Altitudinal migration is a short-distance

animal migration Animal migration is the relatively long-distance movement of individual animals, usually on a seasonal basis. It is the most common form of migration in ecology. It is found in all major animal groups, including birds, mammals, fish, reptile ...

from lower altitudes to higher altitudes and back. Altitudinal migrants change their elevation with the seasons making this form of animal migration seasonal. Altitudinal migration can be most commonly observed in species inhabiting temperate or tropical ecosystems. This behavior is commonly seen among avian species but can also be observed within other vertebrates and some invertebrates. It is commonly thought to happen in response to climate and food availability changes as well as increasingly due to anthropogenic influence. These migrations can occur both during reproductive and non-reproductive seasons. The patterns of altitudinal migration may be affected by climate change resulting in potentially life-threatening situations for some species. Deforestation can affect the migration corridors of altitudinal migrant and could lead to smaller areas for these species to migrate. Changes the environment of altitudinally migrating species can also affect seed dispersal.

Regions

Species that exhibit altitudinal migration can be found in almost every continent on Earth. The only continent in which altitudinal migration cannot be observed is Antarctica. There are many documented examples of migratory range shifts along an elevation gradient among

temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...

species. While these migrations are more understood in temperate regions, and far less understood among

tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

ecosystems and species, there are documented cases. Altitudinal migration is typically seen among

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

found in

montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial ...

areas. Generally, as elevation increases, the species richness decreases. Typical characteristics of tropical altitudinal migrants include: a high rate of

frugivory A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...

or nectarivory; movement between lower elevation areas during non-breeding seasons and higher elevation areas during breeding seasons, or on a consistent annual or seasonal cycle; at least part of the population being migratory with a possible portion of the population residing at breeding sites year-round. This last characteristic can be sex-biased, as it is with

junco A junco , genus ''Junco'', is a small North American bird in the New World sparrow family Passerellidae. Junco systematics are still confusing after decades of research, with various authors accepting between three and twelve species. Despite ...

s, in which the males are less likely to migrate than the females. The white-ruffed manakin provides a good example of an altitudinal migrant by displaying all of these traits. It has a high rate of frugivory, migrates from lower elevations to higher elevations on a predictable breeding-season based cycle, and part of the population is migratory, with a small portion possibly remaining at the breeding sites year-round.

Species

There are many species of animals which exhibit altitudinal migration. While there is many examples of vertebrates which exhibit altitudinal migration, as of 2021 there are few documented examples of invertebrates which exhibit altitudinal migration.

Vertebrates

Birds

In the tropics, altitudinal migrations are most commonly seen among frugivores or nectarivores, such as what is seen among tropical hummingbirds, which migrate altitudinally in response to shifts in food abundance and availability. This migration pattern has been observed in

neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...

birds, but has also been seen in other terrestrial, tropical montane species such as Baird's tapir and

white-lipped peccary The white-lipped peccary (''Tayassu pecari'') is a species of peccary found in Central and South America and the only member of the genus ''Tayassu''. Multiple subspecies have been identified. White-lipped peccaries are similar in appearance to ...

. Tropical avian species that are altitudinal migrants include the white-ruffed manakin,

resplendent quetzal The resplendent quetzal (''Pharomachrus mocinno'') is a small bird found in southern Mexico and Central America, with two recognized subspecies, ''P. m. mocinno'' and ''P. m. costaricensis''. These animals live in tropical forests, particularly ...

, at least 16 species of

raptor Raptor or RAPTOR may refer to: Animals The word "raptor" refers to several groups of bird-like dinosaurs which primarily capture and subdue/kill prey with their talons. * Raptor (bird) or bird of prey, a bird that primarily hunts and feeds on ...

, and many species of hummingbird. The

Hawaiian goose The nene (''Branta sandvicensis''), also known as the nēnē or the Hawaiian goose, is a species of bird Endemism in birds, endemic to the Hawaiian Islands. The nene is exclusively found in the wild on the islands of Oahu, Maui, Kauai, Kauai, Mo ...

