Resumen Pincoya

Resumen Pincoya
Contents
1 Salmon 1
1.1 Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.3 Life cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.4 Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.4.1 Bears and salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.4.2 Beavers and salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.4.3 Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.5 Wild fisheries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.5.1 Commercial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.5.2 Recreational . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.6 Farmed salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.7 Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1.8 As food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1.9 History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1.10 Mythology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1.11 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1.12 Further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
1.13 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2 Atlantic salmon 27
2.1 Nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.2 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.3 Distribution and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
2.4 Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.5 Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
2.6 Life stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
2.6.1 Freshwater phase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.6.2 Saltwater phases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.7 Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
i
ii CONTENTS
2.8 Hybridization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.9 Aquaculture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.9.1 Process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
2.9.2 Controversy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
2.10 Human impact . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
2.11 Recovery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
2.12 Beaver impact . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.13 Legislation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.13.1 England and Wales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.13.2 Scotland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.13.3 United States . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.13.4 Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.14 NASCO . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.15 Sustainable consumption . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.16 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.17 Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.18 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
2.19 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
3 Hydroacoustics 42
3.1 Related Publications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
3.2 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
4 Underwater acoustics 45
4.1 History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.2 Theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.2.1 Sound waves in water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.2.2 Speed of sound, density and impedance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.2.3 Absorption of sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
4.2.4 Sound Reflection and Scattering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
4.2.5 Propagation of sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
4.3 Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
4.3.1 Sound speed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
4.3.2 Absorption . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.3.3 Ambient noise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.3.4 Reverberation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.3.5 Bottom Loss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.4 Underwater hearing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.4.1 Comparison with airborne sound levels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
CONTENTS iii
4.4.2 Hearing sensitivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

4.4.3 Safety thresholds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
4.5 Applications of underwater acoustics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
4.5.1 Sonar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
4.5.2 Underwater communication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
4.5.3 Underwater Navigation and Tracking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.5.4 Seismic exploration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.5.5 Weather and climate observation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.5.6 Oceanography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.5.7 Marine biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.5.8 Particle physics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4.6 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4.7 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4.8 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
5 Bioacoustics 56
5.1 History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
5.2 Methods in bioacoustics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
5.2.1 Acoustic signals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
5.2.2 Sound production, detection, and use in animals . . . . . . . . . . . . . . . . . . . . . . . . . 58
5.2.3 Biomass estimation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
5.3 Animal sounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
5.4 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
5.5 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
5.6 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
5.7 Further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
6 Hydrophone 64
6.1 History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
6.2 Directional hydrophones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
6.2.1 Focused transducers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
6.2.2 Arrays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
6.3 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
6.4 Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
6.5 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
6.6 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
6.7 Text and image sources, contributors, and licenses . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
6.7.1 Text . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
6.7.2 Images . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
iv CONTENTS
6.7.3 Content license . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

Chapter 1
Salmon
For other uses, see Salmon (disambiguation).

This article is about a particular kind of fish. For the food, see Salmon as food.
Pacific salmon leaping at Willamette Falls, Oregon
Commercial production of salmon in million tonnes 1950–

2010[1]
Salmon /ˈsæmən/ is the common name for several species of ray-finned fish in the family Salmonidae. Other fish in the
same family include trout, char, grayling and whitefish. Salmon are native to tributaries of the North Atlantic (genus
Salmo) and Pacific Ocean (genus Oncorhynchus). Many species of salmon have been introduced into non-native envi-
1
2 CHAPTER 1. SALMON
ronments such as the Great Lakes of North America and Patagonia in South America. Salmon are intensively farmed in
many parts of the world.
Typically, salmon are anadromous: they are born in fresh water, migrate to the ocean, then return to fresh water to
reproduce. However, populations of several species are restricted to fresh water through their lives. Various species of
salmon display anadromous life strategies while others display freshwater resident life strategies. Folklore has it that the
fish return to the exact spot where they were born to spawn; tracking studies have shown this to be mostly true. A portion
of a returning salmon run may stray and spawn in different freshwater systems. The percent of straying depends on the
species of salmon.[2] Homing behavior has been shown to depend on olfactory memory.[3][4]
1.1 Species
The term “salmon” comes from the Latin salmo, which in turn may have originated from salire, meaning “to leap”.[5] The
nine commercially important species of salmon occur in two genera. The genus Salmo contains the Atlantic salmon, found
in the north Atlantic, as well as many species commonly named trout. The genus Oncorhynchus contains eight species
which occur naturally only in the North Pacific. As a group, these are known as Pacific salmon. Chinook salmon have
been introduced in New Zealand and Patagonia. Coho, freshwater sockeye, and Atlantic salmon have been established in
Patagonia, as well.[6]
†
Both the Salmo and Oncorhynchus genera also contain a number of species referred to as trout. Within Salmo, additional minor taxa
have been called salmon in English, i.e. the Adriatic salmon (Salmo obtusirostris) and Black Sea salmon (Salmo labrax). The steelhead
anadromous form of the rainbow trout migrates to sea, but it is not termed “salmon”.
Also a number of other species have common names which refer to them as being salmon. Of those listed below, the
Danube salmon or huchen is a large freshwater salmonid related to the salmon above, but others are marine fishes of the
unrelated Perciformes order:
Eosalmo driftwoodensis, the oldest known salmon in the fossil record, helps scientists figure how the different species of
salmon diverged from a common ancestor. The British Columbia salmon fossil provides evidence that the divergence
between Pacific and Atlantic salmon had not yet occurred 40 million years ago. Both the fossil record and analysis of
mitochondrial DNA suggest the divergence occurred by 10 to 20 million years ago. This independent evidence from DNA
analysis and the fossil record rejects the glacial theory of salmon divergence.[39]
1.2 Distribution
Atlantic salmon, Salmo salar
• Atlantic salmon (Salmo salar) reproduce in northern rivers on both coasts of the Atlantic Ocean.
1.3. LIFE CYCLE 3
• Landlocked salmon (Salmo salar m. sebago) live in a number of lakes in eastern North America and in
Northern Europe, for instance in lakes Sebago, Onega, Ladoga, Saimaa, Vänern, and Winnipesaukee. They
are not a different species from the Atlantic salmon, but have independently evolved a non-migratory life
cycle, which they maintain even when they could access the ocean.
• Chinook salmon (Oncorhynchus tshawytscha) are also known in the US as king salmon or blackmouth salmon, and
as spring salmon in British Columbia. Chinook are the largest of all Pacific salmon, frequently exceeding 14 kg (30
lb).[40] The name tyee is used in British Columbia to refer to Chinook over 30 pounds, and in the Columbia River
watershed, especially large Chinook were once referred to as June hogs. Chinook salmon are known to range as
far north as the Mackenzie River and Kugluktuk in the central Canadian arctic,[41] and as far south as the Central
California coast.[42]
• Chum salmon (Oncorhynchus keta) are known as dog, keta, or calico salmon in some parts of the US. This species
has the widest geographic range of the Pacific species:[43] south to the Sacramento River in California in the eastern
Pacific and the island of Kyūshū in the Sea of Japan in the western Pacific; north to the Mackenzie River in Canada
in the east and to the Lena River in Siberia in the west.
• Coho salmon (Oncorhynchus kisutch) are also known in the US as silver salmon. This species is found throughout
the coastal waters of Alaska and British Columbia and as far south as Central California (Monterey Bay).[44] It is
also now known to occur, albeit infrequently, in the Mackenzie River.[41]
• Masu salmon or cherry salmon (Oncorhynchus masou) are found only in the western Pacific Ocean in Japan, Korea,
and Russia. A land-locked subspecies known as the Taiwanese salmon or Formosan salmon (Oncorhynchus masou
formosanus) is found in central Taiwan’s Chi Chia Wan Stream.[45]
• Pink salmon (Oncorhynchus gorbuscha), known as humpies in southeast and southwest Alaska, are found from
northern California and Korea, throughout the northern Pacific, and from the Mackenzie River[41] in Canada to the
Lena River in Siberia, usually in shorter coastal streams. It is the smallest of the Pacific species, with an average
weight of 1.6 to 1.8 kg (3.5 to 4.0 lb).[46]
• Sockeye salmon (Oncorhynchus nerka) are also known in the US as red salmon.[47] This lake-rearing species is
found south as far as the Klamath River in California in the eastern Pacific and northern Hokkaidō island in Japan
in the western Pacific and as far north as Bathurst Inlet in the Canadian Arctic in the east and the Anadyr River
in Siberia in the west. Although most adult Pacific salmon feed on small fish, shrimp, and squid, sockeye feed on
plankton they filter through gill rakers.[48] Kokanee salmon are the land-locked form of sockeye salmon.
• Danube salmon, or huchen (Hucho hucho), are the largest permanent freshwater salmonid species.
1.3 Life cycle
See also: Salmon run and Juvenile salmon
Eggs in different stages of development: In some, only a few cells grow on top of the yolk,
in the lower right, the blood vessels surround the yolk, and in the upper left, the black eyes are visible, even the little lens.
4 CHAPTER 1. SALMON
Life cycle of Pacific salmon
Salmon fry hatching — the baby has grown around the remains of the yolk — visible are
the arteries spinning around the yolk and little old drops, also the gut, the spine, the main caudal blood vessel, the bladder,
and the arcs of the gills
Salmon eggs are laid in freshwater streams typically at high latitudes. The eggs hatch into alevin or sac fry. The fry quickly
develop into parr with camouflaging vertical stripes. The parr stay for six months to three years in their natal stream before
becoming smolts, which are distinguished by their bright, silvery colour with scales that are easily rubbed off. Only 10%
of all salmon eggs are estimated to survive to this stage.[49] The smolt body chemistry changes, allowing them to live in
salt water. Smolts spend a portion of their out-migration time in brackish water, where their body chemistry becomes
accustomed to osmoregulation in the ocean.
1.3. LIFE CYCLE 5
Juvenile salmon, parr, grow up in the relatively protected natal river
The parr lose their camouflage bars and become smolt as they become ready for the tran-
sition to the ocean.
Male ocean-phase adult sockeye
Male spawning-phase adult sockeye
The salmon spend about one to five years (depending on the species) in the open ocean, where they gradually become
sexually mature. The adult salmon then return primarily to their natal streams to spawn. Atlantic salmon spend between
one and four years at sea. (When a fish returns after just one year’s sea feeding, it is called a grilse in Canada, Britain,
and Ireland.) Prior to spawning, depending on the species, salmon undergo changes. They may grow a hump, develop
canine-like teeth, or develop a kype (a pronounced curvature of the jaws in male salmon). All change from the silvery
blue of a fresh-run fish from the sea to a darker colour. Salmon can make amazing journeys, sometimes moving hundreds
of miles upstream against strong currents and rapids to reproduce. Chinook and sockeye salmon from central Idaho, for
example, travel over 1,400 km (900 mi) and climb nearly 2,100 m (7,000 ft) from the Pacific Ocean as they return to
spawn. Condition tends to deteriorate the longer the fish remain in fresh water, and they then deteriorate further after they
spawn, when they are known as kelts. In all species of Pacific salmon, the mature individuals die within a few days or
weeks of spawning, a trait known as semelparity. Between 2 and 4% of Atlantic salmon kelts survive to spawn again, all
females. However, even in those species of salmon that may survive to spawn more than once (iteroparity), postspawning
mortality is quite high (perhaps as high as 40 to 50%.)
To lay her roe, the female salmon uses her tail (caudal fin), to create a low-pressure zone, lifting gravel to be swept
downstream, excavating a shallow depression, called a redd. The redd may sometimes contain 5,000 eggs covering 2.8
m2 (30 sq ft).[50] The eggs usually range from orange to red. One or more males approach the female in her redd,
depositing sperm, or milt, over the roe.[48] 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 may make as many as seven redds before her
supply of eggs is exhausted.[48]
Each year, the fish experiences a period of rapid growth, often in summer, and one of slower growth, normally in winter.
This results in ring formation around an earbone called the otolith, (annuli) analogous to the growth rings visible in a tree
trunk. Freshwater growth shows as densely crowded rings, sea growth as widely spaced rings; spawning is marked by
significant erosion as body mass is converted into eggs and milt.
Freshwater streams and estuaries provide important habitat for many salmon species. They feed on terrestrial and aquatic
insects, amphipods, and other crustaceans while young, and primarily on other fish when older. Eggs are laid in deeper
water with larger gravel, and need cool water and good water flow (to supply oxygen) to the developing embryos. Mor-
tality of salmon in the early life stages is usually high due to natural predation and human-induced changes in habitat,
such as siltation, high water temperatures, low oxygen concentration, loss of stream cover, and reductions in river flow.
Estuaries and their associated wetlands provide vital nursery areas for the salmon prior to their departure to the open
6 CHAPTER 1. SALMON
ocean. Wetlands not only help buffer the estuary from silt and pollutants, but also provide important feeding and hiding
areas.
Salmon not killed by other means show greatly accelerated deterioration (phenoptosis, or “programmed aging”) at the
end of their lives. Their bodies rapidly deteriorate right after they spawn as a result of the release of massive amounts of
corticosteroids.
1.4 Ecology
Bear cub with salmon
1.4.1 Bears and salmon

See also: Salmon run
In the Pacific Northwest and Alaska, salmon are keystone species, supporting wildlife such as birds, bears and otters.[51]
The bodies of salmon represent a transfer of nutrients from the ocean, rich in nitrogen, sulfur, carbon and phosphorus, to
the forest ecosystem.
Grizzly bears function as ecosystem engineers, capturing salmon and carrying them into adjacent wooded areas. There
they deposit nutrient-rich urine and feces and partially eaten carcasses. Bears are estimated to leave up to half the salmon
1.4. ECOLOGY 7
they harvest on the forest floor,[52][53] in densities that can reach 4,000 kilograms per hectare,[54] providing as much as
24% of the total nitrogen available to the riparian woodlands. The foliage of spruce trees up to 500 m (1,600 ft) from a
stream where grizzlies fish salmon have been found to contain nitrogen originating from fished salmon.[55]
1.4.2 Beavers and salmon

