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The Santa Marta montane forests (NT0159) is an ecoregion in the Sierra Nevada de Santa Marta, a massif on the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
coast of northern Colombia. The ecoregion covers altitudes from near sea level up to around , where it gives way to Santa Marta páramo. The isolation of the massif and the range of elevations and climates has resulted in a wide variety of species including many endemics. The lower levels contained tropical rainforest, which has largely been cleared. Higher up, this gives way to cloud forest. Much of this has also been cleared for coffee plantations, pasture for sheep and cattle, and farming.


Geography


Location

The ecoregion covers the slopes of the Sierra Nevada de Santa Marta in the north of Colombia, with an area of . The range rises to snow-covered peaks only from the Caribbean sea. The ecoregion is almost entirely surrounded by the
Sinú Valley dry forests The Sinú Valley dry forests (NT0229) is an ecoregion in the north of Colombia. Geography Location The Sinú Valley is an area of . located within the zone of parallel, north-northeast trending hills that lies between the low-point Magdalena ...
ecoregion. To the northeast and northwest it transitions directly into
Guajira–Barranquilla xeric scrub The Guajira–Barranquilla xeric scrub is a xeric shrubland ecoregion in Colombia, Venezuela, and the ABC Islands (Leeward Antilles), covering an estimated area of . Rainfall varies from , and the median temperature is . Location The ecoregio ...
. In the higher elevations of the mountains the ecoregion gives way to Santa Marta páramo.


Terrain

The triangular Sierra Nevada de Santa Marta massif covers and rises to an elevation of , with peaks that are permanently capped in snow. The northern edge is parallel to the Caribbean coast. The southwestern edge descends to the dry and warm Magdalena River alluvial plain and the
Ciénaga Grande de Santa Marta The Ciénaga Grande de Santa Marta (Spanish for Large Marsh of Saint Martha) is the largest of the swampy marshes located in Colombia between the Magdalena River and the Sierra Nevada de Santa Marta. It has an area of 4280 km2 and belongs to the ...
marshes. The southeastern edge adjoins the valleys of the Cesar River and Ranchería River. Volcanic rocks from the Mesozoic (252–66 Ma) and Tertiary (65–2.58 Ma) eras include
batholith A batholith () is a large mass of intrusive igneous rock (also called plutonic rock), larger than in area, that forms from cooled magma deep in Earth's crust. Batholiths are almost always made mostly of felsic or intermediate rock types, such ...
s of granite, diorite and quartz monzonite, and there are areas of sedimentary rocks. The massif was pushed up to its present height in the Miocene (23–5.33 Ma) and
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
(126,000–11,700 years ago). Glacial lakes at elevations over are the sources of the main rivers that originate in the massif.


Climate

Average air temperatures range from in the plains to or less in the peaks. Rain is carried by the northeast trade winds, with wet seasons from September to December and from May to July. The north side receives the most precipitation, up to annually at above sea level, with less rain higher up and to the south of the massif. At a sample location at the Köppen climate classification is Cwb: warm temperate, winter dry, warm summer. Mean temperatures range from in January to in June. Total annual precipitation is about . Monthly precipitation at the sample location ranges from in February to in October.


Ecology

The ecoregion is in the neotropical realm, in the tropical and subtropical moist broadleaf forests biome. The isolated location surrounded by dry forests has allowed many endemic species of flora and fauna to develop. It is seen as a biological and geographic island, distinct from the Andes. However, there are similarities to the
Serranía del Perijá The Serranía del Perijá, Cordillera de Perijá or Sierra de Perijá is a mountain range, an extension of the eastern Andean branch ( Cordillera Oriental), in northern South America, between Colombia and Venezuela, ending further north in the ...
, to the east, which is normally included in the Cordillera Oriental montane forests. 76% of bird species in Perijá are shared with the Sierra Nevada de Santa Marta. 80% of Perijá's butterflies in the '' Ithomiinae'' and '' Heliconiinae'' families is shared with the Sierra Nevada, but only 40% with the eastern slope of the Cordillera Oriental further to the south. The ecoregion is part of the Northern Andean Montane Forests global ecoregion, which includes the Magdalena Valley montane forests, Venezuelan Andes montane forests, Northwestern Andean montane forests, Cauca Valley montane forests, Cordillera Oriental montane forests, Santa Marta montane forests and Eastern Cordillera Real montane forests terrestrial ecoregions.