, or Nene, is a species of goose native to the Hawaiian islands which has been known to migrate altitudinally. This species of goose can be found in lower elevations during breeding seasons and when molting, and in higher elevations during non-breeding seasons. Temperate avian species which have been known to altitudinally migrate include the American robin,

mountain chickadee The mountain chickadee (''Poecile gambeli'') is a small songbird, a passerine bird in the tit family Paridae. Taxonomy The specific name honors naturalist William Gambel. The mountain chickadee was formerly placed in the genus ''Parus'' with ...

, and

American dipper The American dipper (''Cinclus mexicanus''), also known as a water ouzel, is a stocky dark grey bird with a head sometimes tinged with brown, and white feathers on the eyelids that cause the eyes to flash white as the bird blinks. It is long, ...

Mammals

While less common in avian species in the temperate regions, altitudinal migration still plays a part in migration patterns in montane zones and is seen in most

ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...

s in the

Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico ...

. Ungulates that have been observed to migrate altitudinally include

roe deer The roe deer (''Capreolus capreolus''), also known as the roe, western roe deer, or European roe, is a species of deer. The male of the species is sometimes referred to as a roebuck. The roe is a small deer, reddish and grey-brown, and well-adapt ...

bighorn sheep The bighorn sheep (''Ovis canadensis'') is a species of sheep native to North America. It is named for its large horns. A pair of horns might weigh up to ; the sheep typically weigh up to . Recent genetic testing indicates three distinct subspec ...

, and

mountain goats The mountain goat (''Oreamnos americanus''), also known as the Rocky Mountain goat, is a hoofed mammal endemic to mountainous areas of western North America. A subalpine to alpine species, it is a sure-footed climber commonly seen on cliffs and ...

There are fewer documented examples of tropical mammal altitudinal migrants. While there are less known examples, some tropical bat species have been known to migrate altitudinally. As of 2014, there is not much information as to why tropical species migrate altitudinally, other than that it may be for food resources or reproduction, as it is for temperate bat species. Temperate bat species are also altitudinal migrants. Their migratory patterns are sex-biased altitudinal migrations, with females inhabiting lower elevations during reproductive periods.

Invertebrates

There is little documentation regarding invertebrates that migrate altitudinally compared to avian species, but documentation does exist. The Monarch butterfly a species which has been known to migrate altitudinally, as well as the chestnut tiger butterfly. According to Masahito T. Kimura, invertebrates migrate altitudinally "as a means to escape from unfavorable conditions such as low winter temperature, summer heat, low resource availability, high parasitism, severe microbe infection or overcrowding."

Causes

Altitudinal migration, as a short-distance migration pattern, has been easier to trace than long-distance patterns. Still, while the proximate causes and physiological adaptations for migrations are well understood, determining the ultimate causes have been difficult. This difficulty has been linked to limited success of mark and recapture techniques used to track migratory species. There are many hypotheses for why altitudinal migration may occur, including correlations between food abundance and nutrition-the need to migrate in order to meet specific needs associated with varying abundance and nutrition; reproduction-breeding sites being at elevations different from those of non-breeding sites; anthropogenic-species being increasingly driven to higher altitudes due to human actions.

Food abundance and nutrition

Migration in response to food abundance has been the most accepted hypothesis for why species migrate altitudinally. This hypothesis states that peaks in food abundance along an elevational gradient, such as the slope of a mountain, drive migration patterns as species exploit available food resources. Peaks in food abundance along this gradient often coincide with the breeding season. Some frugivorous birds, such as white-ruffed manakins (''Corapipo altera'') migrate to higher elevations to exploit peaks in fruit abundance. Evidence supports the possibility that migrants have a competitive advantage compared to non-migrant (sedentary) species, due to increased foraging ability over a larger area, resulting in greater food and nutrient uptake. It's been shown that diet differs between non-migratory and migratory species in large-scale analyses and species-pair comparisons of frugivorous tropical birds. While this hypothesis is supported, and has been the most accepted, it fails to explain why altitudinal migrants return to lower elevations, or if it is done in response to shifting food resources. It has been proposed that weather-related resource availability may trigger the elevational migration of some species, such as the white-ruffed manakin during storms.