Beavers also function as ecosystem engineers; in the process of clear-cutting and damming, beavers alter their ecosystems
extensively. Beaver ponds can provide critical habitat for juvenile salmon. An example of this was seen in the years
following 1818 in the Columbia River Basin. In 1818, the British government made an agreement with the U.S. gov-
ernment to allow U.S. citizens access to the Columbia catchment (see Treaty of 1818). At the time, the Hudson’s Bay
Company sent word to trappers to extirpate all furbearers from the area in an effort to make the area less attractive to
U.S. fur traders. In response to the elimination of beavers from large parts of the river system, salmon runs plummeted,
even in the absence of many of the factors usually associated with the demise of salmon runs. Salmon recruitment can
be affected by beavers’ dams because dams can:[56][57][58]
• Slow the rate at which nutrients are flushed from the system; nutrients provided by adult salmon dying throughout
the fall and winter remain available in the spring to newly hatched juveniles
• Provide deeper water pools where young salmon can avoid avian predators
• Increase productivity through photosynthesis and by enhancing the conversion efficiency of the cellulose-powered
detritus cycle
• Create low-energy environments where juvenile salmon put the food they ingest into growth rather than into fighting
currents
• Increase structural complexity with many physical niches where salmon can avoid predators
Beavers’ dams are able to nurture salmon juveniles in estuarine tidal marshes where the salinity is less than 10 ppm.
Beavers build small dams of generally less than 60 cm (2 ft) high in channels in the myrtle zone. These dams can be
overtopped at high tide and hold water at low tide. This provides refuges for juvenile salmon so they do not have to swim
into large channels where they are subject to predation.[59]
1.4.3 Parasites
Main article: Diseases and parasites in salmon
According to Canadian biologist Dorothy Kieser, the myxozoan parasite Henneguya salminicola is commonly found in
the flesh of salmonids. It has been recorded in the field samples of salmon returning to the Haida Gwaii Islands. The fish
responds by walling off the parasitic infection into a number of cysts that contain milky fluid. This fluid is an accumulation
of a large number of parasites.
Henneguya and other parasites in the myxosporean group have complex life cycles, where the salmon is one of two
hosts. The fish releases the spores after spawning. In the Henneguya case, the spores enter a second host, most likely
an invertebrate, in the spawning stream. When juvenile salmon migrate to the Pacific Ocean, the second host releases
a stage infective to salmon. The parasite is then carried in the salmon until the next spawning cycle. The myxosporean
parasite that causes whirling disease in trout has a similar life cycle.[60] However, as opposed to whirling disease, the
Henneguya infestation does not appear to cause disease in the host salmon — even heavily infected fish tend to return to
spawn successfully.
According to Dr. Kieser, a lot of work on Henneguya salminicola was done by scientists at the Pacific Biological Station
in Nanaimo in the mid-1980s, in particular, an overview report[61] which states, “the fish that have the longest fresh water
residence time as juveniles have the most noticeable infections. Hence in order of prevalence coho are most infected
followed by sockeye, chinook, chum and pink.” As well, the report says, at the time the studies were conducted, stocks
8 CHAPTER 1. SALMON
from the middle and upper reaches of large river systems in British Columbia such as Fraser, Skeena, Nass and from
mainland coastal streams in the southern half of B.C., “are more likely to have a low prevalence of infection.” The report
also states, “It should be stressed that Henneguya, economically deleterious though it is, is harmless from the view of
public health. It is strictly a fish parasite that cannot live in or affect warm blooded animals, including man”.
According to Klaus Schallie, Molluscan Shellfish Program Specialist with the Canadian Food Inspection Agency, "Hen-
neguya salminicola is found in southern B.C. also and in all species of salmon. I have previously examined smoked
chum salmon sides that were riddled with cysts and some sockeye runs in Barkley Sound (southern B.C., west coast of
Vancouver Island) are noted for their high incidence of infestation.”
Sea lice, particularly Lepeophtheirus salmonis and various Caligus species, including C. clemensi and C. rogercresseyi, can
cause deadly infestations of both farm-grown and wild salmon.[62][63] Sea lice are ectoparasites which feed on mucus,
blood, and skin, and migrate and latch onto the skin of wild salmon during free-swimming, planktonic nauplii and cope-
podid larval stages, which can persist for several days.[64][65][66] Large numbers of highly populated, open-net salmon
farms can create exceptionally large concentrations of sea lice; when exposed in river estuaries containing large numbers
of open-net farms, many young wild salmon are infected, and do not survive as a result.[67][68] Adult salmon may survive
otherwise critical numbers of sea lice, but small, thin-skinned juvenile salmon migrating to sea are highly vulnerable. On
the Pacific coast of Canada, the louse-induced mortality of pink salmon in some regions is commonly over 80%.[69]
1.5 Wild fisheries
1.5.1 Commercial
As can be seen from the production chart at the left, the global capture reported by different countries to the FAO of
commercial wild salmon has remained fairly steady since 1990 at about one million tonnes per year. This is in contrast
to farmed salmon (below) which has increased in the same period from about 0.6 million tonnes to well over two million
tonnes.[1]
Nearly all captured wild salmon are Pacific salmon. The capture of wild Atlantic salmon has always been relatively small,
and has declined steadily since 1990. In 2011 only 2,500 tonnes were reported.[8] In contrast about half of all farmed
salmon are Atlantic salmon.
1.5.2 Recreational
Recreational salmon fishing can be a technically demanding kind of sport fishing, not necessarily congenial for beginning
fishermen.[70] A conflict exists between commercial fishermen and recreational fishermen for the right to salmon stock
resources. Commercial fishing in estuaries and coastal areas is often restricted so enough salmon can return to their natal
rivers where they can spawn and be available for sport fishing. On parts of the North American west coast sport salmon
fishing completely replaces inshore commercial fishing.[71] The commercial value of a salmon can be several times less
than the value of the same fish caught by a sport fisherman. This is “a powerful economic argument for allocating stock
resources preferentially to sport fishing.”[71]
1.6 Farmed salmon

Main article: Aquaculture of salmon
Salmon aquaculture is a major contributor to the world production of farmed finfish, representing about US$10 billion
annually. Other commonly cultured fish species include: tilapia, catfish, sea bass, carp and bream. Salmon farming is
significant in Chile, Norway, Scotland, Canada and the Faroe Islands; it is the source for most salmon consumed in the
United States and Europe. Atlantic salmon are also, in very small volumes, farmed in Russia and the island of Tasmania,
Australia.
1.7. MANAGEMENT 9
Salmon are carnivorous. They are fed a meal produced from catching other wild fish and other marine organisms. Salmon
farming leads to a high demand for wild forage fish. Salmon require large nutritional intakes of protein, and farmed salmon
consume more fish than they generate as a final product. To produce one pound of farmed salmon, products from several
pounds of wild fish are fed to them. As the salmon farming industry expands, it requires more wild forage fish for feed,
at a time when 75% of the world’s monitored fisheries are already near to or have exceeded their maximum sustainable
yield.[72] The industrial-scale extraction of wild forage fish for salmon farming affects the survivability of the wild predator
fish which rely on them for food.
Work continues on substituting vegetable proteins for animal proteins in the salmon diet. This substitution results in lower
levels of the highly valued omega-3 fatty acid content in the farmed product.
Intensive salmon farming uses open-net cages, which have low production costs. It has the drawback of allowing disease
and sea lice to spread to local wild salmon stocks.[73]
On a dry weight basis, 2–4 kg of wild-caught fish are needed to produce one kg of salmon.[74]
Another form of salmon production, which is safer but less controllable, is to raise salmon in hatcheries until they are
old enough to become independent. They are released into rivers in an attempt to increase the salmon population. This
system is referred to as ranching. It was very common in countries such as Sweden, before the Norwegians developed
salmon farming, but is seldom done by private companies. As anyone may catch the salmon when they return to spawn,
a company is limited in benefiting financially from their investment.
Because of this, the ranching method has mainly been used by various public authorities and nonprofit groups, such as
the Cook Inlet Aquaculture Association, as a way to increase salmon populations in situations where they have declined
due to overharvesting, construction of dams, and habitat destruction or fragmentation. Negative consequences to this
sort of population manipulation include genetic “dilution” of the wild stocks. Many jurisdictions are now beginning to
discourage supplemental fish planting in favour of harvest controls, and habitat improvement and protection.
A variant method of fish stocking, called ocean ranching, is under development in Alaska. There, the young salmon are
released into the ocean far from any wild salmon streams. When it is time for them to spawn, they return to where they
were released, where fishermen can catch them.
An alternative method to hatcheries is to use spawning channels. These are artificial streams, usually parallel to an existing
stream, with concrete or rip-rap sides and gravel bottoms. Water from the adjacent stream is piped into the top of the
channel, sometimes via a header pond, to settle out sediment. Spawning success is often much better in channels than in
adjacent streams due to the control of floods, which in some years can wash out the natural redds. Because of the lack of
floods, spawning channels must sometimes be cleaned out to remove accumulated sediment. The same floods that destroy
natural redds also clean the regular streams. Spawning channels preserve the natural selection of natural streams, as there
is no benefit, as in hatcheries, to use prophylactic chemicals to control diseases.
Farm-raised salmon are fed the carotenoids astaxanthin and canthaxanthin to match their flesh colour to wild salmon[75]
to improve their marketability.[76]
One proposed alternative to the use of wild-caught fish as feed for the salmon, is the use of soy-based products. This
should be better for the local environment of the fish farm, but producing soy beans has a high environmental cost for the
producing region.
Another possible alternative is a yeast-based coproduct of bioethanol production, proteinaceous fermentation biomass.
Substituting such products for engineered feed can result in equal (sometimes enhanced) growth in fish.[77] With its
increasing availability, this would address the problems of rising costs for buying hatchery fish feed.
Yet another attractive alternative is the increased use of seaweed. Seaweed provides essential minerals and vitamins for
growing organisms. It offers the advantage of providing natural amounts of dietary fiber and having a lower glycemic load
than grain-based fish meal.[77] In the best-case scenario, widespread use of seaweed could yield a future in aquaculture
that eliminates the need for land, freshwater, or fertilizer to raise fish.[78]
10 CHAPTER 1. SALMON
1.7 Management
Main article: Environmental issues with salmon
See also: Salmon conservation and Aquaculture of salmon § Issues
The population of wild salmon declined markedly in recent decades, especially North Atlantic populations, which spawn
in the waters of western Europe and eastern Canada, and wild salmon in the Snake and Columbia River systems in
northwestern United States.
Salmon population levels are of concern in the Atlantic and in some parts of the Pacific. Alaska fishery stocks are still
abundant, and catches have been on the rise in recent decades, after the state initiated limitations in 1972.[79][80] Some
of the most important Alaskan salmon sustainable wild fisheries are located near the Kenai River, Copper River, and in
Bristol Bay. Fish farming of Pacific salmon is outlawed in the United States Exclusive Economic Zone, however, there is
a substantial network of publicly funded hatcheries,[81] and the State of Alaska’s fisheries management system is viewed
as a leader in the management of wild fish stocks. In Canada, returning Skeena River wild salmon support commercial,
subsistence and recreational fisheries, as well as the area’s diverse wildlife on the coast and around communities hundreds
of miles inland in the watershed. The status of wild salmon in Washington is mixed. Of 435 wild stocks of salmon and
steelhead, only 187 of them were classified as healthy; 113 had an unknown status, one was extinct, 12 were in critical
condition and 122 were experiencing depressed populations.[82]
The commercial salmon fisheries in California have been either severely curtailed or closed completely in recent years,
due to critically low returns on the Klamath and or Sacramento rivers, causing millions of dollars in losses to commercial
fishermen.[83] Both Atlantic and Pacific salmon are popular sportfish.
Salmon populations have been established in all the Great Lakes. Coho stocks were planted by the state of Michigan in
the late 1960s to control the growing population of non-native alewife. Now Chinook (king), Atlantic, and coho (silver)
salmon are annually stocked in all Great Lakes by most bordering states and provinces. These populations are not self-
sustaining and do not provide much in the way of a commercial fishery, but have led to the development of a thriving
sport fishery.
1.8 As food
Main article: Salmon as food
Salmon is a popular food. Classified as an oily fish,[84] salmon is considered to be healthy due to the fish’s high protein,
high omega-3 fatty acids, and high vitamin D[85] content. Salmon is also a source of cholesterol, with a range of 23–214
mg/100 g depending on the species.[86] According to reports in the journal Science, farmed salmon may contain high
levels of dioxins. PCB (polychlorinated biphenyl) levels may be up to eight times higher in farmed salmon than in wild
salmon,[87] but still well below levels considered dangerous.[88][89] Nonetheless, according to a 2006 study published in the
Journal of the American Medical Association, the benefits of eating even farmed salmon still outweigh any risks imposed
by contaminants.[90] Farmed salmon has a high omega 3 fatty acid content comparable to wild salmon.[91] The type of
omega-3 present may not be a factor for other important health functions.
Salmon flesh is generally orange to red, although white-fleshed wild salmon with white-black skin colour occurs. The
natural colour of salmon results from carotenoid pigments, largely astaxanthin, but also canthaxanthin, in the flesh.[92]
Wild salmon get these carotenoids from eating krill and other tiny shellfish.
The vast majority of Atlantic salmon available around the world are farmed (almost 99%),[93] whereas the majority of
Pacific salmon are wild-caught (greater than 80%). Canned salmon in the US is usually wild Pacific catch, though some
farmed salmon is available in canned form. Smoked salmon is another popular preparation method, and can either be hot
or cold smoked. Lox can refer to either cold-smoked salmon or salmon cured in a brine solution (also called gravlax).
Traditional canned salmon includes some skin (which is harmless) and bone (which adds calcium). Skinless and boneless
canned salmon is also available.
Raw salmon flesh may contain Anisakis nematodes, marine parasites that cause anisakiasis. Before the availability of
refrigeration, the Japanese did not consume raw salmon. Salmon and salmon roe have only recently come into use in
1.9. HISTORY 11
making sashimi (raw fish) and sushi.