Flora

There are warm, wet forests on the north of the massif and part of the west, isolated from similar forests elsewhere. These tropical rain forests at altitudes up to have trees that may reach in height depending on rainfall. The understory contains tree ferns and grasses with large leaves. There are some endemic species, but relatively low diversity compared to other neotropical ecoregions. These forests are considered to be a Pleistocene refuge. There are few species in common with the cloud forests higher up, but there are similarities to wet forests elsewhere in Colombia, in Panama and Venezuela, and at a family level to forests in the Amazon. On the drier western side forests at elevations from above sea level have large trees including '' Poulsenia armata'', red ucuuba (''
Virola sebifera ''Virola sebifera'' is a species of tree in the family Myristicaceae, from North and South America.Markus Wiesenauer, Suzann Kirschner-Brouns: Homöopathie - Das große Handbuch, Gräfe & Unzer Verlag, 2007, Description ''V. sebifera'' is a t ...
''), '' Pterygota colombiana'', muskwood ('' Guarea guidonia''), Panama rubber tree ('' Castilla elastica''), '' Ficus macrosyce'', avocado (''Persea americana'') and Andean royal palm ('' Dictyocaryum lamarckianum''). Above the trees and palms are smaller, and include '' Zygia longifolia'', forest acai ('' Euterpe precatoria''), Súrtuba (''
Geonoma interrupta ''Geonoma'' is a genus of small to medium-sized palms native to the forest understorey of tropical Central and South America. This palm genus is one of the largest in the Neotropics. Its 64 species are distributed from Mexico and Haiti ...
'') and spiny treefern (''
Cyathea pungens ''Cyathea'' is a genus of tree ferns, the type genus of the fern order Cyatheales. The genus name ''Cyathea'' is derived from the Greek ''kyatheion'', meaning "little cup", and refers to the cup-shaped sori on the underside of the fronds. Des ...
''). The forests include vascular
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s such as '' Vriesea elata'' and droophead tufted airplant (''
Guzmania lingulata ''Guzmania lingulata'', the droophead tufted airplant or scarlet star, is a species of flowering plant in the family (biology), family Bromeliaceae, subfamily Tillandsioideae. This evergreen epiphytic perennial plant, perennial is native plant, n ...
''), understory plants with large leaves such as Rattlesnake Plant ('' Calathea insignis'') and dumb cane ('' Dieffenbachia longispatha''), and mosses and liverworts such as ''
Octoblepharum albidum ''Octoblepharum'' is a genus of haplolepideous mosses (Dicranidae) in the monotypic family Octoblepharaceae . The genus ''Octoblepharum'' was previously placed in family Calymperaceae. Species The genus contains the following species: *''Octo ...
'' and '' Leucomium strumosum''. There are cloud forests above , with vegetation related to the forests of the tropical Andes and the higher elevations in the Caribbean, which contain many endemic species. In some areas the cloud forest dips down to lower levels. The trees at elevations up to of altitude form a canopy of in height. The flowering plant families with the largest numbers of genera are Asteraceae, Orchidaceae,
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
and Rubiaceae. The genera ''
Monochaetum ''Monochaetum'' is a neotropical genus of shrubs and subshrubs with about 54 species. It occurs in warm temperate to tropical montane habitats from Mexico and Central America to the South American Andes of Colombia, Venezuela, Ecuador and Peru ...
