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The term p?ramo can refer to a variety of alpine tundra ecosystems. Some ecologists describe the p?ramo broadly as ?all high, tropical, montane vegetation above the continuous timberline?.^1] A more narrow term classifies the p?ramo according to its regional placement - specifically located in ?the northern Andes of South America and adjacent southern Central America?. The p?ramo is the ecosystem of the regions above the continuous forest line, yet below the permanent snowline.^[1] It is a ?Neotropical high mountain biome with a vegetation composed mainly of giant rosette plants, shrubs and grasses?^[2]

Location

In the strictest sense of the term, all p?ramo ecosys tems are located in the Neotropics, namely South and Central America. Scattered throughout the regions between 11N? and 8S? latitudes, these ecosystems are located mainly in the NW corner of South America: specifically in the Cordillera de M?rida mountain range of Venezuela, and in Peru and Ecuador.

P?ramo ecosystems are also found in the mountain ranges of the Andes in Colombia, the Sierra Nevada de Santa Marta, as well as in the regions of Huehuetenango and El Quich? of Guatemala in the Sierra de los Cuchumatanes. The Cordillera de Talamanca of Costa Rica and Panama has a good example of the p?ramo. In northern Ecuador, the Guandera Biological Station is a fairly undisturbed p?ramo ecosystem.

The Sumapaz in the Altiplano Cundiboyacense mountain range of Colombia (about 20km from Bogot?), which is the largest p?ramo in the world. This region was declared a National Park of Colombia in 1977 because of its importance as a biodiversity hotspot and main source of water for the most densely populated area of the country, the Bogot? Savannah.

The 5.7-square-kilometre (1,405-acre) Paramo Wildlife Refuge Park in the San Jose Province of Costa Rica ?protects tropical forest areas in the high elevations of the Talamanca Mountains?^[3]

Cotopaxi National Park contains 329.9 square kilometres (81,524 acres) of protected land in the Cotopaxi Province of Ecuador. Much of this park represents the paramo ecosystem, wherein plants such as ?gentians, lycopodiums, almohadillas and obtrusive valerian, loricarias and chuquiragua grow?^[4]

Climate

P?ramo climates differ slightly depending on the specific location. In Colombia and northern Ecudaor, air masses from the Intertropical Convergence Zone (ITCZ) have a substantial effect on the climate, and these regions tend to be consistently humid (approx. 70-85%)^[2] throughout the year. The Andes also play a key role in the climate of these regions as they cause an orographic uplift in which moist air rises. This creates continuous moisture via rain, clouds, and fog, with many of them receiving over 2000 mm of rain annually^[2]

The p?ramos of the northern-most Andes of Venezuela, northern Colombia, and Costa Rica experience a different climate due to the dry season, which is caused by northeasterly trade winds. Southern Ecuador and northern Peru experience the most severe dryness as they are influenced by an air mass from the Amazon Basin, which releases its moisture on the eastern slopes, as well as another air mass from the west that is influenced by the Humboldt current.

Overall, p?ramo climates are known for their daily fluctuations in temperature and humidity. While they are generally cold and humid ecosystems, they often undergo a sudden and drastic change in weather in which they fluctuate between temperatures from below freezing to as high as 30C. This oscillation often results in a daily freeze-and-thaw cycle, sometimes described as ?summer every day and winter every night.?^[2] Mean annual temperatures of p?ramo ecosystems range from 2C to 10C, with increasingly colder temperatures at higher latitudes.

Soils

Soils in p?ramo ecosystems vary, but most are young and partially weathered. The soil has a relatively low pH because of an abundance of moisture and organic content. Organic content, even within disturbed sites averages very high which contributes to water retention in the soil. During cold and wet weather, there are few nutrients available and productivity is very low in p?ramo soils^[5]. Soils in p?ramo ecosystems have changed because of human activity, especially due to burning vegetation to clear land for grazing.

Soils in the south Ecuadorian p?ramo are characterized broadly into Andisols, Inceptisols, Histosols, Entisols, and Mollisols ^[2]. Recently, there has been an increase in Andisol soils, largely due to more volcanic activity^[6] These soils have a very high water retention rate, which contributes to the rise in cultivation and differential land use. This water supply stored in the soil in the higher elevation p?ramo in the Andes becomes the water supply for Andean settlements in lower altitudes.

Vegetation Zones

P?ramos are divided into separate zones based on altitude and vegetation structure, with the three main types of p?ramo vegetation unequally distributed throughout the different zones.