To the Indigenous peoples of the Pacific Northwest Coast, salmon is considered a vital part of the diet. Specifically, the
indigenous peoples of Haida Gwaii, located near former Queen Charlotte Island in British Columbia, rely on salmon as
one of their main sources of food, although many other bands have fished Pacific waters for centuries.[94] Salmon are not
only ancient and unique, but it is important because it is expressed in culture, art forms, and ceremonial feasts. Annually,
salmon spawn in Haida, feeding on everything on the way upstream and down.[94] In Haida, salmon is referred to as
“tsiin” [94] There are many forms of preparing it including: smoking, baking, frying, making soup, or candied salmon
Historically, there has always been enough salmon, as people would not overfish, and only took what they needed.[95] In
2003, a report on First Nation participation in commercial fisheries, including salmon, commissioned by BC’s Ministry
of Agriculture, Food and Fisheries found that there were 595 First Nation-owned and operated commercial vessels in
the province. Of those vessels, First Nations’ members owned 564.[95] However, employment within the industry has
decreased overall by 50% in the last decade, with 8,142 registered commercial fishermen in 2003. This has affected
employment for many fisherman, who rely on salmon as a source of income.
Black bears also rely on salmon as food. The leftovers the bears leave behind are considered important nutrients for
the forest, such as the soil, trees, and plants. In this sense, the salmon feed the forest and in return receive clean water
and gravel in which to hatch and grow, sheltered from extremes of temperature and water flow in times of high and
low rainfall.[94] However, the condition of the salmon in Haida has been affected in recent decades. Due to logging
and development, much of the salmon’s habitat (i.e.: Ain River) has been destroyed, resulting in the fish being close to
endangered.[94] For residents, this has resulted in limits on catches, in turn, has affected families diets, and cultural events
such as feasts. Some of the salmon systems in danger include: the Davidon, Naden, Mamim, and Mathers.[94] It is clear
that further protection is needed for salmon, such as their habitats, where logging commonly occurs.
1.9 History
See also: Salmon cannery
The salmon has long been at the heart of the culture and livelihood of coastal dwellers. Many people of the northern
Pacific shore had a ceremony to honor the first return of the year. For many centuries, people caught salmon as they swam
upriver to spawn. A famous spearfishing site on the Columbia River at Celilo Falls was inundated after great dams were
built on the river. The Ainu, of northern Japan, trained dogs to catch salmon as they returned to their breeding grounds
en masse. Now, salmon are caught in bays and near shore.
The Columbia River salmon population is now less than 3% of what it was when Lewis and Clark arrived at the river.[96]
Salmon canneries established by settlers beginning in 1866 had a strong negative impact on the salmon population. In
his 1908 State of the Union address, U.S. President Theodore Roosevelt observed that the fisheries were in significant
decline:[97][98]
The salmon fisheries of the Columbia River are now but a fraction of what they were twenty-five years
ago, and what they would be now if the United States Government had taken complete charge of them by
intervening between Oregon and Washington. During these twenty-five years the fishermen of each State have
naturally tried to take all they could get, and the two legislatures have never been able to agree on joint action
of any kind adequate in degree for the protection of the fisheries. At the moment the fishing on the Oregon side
is practically closed, while there is no limit on the Washington side of any kind, and no one can tell what the
courts will decide as to the very statutes under which this action and non-action result. Meanwhile very few
salmon reach the spawning grounds, and probably four years hence the fisheries will amount to nothing; and
this comes from a struggle between the associated, or gill-net, fishermen on the one hand, and the owners of
the fishing wheels up the river.
On the Columbia River the Chief Joseph Dam completed in 1955 completely blocks salmon migration to the upper
Columbia River system.
The Fraser River salmon population was affected by the 1914 slide caused by the Canadian Pacific Railway at Hells Gate.
The 1917 catch was one quarter of the 1913 catch.[99]
1.10 Mythology
The salmon is an important creature in several strands of Celtic mythology and poetry, which often associated them with
wisdom and venerability. In Irish mythology, a creature called the Salmon of Knowledge[100] plays key role in the tale
The Boyhood Deeds of Fionn. In the tale, the Salmon will grant powers of knowledge to whoever eats it, and is sought by
poet Finn Eces for seven years. Finally Finn Eces catches the fish and gives it to his young pupil, Fionn mac Cumhaill,
to prepare it for him. However, Fionn burns his thumb on the salmon’s juices, and he instinctively puts it in his mouth.
In so doing, he inadvertently gains the Salmon’s wisdom. Elsewhere in Irish mythology, the salmon is also one of the
incarnations of both Tuan mac Cairill[101] and Fintan mac Bóchra.[102]
Salmon also feature in Welsh mythology. In the prose tale Culhwch and Olwen, the Salmon of Llyn Llyw is the oldest
animal in Britain, and the only creature who knows the location of Mabon ap Modron. After speaking to a string of other
ancient animals who do not know his whereabouts, King Arthur's men Cai and Bedwyr are led to the Salmon of Llyn
Llyw, who lets them ride its back to the walls of Mabon’s prison in Gloucester.
In Norse mythology, after Loki tricked the blind god Höðr into killing his brother Baldr, Loki jumped into a river and
transformed himself into a salmon to escape punishment from the other gods. When they held out a net to trap him he
attempted to leap over it but was caught by Thor who grabbed him by the tail with his hand, and this is why the salmon’s
tail is tapered.[103]
Salmon are central spiritually and culturally to Native American mythology on the Pacific coast, from the Haida and Coast
Salish peoples, to the Nuu-chah-nulth peoples in British Columbia.[104]
1.11 References
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[60] Crosier, Danielle M.; Molloy, Daniel P.; Bartholomew, Jerri. “Whirling Disease – Myxobolus cerebralis" (PDF). Retrieved 13
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[61] Boyce, N.P.; Kabata, Z.; Margolis, L. (1985). “Investigation of the Distribution, Detection, and Biology of Henneguya salmini-
cola (Protozoa, Myxozoa), a Parasite of the Flesh of Pacific Salmon”. Canadian Technical Report of Fisheries and Aquatic
Sciences (1450): 55.
[62] Sea Lice and Salmon: Elevating the dialogue on the farmed-wild salmon story Watershed Watch Salmon Society, 2004
[63] Bravo, S. (2003). “Sea lice in Chilean salmon farms”. Bull. Eur. Assoc. Fish Pathol. 23: 197–200.
[64] Morton, A.; R. Routledge; C. Peet & A. Ladwig (2004). “Sea lice (Lepeophtheirus salmonis) infection rates on juvenile pink
(Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon in the nearshore marine environment of British Columbia,
Canada”. Canadian Journal of Fisheries and Aquatic Sciences. 61 (2): 147–157. doi:10.1139/f04-016.
[65] Peet, C. R. (2007). Interactions between sea lice (Lepeophtheirus salmonis and Caligus clemensii), juvenile salmon (On-
corhynchus keta and Oncorhynchus gorbuscha) and salmon farms in British Columbia. M.Sc. Thesis, University of Victoria,
Victoria, British Columbia, Canada.
[66] Krkošek, M.; A. Gottesfeld; B. Proctor; D. Rolston; C. Carr-Harris; M.A. Lewis (2007). “Effects of host migration, diversity
and aquaculture on sea lice threats to Pacific salmon populations”. Proceedings of the Royal Society B: Biological Sciences. 274
(1629): 3141–9. doi:10.1098/rspb.2007.1122. PMC 2293942 . PMID 17939989.
[67] Morton, A.; R. Routledge; M. Krkošek (2008). “Sea Louse Infestation in Wild Juvenile Salmon and Pacific Herring Associated
with Fish Farms off the East-Central Coast of Vancouver Island, British Columbia” (PDF). North American Journal of Fisheries
Management. 28 (2): 523–532. doi:10.1577/M07-042.1.
[68] Krkošek, M.; M.A. Lewis; A. Morton; L.N. Frazer; J.P. Volpe (2006). “Epizootics of wild fish induced by farm fish”. Pro-
ceedings of the National Academy of Sciences. 103 (42): 15506–10. doi:10.1073/pnas.0603525103. PMC 1591297 . PMID
17021017.
[69] Krkošek, Martin (2007). “Declining Wild Salmon Populations in Relation to Parasites from Farm Salmon”. Science. 318
(5857): 1772–5. doi:10.1126/science.1148744. PMID 18079401.
[70] Weissglas G and Appelblad H (1997) “Wild-spawning Baltic salmon – A natural resource redefined: From food to toys for
“boys"? In: A-L Toivonen and Tuunainen P (Eds) Socio-economics of recreational fishery, pp. 89–95, Nordic Council of
Ministers. ISBN 9789289301206.
1.11. REFERENCES 15
[71] Shaw, Susan and Muir, James F. (1987) Salmon: Economics and Marketing Page 250, Springer. ISBN 9780709933441.
[72] Seafood Choices Alliance (2005) It’s all about salmon
[73] Wright, Matt. “Fish farms drive wild salmon populations toward extinction”, EurekAlert, 13 December 2007.
[74] Naylor, Rosamond L. “Nature’s Subsidies to Shrimp and Salmon Farming” (PDF). Science; 10/30/98, Vol. 282 Issue 5390,
p883.
[75] “Pigments in Salmon Aquaculture: How to Grow a Salmon-colored Salmon”. Retrieved 26 August 2007. Astaxanthin (3,3'-
hydroxy-β,β-carotene-4,4'-dione) is a carotenoid pigment, one of a large group of organic molecules related to vitamins and
widely found in plants. In addition to providing red, orange, and yellow colours to various plant parts and playing a role in
photosynthesis, carotenoids are powerful antioxidants, and some (notably various forms of carotene) are essential precursors to
vitamin A synthesis in animals.
[76] Guilford, Gwynn (12 March 2015). “Here’s why your farmed salmon has color added to it”. Quartz (publication). Retrieved 12
March 2015.
[77] Rust, Michael B. et al. (November 2010) The Future of Aquafeeds. aquaculture.noaa.gov. p. 56.
[78] Salmon Recovery Planning. nwr.noaa.gov. p. 57.
[79] “1878–2010, Historical Commercial Salmon Catches and Exvessel Values”. Alaska Department of Fish and Game. Retrieved
6 August 2011.
[80] Viechnicki, Joe (3 August 2011). “Pink salmon numbers record setting in early season”. KRBD Public Radio in Ketchikan,
Alaska. Retrieved 6 August 2011.
[81] media.aprn.org|low fish returns in Southeast this summer have been tough on the region’s hatcheries
[82] Johnson, Thom H.; Lincoln, Rich; Graves, Gary R. & Gibbons, Robert G. “Status of Wild Salmon and Steelhead Stocks in
Washington State”. In Stouder, Deanna J.; Bisson, Peter A. & Naiman, Robert J. Pacific Salmon and Their Ecosystems: Status
and Future Options. Springer. pp. 127–144. doi:10.1007/978-1-4615-6375-4_11. ISBN 978-1-4615-6375-4.
[83] Hackett, S. & D. Hansen. “Cost and Revenue Characteristics of the Salmon Fisheries in California and Oregon”. Retrieved 1
June 2009.
[84] “What’s an oily fish?". Food Standards Agency. 24 June 2004. Archived from the original on 19 October 2014.
[85] “Dietary Supplement Fact Sheet: Vitamin D”. National Institutes of Health. Archived from the original on 13 December 2007.
Retrieved 13 December 2007.
[86] “Cholesterol: Cholesterol Content in Seafoods (Tuna, Salmon, Shrimp)". Retrieved 13 December 2007.
[87] Hites, R. A.; Foran, J. A.; Carpenter, D. O.; Hamilton, M. C.; Knuth, B. A.; Schwager, S. J. (2004). “Global Assessment of
Organic Contaminants in Farmed Salmon”. Science. 303 (5655): 226–9. doi:10.1126/science.1091447. PMID 14716013.
[88] “Farmed vs. wild salmon – which is better?". CTV News. Retrieved 28 April 2013.
[89] Foran, J. A.; Carpenter, D. O.; Hamilton, M. C.; Knuth, B. A.; Schwager, S. J. (2005). “Risk-Based Consumption Advice
for Farmed Atlantic and Wild Pacific Salmon Contaminated with Dioxins and Dioxin-like Compounds”. Environmental Health
Perspectives. 113 (5): 552–556. doi:10.1289/ehp.7626. PMC 1257546 . PMID 15866762.
[90] Mozaffarian, Dariush; Rimm, Eric B. (2006). “Fish Intake, Contaminants, and Human Health”. JAMA. 296 (15): 1885–99.
doi:10.1001/jama.296.15.1885. PMID 17047219.
[91] Raatz, S. K.; Rosenberger, T. A.; Johnson, L. K.; Wolters, W. W.; Burr, G. S.; Picklo Mj, Sr (2013). “Dose-Dependent Con-
sumption of Farmed Atlantic Salmon (Salmo salar) Increases Plasma Phospholipid n-3 Fatty Acids Differentially”. Journal of
the Academy of Nutrition and Dietetics. 113 (2): 282–7. doi:10.1016/j.jand.2012.09.022. PMC 3572904 . PMID 23351633.
[92] “Opinion of the Scientific Committee on Animal Nutrition on the use of canthaxanthin in feedingstuffs for salmon and trout, lay-
ing hens, and other poultry.” (PDF). European Commission — Health & Consumer Protection Directorate. pp. 6–7. Retrieved
13 November 2006.
[93] Montaigne, Fen. “Everybody Loves Atlantic Salmon: Here’s the Catch...”. National Geographic. Retrieved 17 November 2006.
[94] Haida Gwaii Strategic Land Use Agreement. haidanation.ca (September 2007)
[95] Garner, Kerri and Parfitt, Ben (April 2006) First Nations, Salmon Fisheries and the Rising Importance of Conservation. Van-
couver, BC: Prepared for the Pacific Fisheries Resource Conservation Council. ISBN 1-897110-28-6
[96] “Endangered Salmon”. U.S. Congressman Jim McDermott. Archived from the original on 15 November 2006. Retrieved 17
November 2006.
[97] “Columbia River History: Commercial Fishing”. Northwest Power and Conservation Council. 2010. Retrieved 26 January
2012.
[98] Roosevelt, Theodore (8 December 1908). “State of the Union Address Part II by Theodore Roosevelt”. Retrieved 31 January
2012.
[99] Babcock, John P. (1920). Fraser River Salmon Situation a Reclamation Project. Victoria, B.C.: W. H. Cullin. p. 5
[100] “The Salmon of Knowledge. Celtic Mythology, Fairy Tale”. Luminarium.org. 18 January 2007. Retrieved 1 June 2010.
[101] “The Story of Tuan mac Cairill”. Maryjones.us. Retrieved 18 March 2010.
[102] “The Colloquy between Fintan and the Hawk of Achill”. Ucc.ie. Retrieved 18 March 2010.
[103] “The Poetic Edda,”. Translated by Henry Adams Bellows. Retrieved 27 April 2011.
[104] Salmon Culture of the Pacific Northwest Tribes. Columbia River Inter-Tribal Fish Commission
1.12 Further reading

• Atlas of Pacific Salmon, Xanthippe Augerot and the State of the Salmon Consortium, University of California Press,
2005, hardcover, 152 pages, ISBN 0-520-24504-0
• Making Salmon: An Environmental History of the Northwest Fisheries Crisis, Joseph E. Taylor III, University of
Washington Press, 1999, 488 pages, ISBN 0-295-98114-8
• Trout and Salmon of North America, Robert J. Behnke, Illustrated by Joseph R. Tomelleri, The Free Press, 2002,
hardcover, 359 pages, ISBN 0-7432-2220-2
• Come back, salmon, By Molly Cone, Sierra Club Books, 48 pages, ISBN 0-87156-572-2 – A book for juveniles
describes the restoration of 'Pigeon Creek'.
• The salmon: their fight for survival, By Anthony Netboy, 1973, Houghton Mifflin Co., 613 pages, ISBN 0-395-
14013-7
• A River Lost, by Blaine Harden, 1996, WW Norton Co., 255 pages, ISBN 0-393-31690-4. (Historical view of the
Columbia River system).
• River of Life, Channel of Death, by Keith C. Peterson, 1995, Confluence Press, 306 pages, ISBN 978-0-87071-
496-2. (Fish and dams on the Lower Snake River.)
• Salmon, by Dr Peter Coates, 2006, ISBN 1-86189-295-0
• Lackey, Robert T (2000) “Restoring Wild Salmon to the Pacific Northwest: Chasing an Illusion?" In: Patricia Koss
and Mike Katz (Eds) What we don't know about Pacific Northwest fish runs: An inquiry into decision-making under
uncertainty, Portland State University, Portland, Oregon. Pages 91–143.
• Mills D (2001) “Salmonids” In: pp. 252–261, Steele JH, Thorpe SA and Turekian KK (2010) Marine Biology: A
Derivative of the Encyclopedia of Ocean Sciences, Academic Press. ISBN 978-0-08-096480-5.
• NEWS January 31, 2007: U.S. Orders Modification of Klamath River – Dams Removal May Prove More Cost-
Effective for allowing the passage of Salmon
1.13. EXTERNAL LINKS 17
• Salmon age and sex composition and mean lengths for the Yukon River area, 2004 / by Shawna Karpovich and
Larry DuBois. Hosted by Alaska State Publications Program.
• "Studies in the Natural History of the Sacramento Salmon". Popular Science Monthly. 61. July 1902.
• Trading Tails: Linkages Between Russian Salmon Fisheries and East Asian Markets. Shelley Clarke. (November
2007). 120pp. ISBN 978-1-85850-230-4.
• The Salmons Tale one of the twelve Ionan Tales by Jim MacCool
1.13 External links

• “Last Stand of the American Salmon,” G. Bruce Knecht for Men’s Journal
• Plea for the Wanderer, an NFB documentary on West Coast salmon
• Genetic Status of Atlantic Salmon in Maine: Interim Report (2002) Online book
• Arctic Salmon on Facebook research project studying Pacific salmon in the Arctic and potential links to climate
change
• University of Washington Libraries Digital Collections – Salmon Collection A collection of documents describing
salmon of the Pacific Northwest.
• Salmon Nation A movement to create a bioregional community, based on the historic spawning area of Pacific
salmon (CA to AK).
• Arctic Salmon Pacific salmon distribution and abundance seems to be increasing in the Arctic. Links to a Canadian
research project documenting changes in Pacific salmon and studying Pacific salmon ecology in the Arctic.
Henneguya salminicola, a myxozoan parasite commonly found in the flesh of salmonids on the West Coast of Canada, in coho salmon
Wild fisheries – commercial capture of all true wild salmon species 1950–2010, as reported by the FAO[1]
Angler and gillie land a salmon, Scotland
Aquaculture production of all true salmon species 1950–2010, as reported by the FAO [1]
Artificially incubated chum salmon