'' and '' Tillandsia'' each have five endemic species. The family Melastomataceae has 13 endemic species. Andean royal palms (''Dictyocarium lamarckianum'') emerge from the cloud forest canopy, and dominate wet and sloping areas. They are common from , and the wax palm (''
Ceroxylon ceriferum ''Ceroxylon ceriferum'', also known as the Sacred wax palm is a species of ''Ceroxylon ''Ceroxylon'' is a genus of flowering plants in the family Arecaceae, native to the Andes in Venezuela, Colombia, Ecuador, Peru, and Bolivia, known as Ande ...
'') is common from . Other cloud forest tree species include palo azul ('' Calatola costaricensis''), ''
Cavendishia callista ''Cavendishia'' is a genus of about 100 species of woody perennial plants, many of which are epiphytic. The genus is native to tropical South America and Central America Central America ( es, América Central or ) is a subregion of the ...
'', '' Graffenrieda santamartensis'', '' Gustavia speciosa'' and ''
Tovomita weddelliana ''Tovomita'' is a genus of flowering plants in the family Clusiaceae. They are noted for having white-yellow latex and containing xanthones. The genus is distributed in the tropical Americas, with many occurring in Venezuela. Most are native to t ...
''. The understory includes tree ferns, palms, prop-root plants, vascular epiphytes and woody
liana A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...
s. There are many '' Bryophytes'' such as the thallose liverwort '' Dumortiera hirsuta'' and the moss '' Phyllogonium fulgens''. Cloud forests from high are found in areas with heavy fog and mist between elevations of , although there is less rainfall than on the lower slopes. Typical tree species are wax palm (''
Ceroxylon ceriferum ''Ceroxylon ceriferum'', also known as the Sacred wax palm is a species of ''Ceroxylon ''Ceroxylon'' is a genus of flowering plants in the family Arecaceae, native to the Andes in Venezuela, Colombia, Ecuador, Peru, and Bolivia, known as Ande ...
''), ''
Chaetolepis santamartensis ''Chaetolepis'' is a genus of flowering plants belonging to the family Melastomataceae. Its native range is Costa Rica to Guyana. Species: *''Chaetolepis alpina'' *''Chaetolepis anisandra'' *''Chaetolepis cufodontisii'' *''Chaetolepis lin ...
'', '' Chusquea tuberculosa'', ''
Hesperomeles ferruginea ''Hesperomeles ferruginea'' is an evergreen shrub or tree in the family Rosaceae, native to montane forests on the Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming ...
'', ''
Monochaetum uberrimum ''Monochaetum'' is a neotropical genus of shrubs and subshrubs with about 54 species. It occurs in warm temperate to tropical montane habitats from Mexico and Central America to the South American Andes of Colombia, Venezuela, Ecuador and ...
'', '' Myrcianthes myrsinoides'', ''
Myrsine coriacea ''Myrsine'' is a genus of flowering plants in the family Primulaceae. It was formerly placed in the family Myrsinaceae before this was merged into the Primulaceae. It is found nearly worldwide, primarily in tropical and subtropical areas. It cont ...
'', '' Paragynoxys undatifolia'', Pino de Pasto (''
Podocarpus oleifolius ''Podocarpus oleifolius'' is a species of conifer in the family Podocarpaceae. It is found in Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Panama, Peru, and Venezuela. ''Podocarpus oleifolius'' contains one sub ...
'') and '' Weinmannia pinnata''. Bryophytes and lichens include '' Anastrophyllum auritum'', Musgo ('' Campylopus benedictii''), bonfire moss (''
Funaria hygrometrica ''Funaria hygrometrica'', the bonfire moss or common cord-moss, is a type of water moss which grows on shady, moist soil. It can also be found on moist walls and the crevices of rocks and places where recent fires have taken place. It has been re ...
'') and '' Hypotrachyna gigas''.