Superp?ramo is at the highest elevation and is usually considered to be the transition zone between the higher, permanent snow region and the lower grass p?ramo zone. The superp?ramo zone is generally narrow and exists atop loose stones and sandy soils at about 4500-4800 m. It has the lowest air temperature, precipitation level, soil water-holding capacity, and nutrient content of all the zones. Being the highest in altitude, it also has the highest levels of solar radiation and night frost. For this reason, vegetation in the superp?ramo must be highly resistant to such severe fluctuations in weather.1 Air temperatures are low- cold at night and cool during the day with daily oscillations larger than the oscillations of monthly averages^[2]. Due to its localization on high mountains, this area is the least disturbed by humans and contains the most endemic species of all the zones.This p?ramo often has so much visible sand that it?s easily mistaken for a beach or desert, however there are many tiny, clumped or scattered plants that persist in this zone. Some examples include Azorella pedunculat from the Apiaceae family, as well as plants from the Asteraceae, Fabaceae, and Ericaceae families.

The most broadly described zone is that of the grass p?ramo. Grass p?ramos cover large areas of mountain ranges, while others are limited to small areas on slopes and summits of very high mountains^[7]. This is often what people are referring to when they use the term ?p?ramo.? It has continuous vegetation and plant cover with a ?yellowish to olive?brown? look due to the combination of dead grasses amongst the living^[2]. The grass p?ramo exists from approximately 3500 - 4100 m, and is composed of mostly tussock-grasses or bunch-grasses^[2]. Plant species of the Calamagrostis and Festuca genera tend to dominate this zone. Other common vegetation includes large and small shrubs, stunted trees, cushion plants, herbs, graminoids, and rosette plants^[2]. While these are the dominant species, grass p?ramos can also contain tall- and short-grass communities, including herbaceous and woody vegetation. Due to its easier access and high levels of grass, this zone is more impacted by humans and suffers from both burning and grazing activities.

Subp?ramo is the lowest and most diverse zone. At 3000 ? 3500 m, it is a shrub-dominated zone that combines aspects of both the grass p?ramo above and the forest below. Along with shrubs, this zone also contains small, scattered trees which gradually transition into the grasses and herbs of the grass p?ramo above. Plant communities in this vegetation zone are also known to include thickets that are mainly composed of shrubby or woody vegetation, including species from the Ilex, Ageratina, and Baccharis genera. Fragmented forests can appear in the subp?ramo due to microclimatic or edaphic conditions, yet the more abrupt changes are generally due to anthropogenic disruption such as cutting, burning, and grazing activities. Because of these high levels of disruption, it is believed that subp?ramos are made up of largely secondary-growth communities. The high levels of disruption also make this zone particularly difficult to define, as humans typically extend and expand the zone for their own purposes, sometimes over hundreds or thousands of years. This has altered forest lines, often lowering them by several hundred meters, which has also affected the zoning habitats for many animals.

Fauna

The vegetation of the p?ramo provides shelter and habitat for a variety of mamm als, birds, insects, amphibians, and reptiles. Some animals commonly found in p?ramo ecosystems include the Andean Fox (also referred to as the ?P?ramo Wolf?, the White-Tailed Deer, and the Spectacled Bear which occasionally forages in the high p?ramo for its favorite snack, Puya bromeliads^[8]. Invertebrates such as grasshoppers, cockroaches, beetles, and flies are found mostly in the subp?ramo. Amphibians have been well documented in p?ramo ecosystems, including the salamander (Bolitoglossa spp.), and the frog (Eleutherodactylus spp.). Reptiles commonly sighted include lizards (Stenocercus spp,. Phenacosaurus spp., and Proctoporus spp.)

In all, 69 species of birds are listed as ?total users? of p?ramo habitats, with ?41 species making it their primary habitat and 16 as indicator species?^[2]. The Condor, also called ?King of the Andes? is one of the most impressive birds in the Andes due to its wide wingspan and majestic flight, but is no longer frequently seen. The most numerous bird families in the p?ramo, however, include ?the hummingbirds, furnariids, finches and flycatchers?, which tolerate the cold climate by going into ?a kind of nightly hibernation.?^[8] . Other commonly sighted birds include the turkey buzzard (Cathartes aura), eagle, hawk, falcon (Phalcoboenus carunculatus), and the rufous-fronted parakeet (Bolborhynchus ferrugineifrons)^[2].

Hummingbirds, bees and flies are all important pollinators in the p?ramo, while birds and smaller mammals such as rabbits and guinea pigs are important seed dispersers. Many of the larger mammals of the p?ramo are now rarely seen as their numbers have continuously dwindled due to being hunted excessively by man.

Human Impact and Climate Change

Humans have inhabited the p?ramo of the Andes for approximately the past 15,000 years. Deforestation has been extensive and in some cases, like the northern Andes, 90-95% of forests have been cleared. Other sites in Venezuela and Colombia show evidence that humans settled there at least 500-800 years ago and used the land for agriculture and hunting^[2].

