}}.
* Ueda, Hiroshi (2013) [https://books.google.com/books?hl=en&lr=&id=ohwbAAAAQBAJ&oi=fnd&pg=PP1 "Physiology of imprinting and homing migration in Pacific salmon"] In: H Ueda and K Tsukamoto (eds),''Physiology and Ecology of Fish Migration'', CRC Press, pp.&nbsp;1–27. {{ISBN|9781466595132}}.
* {{cite journal|vauthors=Walker MM, Diebel CE, Haugh CV, Pankhurst PM, Montgomery JC, Green CR |title=Structure and function of the vertebrate magnetic sense|journal=Nature|volume=390|issue=6658|pages=371–6 |doi=10.1038/37057 |pmid=20358649|year=1997|bibcode=1997Natur.390..371W|s2cid=4386772}}
* [[Wired (magazine)|''Wired'']]. [https://www.wired.com/wiredscience/2008/12/to-save-threate/ Hacking Salmon's Mental Compass to Save Endangered Fish] 2 December 2008.

;Nitrogen
* {{cite journal | vauthors = Cederholm CJ, Kunze MD, Murota T, Sibatani T | year = 1999 | title = Pacific salmon carcasses: essential contributions of nutrients and energy for aquatic and terrestrial ecosystems | url = http://www.nativefishsociety.org/conservation/wild_population/annotated_bib_salmonids_hatcheries/nutrient_enrichment/Pacific.pdf | journal = Fisheries | volume = 24 | issue = 10 | pages = 6–15 | doi = 10.1577/1548-8446(1999)024<0006:psc>2.0.co;2 | bibcode = 1999Fish...24j...6C | access-date = 21 December 2011 | archive-date = 5 March 2016 | archive-url = https://web.archive.org/web/20160305080342/http://www.nativefishsociety.org/conservation/wild_population/annotated_bib_salmonids_hatcheries/nutrient_enrichment/Pacific.pdf }}
* {{cite journal | vauthors = Gresh T, Lichatowich J, Schoonmaker P | year = 2000 | title = Salmon Decline Creates Nutrient Deficit in Northwest Streams | url = http://www.inforain.org/reports/salmon_decline.html | archive-url = https://web.archive.org/web/20080509143932/http://www.inforain.org/reports/salmon_decline.html | archive-date = 2008-05-09 | journal = Fisheries | volume = 15 | issue = 1 | pages = 15–21 | doi = 10.1577/1548-8446(2000)025<0015:AEOHAC>2.0.CO;2 | url-access = subscription }}
* {{cite journal | vauthors = Hocking MD, Reynolds JD | year = 2011 | title = Impacts of salmon on riparian plant diversity | url = https://www.sfu.ca/~mhocking/Morgan_D._Hocking_files/Hocking%20and%20Reynolds%202011.pdf | journal = Science | volume = 331 | issue = 6024 | pages = 1609–1612 | doi = 10.1126/science.1201079 | pmid = 21442794 | bibcode = 2011Sci...331.1609H | s2cid = 30725341 }}{{dead link|date=May 2018 |bot=InternetArchiveBot |fix-attempted=yes }}
* {{cite journal |vauthors=Naiman RJ, Bilby RE, Schindler DE, Helfield JM |url=http://www.fish.washington.edu/people/naiman/CV/reprints/naiman_ecosys_salmon_2002.pdf |title=Pacific salmon, nutrients, and the dynamics of freshwater and riparian ecosystems |journal=Ecosystems |volume=5 |issue=4 |pages=399–417 |doi=10.1007/s10021-001-0083-3 |year=2002 |bibcode=2002Ecosy...5..399N |s2cid=5607299 |archive-url=https://web.archive.org/web/20120424224623/http://www.fish.washington.edu/people/naiman/CV/reprints/naiman_ecosys_salmon_2002.pdf |archive-date=2012-04-24 }}
* {{cite journal | vauthors = Ruckelshaus MH, Levin P, Johnson JB, Kareiva PM | year = 2002 | title = The Pacific salmon wars: what science brings to the challenge of recovering species | url = http://www.olemiss.edu/courses/bisc677/Ruckelshaus2002.pdf | journal = Annu. Rev. Ecol. Syst. | volume = 33 | pages = 665–706 | doi = 10.1146/annurev.ecolsys.33.010802.150504 | archive-date = 24 April 2023 | access-date = 21 December 2011 | archive-url = https://web.archive.org/web/20230424033602/https://olemiss.edu/courses/bisc677/Ruckelshaus2002.pdf }}

;Resilience
* {{cite journal | vauthors = Bottom DL, Jones KK, Simenstad CA, Smith CL | year = 2009 | title = Reconnecting social and ecological resilience in salmon ecosystems | journal = Ecology and Society | volume = 14 | issue = 1 | page = 5 | article-number = art5 | doi = 10.5751/es-02734-140105 | url = http://dlc.dlib.indiana.edu/dlc/bitstream/handle/10535/2644/Reconnecting_Social_and_Ecological_Resilience_in_Salmon_Ecosystems.pdf?sequence=1&isAllowed=y | doi-access = free }}
* Bottom DL, Jones KK, Simenstad CA and Smith CL (Eds.) (2010) [http://www.ecologyandsociety.org/issues/view.php?sf=34 Pathways to Resilient Salmon Ecosystems] {{Webarchive|url=https://web.archive.org/web/20210419002239/https://www.ecologyandsociety.org/issues/view.php?sf=34 |date=19 April 2021 }} ''Ecology and Society'', Special Feature.
{{Refend}}

==External links==
* [https://web.archive.org/web/20130508081257/http://trueslant.com/hivemind/2009/07/09/putting-a-price-on-salmon/ Putting a Price on Salmon] ''True Slant'', 9 July 2009.
* [http://www.nwcouncil.org/history/fishpassage.asp Fish passage at dams] ''[[Northwest Power and Conservation Council]]''. Retrieved 17 December 2011.
* [https://www.wired.com/wiredscience/2011/01/salmon-disease/#more-48063 Mystery Disease Found in Pacific Salmon] [[Wired (magazine)|''Wired'']], 13 January 2011.
* [https://www.nps.gov/olym/learn/education/upload/Lesson9-SalmonAnadromousLifestyles.pptx Pacific Salmon: Anadromous Lifestyles] ''US National Park Service''.
* [https://web.archive.org/web/20120512173410/http://www.mtshastanews.com/news/x1991994365/Study-takes-long-term-diversified-view-of-salmon-issues Study takes long-term, diversified view of salmon issues] ''Mount Shasta News'', 30 September 2009.

{{salmon|state=expanded}}
{{swarming}}

[[Category:Marine biology]]
[[Category:Fishing]]
[[Category:Salmon]]
[[Category:Fish migrations]]