Fauna

Mammal species include the
Central American agouti The Central American agouti (''Dasyprocta punctata'') is a species of agouti from the family Dasyproctidae. The main portion of its range is from Chiapas and the Yucatan Peninsula (southern Mexico), through Central America, to northwestern Ecuad ...
(''Dasyprocta punctata''),
jaguar The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus '' Panthera'' native to the Americas. With a body length of up to and a weight of up to , it is the largest cat species in the Americas and the th ...
(''Panthera onca''), lowland paca (Cuniculus paca).
neotropical otter The Neotropical otter or Neotropical river otter (''Lontra longicaudis'') is an otter species found in Mexico, Central America, South America, and the island of Trinidad. It is physically similar to the northern and southern river otter, which o ...
(''Lontra longicaudis''),
ocelot The ocelot (''Leopardus pardalis'') is a medium-sized spotted wild cat that reaches at the shoulders and weighs between on average. It was first described by Carl Linnaeus in 1758. Two subspecies are recognized. It is native to the southwes ...
(''Leopardus pardalis''), South American tapir (''Tapirus terrestris''), Venezuelan red howler (''Alouatta seniculus'') and white-lipped peccary (''Tayassu pecari''). Endemic mammals include the red-tailed squirrel (''Sciurus granatensis''), Tomes's rice rat (''
Nephelomys albigularis ''Nephelomys albigularis'', also known as the white-throated oryzomys or Tomes's rice rat, is a species of rodent in the genus '' Nephelomys'' of family Cricetidae. Described in 1860, it was the first ''Nephelomys'' species to be discovered. It w ...
'') and
unicolored Oldfield mouse The unicolored Oldfield mouse (''Thomasomys monochromos'') is a species of rodent in the family Cricetidae. It is found only in Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular r ...
(''Thomasomys monochromos''). Endangered mammals include the red-crested tree-rat (''Santamartamys rufodorsalis''). Bird species include
bay-headed tanager The bay-headed tanager (''Tangara gyrola'') is a medium-sized passerine bird. This tanager is a resident breeder in Costa Rica, Panama, South America south to Ecuador, Bolivia and north-western Brazil, and on Trinidad. Taxonomy The bay-headed ta ...
(''Tangara gyrola''),
black-chested jay The black-chested jay (''Cyanocorax affinis'') is a species of bird in the family Corvidae. Measuring long, this jay is easily recognized from its distinctive facial pattern and yellow eye. The head, face, and chest are mostly black with violet ...
(''Cyanocorax affinis''),
blue-billed curassow The blue-billed curassow (''Crax alberti'') is a species of bird in the family Cracidae, the chachalacas, Guan (bird), guans, and curassows. It is endemic to Colombia. Taxonomy and systematics The blue-billed curassow is monotypic. However, i ...
(''Crax alberti''),
coppery emerald The coppery emerald (''Chlorostilbon russatus'') is a species of hummingbird in the "emeralds", tribe Trochilini of subfamily Trochilinae. It is found in Colombia and Venezuela.HBW and BirdLife International (2020) ''Handbook of the Birds of ...
(''Chlorostilbon russatus''),
crested guan The crested guan (''Penelope purpurascens'') is a member of an ancient group of birds of the family Cracidae, which are related to the Australasian megapodes or mound builders (Megapodiidae). It is found in the Neotropics, in lowlands forest ...
(''Penelope purpurascens''),
keel-billed toucan The keel-billed toucan (''Ramphastos sulfuratus''), also known as sulfur-breasted toucan or rainbow-billed toucan, is a colorful Latin American member of the toucan family. It is the national bird of Belize. The species is found in tropical jungl ...
(''Ramphastos sulfuratus''),
Santa Marta blossomcrown The Santa Marta blossomcrown (''Anthocephala floriceps'') is a Vulnerable species of hummingbird in the "emeralds", tribe Trochilini of subfamily Trochilinae. It endemic to the Sierra Nevada de Santa Marta of Colombia.HBW and BirdLife Internati ...
(''Anthocephala floriceps''),
scarlet-fronted parakeet The scarlet-fronted parakeet (''Psittacara wagleri''), also known as the scarlet-fronted conure'','' red-fronted conure or Wagler's conure, is a long-tailed South American species of parrot. It is found in Colombia and Venezuela. Its natural habi ...
(''Psittacara wagleri'') and white-tipped quetzal (''Pharomachrus fulgidus''). Birds with restricted ranges include black-backed thornbill (''Ramphomicron dorsale''),
green-bearded helmetcrest The green-bearded helmetcrest (''Oxypogon guerinii'') is a species of hummingbird in the "coquettes", tribe Lesbiini of subfamily Lesbiinae. It is endemic to Colombia.HBW and BirdLife International (2020) ''Handbook of the Birds of the World ...
(''Oxypogon guerinii''),
montane woodcreeper The montane woodcreeper (''Lepidocolaptes lacrymiger'') is a perching bird species in the subfamily Dendrocolaptinae of the ovenbird family ( Furnariidae). It is found in Bolivia, Colombia, Ecuador, Peru, and Venezuela. Its natural habitat is su ...