Rainbow trout farm in an archipelago of Finland

Spawning sockeye salmon in Becharof Creek, Becharof Wilderness, Alaska

Salmon sashimi
Seine fishing for salmon – Wenzel Hollar, 1607–1677

Scales on the “Big Fish” or “Salmon of Knowledge” celebrates the return of fish to the River Lagan
Chapter 2
Atlantic salmon
This article is about a particular species of fish. For the food, see Salmon as food.
The Atlantic salmon (Salmo salar) is a species of ray-finned fish in the family Salmonidae. It is found in the northern
Atlantic Ocean, in rivers that flow into the north Atlantic and, due to human introduction, in the north Pacific Ocean.[2][3]
Atlantic salmon have long been the target of recreational and commercial fishing, and this, as well as habitat destruction,
has reduced their numbers significantly; the species is the subject of conservation efforts in several countries.
2.1 Nomenclature
The Atlantic salmon was given its scientific binomial name by 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,[4]
but more likely meaning “resident of salt water”. 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.
Other names used to reference Atlantic salmon are: bay salmon, black salmon, caplin-scull salmon, Sebago salmon, silver
salmon, fiddler, or outside salmon. 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 or
ouananiche.
2.2 Description
The species is the longest and heaviest in 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.[5] But specimens can be much larger. An Atlantic salmon netted in
1960 in Scotland, in the estuary of the river Hope, was weighed in at 49.44 kg (109.0 lb), while another netted in 1925
in Norway measured 160.65 cm (63.25 in) in length.[6]
The colouration 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 colouration. The salmon has a fusiform body, and well-developed teeth. All fins, save the
adipose, are bordered with black.
27
28 CHAPTER 2. ATLANTIC SALMON
Atlantic salmon are among the largest salmon species
2.3 Distribution and habitat
The distribution of Atlantic salmon depends on water temperature. Because of climate change, some of the species’
southern populations, in Spain and other warm countries, are growing smaller[8][9] and are expected to be extirpated
soon. Before human influence, the natural breeding grounds of Atlantic salmon were rivers in Europe and the eastern
coast of North America. When North America was settled by Europeans, eggs were brought on trains to the west coast
and introduced into the rivers there. Other attempts to bring Atlantic salmon to new settlements were made; e.g. New
Zealand. But since there are no suitable ocean currents on New Zealand, most of these introductions failed. There is at
least one landlocked population of Atlantic salmon on New Zealand, where the fish never go out to sea.
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
behavioral 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.
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 kilometers at sea; instead they surf through sea
surface currents. When they reach their natal river they find it by smell; only 5% of Atlantic salmon go up the wrong river.
Thus, the habitat of Atlantic salmon is the river where they are born and the sea surface currents that are connected to
that river in a circular path.
Wild salmon disappeared from many rivers during the twentieth century due to overfishing and habitat change.[2] By 2000
the numbers of Atlantic salmon had dropped to critically low levels.[10]
2.4. DIET 29
Ocean migration of Atlantic salmon from the Connecticut River[7]
2.4 Diet
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. The most commonly eaten
foods include caddisflies, blackflies, mayflies, and stoneflies.[2]
As adults, the fish feed on much larger food: Arctic squid, sand eels, amphipods, Arctic shrimp, and sometimes herring,
and the fishes’ size increases dramatically.[2]
2.5 Behavior
Fry and parr have been said to be territorial, but evidence showing them to guard territories is inconclusive. While they
may occasionally be aggressive towards each other, the social hierarchy is still unclear. Many have been found to school,
especially when leaving the estuary.
Adult Atlantic salmon are considered much more aggressive than other salmon, and are more likely to attack other fish
than others. A matter of concern is where they have become an invasive threat, attacking native salmon, such as Chinook
salmon and coho salmon.[2]
2.6 Life stages
Life cycle of the Atlantic salmon
See also: Juvenile salmon
Most Atlantic salmon follow an anadromous fish migration pattern,[3] 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.
Atlantic salmon do not require saltwater. Numerous examples of fully freshwater (i.e., “landlocked”) populations of the
species exist throughout the Northern Hemisphere,[3] including a now extinct population in Lake Ontario, which have been
2.6. LIFE STAGES 31
shown in recent studies to have spent their entire life cycle in watershed of the lake.[11] In North America, the landlocked
strains are frequently known as ouananiche.
2.6.1 Freshwater phase

The freshwater phases of Atlantic salmon vary between one and eight years, according to river location.[12] 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.[12] The average age correlates to temperature exceeding 7 °C (45 °F).[2]
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.
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.
2.6.2 Saltwater phases

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.
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.
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%.[2]
• Very young fertilized salmon eggs, notice the developing eyes and neural tube
• Newly hatched alevin feed on their yolk sacs

• When the alevin or sac fry have depleted their yolk sac or “lunch box”, they
emerge from the gravel habitat of their redd (nest) to look for food as fry.
• The fry become parr, and pick home rocks or plants in the streambed from
which they dart out to capture insect larvae and other passing food
• When the parr are ready for migration to the ocean, they become smolt
2.7 Breeding
See also: Salmon run
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.
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.
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.[3][13] Migration and spawning exact an enormous physiological
toll on individuals, such that repeat spawners are the exception rather than the norm.[13] 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.[12]
2.8. HYBRIDIZATION 33
Fish ladder for Atlantic salmon constructed in the middle of a large weir
2.8 Hybridization
When in shared breeding habitats, Atlantic salmon will hybridize with brown trout (Salmo trutta).[14][15][16] Hybrids be-
tween 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 they were not significantly heterogeneous 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.[17]
2.9 Aquaculture
See also: Salmon in aquaculture
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 and a supplemental food fishery. However, the wild
Atlantic salmon fishery is commercially dead; after extensive habitat damage and overfishing, wild fish make up only 0.5%
of the Atlantic salmon available in world fish markets. The rest are farmed, predominantly from aquaculture in Norway,
Chile, Canada, the UK, Ireland, Faroe Islands, Russia and Tasmania in Australia. Sport fishing communities, mainly from
Iceland and Scandinavia, have joined in the North Atlantic Salmon Fund to buy away commercial quotas in an effort to
save the wild species of Salmo salar.[13]
Atlantic salmon marine cages in the Faroe Islands
2.9.1 Process
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.[18]
Some researchers have even studied cryopreservation of their eggs.[19]
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 USA, or parts of Europe.[13]
Generally, cages are made of two nets. Inner nets, which wrap around the cages, hold the salmon. Outer nets, which are
held by floats, keep predators out.[18]
2.9.2 Controversy
Some Atlantic salmon escape from cages at sea. These salmon tend to lessen the genetic diversity of the species leading
to lower survival rates, and lower catch rates. On the west coast of North America, the non-native salmon can be an
invasive threat, especially in Alaska and parts of Canada. This causes them to compete with native salmon for resources.
Extensive efforts are underway to prevent the spread of Atlantic salmon in the Pacific and elsewhere.[20]
On the west coast of northern America, aquaculturists have taken care to ensure the non-native salmon cannot escape
from their open-net pens, and escape is no longer considered a major concern. Evidence of Atlantic salmon surviving and
establishing wild populations in the Pacific is lacking.
From 1905 until 1935, 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
2.10. HUMAN IMPACT 35
never materialized. 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.[21]
2.10 Human impact
Seine fishing for salmon – Wenzel Hollar, 1607–1677
Atlantic salmon were once abundant throughout the North Atlantic. European fishermen gillnetted for Atlantic salmon in
rivers using hand-made nets for at least several centuries.[22] Wood and stone weirs along streams and ponds were used for
millennia to harvest salmon in the rivers of Maine and New England,[23] and gillnetting was an early fishing technology
in colonial America.[24]
Human activities have heavily damaged salmon populations across their range. The major impacts were from overfishing
and habitat change, and the new threat from competitive farmed fish. 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.[25][11] When dams were constructed on the Oswego River, their spawning areas were
cut off and they went extinct locally.
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.[26] Overfishing at sea is generally considered the primary factor.
Beginning around 1990, the rates of Atlantic salmon mortality at sea more than doubled. In the western Atlantic, fewer
than 100,000 of the important multiple sea-winter salmon were returning. Rivers of the coast of Maine, southern New
Brunswick and much of mainland Nova Scotia saw runs drop precipitously, and even disappear. To find out more about the
increased mortality rate, a concerted international effort has been organized by the North Atlantic Salmon Conservation
Organization.[27]
Possibly because of improvements in ocean feeding grounds, returns in 2008 were 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.[28]
2.11 Recovery
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 introduced successfully, but the
percentage of salmon reproducing naturally is very low. Most are stocked annually. Atlantic salmon were native to Lake
Ontario, but were extirpated by habitat loss and overfishing in the late 19th century. The state of New York has since
stocked its adjoining rivers and tributaries, and in many cases does not allow active fishing.[3][20]
Historically, the Housatonic River, and its Naugatuck River tributary, hosted the southernmost Atlantic salmon spawning
runs in the United States.[29][30] However, there are historical accounts as early as 1609 from Henry Hudson that Atlantic
salmon once ran up the Hudson River.[31]
In the early 1990s, Carlson challenged the notion that Atlantic salmon were prehistorically abundant in New England,
when the climate was warmer as it is now. This idea was based on a paucity of bone data in archaeological sites relative to
other fish species and claimed that historical observer records were exaggerated.[32][33] However, arguments that lack of
archaeological bone fragments rule out historic abundance are more recently disputed because salmon bones are rare at
sites that still have large salmon runs, salmonid bones in general are poorly recovered relative to other fish species, and that
salmon remains may have been diluted by the large numbers of other anadromous fishes using northeastern streams.[34][35]
In addition, fish scale evidence dating to 10,000 years BP places Atlantic salmon in a coastal New Jersey pond.[36]
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. In Ontario, the Atlantic Salmon Restoration Program[37] was started in 2006, and is one of the largest freshwater
conservation programs in North America. It has stocked Lake Ontario with over 700,000 young Atlantic salmon. 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.[25]
• A migrating salmon was observed in Ontario’s Credit River in November 2007.[25]
• As of 2013, there has been some success in establishing Atlantic salmon in Fish Creek, a tributary of Oneida Lake
in central New York.[38]
• In November 2015, salmon nests were observed in Connecticut in the Farmington River, a tributary of the Con-
necticut River where Atlantic salmon had not been observed spawning since “probably the Revolutionary War".[39]
A 45-year, $25 million federal government effort to restore wild Atlantic salmon to the Connecticut River water-
shed was discontinued in 2012, but now appears to have been successful.[40]
Atlantic salmon still remains a popular fish for human consumption.[3] It is commonly sold fresh, canned, or frozen.
2.12. BEAVER IMPACT 37
2.12 Beaver impact

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 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 Salmo salar upstream from the dams suggests the dams are
penetrated by parr.[41] Downstream migration of Atlantic salmon smolts was similarly unaffected by beaver dams, even
in periods of low flows.[41]
In a 2003 study, Atlantic salmon and sea-run brown trout/sea trout spawning in the Numedalslågen River and 51 of its
tributaries in southeastern Norway were unhindered by beavers.[42] 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.[43]
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.[41] 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.[44]
2.13 Legislation
The first laws regarding the Atlantic salmon were started nearly 800 years ago.
2.13.1 England and Wales
Edward I instituted a penalty for collecting salmon during certain times of the year. His son Edward II continued, regu-
lating 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.
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.
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 were reduced to 10 regional water authorities (RWAs). Although only the Northumbrian, Welsh,
northwest and southwest RWA’s had considerable salmon populations, all ten also cared for trout and freshwater eels.
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.[2]
2.13.2 Scotland
Legislation in Scotland to help Atlantic salmon began in 1318 by Alexander II. It prohibited certain types of traps in
rivers.
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. Because
the fish were held in such high regard, poachers were severely punished.
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.[2][18]
2.13.3 United States

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. Returns in 2008 have been around 2,000, more than double the 2007
return of 940.
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”.[45]
2.13.4 Canada
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, the government has attempted to
pass a new version of the century-old Fisheries Act through Parliament.
Federal legislation regarding at-risk populations is weak. Inner Bay of Fundy Atlantic salmon runs were declared endan-
gered in 2000. As of 2008, no recovery plan is in place.
It takes constant pressure from nongovernmental organizations, such as the Atlantic Salmon Federation, for improvements
in management, and for initiatives to be considered. For example, the technology for mitigation of acid rain-affected rivers
used in Norway is needed in 54 Nova Scotia rivers. Yet, an initiative of the ASF and the Nova Scotia Salmon Association
raised the funds to get a project in place, in West River-Sheet Harbour.
In Quebec, the daily catch limit for Atlantic salmon is one fish over 63 cm (25 in), two fish under 63 cm (25 in) or one
fish over and one under 63 cm (25 in), provided the smaller fish was the first one caught (a provision designed to prevent
an angler from continuing to fish if a large fish is already in possession). The annual catch limit is seven Atlantic salmon
of any size.
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.[46][47]
2.14 NASCO
The North Atlantic Salmon Conservation Organization is an international council made up of Canada, the European
Union, Iceland, Norway, the Russian Federation, and the United States, with its headquarters in Edinburgh.[48] 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.
2.15 Sustainable consumption

In 2010, Greenpeace International has added the Atlantic salmon to its seafood red list. “The Greenpeace International
seafood red list is a list of fish that are commonly sold in supermarkets around the world, and which have a very high risk
2.16. SEE ALSO 39
of being sourced from unsustainable fisheries”.[49]
2.16 See also

