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Last Update: 2 November 2017 Part I

M - 43
Migration of Fish
In ethology, the regular, usually seasonal, movement of all or part of an animal population to and from a
given area. Familiar migrants include many birds; hoofed animals, especially in East Africa and in the Arctic
tundra; bats; whales and porpoises; seals; and fishes, such as salmon.

Migration can be contrasted with emigration, which involves a change in location not necessarily followed
by a return journey; invasion or interruption, both of which involve the appearance and subsequent disappearance
of great numbers of animals at irregular times and locations; and range expansion, which tends to enlarge the
distribution of a species, particularly its breeding area.

The migration cycle is often annual and thus closely linked with the cyclic pattern of the seasons. The
migration of most birds and mammals and many of the fishes are on a yearly cycle. In many cases (e.g., salmon
and eels) animals with a relatively long life-span return to the place of birth in order to reproduce and eventually
die. In other cases, as in certain invertebrates, where the animal has a relatively brief life-span and reproduces
rapidly, migrations may not occur in every generation. The daily movements of certain fishes and invertebrates
have also been called migrations because of their regular occurrence. This type of movement, however, is not to be
confused with migration in the strict sense.

Most migrations involve horizontal travel. The distance traversed may be a few miles or several thousands
of miles.

Some migrations take a vertical direction and involve no appreciable horizontal movement. Certain aquatic
animals, for example, move from deep water to the surface according to the season. Certain birds, mammals, and
insects migrate altitudinally in mountainous areas, going from the upper zones, where they breed, to the foothills or
plains during seasons when the weather is severe and unfavourable. Such vertical travels involve essentially the
same type of environmental change as horizontal, or latitudinal, migrations over long distances.