(''Lepidocolaptes lacrymiger''),
mountain velvetbreast The mountain velvetbreast (''Lafresnaya lafresnayi'') is a species of hummingbird in the "brilliants", tribe Heliantheini in subfamily Lesbiinae. It is found in Colombia, Ecuador, Peru, and Venezuela.HBW and BirdLife International (2020) ''Handb ...
(''Lafresnaya lafresnayi''), Santa Marta parakeet (''Pyrrhura viridicata''),
strong-billed woodcreeper The strong-billed woodcreeper (''Xiphocolaptes promeropirhynchus'') is a species of bird in the Dendrocolaptinae subfamily. It is one of the largest woodcreepers and the largest furnariids, though the slender long-billed woodcreeper is longer an ...
(''Xiphocolaptes promeropirhynchus''), white-tipped quetzal (''Pharomachrus fulgidus'') and yellow-crowned whitestart (''Myioborus flavivertex''). Endemic birds include the Santa Marta parakeet (''Pyrrhura viridicata'') and
white-tailed starfrontlet The white-tailed starfrontlet (''Coeligena phalerata'') is a species of hummingbird in the "brilliants", tribe Heliantheini in subfamily Lesbiinae. It is endemic to the Sierra Nevada de Santa Marta of northeastern Colombia.HBW and BirdLife Int ...
(Coeligena phalerata). Endangered birds include the black-and-chestnut eagle (''Spizaetus isidori''), black-backed thornbill (''Ramphomicron dorsale''),
blue-billed curassow The blue-billed curassow (''Crax alberti'') is a species of bird in the family Cracidae, the chachalacas, Guan (bird), guans, and curassows. It is endemic to Colombia. Taxonomy and systematics The blue-billed curassow is monotypic. However, i ...
(''Crax alberti''), Santa Marta bush tyrant (''Myiotheretes pernix''), Santa Marta parakeet (''Pyrrhura viridicata''),
Santa Marta sabrewing The Santa Marta sabrewing (''Campylopterus phainopeplus'') is a Critically Endangered species of hummingbird in the "emeralds", tribe Trochilini of subfamily Trochilinae. It is endemic to the Guatapurí dry enclave on the eastern side of the S ...
(''Campylopterus phainopeplus'') and Santa Marta wren (''Troglodytes monticola''). There are many species of reptiles and amphibians. Endemic sauria include the Colombian Clawed Gecko (''Pseudogonatodes furvus''), '' Ptychoglossus romaleos'', '' Proctoporus specularis'' and Santa Marta Anole (''Anolis santamartae''), Amphibians include several ''
Pristimantis ''Pristimantis'' is a very large genus of frogs distributed in the southern Caribbean islands ( Lesser Antilles) and in Central and South America from Honduras to northern Argentina and southern Brazil. With 596 described species (as of October ...
'' species, mostly at the higher elevations, including Cebolleta Robber Frog (''
Pristimantis tayrona ''Tachiramantis tayrona'' is a species of frog in the family Strabomantidae. It is endemic to the north-western Sierra Nevada de Santa Marta, Colombia. The specific name ''tayrona'' refers to the pre-Columbian Tairona culture. Lynch and Ru� ...
''), Cristina's Robber Frog ('' Pristimantis cristinae''), Ruthven's Robber Frog ('' Pristimantis ruthveni'') and Santa Marta robber frog ('' Pristimantis sanctaemartae''). The only endemic amphibian genus of the Sierra Nevada includes Walker's Sierra frog (''
Geobatrachus walkeri ''Geobatrachus walkeri'', commonly known as Walker's Sierra frog, is a species of frog in the family Craugastoridae. It is the only species in the monotypic genus ''Geobatrachus''. It is endemic to Sierra Nevada de Santa Marta, Colombia. Its nat ...
''). Endangered amphibians include '' Atelopus arsyecue'',
Guajira stubfoot toad The Guajira stubfoot toad or Carrikeri harlequin frog (''Atelopus carrikeri'') is a species of toad in the family Bufonidae. It is about 5 cm (2.0 in) long and typically black, though some populations have orange coloration. This specie ...
(''Atelopus carrikeri''), Harlequin frog (''
Atelopus laetissimus ''Atelopus laetissimus'' is a species of toad in the family Bufonidae. It is endemic to Colombia and only known from the area of its type locality in the northwestern part of the Sierra Nevada de Santa Marta, in the Magdalena Department. ...
''), Nahumae stubfoot toad (''
Atelopus nahumae ''Atelopus nahumae'' (San Lorenzo harlequin frog) is a species of toad in the family Bufonidae. It is endemic to Colombia and only known from humid montane forest of Sierra Nevada de Santa Marta in the Magdalena, La Guajira, and Cesar Depa ...
''), Walker's stubfoot toad ('' Atelopus walkeri''), Santa Marta poison arrow frog ('' Colostethus ruthveni''), Boulenger's backpack frog ('' Cryptobatrachus boulengeri''), Walker's sierra frog (''
Geobatrachus walkeri ''Geobatrachus walkeri'', commonly known as Walker's Sierra frog, is a species of frog in the family Craugastoridae. It is the only species in the monotypic genus ''Geobatrachus''. It is endemic to Sierra Nevada de Santa Marta, Colombia. Its nat ...
''), Ground robber frog ('' Pristimantis insignitus'') and Ruthven's robber frog ('' Pristimantis ruthveni'').