• AquAdvantage salmon, a genetically modified Atlantic salmon
• Atlantic Salmon Federation (ASF)
• Salmon as food
2.17 Notes
[1] Baillie, J. and Groombridge, B. (1996). "Salmo salar". IUCN Red List of Threatened Species. IUCN. 1996: e.T14144A4408913.
doi:10.2305/iucn.uk.1996.rlts.t19855a9026693.en. Retrieved 26 August 2016.
[2] Shearer, W. (1992). The Atlantic Salmon. Halstead Press.
[3] The Audubon Society Field Guide to North American Fishes, Whales & Dolphins. Chanticleer Press. 1983. p. 395.
[4] Barton, M.: “Biology of Fishes.”, pages 198–202 Thompson Brooks/Cole 2007
[5] “Atlantic salmon (Salmo salar)". NOAA Fisheries - Office of Protected Resources.
[6] “Buller F, The Domesday Book of Giant Salmon Volume 1 & 2. Constable (2007) & Constable (2010)
[7] Atlantic Salmon Life Cycle Connecticut River Coordinator’s Office, U.S. Fish and Wildlife Service. Updated: 13 September
2010.
[8] “Same number of fishermen, but less salmon in Spanish rivers”. science daily. August 26, 2011. Retrieved April 30, 2016.
[9] J. L. Horreo, G. Machado-Schiaffino, A. M. Griffiths, D. Bright, J. R. Stevens, E. Garcia-Vazquez. (2011). “Atlantic Salmon at
Risk: Apparent Rapid Declines in Effective Population Size in Southern European Populations.”. Transactions of the American
Fisheries Society, 2011; 140 (3): 605 DOI: 10.1080/00028487.2011.585574.
[10] B. Dempson, C. J. Schwarz, D. G. Reddin, M. F. O’Connell, C. C. Mullins, and C. E. Bourgeois (2001). “Estimation of marine
exploitation rates on Atlantic salmon (Salmo salar L.) stocks in Newfoundland, Canada” (PDF). ICES Journal of Marine Science:
331–341. doi:10.1006/jmsc.2000.1014. Retrieved 7 May 2011.
[11] “Study sheds light on extinct Lake Ontario salmon”. Toronto Star, November 9, 2016, page GT1.
[12] Klemetsen A, Amundsen P-A, Dempson JB, Jonsson B, Jonsson N, O’Connell MF, Mortensen E (2003). “Atlantic salmon
Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories”.
Ecology of Freshwater Fish. doi:10.1034/j.1600-0633.2003.00010.x.
[13] Heen, K. (1993). Salmon Aquaculture. Halstead Press.
[14] Youngson, A. F., Webb, J. H., Thompson, C. E., and Knox, D. 1993. Spawning of escaped farmed Atlantic salmon (Salmo
salar): hybridization of females with brown trout (Salmo trutta). Canadian Journal of Fisheries and Aquatic Sciences, 50:1986-
1990.
[15] Matthews, M. A., Poole, W. R., Thompson, C. E., McKillen, J., Ferguson, A., Hindar, K., and Wheelan, K. F. 2000. Incidence
of hybridization between Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in Ireland. Fisheries Management
and Ecology, 7:337-347.
[16] Seawater tolerance in Atlantic salmon, Salmo salar L., brown trout, Salmo trutta L., and S. salar × S. trutta hybrids smolt. Urke
HA, Koksvik J, Arnekleiv JV, Hindar K, Kroglund F, Kristensen T. Source Norwegian Institute of Water Research, 7462,
Trondheim, Norway. henning.urke(@)niva.no
[17] Natural hybridization between Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in northern Spain by Carlos Garcia
de Leaniz
[18] Sedgwick, S. (1988). Salmon Farming Handbook. Fishing News Books LTD.
[19] N. Bromage (1995). Broodstock Management and Egg and Larval Quality. Blackwell Science.
[20] Mills, D. (1989). Ecology and Management of Atlantic Salmon. Springer-Verlag.
[21] R. M. J. Ginetz (May 2002). “On the Risk of Colonization by Atlantic Salmon in BC waters” (PDF). B.C. Salmon Farmers
Association.
[22] Jenkins, J. Geraint (1974). Nets and Coracles, p. 68. London, David and Charles.
[23] “The River”. The Penobscot River Restoration Trust. Retrieved 19 November 2013.
[24] Netboy, Anthony (1973) The Salmon: Their Fight for Survival, pp. 181-182. Boston, Houghton Mifflin.
[25] Harb, M. “Upstream Battle”, Canadian Geographic Magazine, June 2008, p. 24
[26] “Salmon campaigner lands top award”. BBC News. 22 April 2007.
[27] “Atlantic Salmon”. animallist.weebly.com. Retrieved 19 November 2013.
[28] Carpenter, Murray (26 December 2011). “Shiny Patches in Maine’s Streambeds Are Bright Sign for Salmon”. The New York
Times. Retrieved February 2012. Check date values in: |access-date= (help)
[29] Fay, C., M. Bartron, S. Craig, A. Hecht, J. Pruden, R. Saunders, T. Sheehan, and J. Trial (2006). Status Review for Anadromous
Atlantic Salmon (Salmo salar) in the United States. Report to the National Marine Fisheries Service and U.S. Fish and Wildlife
Service (Report). p. 294. Retrieved July 3, 2016.
[30] Kendall, W. C. (1935). The fishes of New England: the salmon family. Part 2 - the salmons. Boston, Massachusetts: Memoirs
of the Boston Society of Natural History: monographs on the natural history of New England. p. 90. Retrieved July 3, 2016.
[31] W.C. Kendall (1935). The fishes of New England- the salmon family. Part 2 - the salmons. Memoirs of the Boston Society of
Natural History- monographs on the natural history of New England. 9. pp. 1–166. Retrieved July 3, 2016.
[32] Catherine C. Carlson (1988). GP Nicholas, ed. Where’s the salmon? A reevaluation of the role of anadromous fisheries in
aboriginal New England in Holocene human ecology in Northeastern North America. New York: Plenum Press. ISBN 978-
0306428692.
[33] Catherine C. Carlson (1996). “The [In]Significance of Atlantic Salmon”. History Through a Pinhole. 8(3/4 (Fall/Winter).
Retrieved July 3, 2016.
[34] Stephen F. Jane, Keith H. Nislow, Andrew R. Whiteley (September 2014). “The use (and misuse) of archaeological salmon
data to infer historical abundance in North America with a focus on New England”. Reviews in Fish Biology and Fisheries. 24
(3): 943–954. doi:10.1007/s11160-013-9337-3. Retrieved July 3, 2016.
[35] Brian S. Robinson, George L. Jacobson, Martin G. Yates, Arthur E. Spiess, Ellen R. Cowie (October 2009). “Atlantic salmon,
archaeology and climate change in New England”. Journal of Archaeological Science. 36 (10): 2184–2191. doi:10.1016/j.jas.2009.06.001.
Retrieved July 3, 2016.
[36] Robert A. Daniels and Doroty Peteet (November 1998). “Fish scale evidence for rapid post-glacial colonization of an Atlantic
coastal pond”. Global Ecology & Biogeography Letters. 7 (6): 467–476. doi:10.2307/2997716. Retrieved July 3, 2016.
[37] Atlantic Salmon Restoration Program
[38] Figura, David. “Cicero angler lands 27-inch Atlantic salmon in Oneida Lake”. Syracuse.com. Syracuse Media Group. Retrieved
4 February 2016.
[39] Hladky, Gregory B. (25 Dec 2015). “Salmon Found Spawning In Farmington River Watershed For First Time in Centuries”.
Hartford Courant. Tribune Company. Retrieved 4 February 2016.
[40] Jaymi Heimbuch (March 11, 2016). “Wild Atlantic salmon are spawning in Connecticut River for the first time in 200 years”.
Mother Nature Network. Retrieved July 3, 2016.
[41] P. Collen & R. J. Gibson (2001). “The general ecology of beavers (Castor spp.), as related to their influence on stream ecosystems
and riparian habitats, and the subsequent effects on fish – a review” (PDF). Reviews in Fish Biology and Fisheries. 10 (4): 439–
461. doi:10.1023/A:1012262217012.
2.18. REFERENCES 41
[42] Howard Park & Øystein Cock Rønning (2007). “Low potential for restraint of anadromous salmonid reproduction by beaver
Castor fiber in the Numedalslågen river catchment, Norway”. River Research and Applications. 23 (7): 752–762. doi:10.1002/rra.1008.
[43] Barry A. Taylor, Charles MacInnis, Trevor A. Floyd (2010). “Influence of Rainfall and Beaver Dams on Upstream Move-
ment of Spawning Atlantic Salmon in a Restored Brook in Nova Scotia, Canada”. River Research and Applications: 183–193.
doi:10.1002/rra.1252.
[44] Douglas B. Sigourney, Benjamin H. Letcher & Richard A. Cunjak (2006). “Influence of beaver activity on summer growth and
condition of age-2 Atlantic salmon parr”. Transactions of the American Fisheries Society. 135 (4): 1068–1075. doi:10.1577/T05-
159.1.
[45] (16 U.S.C. 1532(19)) http://www.epa.gov/EPA-SPECIES/1998/May/Day-01/e11668.htm[]
[46] “Frequently Asked Questions (FAQ)". Bring Back the Salmon Lake Onterio. Retrieved 17 September 2015.
[47] “Endangered Populations”. Atlantic Salmon Federation. Retrieved 17 September 2015.
[48] “NASCO ~ The North Atlantic Salmon Conservation Organization”. Nasco.int. Retrieved 11 February 2012.
[49] Greenpeace International Seafood Red list Archived 10 April 2010 at the Wayback Machine.
2.18 References
• Atlantic salmon NOAA FishWatch. Retrieved 4 November 2012.
2.19 External links

• Atlantic Salmon Conservation Foundation
• NOAA Fisheries Atlantic salmon page

• Atlantic Salmon Federation
• Issues salmon face – Smithsonian Ocean Portal

• State of Maine, Atlantic Salmon Commission
• University of Massachusetts information regarding Atlantic salmon
• Atlantic salmon’s page on World Wide Fund for Nature’s website

• Alaska Fish and Game Atlantic salmon page
• Atlantic Salmon Trust, an organization working to protect the Atlantic salmon

• Atlantic Salmon Fishing, Information about Atlantic Salmon Fishing
• Ontario Ministry of Natural Resources, Information About Atlantic Salmon Restoration Program
• Atlantic Salmon Fishing Reports & Images
• Salmon Ireland, information on the salmon rivers

Chapter 3
Hydroacoustics
Hydroacoustics is a general term for the study and application of sound in water. The term comes from Greek ὕδωρ,
water,[1] and ακουστική, acoustics. Hydroacoustics, utilizing sonar technology, is most commonly used for detection,
assessment, and monitoring of underwater physical and biological characteristics.
Collecting Multibeam Sonar Data
Hydroacoustics can be utilized to detect the depth of a water body (bathymetry), as well as the presence or absence,
abundance, distribution, size, and behavior of underwater plants and animals. Hydroacoustic sensing involves "passive
acoustics" (listening for sounds) or active acoustics making a sound and listening for the echo, hence the common name
for the device, echo sounder or echosounder.
There are a number of different causes of noise from shipping. These can be subdivided into those caused by the propeller,
those caused by machinery, and those caused by the movement of the hull through the water. The relative importance of
these three different categories will depend, amongst other things, on the ship type [2] One of the main causes of hydro
acoustic noise from fully submerged lifting surfaces is the unsteady separated turbulent flow near the surface’s trailing
42
3.1. RELATED PUBLICATIONS 43
edge that produces pressure fluctuations on the surface and unsteady oscillatory flow in the near wake.The relative motion
between the surface and the ocean creates a turbulent boundary layer (TBL) that surrounds the surface. The noise is
generated by the fluctuating velocity and pressure fields within this TBL.
Specific Applications
• Surveillance Towed Array Sensor System
• Fisheries Acoustics
• Acoustic Doppler Current Profiler for water speed measurement
• Acoustic Camera
• Passive Acoustic Monitoring
3.1 Related Publications

• Quality assurance of hydroacoustic surveys: the repeatability of fish-abundance and biomass estimates in lakes
within and between hydroacoustic systems (free link to document)
• Hydroacoustics as a tool for assessing fish biomass and size distribution associated with discrete shallow water
estuarine habitats in Louisiana
• Acoustic assessment of squid stocks
• Summary of the use of hydroacoustics for quantifying the escapement of adult salmonids (Oncorhynchus and Salmo
spp.) in rivers. Ransom, B.H., S.V. Johnston, and T.W. Steig. 1998. Presented at International Symposium and
Workshop on Management and Ecology of River Fisheries, University of Hull, England, 30 March-3 April 1998
• Multi-frequency acoustic assessment of fisheries and plankton resources. Torkelson,T.C., T.C. Austin, and P.H.
Weibe. 1998. Presented at the 135th Meeting of the Acoustical Society of America and the 16th Meeting of the
International Congress of Acoustics, Seattle, Washington.
3.2 References
• Acoustics Unpacked A great reference for freshwater hydroacoustics for resource assessment
• Inter-Calibration of Scientific Echosounders in the Great Lakes
• Hydroacoustic Evaluation of Spawning Red Hind Aggregations Along the Coast of Puerto Rico in 2002 and 2003
• Feasibility Assessment of Split-Beam Hydroacoustic Techniques for Monitoring Adult Shortnose Sturgeon in the
Delaware River
• Categorising Salmon Migration Behaviour Using Characteristics of Split-beam Acoustic Data
• Evaluation of Methods to Estimate Lake Herring Spawner Abundance in Lake Superior
• Estimating Sockeye Salmon Smolt Flux and Abundance with Side-Looking Sonar
• Herring Research: Using Acoustics to Count Fish.
• Hydroacoustic Applications in Lake, River and Marine environments for study of plankton, fish, vegetation, sub-
strate or seabed classification, and bathymetry.
• Hydroacoustics: Rivers (in: Salmonid Field Protocols Handbook: Chapter 4)

44 CHAPTER 3. HYDROACOUSTICS
• Hydroacoustics: Lakes and Reservoirs (in: Salmonid Field Protocols Handbook: Chapter 5)
• PAMGUARD: An Open-Source Software Community Developing Marine Mammal Acoustic Detection and Lo-
calisation Software to Benefit the Marine Environment; http://www.pamguard.org/home.shtml
[1] Liddell, H.G. & Scott, R. (1940). A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with
the assistance of. Roderick McKenzie. Oxford: Clarendon Press.
[2] reducing underwater noise pollution from large commercial vessels