Origin and evolution of migration

The origins of migration remain in the realm of pure conjecture; neither observation nor experiment has
resolved the matter. The explanation, however, must be related to geographical and climatological factors that have
prevailed since the Tertiary Period, which ended some 2,500,000 years ago. The great Quaternary ice ages, which
came later, were very important in altering the distribution of animals over a large part of the world, but migrations
occurred long before.
Migration, as it is now known among modern birds and mammals, probably appeared gradually by stages.
Some animals changed their habitat only slightly, never leaving the same general region. The movements of other
animals were more erratic, their dispersal being oriented toward the most favourable places. Such movements are
the first stages of true migration--a phenomenon characterized by elaborate mechanisms--which gradually acquired
stability through natural selection. At first, many populations must have perished rather than attempt to flee from
unfavourable conditions. Only a fraction of such populations probably sought more favourable conditions
elsewhere, but natural selection favoured the "migrants," and migratory tendencies were retained.
In some cases, original habitats were in present-day wintering areas, and animals developed a tendency to
leave in spring in order to breed in other territories. Seasonal changes of weather and food supply in these newly
settled regions forced the animals to migrate in fall, and they thus retreated to their former range. Among birds
nesting in the Northern Hemisphere, hummingbirds, tyrant flycatchers, tanagers, orioles, bee-eaters, and swifts
have distinct tropical affinities; in recent geological times these birds gradually spread northward as glacial ice
receded and the continent became warmer. Other birds, such as plover, ducks, and geese, originally lived in what is
now their breeding area. Gradual climatic changes forced them to spend their winters in regions far to the south.
Migrations thus appear to be the consequence of invasions or emigrations, during which animals settle in new areas
during a segment of the annual cycle.
Migratory birds use the routes by which their ancestors first invaded new regions after the glacial recession.
The yellow wagtail (Motacilla flava) and the wheatear (Oenanthe oenanthe) settled in Alaska; they migrate
annually into other parts of the Western Hemisphere but spend their winters in the warm regions of southeastern
Asia and even Africa, probably following the migratory route of their ancestors. A typically North American
species, the gray-cheeked thrush (Hylocichla minima), which has extended its breeding area to northeastern
Siberia, returns to spend the winter in the central regions of South America.
Migration of Fish
Many species of fish wander annually through a particular area of the ocean. Some are true migrants,
travelling regularly over great distances. Young fish usually leave the spawning grounds for areas where they
develop into juveniles, before joining the adult stock at the feeding grounds. Adults move to the spawning grounds,
then return to the feeding grounds. Migratory patterns of fish are related to oceanographic factors and to currents.
Eggs, larvae, and young drift passively with the current, although migration of adult fish toward breeding grounds is
usually against the current. Adult movements thus are directional rather than passive, and the fish respond to
environmental conditions--e.g., climate.
Three categories of migratory fishes can be distinguished: oceanodromous, anadromous, and catadromous.
Oceanodromous fish
Oceanodromous fish, which occur widely throughout the world's oceans, live and migrate wholly in the sea.
They differ mainly from one another by the method and extent of their migration.
Herring (Clupea harengus), extensively studied because of their economic importance, are the best known
of the oceanodromous type and can be classified into several populations, or local races, which do not mix freely.
In addition, each has a particular migratory behaviour. In the North Sea, distinct groups spawn in different seasons
and on different grounds: Buchan herring spawn in August and September off the coast of Scotland and migrate to
the coast of southwestern Norway; Dogger Bank herring spawn in September and October in the central part of the
North Sea and along the English coast and then migrate to the Skagerrak, an arm of the North Sea between
Denmark and Norway; Downs herring spawn from November to January off the French coast, mainly between
Dunkirk and Fcamp, then feed in summer in the middle and northern parts of the North Sea, sharing the feeding
grounds with other populations. The diversity of migration and of reproductive seasons is closely connected with
the annual cycle of oceanographic conditions in the North Sea.
Cod (Gadus morhua) have migration patterns similar to those of herring. The migrations of other fish cover
even greater distances; in the Atlantic, for example, white tuna (Germo alalunga) are found in winter around the
Azores and the Canary Islands, where they spawn in spring. They then migrate northward to the Gulf of Gascogne
and afterward to the waters around Iceland, arriving there in July. Populations of red tuna (Thunnus thynnus) occur
throughout the Mediterranean Sea and the eastern Atlantic. In May and June they spawn in the western
Mediterranean. During summer they spread northward to feed, finally reaching the Arctic Ocean. Similar
migrations occur along the North American coast in the Atlantic and throughout the Pacific.
Anadromous fish
Anadromous fish live in the sea and migrate to fresh water to breed. Their adaptations to conditions of
different habitats are precise, particularly with regard to salinity of the water.
Salmon (Salmo, Oncorhynchus) spawn in the cold, clear waters of lakes or upper streams. Eggs are laid in
gravel beds. The young of the Atlantic salmon remain in fresh water for two to three years, sometimes as long as
six; Pacific salmon sometimes migrate to the sea in their first year. Adult fish usually remain in the sea for two or
three winters, but sometimes only one. Then, as grilse (adolescents) or as adults, they return to fresh water and
spawn, after changes occur in colour and other external features. Some Atlantic salmon die in fresh water after a
single spawning; others return to the sea.
The tagging of salmon has shown that European types may cross from Norway to Scotland, as well as the
reverse. Pacific salmon are probably distributed over the Pacific Ocean and Bering Sea, between latitudes 45 N
and 65 N with surface waters of 2 to 11 C (36 to 52 F).
Experiments in Canada and the United States, in which young salmon migrating to the Pacific have been
tagged, have shown that a high proportion of the fish return to the river where they hatched. Tagging of Atlantic
salmon has shown that a few survivors have migrated two or even three times to a particular river in successive
years. Adults reared from experimentally transplanted eggs return to the stream where they were hatched or grew,
not to the stream where the eggs were laid. Aside from other means of orientation, such as reference to celestial
features, topographical features are believed to play an important part in recognition of the original habitat. The
sense of smell, or olfaction, however, has the most important role. Experiments have shown that migrating salmon
are attracted to the waters of the stream in which they are going to spawn. Experiential imprinting at an early stage
of development enables a grown fish to respond to waters that contain substances with a particular odour or that
have a characteristic temperature.
Catadromous fish
Catadromous fish spend most of their lives in fresh water, then migrate to the sea to breed. This type is