Reproduction

A number of species engage in movement that could be defined as altitudinal migration as part of their mating or reproductive behaviors. For example, in male white-ruffed manakins, migratory behavior has shown to lessen social status and mating success at

leks A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...

the following breeding season. Most hummingbird species at

Monteverde Monteverde is the twelfth canton of the Puntarenas province of Costa Rica. It is located in the Cordillera de Tilarán mountain range. Roughly a four-hour drive from the Central Valley, Monteverde is one of the country's major ecotourism des ...

increase altitude during the wet season in order to breed. Of the 16 species of neotropical raptors (including the

Andean condor The Andean condor (''Vultur gryphus'') is a giant South American Cathartid vulture and is the only member of the genus ''Vultur''. Found in the Andes mountains and adjacent Pacific coasts of western South America, the Andean condor is the larg ...

''Vultur gryphus''), that are known to be altitudinal migrants, most breed in the high

Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...

and migrate to lowland areas during non-breeding seasons.

Nest predation

Studies have shown a decreased risk of nest predation at higher altitudes, which may explain the seasonal (breeding season—non-breeding season) altitudinal migration of some passerine birds. An experiment using 385 nests at varying locations on the Atlantic slope of Costa Rica showed decreased predation at increasing altitudes, with predation highest at intermediate altitudes. This hypothesis proposes that altitudinal migration may have evolved among some species as a response to nest predation, as a way lower the risk. Studies have also shown that elevation of home range influences breeding time.

Anthropogenic

The

walia ibex The walia ibex (''Capra walie'', Amharic: ዋልያ ''wālyā'') is an endangered species of ibex. It is sometimes considered a subspecies of the Alpine ibex. Threats against the species include habitat loss, poaching, and restricted range; only ...

(''Capra walie'') has increasingly been driven to higher altitudes in Ethiopian mountain ranges. This has occurred because of human activity impacting their native range, including war, expanding human settlement, and cultivation.

Conservation implications

Climate change

Climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...

could be causing migration patterns to shift into an earlier time frame, coinciding with an earlier start of the growing period. This means that migratory species may leave lower altitudes for higher-altitude breeding sites while those breeding sites still lack the necessary resources. Some species that have shorter migratory paths may be able to return to the lower elevations and wait, but run the risk of running out of the resources in that lower altitude, such as food and cover, that may only be available for a short, set period of time. Over 30% of birds and other species in montane forests show altitudinal migration patterns. Because of this, changes in climate and seasonality (decrease or increase) would affect a large portion of tropical species and have the potential to cause a

trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...

on the community-level. Furthermore, climate change may cause seasonal storms and rainfall patterns to change, shifting the timing and/or need for altitudinal migration in the future by shifting availability of resources, which is believed to be a driving cause of altitudinal migration. The upward shift of species caused by climate change also holds the potential to cause both mountaintop

extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

and lowland biotic attrition. This is because the lowland tropics lack species that can cope with increasing temperatures. An overall loss of species richness can occur due to there being fewer migratory species to replace lost ones. This has been seen occurring by looking at the average time of arrival and departure at high altitude areas for the American robin (''Turdus migratorius''). The average time interval has shifted by as much as two weeks due to the change in seasonal patterns of resource abundance and temperature.

Migration corridors

Migration corridors, connecting lowland and montane habitats are essential for the upkeep and survival of migratory species. Some species can cross cleared lands, such as

pasture Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep, or sw ...

s, but many require closed forest areas, such as what is provided by these migration corridors. Deforestation can disrupt these corridors and affect the migration pattern of species which do exhibit altitudinal migration. Deforestation can limit the amount of space available for migration, leading to a more narrow and limited migration pattern.

Seed dispersal

Tropical frugivorous birds have complex elevational migrations and are responsible for the dispersal of many species of seeds across the different

Holdridge life zones The Holdridge life zones system is a global bioclimatic scheme for the classification of land areas. It was first published by Leslie Holdridge in 1947, and updated in 1967. It is a relatively simple system based on few empirical data, giving ob ...

and transitional regions, causing wide dispersal of plant species and significant ecological linkage. A change in migration patterns can cause a decrease in the efficiency, and ability of these species as seed dispersal agents.

References

{{reflist, 26em Bird migration Animal migration Transhumance