Chapter 4
Underwater acoustics
Output of a computer model of underwater acoustic propagation in a simplified ocean environment.
Underwater acoustics is the study of the propagation of sound in water and the interaction of the mechanical waves that
constitute sound with the water and its boundaries. The water may be in the ocean, a lake or a tank. Typical frequencies
associated with underwater acoustics are between 10 Hz and 1 MHz. The propagation of sound in the ocean at frequencies
lower than 10 Hz is usually not possible without penetrating deep into the seabed, whereas frequencies above 1 MHz are
rarely used because they are absorbed very quickly. Underwater acoustics is sometimes known as hydroacoustics.
The field of underwater acoustics is closely related to a number of other fields of acoustic study, including sonar, transduction,
acoustic signal processing, acoustical oceanography, bioacoustics, and physical acoustics.
45
46 CHAPTER 4. UNDERWATER ACOUSTICS
4.1 History
Underwater sound has probably been used by marine animals for millions of years. The science of underwater acoustics
began in 1490, when Leonardo da Vinci wrote the following,[1]
“If you cause your ship to stop and place the head of a long tube in the water and place the outer extremity
to your ear, you will hear ships at a great distance from you.”
In 1687 Isaac Newton wrote his Mathematical Principles of Natural Philosophy which included the first mathematical
treatment of sound. The next major step in the development of underwater acoustics was made by Daniel Colladon, a
Swiss physicist, and Charles Sturm, a French mathematician. In 1826, on Lake Geneva, they measured the elapsed
time between a flash of light and the sound of a submerged ship’s bell heard using an underwater listening horn.[2] They
measured a sound speed of 1435 metres per second over a 17 kilometre(Km) distance, providing the first quantitative
measurement of sound speed in water.[3] The result they obtained was within about 2% of currently accepted values. In
1877 Lord Rayleigh wrote the Theory of Sound and established modern acoustic theory.
The sinking of Titanic in 1912 and the start of World War I provided the impetus for the next wave of progress in
underwater acoustics. Systems for detecting icebergs and U-boats were developed. Between 1912 and 1914, a number
of echolocation patents were granted in Europe and the U.S., culminating in Reginald A. Fessenden's echo-ranger in
1914. Pioneering work was carried out during this time in France by Paul Langevin and in Britain by A B Wood and
associates.[4] The development of both active ASDIC and passive sonar (SOund Navigation And Ranging) proceeded
apace during the war, driven by the first large scale deployments of submarines. Other advances in underwater acoustics
included the development of acoustic mines.
In 1919, the first scientific paper on underwater acoustics was published,[5] theoretically describing the refraction of sound
waves produced by temperature and salinity gradients in the ocean. The range predictions of the paper were experimentally
validated by transmission loss measurements.
The next two decades saw the development of several applications of underwater acoustics. The fathometer, or depth
sounder, was developed commercially during the 1920s. Originally natural materials were used for the transducers, but
by the 1930s sonar systems incorporating piezoelectric transducers made from synthetic materials were being used for
passive listening systems and for active echo-ranging systems. These systems were used to good effect during World
War II by both submarines and anti-submarine vessels. Many advances in underwater acoustics were made which were
summarised later in the series Physics of Sound in the Sea, published in 1946.
After World War II, the development of sonar systems was driven largely by the Cold War, resulting in advances in the
theoretical and practical understanding of underwater acoustics, aided by computer-based techniques.
4.2 Theory
4.2.1 Sound waves in water
A sound wave propagating underwater consists of alternating compressions and rarefactions of the water. These compres-
sions and rarefactions are detected by a receiver, such as the human ear or a hydrophone, as changes in pressure. These
waves may be man-made or naturally generated.
4.2.2 Speed of sound, density and impedance
The speed of sound c (i.e., the longitudinal motion of wavefronts) is related to frequency f and wavelength λ of a wave
by c = f · λ .
This is different from the particle velocity u , which refers to the motion of molecules in the medium due to the sound,
and relates the plane wave pressure p to the fluid density ρ and sound speed c by p = c · u · ρ .
4.2. THEORY 47
The product of c and ρ from the above formula is known as the characteristic acoustic impedance. The acoustic power
(energy per second) crossing unit area is known as the intensity of the wave and for a plane wave the average intensity is
given by I = q 2 /(ρc) , where q is the root mean square acoustic pressure.
At 1 kHz, the wavelength in water is about 1.5 m. Sometimes the term “sound velocity” is used but this is incorrect as
the quantity is a scalar.
The large impedance contrast between air and water (the ratio is about 3600) and the scale of surface roughness means
that the sea surface behaves as an almost perfect reflector of sound at frequencies below 1 kHz. Sound speed in water
exceeds that in air by a factor of 4.4 and the density ratio is about 820.
4.2.3 Absorption of sound
Absorption of low frequency sound is weak.[6] (see Technical Guides - Calculation of absorption of sound in seawater
for an on-line calculator). The main cause of sound attenuation in fresh water, and at high frequency in sea water (above
100 kHz) is viscosity. Important additional contributions at lower frequency in seawater are associated with the ionic
relaxation of boric acid (up to c. 10 kHz)[6] and magnesium sulfate (c. 10 kHz-100 kHz).[7]
Sound may be absorbed by losses at the fluid boundaries. Near the surface of the sea losses can occur in a bubble layer
or in ice, while at the bottom sound can penetrate into the sediment and be absorbed.
4.2.4 Sound Reflection and Scattering
Boundary interactions
Both the water surface and bottom are reflecting and scattering boundaries.
Surface For many purposes the sea-air surface can be thought of as a perfect reflector. The impedance contrast is
so great that little energy is able to cross this boundary. Acoustic pressure waves reflected from the sea surface expe-
rience a reversal in phase, often stated as either a “pi phase change” or a “180 deg phase change”. This is represented
mathematically by assigning a reflection coefficient of minus 1 instead of plus one to the sea surface.
At high frequency (above about 1 kHz) or when the sea is rough, some of the incident sound is scattered, and this is
taken into account by assigning a reflection coefficient whose magnitude is less than one. For example, close to normal
incidence, the reflection coefficient becomes R = −e−2k h sin A , where h is the rms wave height.[8]
2 2 2
A further complication is the presence of wind generated bubbles or fish close to the sea surface.[9] The bubbles can also
form plumes that absorb some of the incident and scattered sound, and scatter some of the sound themselves.[10]
Seabed The acoustic impedance mismatch between water and the bottom is generally much less than at the surface and
is more complex. It depends on the bottom material types and depth of the layers. Theories have been developed for
predicting the sound propagation in the bottom in this case, for example by Biot [11] and by Buckingham.[12]
At Target
The reflection of sound at a target whose dimensions are large compared with the acoustic wavelength depends on its size
and shape as well as the impedance of the target relative to that of water. Formulae have been developed for the target
strength of various simple shapes as a function of angle of sound incidence. More complex shapes may be approximated
by combining these simple ones.[1]
4.2.5 Propagation of sound
Underwater acoustic propagation depends on many factors. The direction of sound propagation is determined by the sound
speed gradients in the water.This is a imortant thing that happen in water, becouse the speed of sound travel in water with
velocity regular. In the sea the vertical gradients are generally much larger than the horizontal ones. Combining this with
a tendency towards increasing sound speed at increasing depth, due to the increasing pressure in the deep sea, causes a
reversal of the sound speed gradient in the thermocline, creating an efficient waveguide at the depth, corresponding to the
minimum sound speed. The sound speed profile may cause regions of low sound intensity called “Shadow Zones,” and
regions of high intensity called “Caustics”. These may be found by ray tracing methods.
At equator and temperate latitudes in the ocean, the surface temperature is high enough to reverse the pressure effect,
such that a sound speed minimum occurs at depth of a few hundred metres. The presence of this minimum creates a
special channel known as Deep Sound Channel, previously known as the SOFAR (sound fixing and ranging) channel,
permitting guided propagation of underwater sound for thousands of kilometres without interaction with the sea surface
or the seabed. Another phenomenon in the deep sea is the formation of sound focusing areas, known as Convergence
Zones. In this case sound is refracted downward from a near-surface source and then back up again. The horizontal
distance from the source at which this occurs depends on the positive and negative sound speed gradients. A surface
duct can also occur in both deep and moderately shallow water when there is upward refraction, for example due to cold
surface temperatures. Propagation is by repeated sound bounces off the surface.
In general, as sound propagates underwater there is a reduction in the sound intensity over increasing ranges, though in
some circumstances a gain can be obtained due to focusing. Propagation loss (sometimes referred to as transmission loss)
is a quantitative measure of the reduction in sound intensity between two points, normally the sound source and a distant
receiver. If Is is the far field intensity of the source referred to a point 1 m from its acoustic centre and Ir is the intensity
at the receiver, then the propagation loss is given by[1] P L = 10log(Is /Ir ) . In this equation Ir is not the true acoustic
intensity at the receiver, which is a vector quantity, but a scalar equal to the equivalent plane wave intensity (EPWI) of
the sound field. The EPWI is defined as the magnitude of the intensity of a plane wave of the same RMS pressure as the
true acoustic field. At short range the propagation loss is dominated by spreading while at long range it is dominated by
absorption and/or scattering losses.
An alternative definition is possible in terms of pressure instead of intensity,[13] giving P L = 20log(ps /pr ) , where ps
is the RMS acoustic pressure in the far-field of the projector, scaled to a standard distance of 1 m, and pr is the RMS
pressure at the receiver position.
These two definitions are not exactly equivalent because the characteristic impedance at the receiver may be different from
that at the source. Because of this, the use of the intensity definition leads to a different sonar equation to the definition
based on a pressure ratio.[14] If the source and receiver are both in water, the difference is small.
Propagation modelling
The propagation of sound through water is described by the wave equation, with appropriate boundary conditions. A
number of models have been developed to simplify propagation calculations. These models include ray theory, normal
mode solutions, and parabolic equation simplifications of the wave equation.[15] Each set of solutions is generally valid and
computationally efficient in a limited frequency and range regime, and may involve other limits as well. Ray theory is more
appropriate at short range and high frequency, while the other solutions function better at long range and low frequency.[16]
Various empirical and analytical formulae have also been derived from measurements that are useful approximations.[17]
Reverberation
Transient sounds result in a decaying background that can be of much larger duration than the original transient signal.
The cause of this background, known as reverberation, is partly due to scattering from rough boundaries and partly due
to scattering from fish and other biota. For an acoustic signal to be detected easily, it must exceed the reverberation level
as well as the background noise level.
4.3. MEASUREMENTS 49
Doppler Shift
If an underwater object is moving relative to an underwater receiver, the frequency of the received sound is different
from that of the sound radiated (or reflected) by the object. This change in frequency is known as a Doppler shift. The
shift can be easily observed in active sonar systems, particularly narrow-band ones, because the transmitter frequency is
known, and the relative motion between sonar and object can be calculated. Sometimes the frequency of the radiated
noise (a tonal) may also be known, in which case the same calculation can be done for passive sonar. For active systems
the change in frequency is 0.69 Hz per knot per kHz and half this for passive systems as propagation is only one way. The
shift corresponds to an increase in frequency for an approaching target.
Intensity Fluctuations
Though acoustic propagation modelling generally predicts a constant received sound level, in practice there are both
temporal and spatial fluctuations. These may be due to both small and large scale environmental phenomena. These can
include sound speed profile fine structure and frontal zones as well as internal waves. Because in general there are multiple
propagation paths between a source and receiver, small phase changes in the interference pattern between these paths can
lead to large fluctuations in sound intensity.
Non-linearity
In water, especially with air bubbles, the change in density due to a change in pressure is not exactly linearly proportional.
As a consequence for a sinusoidal wave input additional harmonic and subharmonic frequencies are generated. When two
sinusoidal waves are input, sum and difference frequencies are generated. The conversion process is greater at high source
levels than small ones. Because of the non-linearity there is a dependence of sound speed on the pressure amplitude so
that large changes travel faster than small ones. Thus a sinusoidal waveform gradually becomes a sawtooth one with a
steep rise and a gradual tail. Use is made of this phenomenon in parametric sonar and theories have been developed to
account for this, e.g. by Westerfield.
4.3 Measurements
Sound in water is measured using a hydrophone, which is the underwater equivalent of a microphone. A hydrophone
measures pressure fluctuations, and these are usually converted to sound pressure level (SPL), which is a logarithmic
measure of the mean square acoustic pressure.
Measurements are usually reported in one of three forms :-
• RMS acoustic pressure in micropascals (or dB re 1 μPa)
• RMS acoustic pressure in a specified bandwidth, usually octaves or thirds of octave (dB re 1 μPa)
• spectral density (mean square pressure per unit bandwidth) in micropascals-squared per hertz (dB re 1 μPa²/Hz)
The scale for acoustic pressure in water differs from that used for sound in air. In air the reference pressure is 20 μPa
rather than 1 μPa. For the same numerical value of SPL, the intensity of a plane wave (power per unit area, proportional
to mean square sound pressure divided by acoustic impedance) in air is about 202 ×3600 = 1 440 000 times higher than
in water. Similarly, the intensity is about the same if the SPL is 61.6 dB higher in the water.
4.3.1 Sound speed

Approximate values for fresh water and seawater, respectively, at atmospheric pressure are 1450 and 1500 m/s for the
sound speed, and 1000 and 1030 kg/m³ for the density.[18] The speed of sound in water increases with increasing pressure,
temperature and salinity.[19][20] The maximum speed in pure water under atmospheric pressure is attained at about 74 °C;
sound travels slower in hotter water after that point; the maximum increases with pressure.[21] On-line calculators can be
found at Technical Guides - Speed of Sound in Sea-Water and Technical Guides - Speed of Sound in Pure Water.
4.3.2 Absorption
Many measurements have been made of sound absorption in lakes and the ocean [6][7] (see Technical Guides - Calculation
of absorption of sound in seawater for an on-line calculator).
4.3.3 Ambient noise
Measurement of acoustic signals are possible if their amplitude exceeds a minimum threshold, determined partly by the
signal processing used and partly by the level of background noise. Ambient noise is that part of the received noise that
is independent of the source, receiver and platform characteristics. This it excludes reverberation and towing noise for
example.
The background noise present in the ocean, or ambient noise, has many different sources and varies with location and
frequency.[22] At the lowest frequencies, from about 0.1 Hz to 10 Hz, ocean turbulence and microseisms are the primary
contributors to the noise background.[23] Typical noise spectrum levels decrease with increasing frequency from about
140 dB re 1 μPa²/Hz at 1 Hz to about 30 dB re 1 μPa²/Hz at 100 kHz. Distant ship traffic is one of the dominant noise
sources in most areas for frequencies of around 100 Hz, while wind-induced surface noise is the main source between 1
kHz and 30 kHz. At very high frequencies, above 100 kHz, thermal noise of water molecules begins to dominate. The
thermal noise spectral level at 100 kHz is 25 dB re 1 μPa²/Hz. The spectral density of thermal noise increases by 20 dB
per decade (approximately 6 dB per octave).[24]
Transient sound sources also contribute to ambient noise. These can include intermittent geological activity, such as
earthquakes and underwater volcanoes,[25] rainfall on the surface, and biological activity. Biological sources include
cetaceans (especially blue, fin and sperm whales),[26][27] certain types of fish, and snapping shrimp.
Rain can produce high levels of ambient noise. However the numerical relationship between rain rate and ambient noise
level is difficult to determine because measurement of rain rate is problematic at sea.
4.3.4 Reverberation
Many measurements have been made of sea surface, bottom and volume reverberation. Empirical models have sometimes
been derived from these. A commonly used expression for the band 0.4 to 6.4 kHz is that by Chapman and Harris.[28] It is
found that a sinusoidal waveform is spread in frequency due to the surface motion. For bottom reverberation a Lambert’s
Law is found often to apply approximately, for example see Mackenzie.[29] Volume reverberation is usually found to occur
mainly in layers, which change depth with the time of day, e.g., see Marshall and Chapman.[30] The under-surface of ice
can produce strong reverberation when it is rough, see for example Milne.[31]
4.3.5 Bottom Loss
Bottom loss has been measured as a function of grazing angle for many frequencies in various locations, for example
those by the US Marine Geophysical Survey.[32] The loss depends on the sound speed in the bottom (which is affected
by gradients and layering) and by roughness. Graphs have been produced for the loss to be expected in particular cir-
cumstances. In shallow water bottom loss often has the dominant impact on long range propagation. At low frequencies
sound can propagate through the sediment then back into the water.
4.4 Underwater hearing

4.5. APPLICATIONS OF UNDERWATER ACOUSTICS 51
4.4.1 Comparison with airborne sound levels

As with airborne sound, sound pressure level underwater is usually reported in units of decibels, but there are some
important differences that make it difficult (and often inappropriate) to compare SPL in water with SPL in air. These
differences include:[33]
• difference in reference pressure: 1 μPa (one micropascal, or one millionth of a pascal) instead of 20 μPa.[13]
• difference in interpretation: there are two schools of thought, one maintaining that pressures should be compared
directly, and the other that one should first convert to the intensity of an equivalent plane wave.
• difference in hearing sensitivity: any comparison with (A-weighted) sound in air needs to take into account the
differences in hearing sensitivity, either of a human diver or other animal.[34]
4.4.2 Hearing sensitivity

The lowest audible SPL for a human diver with normal hearing is about 67 dB re 1 μPa, with greatest sensitivity occurring
at frequencies around 1 kHz.[35] This corresponds to a sound intensity 5.4 dB, or 3.5 times, higher than the threshold
in air (see #Measurements above). Dolphins and other toothed whales are renowned for their acute hearing sensitivity,
especially in the frequency range 5 to 50 kHz.[34][36] Several species have hearing thresholds between 30 and 50 dB re 1
μPa in this frequency range. For example, the hearing threshold of the killer whale occurs at an RMS acoustic pressure
of 0.02 mPa (and frequency 15 kHz), corresponding to an SPL threshold of 26 dB re 1 μPa.[37] By comparison the most
sensitive fish is the soldier fish, whose threshold is 0.32 mPa (50 dB re 1 μPa) at 1.3 kHz, whereas the lobster has a hearing
threshold of 1.3 Pa at 70 Hz (122 dB re 1 μPa).[37]
4.4.3 Safety thresholds

High levels of underwater sound create a potential hazard to marine and amphibious animals as well as to human
divers.[34][38] Guidelines for exposure of human divers and marine mammals to underwater sound are reported by the
SOLMAR project of the NATO Undersea Research Centre.[39] Human divers exposed to SPL above 154 dB re 1 μPa in
the frequency range 0.6 to 2.5 kHz are reported to experience changes in their heart rate or breathing frequency. Diver
aversion to low frequency sound is dependent upon sound pressure level and center frequency.[40]
4.5 Applications of underwater acoustics
4.5.1 Sonar
Main article: Sonar
Sonar is the name given to the acoustic equivalent of radar. Pulses of sound are used to probe the sea, and the echoes are
then processed to extract information about the sea, its boundaries and submerged objects. An alternative use, known as
passive sonar, attempts to do the same by listening to the sounds radiated by underwater objects.
4.5.2 Underwater communication

