Animal migration

Wildebeest on the Serengeti 'great migration'

Migration can take very different forms in different species, and has a variety of causes.[1][2][3] As such, there is no simple accepted definition of migration.[4] One of the most commonly used definitions, proposed by the zoologist J. S. Kennedy[5] is

Migratory behavior is persistent and straightened-out movement effected by the animal's own locomotory exertions or by its active embarkation on a vehicle. It depends on some temporary inhibition of station-keeping responses, but promotes their eventual disinhibition and recurrence.[5]

Migration encompasses four related concepts: persistent straight movement; relocation of an individual on a greater scale (in both space and time) than its normal daily activities; seasonal to-and-fro movement of a population between two areas; and movement leading to the redistribution of individuals within a population.[4] Migration can be either obligate, meaning individuals must migrate, or facultative, meaning individuals can "choose" to migrate or not. Within a migratory species or even within a single population, often not all individuals migrate. Complete migration is when all individuals migrate, partial migration is when some individuals migrate while others do not, and differential migration is when the difference between migratory and non-migratory individuals is based on discernible characteristics like age or sex.[4] Irregular (non-cyclical) migrations such as irruptions can occur under pressure of famine, overpopulation of a locality, or some more obscure influence.[6]

Seasonal migration is the movement of various species from one habitat to another during the year. Resource availability changes depending on seasonal fluctuations, which influence migration patterns. Some species such as Pacific salmon migrate to reproduce; every year, they swim upstream to mate and then return to the ocean.[7] Temperature is a driving factor of migration that is dependent on the time of year. Many species, especially birds, migrate to warmer locations during the winter to escape poor environmental conditions.[8]

Circadian migration is where birds utilise circadian rhythm (CR) to regulate migration in both fall and spring. In circadian migration, clocks of both circadian (daily) and circannual (annual) patterns are used to determine the birds' orientation in both time and space as they migrate from one destination to the next. This type of migration is advantageous in birds that, during the winter, remain close to the equator, and also allows the monitoring of the auditory and spatial memory of the bird's brain to remember an optimal site of migration. These birds also have timing mechanisms that provide them with the distance to their destination.[9]

Tidal migration is the use of tides by organisms to move periodically from one habitat to another. This type of migration is often used in order to find food or mates. Tides can carry organisms horizontally and vertically for as little as a few nanometres to even thousands of kilometres.[10] The most common form of tidal migration is to and from the intertidal zone during daily tidal cycles.[10] These zones are often populated by many different species and are rich in nutrients. Organisms like crabs, nematodes, and small fish move in and out of these areas as the tides rise and fall, typically about every twelve hours. The cycle movements are associated with foraging of marine and bird species. Typically, during low tide, smaller or younger species will emerge to forage because they can survive in the shallower water and have less chance of being preyed upon. During high tide, larger species can be found due to the deeper water and nutrient upwelling from the tidal movements. Tidal migration is often facilitated by ocean currents.[11][12][13]

While most migratory movements occur on an annual cycle, some daily movements are also described as migration. Many aquatic animals make a diel vertical migration, travelling a few hundred metres up and down the water column,[14] while some jellyfish make daily horizontal migrations of a few hundred metres.[15]

Flocks of birds assembling before migration southwards

Approximately 1,800 of the world's 10,000 bird species migrate long distances each year in response to the seasons.[16] Many of these migrations are north-south, with species feeding and breeding in high northern latitudes in the summer and moving some hundreds of kilometres south for the winter.[17] Some species extend this strategy to migrate annually between the Northern and Southern Hemispheres. The Arctic tern has the longest migration journey of any bird: it flies from its Arctic breeding grounds to the Antarctic and back again each year, a distance of at least 19,000 km (12,000 mi), giving it two summers every year.[18]

Bird migration is controlled primarily by day length, signalled by hormonal changes in the bird's body.[19] On migration, birds navigate using multiple senses. Many birds use a sun compass, requiring them to compensate for the sun's changing position with time of day.[20] Navigation involves the ability to detect magnetic fields.[21]

Many species of salmon migrate up rivers to spawn

Most fish species are relatively limited in their movements, remaining in a single geographical area and making short migrations to overwinter, to spawn, or to feed. A few hundred species migrate long distances, in some cases of thousands of kilometres. About 120 species of fish, including several species of salmon, migrate between saltwater and freshwater (they are 'diadromous').[22][23]

Forage fish such as herring and capelin migrate around substantial parts of the North Atlantic ocean. The capelin, for example, spawn around the southern and western coasts of Iceland; their larvae drift clockwise around Iceland, while the fish swim northwards towards Jan Mayen island to feed and return to Iceland parallel with Greenland's east coast.[24]

In the 'sardine run', billions of Southern African pilchard Sardinops sagax spawn in the cold waters of the Agulhas Bank and move northward along the east coast of South Africa between May and July.[25]

An aggregation of migratory Pantala flavescens dragonflies, known as globe skimmers, in Coorg, India