When Europeans came to the Americas they introduced exotic plants and animals that greatly affected the land, especially the cattle which were introduced to the p?ramos in the early 18th century. By the 20th century, the growing population of settlers led to an increased demand for land, and p?ramo ecosystems suffered accordingly. As more land was needed for cattle, fire became a very popular resource for clearing land and eventually p?ramos became excessively burned and overgrazed.Both burning and grazing have had a significant impact on the vegetation, soils, species diversity, and water storage capacity of the p?ramos. In burned and disturbed sites that were studied in the Andes, the pH and phosphorus concentration in the soil are higher than in non-burned sites^[9].

Growing populations in Colombia, Venezuela, and Ecuador have forced settlements in higher elevations covering more p?ramo. Recent developments such as construction of aqueducts, drainage systems, and roads, mining, and afforestaion have been a huge additional p?ramo disturbance.

Climate change is becoming an increasingly pressing issue for p?ramo ecosystems. Increases in temperature extremes are forcing many fauna and flora species to higher grounds, and eventually they could face extinction. The flora of p?ramos is adapted to specific conditions and is thus vulnerable to even small climatic change. Climate change in the Andes is also causing glaciers in the p?ramo to disappear as well as decreasing rainfall, virtually drying up p?ramo and in turn, drying up the water supply for cities such as Quito, Ecuador and Bogot?, Colombia.

nced by an air mass from the Amazon Basin, which releases its moisture on the eastern slopes, as well as another air mass from the west that is influenced by the Humboldt current.

Overall, p?ramo climates are known for their daily fluctuations in temperature and humidity. While they are generally cold and humid ecosystems, they often undergo a sudden and drastic change in weather in which they fluctuate between temperatures from below freezing to as high as 30C. This oscillation often results in a daily freeze-and-thaw cycle, sometimes described as ?summer every day and winter every night.?^[2] Mean annual temperatures of p?ramo ecosystems range from 2C to 10C, with increasingly colder temperatures at higher latitudes.

Soils

Soils in p?ramo ecosystems vary, but most are young and partially weathered. The soil has a relatively low pH because of an abu ndance of moisture and organic content. Organic content, even within disturbed sites averages very high which contributes to water retention in the soil. During cold and wet weather, there are few nutrients available and productivity is very low in p?ramo soils^[5]. Soils in p?ramo ecosystems have changed because of human activity, especially due to burning vegetation to clear land for grazing.

Soils in the south Ecuadorian p?ramo are characterized broadly into Andisols, Inceptisols, Histosols, Entisols, and Mollisols ^[2]. Recently, there has been an increase in Andisol soils, largely due to more volcanic activity^[6] These soils have a very high water retention rate, which contributes to the rise in cultivation and differential land use. This water supply stored in the soil in the higher elevation p?ramo in the Andes becomes the water supply for Andean settlements in lower altitudes.

Vegetation Zones

P?ramos are divided into separate zones based on altitude and vegetation structure, with the three main types of p?ramo vegetation unequally distributed throughout the different zones.

Superp?ramo is at the highest elevation and is usually considered to be the transition zone between the higher, permanent snow region and the lower grass p?ramo zone. The superp?ramo zone is generally narrow and exists atop loose stones and sandy soils at about 4500-4800 m. It has the lowest air temperature, precipitation level, soil w ater-holding capacity, and nutrient content of all the zones. Being the highest in altitude, it also has the highest levels of solar radiation and night frost. For this reason, vegetation in the superp?ramo must be highly resistant to such severe fluctuations in weather.1 Air temperatures are low- cold at night and cool during the day with daily oscillations larger than the oscillations of monthly averages^[2]. Due to its localization on high mountains, this area is the least disturbed by humans and contains the most endemic species of all the zones.This p?ramo often has so much visible sand that it?s easily mistaken for a beach or desert, however there are many tiny, clumped or scattered plants that persist in this zone. Some examples include Azorella pedunculat from the Apiaceae family, as well as plants from the Asteraceae, Fabaceae, and Ericaceae families.

The most broadly described zone is that of the grass p?ramo. Grass p?ramos cover large areas of mountain ranges, while others are limited to small areas on slopes and summits of very high mountains^[7]. This is often what people are referring to when they use the term ?p?ramo.? It has continuous vegetation and plant cover with a ?yellowish to olive?brown? look due to the combination of dead grasses amongst the living^[2]. The grass p?ramo exists from approximately 3500 - 4100 m, and is composed of mostly tussock-grasses or bunch-grasses^[2]. Plant species of the Calamagrostis and Festuca genera tend to dominate this zone. Other common vegetation includes large and small shrubs, stunted trees, cushion plants, herbs, graminoids, and rosette plants^[2]. While these are the dominant species, grass p?ramos can also contain tall- and short-grass communities, including herbaceous and woody vegetation. Due to its easier access and high levels of grass, this zone is more impacted by humans and suffers from both burning and grazing activities.