Main article: Underwater acoustic communication
The need for underwater acoustic telemetry exists in applications such as data harvesting for environmental monitoring,
communication with and between manned and unmanned underwater vehicles, transmission of diver speech, etc. A related
application is underwater remote control, in which acoustic telemetry is used to remotely actuate a switch or trigger an
event. A prominent example of underwater remote control are acoustic releases, devices that are used to return sea floor
deployed instrument packages or other payloads to the surface per remote command at the end of a deployment. Acoustic
communications form an active field of research [41][42] with significant challenges to overcome, especially in horizontal,
shallow-water channels. Compared with radio telecommunications, the available bandwidth is reduced by several orders
of magnitude. Moreover, the low speed of sound causes multipath propagation to stretch over time delay intervals of tens
or hundreds of milliseconds, as well as significant Doppler shifts and spreading. Often acoustic communication systems
are not limited by noise, but by reverberation and time variability beyond the capability of receiver algorithms. The fidelity
of underwater communication links can be greatly improved by the use of hydrophone arrays, which allow processing
techniques such as adaptive beamforming and diversity combining.
4.5.3 Underwater Navigation and Tracking
Main article: Underwater Acoustic Positioning System
Underwater navigation and tracking is a common requirement for exploration and work by divers, ROV, autonomous
underwater vehicles (AUV), manned submersibles and submarines alike. Unlike most radio signals which are quickly
absorbed, sound propagates far underwater and at a rate that can be precisely measured or estimated.[43] It can thus
be used to measure distances between a tracked target and one or multiple reference of baseline stations precisely, and
triangulate the position of the target, sometimes with centimeter accuracy. Starting in the 1960s, this has given rise to
underwater acoustic positioning systems which are now widely used.
4.5.4 Seismic exploration
Main article: Reflection seismology
Seismic exploration involves the use of low frequency sound (< 100 Hz) to probe deep into the seabed. Despite the
relatively poor resolution due to their long wavelength, low frequency sounds are preferred because high frequencies are
heavily attenuated when they travel through the seabed. Sound sources used include airguns, vibroseis and explosives.
4.5.5 Weather and climate observation
Acoustic sensors can be used to monitor the sound made by wind and precipitation. For example, an acoustic rain gauge
is described by Nystuen.[44] Lightning strikes can also be detected.[45] Acoustic thermometry of ocean climate (ATOC)
uses low frequency sound to measure the global ocean temperature.
4.5.6 Oceanography
Main article: Acoustical oceanography
Large scale ocean features can be detected by acoustic tomography. Bottom characteristics can be measured by side-scan
sonar and sub-bottom profiling.
4.5.7 Marine biology
Main article: Bioacoustics
Due to its excellent propagation properties, underwater sound is used as a tool to aid the study of marine life, from
microplankton to the blue whale. Echo sounders are often used to provide data on marine life abundance, distribution,
4.6. SEE ALSO 53
and behavior information. Echo sounders, also referred to as hydroacoustics is also used for fish location, quantity, size,
and biomass.
Acoustic telemetry is also used for monitoring fishes and marine wildlife. An acoustic transmitter is attached to the fish
(sometimes internally) while an array of receivers listen to the information conveyed by the sound wave. This enables the
researchers to track the movements of individuals in a small-medium scale.[46]
Pistol shrimp create sonoluminescent cavitation bubbles that reach up to 5,000 K (4,700 °C) [47]
4.5.8 Particle physics

A neutrino is a fundamental particle that interacts very weakly with other matter. For this reason, it requires detection
apparatus on a very large scale, and the ocean is sometimes used for this purpose. In particular, it is thought that ultra-high
energy neutrinos in seawater can be detected acoustically.[48]
4.6 See also

• Acoustic Tags (Acoustic Telemetry)
• Bioacoustics
• Hydroacoustics
• Ocean Tracking Network
• Refraction (sound)
• Sonar
• Underwater Acoustic Positioning System
• SOFAR channel
• Underwater acoustic wireless communication system
4.7 References
[1] Urick, Robert J. Principles of Underwater Sound, 3rd Edition. New York. McGraw-Hill, 1983.
[2] C. S. Clay & H. Medwin, Acoustical Oceanography (Wiley, New York, 1977)
[3] Annales de Chimie et de Physique 36 [2] 236 (1827)
[4] A. B. Wood, From the Board of Invention and Research to the Royal Naval Scientific Service, Journal of the Royal Naval Scientific
Service Vol 20, No 4, pp 1-100 (185-284).
[5] H. Lichte (1919). “On the influence of horizontal temperature layers in sea water on the range of underwater sound signals”.
Physik. Z. 17 (385).
[6] R. E. Francois & G. R. Garrison, Sound absorption based on ocean measurements. Part II: Boric acid contribution and equation
for total absorption, J. Acoust. Soc. Am. 72, 1879-1890 (1982).
[7] R. E. Francois and G. R. Garrison, Sound absorption based on ocean measurements. Part I: Pure water and magnesium sulfate
contributions, J. Acoust. Soc. Am. 72, 896-907 (1982).
[8] H. Medwin & C. S. Clay, Fundamentals of Acoustical Oceanography (Academic, Boston, 1998).
[9] D. E. Weston & P. A. Ching, Wind effects in shallow-water transmission, J. Acoust. Soc. Am. 86, 1530-1545 (1989).
[10] G. V. Norton & J. C. Novarini, On the relative role of sea-surface roughness and bubble plumes in shallow-water propagation
in the low-kilohertz region, J. Acoust. Soc. Am. 110, 2946-2955 (2001)
[11] N Chotiros, Biot Model of Sound Propagation in Water Saturated Sand. J. Acoust. Soc. Am. 97, 199 (1995)
[12] M. J. Buckingham, Wave propagation, stress relaxation, and grain-to-grain shearing in saturated, unconsolidated marine sedi-
ments, J. Acoust. Soc. Am. 108, 2796-2815 (2000).
[13] C. L. Morfey, Dictionary of Acoustics (Academic Press, San Diego, 2001).
[14] M. A. Ainslie, The sonar equation and the definitions of propagation loss, J. Acoust. Soc. Am. 115, 131-134 (2004).
[15] F. B. Jensen, W. A. Kuperman, M. B. Porter & H. Schmidt, Computational Ocean Acoustics (AIP Press, NY, 1994).
[16] C. H. Harrison, Ocean propagation models, Applied Acoustics 27, 163-201 (1989).
[17] L. M. Brekhovskikh & Yu. P. Lysanov, Fundamentals of Ocean Acoustics, 3rd edition (Springer-Verlag, NY, 2003).
[18] A. D. Pierce, Acoustics: An Introduction to its Physical Principles and Applications (American Institute of Physics, New York,
1989).
[19] Mackenzie, Nine-term equation for sound speed in the oceans, J. Acoust. Soc. Am. 70, 807-812 (1982).
[20] C. C. Leroy, The speed of sound in pure and neptunian water, in Handbook of Elastic Properties of Solids, Liquids and Gases,
edited by Levy, Bass & Stern, Volume IV: Elastic Properties of Fluids: Liquids and Gases (Academic Press, 2001)
[21] Wilson, Wayne D. (26 Jan 1959). “Speed of Sound in Distilled Water as a Function of Temperature and Pressure”. J. Acoust.
Soc. Am. 31 (8): 1067–1072. Bibcode:1959ASAJ...31.1067W. doi:10.1121/1.1907828. Retrieved 11 February 2012.
[22] G. M. Wenz, Acoustic ambient noise in the ocean: spectra and sources, J. Acoust. Soc. Am. 34, 1936-1956 (1962).
[23] S. C. Webb, The equilibrium oceanic microseism spectrum, J. Acoust. Soc. Am. 92, 2141-2158 (1992).
[24] R. H. Mellen, The Thermal-Noise Limit in the Detection of Underwater Acoustic Signals, J. Acoust. Soc. Am. 24, 478-480
(1952).
[25] R. S. Dietz and M. J. Sheehy, Transpacific detection of myojin volcanic explosions by underwater sound. Bulletin of the
Geological Society 2 942-956 (1954).
[26] M. A. McDonald, J. A. Hildebrand & S. M. Wiggins, Increases in deep ocean ambient noise in the Northeast Pacific west of
San Nicolas Island, California, J. Acoust. Soc. Am. 120, 711-718 (2006).
[27] Ocean Noise and Marine Mammals, National Research Council of the National Academies (The National Academies Press,
Washington DC, 2003).
[28] R Chapman and J Harris, Surface backscattering Strengths Measured with Explosive Sound Sources. J. Acoust. Soc. Am. 34,
547 (1962)
[29] K Mackenzie, Bottom Reverberation for 530 and 1030 cps Sound in Deep Water. J. Acoust. Soc. Am. 36, 1596 (1964)
[30] J. R. Marshall and R. P. Chapman, Reverberation from a Deep Scattering Layer Measured with Explosive Sound Sources. J.
Acoust. Soc. Am. 36, 164 (1964)
[31] A. Milne, Underwater Backscattering Strengths of Arctic Pack Ice. J. Acoust. Soc. Am. 36, 1551 (1964)
[32] MGS Station Data Listing and Report Catalog, Nav Oceanog Office Special Publication 142, 1974
[33] D.M.F. Chapman, D.D. Ellis, The elusive decibel - thoughts on sonars and marine mammals, Can. Acoust. 26(2), 29–31 (1996)
[34] W. J. Richardson, C. R. Greene, C. I. Malme and D. H. Thomson, Marine Mammals and Noise (Academic Press, San Diego,
1995).
[35] S. J. Parvin, E. A. Cudahy & D. M. Fothergill, Guidance for diver exposure to underwater sound in the frequency range 500 to
2500 Hz, Underwater Defence Technology (2002).
[36] W. W. L. Au, The Sonar of Dolphins (Springer, NY, 1993).

[37] D. Simmonds & J. MacLennan, Fisheries Acoustics: Theory and Practice, 2nd edition (Blackwell, Oxford, 2005)
[38] Steevens CC, Russell KL, Knafelc ME, Smith PF, Hopkins EW, Clark JB (1999). “Noise-induced neurologic disturbances in
divers exposed to intense water-borne sound: two case reports”. Undersea Hyperb Med. 26 (4): 261–5. PMID 10642074.
Retrieved 2009-03-31.
[39] NATO Undersea Research Centre Human Diver and Marine Mammal Risk Mitigation Rules and Procedures, NURC Special
Publication NURC-SP-2006-008, September 2006
[40] Fothergill DM, Sims JR, Curley MD (2001). “Recreational scuba divers’ aversion to low-frequency underwater sound”. Under-
sea Hyperb Med. 28 (1): 9–18. PMID 11732884. Retrieved 2009-03-31.
[41] D. B. Kilfoyle and A. B. Baggeroer, “The state of the art in underwater acoustic telemetry,” IEEE J. Oceanic Eng. 25, 4-27
(2000).
[42] M.Stojanovic, “Acoustic (Underwater) Communications,” entry in Encyclopedia of Telecommunications, John G. Proakis, Ed.,
John Wiley & Sons, 2003.
[43] Underwater Acoustic Positioning Systems, P.H. Milne 1983, ISBN 0-87201-012-0
[44] J. A. Nystuen, Listening to raindrops from underwater: An acoustic disdrometer, J Atmospheric and Oceanic Technology,
18(10), 1640-1657 (2001).
[45] R. D. Hill, Investigation of lightning strikes to water surfaces, J. Acoust. Soc. Am. 78, 2096-2099 (1985).
[46] Moore, A., T. Storeton-West, I. C. Russell, E. C. E. Potter, and M. J. Challiss. 1990. A technique for tracking Atlantic salmon
(Salmo salar L.) smolts through estuaries. International Council for the Ex- ploration of the Sea, C.M. 1990/M: 18, Copenhagen.
[47] D. Lohse, B. Schmitz & M. Versluis (2001). “Snapping shrimp make flashing bubbles”. Nature. 413 (6855): 477–478.
Bibcode:2001Natur.413..477L. doi:10.1038/35097152. PMID 11586346.
[48] S. Bevan, S. Danaher, J. Perkin, S. Ralph, C. Rhodes, L. Thompson, T. Sloane, D. Waters and The ACoRNE Collaboration,
Simulation of ultra high energy neutrino induced showers in ice and water, Astroparticle Physics Volume 28, Issue 3, November
2007, Pages 366-379
4.8 External links

• Ocean Acoustics Library
• Ultrasonics and Underwater Acoustics
• Monitoring the global ocean through underwater acoustics

• Underwater Acoustics Research
• ASA Underwater Acoustics Technical Committee

• An Ocean of Sound
• Underwater Acoustic Communications

• Acoustic Communications Group at the Woods Hole Oceanographic Institution
• Sound in the Sea

• SFSU Underwater Acoustics Research Group
• Discovery of Sound in the Sea

• PAMBuoy Passive Acoustic Monitoring
Chapter 5
Bioacoustics
Bioacoustics is a cross-disciplinary science that combines biology and acoustics. Usually it refers to the investigation
of sound production, dispersion and reception in animals (including humans).[1] This involves neurophysiological and
anatomical basis of sound production and detection, and relation of acoustic signals to the medium they disperse through.
The findings provide clues about the evolution of acoustic mechanisms, and from that, the evolution of animals that employ
them.
In underwater acoustics and fisheries acoustics the term is also used to mean the effect of plants and animals on sound
propagated underwater, usually in reference to the use of sonar technology for biomass estimation.[2][3]
5.1 History
For a long time humans have employed animal sounds to recognise and find them. Bioacoustics as a scientific discipline
was established by the Slovene biologist Ivan Regen who began systematically to study insect sounds. In 1925 he used a
special stridulatory device to play in a duet with an insect. Later he put a male cricket behind a microphone and female
crickets in front of a loudspeaker. The females were not moving towards the male but towards the loudspeaker.[4] Regen’s
most important contribution to the field apart from realization that insects also detect airborne sounds was the discovery
of tympanal organ's function.[5]
Relatively crude electro-mechanical devices available at the time (such as phonographs) allowed only for crude appraisal
of signal properties. More accurate measurements were made possible in the second half of the 20th century by advances
in electronics and utilization of devices such as oscilloscopes and digital recorders.
The most recent advances in bioacoustics concern the relationships among the animals and their acoustic environment
and the impact of anthropogenic noise. Bioacoustic techniques have recently been proposed as a non-invasive method for
estimating biodiversity.[6]
5.2 Methods in bioacoustics

Listening is still one of the main methods used in bioacoustical research. Little is known about neurophysiological pro-
cesses that play a role in production, detection and interpretation of sounds in animals, so animal behaviour and the signals
themselves are used for gaining insight into these processes.
5.2.1 Acoustic signals

An experienced observer can use animal sounds to recognize a “singing” animal species, its location and condition in
nature. Investigation of animal sounds also includes signal recording with electronic recording equipment. Due to the
56
5.2. METHODS IN BIOACOUSTICS 57
The sonograms of Thrush nightingale (Luscinia luscinia) and Common nightingale (Luscinia megarhynchos) singing help to distinguish
these two species by voice definitely.
wide range of signal properties and media they propagate through, specialized equipment may be required instead of the
usual microphone, such as a hydrophone (for underwater sounds), detectors of ultrasound (very high-frequency sounds) or
infrasound (very low-frequency sounds), or a laser vibrometer (substrate-borne vibrational signals). Computers are used
for storing and analysis of recorded sounds. Specialized sound-editing software is used for describing and sorting signals
according to their intensity, frequency, duration and other parameters.
Animal sound collections, managed by museums of natural history and other institutions, are an important tool for sys-
tematic investigation of signals. Many effective automated methods involving signal processing, data mining and machine
learning techniques have been developed to detect and classify the bioacoustic signals.[7]
58 CHAPTER 5. BIOACOUSTICS
5.2.2 Sound production, detection, and use in animals