Some winged insects such as locusts and certain butterflies and dragonflies with strong flight migrate long distances. Among the dragonflies, species of Libellula and Sympetrum are known for mass migration, while Pantala flavescens, known as the globe skimmer or wandering glider dragonfly, makes the longest ocean crossing of any insect: between India and Africa.[26] Exceptionally, swarms of the desert locust, Schistocerca gregaria, flew westwards across the Atlantic Ocean for 4,500 kilometres (2,800 mi) during October 1988, using air currents in the Inter-Tropical Convergence Zone.[27]

In some migratory butterflies, such as the monarch butterfly and the painted lady, no individual completes the whole migration. Instead, the butterflies mate and reproduce on the journey, and successive generations continue the migration.[28]

Some mammals undertake exceptional migrations; reindeer have one of the longest terrestrial migrations on the planet, reaching as much as 4,868 kilometres (3,025 mi) per year in North America. However, over the course of a year, grey wolves move the most. One grey wolf covered a total cumulative annual distance of 7,247 kilometres (4,503 mi).[29]

High-mountain shepherds in Lesotho practice transhumance with their flocks.

Mass migration occurs in mammals such as the Serengeti 'great migration', an annual circular pattern of movement with some 1.7 million wildebeest and hundreds of thousands of other large game animals, including gazelles and zebra.[30][31] More than 20 such species engage, or used to engage, in mass migrations.[32] Of these migrations, those of the springbok, black wildebeest, blesbok, scimitar-horned oryx, and kulan have ceased.[33] Long-distance migrations occur in some bats – notably the mass migration of the Mexican free-tailed bat between Oregon and southern Mexico.[34] Migration is important in cetaceans, including whales, dolphins and porpoises; some species travel long distances between their feeding and their breeding areas.[35]

Humans are mammals, but human migration, as commonly defined, is when individuals often permanently change where they live, which does not fit the patterns described here. An exception is some traditional migratory patterns such as transhumance, in which herders and their animals move seasonally between mountains and valleys, and the seasonal movements of nomads.[36][37]

Among the reptiles, adult sea turtles migrate long distances to breed, as do some amphibians. Hatchling sea turtles, too, emerge from underground nests, crawl down to the water, and swim offshore to reach the open sea.[38] Juvenile green sea turtles make use of Earth's magnetic field to navigate.[39]

Christmas Island red crabs on annual migration

Some crustaceans migrate, such as the largely-terrestrial Christmas Island red crab, which moves en masse each year by the millions. Like other crabs, they breathe using gills, which must remain wet, so they avoid direct sunlight, digging burrows to shelter from the sun. They mate on land near their burrows. The females incubate their eggs in their abdominal brood pouches for two weeks. Then they return to the sea to release their eggs at high tide in the moon's last quarter. The larvae spend a few weeks at sea and then return to land.[40][41]

• Aidley, D. J. (1981). Animal migration. Cambridge University Press. ISBN 978-0-52123-274-6.
• Baker, R. R. (1978) The Evolutionary Ecology of Animal Migration. Holmes & Meier. ISBN 978-0-34019-409-6.
• Dingle, H. (1996) Migration: The Biology of Life on the Move. Oxford University Press. ISBN 978-0-19802-577-1.
• Gauthreaux, S. A. (1980) Animal Migration, Orientation, and Navigation. Academic Press. ISBN 978-0-12277-750-9.
• Milner-Gulland, E. J., Fryxell, J. M., and Sinclair, A. R. E. (2011) Animal Migration: A Synthesis. Oxford University Press. ISBN 978-0-19956-899-4.
• Rankin, M. (1985) Migration: Mechanisms and Adaptive Significance: Contributions in Marine Science. Marine Science Institute. OCLC 747358527.
• Riede, K. (2002) Global Register of Migratory Species. With database and GIS maps on CD. ISBN 978-3-78433-826-2.

• Drake, V. A. and Gatehouse, A. G. (1995) Insect migration: tracking resources through space and time. Cambridge University Press. ISBN 978-0-52101-853-1
• Elphick, J. (1995) The atlas of bird migration: tracing the great journeys of the world's birds. Random House. ISBN 978-1-55407-971-1
• Greenberg, R. and Marra, P. P. (2005) Birds of Two Worlds: The Ecology and Evolution of Migration. Johns Hopkins University Press. ISBN 978-0-80188-107-7
• Lucas, M. C. and Baras, E. (2001) Migration of freshwater fishes. Blackwell Science. ISBN 978-0-47099-964-6
• MacKeown, B. A. (1984) Fish migration. Timber Press. ISBN 978-0-91730-499-6
• Sonnenschein, E; Berthold, P. (2003) Avian migration. Springer. ISBN 978-3-54043-408-5

• Gans, R. and Mirocha, P. How do Birds Find their Way? HarperCollins. (Stage 2) ISBN 978-0-43969-940-2
• Marsh, L. (2010) Amazing Animal Journeys. National Geographic Society. (Level 3) ISBN 978-0-00826-686-8