Subp?ramo is the lowest and most diverse zone. At 3000 ? 3500 m, it is a shrub-dominated zone that combines aspects of both the grass p?ramo above and the forest below. Along with shrubs, this zone also contains small, scattered trees which gradually transition into the grasses and herbs of the grass p?ramo above. Plant communities in this vegetation zone are also known to include thickets that are mainly composed of shrubby or woody vegetation, including species from the Ilex, Ageratina, and Baccharis genera. Fragmented forests can appear in the subp?ramo due to microclimatic or edaphic conditions, yet the more abrupt changes are generally due to anthropogenic disruption such as cutting, burning, and grazing activities. Because of these high levels of disruption, it is believed that subp?ramos are made up of largely secondary-growth communities. The high levels of disruption also make this zone particularly difficult to define, as humans typically extend and expand the zone for their own purposes, sometimes over hundreds or thousands of years. This has altered forest lines, often lowering them by several hundred meters, which has also affected the zoning habitats for many animals.

Fauna

The vegetation of the p?ramo provides shelter and habitat for a variety of mammals, birds, insects, amphibians, and reptiles. Some animals commonly found in p?ramo ecosystems include the Andean Fox (also referred to as the ?P?ramo Wolf?, the White-Tailed Deer, and the Spectacled Bear which occasionally forages in the high p?ramo for its favorite snack, Puya bromeliads^[8]. Invertebrates such as grasshoppers, cockroaches, beetles, and flies are found mostly in the subp?ramo. Amphibians have been well documented in p?ramo ecosystems, including the salamander (Bolitoglossa spp.), and the frog (Eleutherodactylus spp.). Reptiles commonly sighted include lizards (Stenocercus spp,. Phenacosaurus spp., and Proctoporus spp.)

In all, 69 species of birds are listed as ?total users? of p?ramo habitats, with ?41 species making it their primary habitat and 16 as indicator species?^[2]. The Condor, also called ?King of the Andes? is one of the most impressive birds in the Andes due to its wide wingspan and majestic flight, but is no longer frequently seen. The most numerous bird families in the p?ramo, however, include ?the hummingbirds, furnariids, finches and flycatchers?, which tolerate the cold climate by going into ?a kind of nightly hibernation.?^[8] . Other commonly sighted birds include the turkey buzzard (Cathartes aura), eagle, hawk, falcon (Phalcoboenus carunculatus), and the rufous-fronted parakeet (Bolborhynchus ferrugineifrons)^[2].

Hummingbirds, bees and flies are all important pollinators in the p?ramo, while birds and smaller mammals such as rabbits and guinea pigs are important seed dispersers. Many of the larger mammals of the p?ramo are now rarely seen as their numbers have continuously dwindled due to being hunted excessively by man.

Human Impact and Climate Change

Humans have inhabited the p?ramo of the Andes for approximately the past 15,000 years. Deforestation has been extensive and in some cases, like the northern Andes, 90-95% of forests have been cleared. Other sites in Venezuela and Colombia show evidence that humans settled there at least 500-800 years ago and used the land for agriculture and hunting^[2].

When Europeans came to the Americas they introduced exotic plants and animals that greatly affected the land, especially the cattle which were introduced to the p?ramos in the early 18th century. By the 20th century, the growing population of settlers led to an increased demand for land, and p?ramo ecosystems suffered accordingly. As more land was needed for cattle, fire became a very popular resource for clearing land and eventually p?ramos became excessively burned and overgrazed.Both burning and grazing have had a significant impact on the vegetation, soils, species diversity, and water storage capacity of the p?ramos. In burned and disturbed sites that were studied in the Andes, the pH and phosphorus concentration in the soil are higher than in non-burned sites^[9].

Growing populations in Colombia, Venezuela, and Ecuador have forced settlements in higher elevations covering more p?ramo. Recent developments such as construction of aqueducts, drainage systems, and roads, mining, and afforestaion have been a huge additional p?ramo disturbance.

Climate change is becoming an increasingly pressing issue for p?ramo ecosystems. Increases in temperature extremes are forcing many fauna and flora species to higher grounds, and eventually they could face extinction. The flora of p?ramos is adapted to specific conditions and is thus vulnerable to even small climatic change. Climate change in the Andes is also causing glaciers in the p?ramo to disappear as well as decreasing rainfall, virtually drying up p?ramo and in turn, drying up the water supply for cities such as Quito, Ecuador and Bogot?, Colombia.

References

External links