Scientists in the field of bioacoustics are interested in anatomy and neurophysiology of organs involved in sound production
and detection, including their shape, muscle action, and activity of neuronal networks involved. Of special interest is
coding of signals with action potentials in the latter.
But since the methods used for neurophysiological research are still fairly complex and understanding of relevant processes
is incomplete, more trivial methods are also used. Especially useful is observation of behavioural responses to acoustic
signals. One such response is phonotaxis – directional movement towards the signal source. By observing response to well
defined signals in a controlled environment, we can gain insight into signal function, sensitivity of the hearing apparatus,
noise filtering capability, etc.
5.2.3 Biomass estimation

Main article: Hydroacoustics
Biomass estimation is a method of detecting and quantifying fish and other marine organisms using sonar technology.[3]
As the sound pulse travels through water it encounters objects that are of different density than the surrounding medium,
such as fish, that reflect sound back toward the sound source. These echoes provide information on fish size, location,
and abundance. The basic components of the scientific echo sounder hardware function is to transmit the sound, receive,
filter and amplify, record, and analyze the echoes. While there are many manufacturers of commercially available “fish-
finders,” quantitative analysis requires that measurements be made with calibrated echo sounder equipment, having high
signal-to-noise ratios.
5.3 Animal sounds

Sounds used by animals that fall within the scope of bioacoustics include a wide range of frequencies and media, and are
often not "sound" in the narrow sense of the word (i.e. compression waves that propagate through air and are detectable
by the human ear). Katydid crickets, for example, communicate by sounds with frequencies higher than 100 kHz, far into
the ultrasound range.[8] Lower, but still in ultrasound, are sounds used by bats for echolocation. On the other side of the
frequency spectrum are low frequency-vibrations, often not detected by hearing organs, but with other, less specialized
sense organs. The examples include ground vibrations produced by elephants whose principal frequency component is
around 15 Hz, and low- to medium-frequency substrate-borne vibrations used by most insect orders.[9] Many animal
sounds, however, do fall within the frequency range detectable by a human ear, between 20 and 20,000 Hz. Mechanisms
for sound production and detection are just as diverse as the signals themselves.
5.4 See also

• Acoustic ecology
• Acoustical oceanography
• Animal communication
• Animal language
• Biomusic
• Diffusion (acoustics)
• Field recording
• Frog hearing and communication

5.5. REFERENCES 59
• List of animal sounds

• List of Bioacoustics Software
• Music therapy
• Natural sounds
• Underwater acoustics
• Vocal learning
• Whale sound
• Zoomusicology
5.5 References
[1] “Bioacoustics - the International Journal of Animal Sound and its Recording”. Taylor & Francis. Retrieved 31 July 2012.
[2] Medwin H. & Clay C.S. (1998). Fundamentals of Acoustical Oceanography, Academic Press
[3] Simmonds J. & MacLennan D. (2005). Fisheries Acoustics: Theory and Practice, second edition. Blackwell
[4] Kočar T. (2004). Kot listja in kobilic (As many as leaves and grasshoppers). GEA, october 2004. Mladinska knjiga, Ljubljana
(Slovene)
[5] Glen Wever, Ernest (2008). “Sound reception: Evidence of hearing and communication in insects”. Britannica online. Retrieved
2008-09-25.
[6] Sueur J.; Pavoine S.; Hamerlynck O.; Duvail S. (December 30, 2008). Reby, David, ed. “Rapid Acoustic Survey for Biodiversity
Appraisal”. PLoS ONE. 3 (12): e4065. Bibcode:2008PLoSO...3.4065S. doi:10.1371/journal.pone.0004065. PMC 2605254 .
PMID 19115006. Retrieved 2009-01-19.
[7] M. Pourhomayoun, P. Dugan, M. Popescu, and C. Clark, “Bioacoustic Signal Classification Based on Continuous Region Fea-
tures, Grid Masking Features and Artificial Neural Network,” International Conference on Machine Learning (ICML), 2013.
[8] Mason A.C., Morris G.K., Wall P. (1991): High Ultrasonic Hearing and Tympanal Slit Function in Rainforest Katydids. Natur-
wissenschaften 78: 365-367.
[9] Virant-Doberlet M. & Čokl A. (2004): Vibrational communication in insects. Neotropical Entomology 33(2): 121-134
5.6 External links

• BioAcoustica: Wildlife Sounds Database
• The British Library Sound Archive has 150,000 recordings of over 10,000 species.
• International Bioacoustics Council links to many bioacoustics resources.
• Borror Laboratory of Bioacoustics at The Ohio State University has a large archive of animal sound recordings.
• Listen to Nature 400 examples of animal songs and calls
• Wildlife Sound Recording Society
• Bioacoustic Research Program at the Cornell Lab of Ornithology distributes a number of different free bioacoustics
synthesis & analysis programs.
• Macaulay Library at the Cornell Lab of Ornithology is the world’s largest collection of animal sounds and associated
video.
5.7 Further reading

• Ewing A.W. (1989): Arthropod bioacoustics: Neurobiology and behaviour. Edinburgh: Edinburgh University Press.
ISBN 0-7486-0148-1
• Fletcher N. (2007): Animal Bioacoustics. IN: Rossing T.D. (ed.): Springer Handbook of Acoustics, Springer. ISBN
978-0-387-33633-6
5.7. FURTHER READING 61
Spectrogram (above) and oscillogram (below) of the humpback whale's calls

5.7. FURTHER READING 63
European starling singing

Chapter 6
Hydrophone
Not to be confused with hydraulophone, a musical instrument.
A hydrophone (Ancient Greek ὕδωρ = water[1] and φωνή = sound[1] ) is a microphone designed to be used underwater
for recording or listening to underwater sound. Most hydrophones are based on a piezoelectric transducer that generates
electricity when subjected to a pressure change. Such piezoelectric materials, or transducers, can convert a sound signal
into an electrical signal since sound is a pressure wave. Some transducers can also serve as a projector, but not all have
this capability, and some may be destroyed if used in such a manner.
A hydrophone can “listen” to sound in air but will be less sensitive due to its design as having a good acoustic impedance
match to water, which is a denser fluid than air. Likewise, a microphone can be buried in the ground, or immersed in
water if it is put in a waterproof container, but will give similarly poor performance due to the similarly bad acoustic
impedance match.
6.1 History
The earliest widely used design was the Fessenden oscillator, an electrodynamically driven clamped-edge circular plate
transducer (not actually an oscillator) operating at 500, 1000, and later 3000 Hz. It was originally marketed as an un-
derwater telegraph, rather than as sonar, but was later very successful, its Canadian inventor, Reginald Fessenden, was
awarded the “Scientific American Magazine Gold Medal of Safety” in 1929 from the American Museum of Safety, an
organization for ship captains;[2] some were still in use during World War II.
Ernest Rutherford, in England, led pioneer research in hydrophones using piezoelectric devices, and his only patent was for
a hydrophone device. The acoustic impedance of piezoelectric materials facilitated their use as underwater transducers.
The piezoelectric hydrophone was used late in World War I, by convoy escorts detecting U-boats, greatly impacting the
effectiveness of submarines.
From late in World War I until the introduction of active sonar, hydrophones were the sole method for submarines to
detect targets while submerged, and remain useful today.
6.2 Directional hydrophones
A small single cylindrical ceramic transducer can achieve near perfect omnidirectional reception. Directional hydrophones
increase sensitivity from one direction using two basic techniques:
64
6.3. SEE ALSO 65
6.2.1 Focused transducers
This device uses a single transducer element with a dish or conical-shaped sound reflector to focus the signals, in a similar
manner to a reflecting telescope. This type of hydrophone can be produced from a low-cost omnidirectional type, but
must be used while stationary, as the reflector impedes its movement through water. A new way to direct is to use a
spherical body around the hydrophone. The advantage of directivity spheres is that the hydrophone can be moved within
the water, ridding it of the interferences produced by a conical-shaped element
6.2.2 Arrays
Multiple hydrophones can be arranged in an array so that it will add the signals from the desired direction while subtracting
signals from other directions. The array may be steered using a beamformer. Most commonly, hydrophones are arranged
in a “line array” but may be in two- or three-dimensional arrangements.
SOSUS hydrophones, laid on the seabed and connected by underwater cables, were used, beginning in the 1950s, by the
U.S. Navy to track movement of Soviet submarines during the Cold War along a line from Greenland, Iceland and the
United Kingdom known as the GIUK gap.[3] These are capable of clearly recording extremely low frequency infrasound,
including many unexplained ocean sounds.
6.3 See also

• Communication with submarines
• Underwater acoustics
• Sonar
• Reflection seismology
6.4 Notes
[1] Liddell, H.G. & Scott, R. (1940). A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with
the assistance of. Roderick McKenzie. Oxford: Clarendon Press.
[2] Frost, Gary Lewis (July 2001). “Inventing Schemes and Strategies: The Making and Selling of the Fessenden Oscillator”.
Technology and Culture. Project MUSE. 42 (3): 462–488. doi:10.1353/tech.2001.0109. External link in |publisher= (help)
[3] Mackay, D.G. "Scotland the Brave? US Strategic Policy in Scotland 1953-1974". Glasgow University, Masters Thesis (research).
2008. Accessed 12 October 2009.
6.5 References
• Pike, John (1999). SOSUS. Retrieved January 28, 2005.
• Watlington, Frank (1979). How to build & use low-cost hydrophones. (ISBN 0830610790)
• Unknown. hydrophone. Retrieved January 28, 2005.
• Unknown. (2005) Schlumberger Oilfield Glossary: Term 'hydrophone'. Retrieved January 28, 2005.
• Onda Corporation (2015). 'Hydrophone Handbook'.

66 CHAPTER 6. HYDROPHONE
6.6 External links

• DOSITS—Hydrophone introduction at Discovery of Sound in the Sea
• orcasound.net—Live hydrophone streams from killer whale habitat

• Passive Acoustic Monitoring—Using hydrophones to monitor underwater sounds
• Build your own hydrophone—free instructions
• Precision Acoustics—useful resource on hydrophones

• The British Library Sound Archive—contains many wildlife and atmospheric recordings made using hydrophones.
• High Quality Hydrophones— High quality manufacturer of Hydrophones.

A hydrophone being lowered into the North Atlantic

6.7. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES 69
6.7 Text and image sources, contributors, and licenses
6.7.1 Text
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ner, Camembert, Sfdan, K.lee, Fred Bauder, Vagemulo~enwiki, Dan Koehl, Ixfd64, Pcb21, Ahoerstemeier, Cyp, DavidWBrooks, Stan Shebs,
Docu, Muriel Gottrop~enwiki, Angela, Glenn, Kils, Nikai, Big iron, Evercat, Samw, David Thrale, Daniel Quinlan, Peregrine981, Tpbrad-
bury, Nv8200pa, Indefatigable, Fvw, Robbot, Dale Arnett, Pigsonthewing, Yas~enwiki, Fredrik, Nurg, Seglea, Naddy, Smallweed, Babbage,
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son42, MPF, Ævar Arnfjörð Bjarmason, Lupin, Zigger, Everyking, Ceejayoz, Jason Quinn, Macrakis, Solipsist, Bobblewik, Neilc, James
Crippen, Andycjp, Toytoy, R. fiend, Slowking Man, Antandrus, OverlordQ, PDH, Vina, Yggdræsil, Daniel11, Starx, Neutrality, Ukexpat, Rak-
ista, WikiDon, Pinnerup, Trevor MacInnis, Freakofnurture, Eyrian, Jiy, Nep, RossPatterson, Discospinster, Supercoop, Vsmith, Gronky, Ben-
der235, Mykhal, ESkog, Dora dora, Kaisershatner, MisterSheik, Ben Webber, Mwanner, Cacophony, Bendono, Bobo192, Robotje, Goblim,
Giraffedata, MarkHab, Rockhopper10r, Kjkolb, Nk, Thewayforward, (aeropagitica), Orangemarlin, Danski14, Alansohn, Carbon Caryatid,
Muugokszhiion, Bart133, DreamGuy, Snowolf, Wtmitchell, Velella, Helixblue, Yuckfoo, Evil Monkey, Max Naylor, RainbowOfLight, TenO-
fAllTrades, Bsadowski1, Sfacets, Versageek, Gene Nygaard, Ghirlandajo, HunterAmor, Luigizanasi, Stemonitis, Richard Arthur Norton (1958-
), Pekinensis, Woohookitty, TigerShark, Scriberius, Thorpe, Benbest, Before My Ken, Riumplus, MONGO, Bennetto, GregorB, John Hill,
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4.4.2 Hearing sensitivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

6.7.3 Content license . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

For other uses, see Salmon (disambiguation).

Paciﬁc salmon leaping at Willamette Falls, Oregon

Commercial production of salmon in million tonnes 1950–

Atlantic salmon, Salmo salar

1.3 Life cycle

See also: Salmon run and Juvenile salmon

Life cycle of Paciﬁc salmon

Juvenile salmon, parr, grow up in the relatively protected natal river

Male ocean-phase adult sockeye

Male spawning-phase adult sockeye

Bear cub with salmon

1.4.1 Bears and salmon

1.4.2 Beavers and salmon

1.5 Wild ﬁsheries

1.6 Farmed salmon

making sashimi (raw ﬁsh) and sushi.

[5] Salmon etymonline.com, Online Etymology Dictionary. Retrieved 25 April 2012.

[9] “Salmo salar”. Integrated Taxonomic Information System.

[13] “Oncorhynchus tshawytscha”. Integrated Taxonomic Information System.

[16] “Oncorhynchus keta”. Integrated Taxonomic Information System.

[19] “Oncorhynchus kisutch”. Integrated Taxonomic Information System.

[21] “Oncorhynchus masou”. Integrated Taxonomic Information System.

[24] “Oncorhynchus gorbuscha”. Integrated Taxonomic Information System.

[27] “Oncorhynchus nerka”. Integrated Taxonomic Information System.

[30] “Arripis trutta”. Integrated Taxonomic Information System.

[32] “Hucho hucho”. Integrated Taxonomic Information System.

[36] “Elagatis bipinnulata”. Integrated Taxonomic Information System.

[38] “Eleutheronema tetradactylum”. Integrated Taxonomic Information System.

[45] “Formosan salmon”. Taiwan Journal. Retrieved 13 December 2006.

[72] Seafood Choices Alliance (2005) It’s all about salmon

[78] Salmon Recovery Planning. nwr.noaa.gov. p. 57.

1.12 Further reading

• Salmon, by Dr Peter Coates, 2006, ISBN 1-86189-295-0

1.13 External links

Angler and gillie land a salmon, Scotland

Artiﬁcially incubated chum salmon

Rainbow trout farm in an archipelago of Finland

Spawning sockeye salmon in Becharof Creek, Becharof Wilderness, Alaska

Seine ﬁshing for salmon – Wenzel Hollar, 1607–1677

Atlantic salmon are among the largest salmon species

2.3 Distribution and habitat

Ocean migration of Atlantic salmon from the Connecticut River[7]

2.6 Life stages

Life cycle of the Atlantic salmon

See also: Juvenile salmon

2.6.1 Freshwater phase

2.6.2 Saltwater phases

• Newly hatched alevin feed on their yolk sacs

See also: Salmon in aquaculture

Atlantic salmon marine cages in the Faroe Islands

2.10 Human impact

Seine ﬁshing for salmon – Wenzel Hollar, 1607–1677

• A migrating salmon was observed in Ontario’s Credit River in November 2007.[25]

2.12 Beaver impact

2.13.1 England and Wales

2.13.3 United States