exemplified by eels of the genus Anguilla, numbering 16 species, the best-known of which are the North American
eel (A. rostrata) and the European eel (A. anguilla).
European eels and North American eels spawn in warm saline waters of the Atlantic, at depths of 400 to 700
metres (about 1,300 to 2,300 feet), in an area centred near latitude 26 N longitude 55 W called the Sargasso Sea.
The pelagic eggs develop into leptocephali--transparent, leaflike forms with relatively small heads--that are carried
by the Gulf Stream to the shallow waters of the continental shelves. When they are about two and one-half years
old and about eight centimetres long (a little more than three inches), a metamorphosis occurs. The leptocephali are
transformed into so-called elvers, which are bottom-dwelling, pigmented, and cylindrical in form. They arrive in
coastal waters as glass eels and begin to swim upstream in freshwater streams in spring. Their migration upstream
is spectacular, as the young fish gather by millions, forming a dense mass several miles long. In freshwater the eels
grow to full size, becoming yellow eels. They live as such for 10 to 15 years before changing into silver eels, with
enlarged eyes; they swim downstream to the sea, return to the spawning grounds (Sargasso Sea), and die.
The migration of these eels is not entirely understood, particularly their return to the Sargasso Sea. It may be
that European eels and North American eels belong to the same species.
Physiological stimulus of migration
Migration, like reproduction and other phases (as molting in birds), is part of the life cycle and depends on a
complex internal rhythm that affects the whole organism, particularly the endocrine glands (glands of internal
secretion) and the gonads. Migration must thus be viewed in relation to the entire annual cycle.
Each year birds return to particular areas to breed, and remain there until the members of the brood can care
for themselves. There is no relation between the reproductive and migratory stimuli, yet the two phenomena,
although independent, are nevertheless stimulated by the same factor.
A physiological study of certain migrants has revealed that metabolic patterns usually change prior to
migration, and fats accumulate in the body tissues. The whitethroat (Sylvia communis) weighs an average of 12 to
13 grams (about 0.4 ounce) during the breeding season, 16 to 19 grams (about 0.6 ounce) in the autumn, and 20 to
24 grams (about 0.8 ounce) in the winter. Food consumption increases with the autumn molt, reaching a peak at the
beginning of the migration season. These fundamental physiological changes, chiefly under the control of the
thyroid gland, are correlated with migratory activity. Such fluctuations are not observed in nonmigratory species.
Variations in metabolism and related phenomena are controlled by an endocrine gland, namely the pituitary
gland, which is located in the lower part of the brain and acts as a command post, sending out instructions in the
form of secretions called hormones. That the pituitary has a cycle independent of environmental factors is
demonstrated by the regularity with which phases such as reproduction occur from year to year in the lives of some
birds, and by the diverse response of various species and populations to the same environmental factors. That the
pituitary is, however, influenced by environmental factors, such as variations in day length and the intensity of the
Sun, has been demonstrated experimentally.
Gonadal development and the deposition of fat, for example, are influenced by the pituitary, which responds
to increasing day length in springtime by accelerating the rate of gonadal development. The pituitary thus governs
the development of gonads and, in addition, affects all metabolic processes, including development of the thyroid
gland, so as to prepare the animal physiologically for migration. If only the pituitary and variations in day length
were involved, migration would be triggered at definite times, because the pituitary cycle is fixed, and
photoperiodism is a highly predictable phenomenon; such a lack of flexibility, however, would inevitably cause
migrant populations to suffer catastrophes because ecological conditions are irregular--meteorological events, such
as the arrival of spring, and biological phenomena, such as flowering, foliation, hatching of insects, and availability
of food, are highly variable from year to year. The pituitary thus serves only to prepare the bird for flight; the
proper ecological conditions, on the other hand, are necessary to initiate it. The availability of food is an important
factor. Temperature and weather conditions also have an influence--a sudden period of cold weather during autumn
may induce the immediate departure of many migrants.
Sensitivity to changes in the weather and other environmental conditions varies markedly among species.
Some, such as the woodcock, snipe, lapwing, starling, and lark, rely on surrounding conditions to initiate their
spring and autumn migrations, and the patterns of their flight depend on temperature and barometric pressure.
Others, such as the swift, cliff swallow, Baltimore oriole, and short-tailed petrel, are less weather dependent, and,
since the dates of their arrival and departure are not regulated by the weather, they occur with remarkable
regularity each year.
The factors that stimulate migration in animals other than birds are not yet well understood. Ecological
conditions play a great part in the migratory activity of mammals, who react to general food shortage by moving to
another region. Whale, for example, leave the Antarctic region as winter modifies the oceanographic conditions.
Seals disperse when the food supply in the area of their breeding colonies is depleted. Environmental factors are of
primary importance in the migration of fishes and marine invertebrates. Annual movements of water masses
change physical conditions such as temperature and salinity; biotic conditions are influenced accordingly.
Ecological significance of migration
There are many ecological implications of migration. The food resources of some regions would not be
adequately exploited without moving populations. The sequence of migratory movement is closely integrated in
the annual cycle of ecosystems characterized by productivity fluctuations. Migratory behaviour concerns only
species located at specific trophic levels (zones of food availability) where maximal fluctuations occur both in
breeding areas and in wintering regions. Migrant birds avoid equatorial forests where productivity is constant
throughout the year, and food surpluses do not occur. They do congregate, on the other hand, in savannas where
productivity varies with the seasons.
Such a coordinated sequence is particularly apparent in the case of birds migrating from the northern Arctic
regions to tropical winter regions; both life zones are characterized by broad fluctuations in productivity. In the
Arctic, vegetal and animal production is very high during the summer; ducks and waders nest in great numbers,
exploiting these resources. As winter comes, food becomes scarce, and water birds migrate to the tropics, where
the rainy season has caused food production to increase to optimal levels. Ducks and wading birds concentrate in
the most favourable areas, remaining until spring, when productivity is lowest. By then the condition of breeding
areas is again favourable for the birds. The life cycles of these birds are closely attuned with the cycles of their
various habitats, and the sizes of bird populations are controlled by the capacity of both areas to sustain them.
Migration, then, has considerable ecological significance. It enables fast-moving animals to exploit
fluctuating resources and to settle in areas where life would not be tenable for animals incapable of rapid travel. On
the other hand, peaks of food production would be unexploited without the periodic presence of migratory