Status

The World Wide Fund for Nature (WWF) gives the region the status of "Vulnerable". The massif was settled in the Pre-Columbian era, and traces of these communities remain. Areas of the ecoregion have been sporadically occupied since the end of the 19th century. From 70% to 80% of the original forest has been cleared in the last 50 years. The wet forest of the lower levels has been reduced to thinned-out fragments by settlements, marijuana and coca cultivation, logging, extraction of firewood and conversion to pasture. Deforestation causes erosion, mud slides in the rainy seasons and silting of the rivers. In the late 19th century large belts of forest at the middle levels were converted to coffee cultivation. Large areas of forest have also been cleared to create pasture, kept clear by annual burns. Higher up, the Ancho and Frío river basins and other parts of the cloud forest have been modified to make space for sheep and cattle rearing, farming of potatoes and fruit, and extraction of wood. Forests clearance has threatened the populations of large animals, which are also hunted for food, for their skins, or as perceived threats to cattle. Roads opened to the higher areas have led to looting of pre-Columbian sites, hunting and extraction of firewood. A 2006 book reported that of the Santa Marta montane forests 33.4% had been transformed by human activity. Parts of the ecoregion are theoretically protected by the
Sierra Nevada de Santa Marta National Park The Sierra Nevada de Santa Marta (English: ''Snow-Covered Mountain Range of Saint Martha'') is an isolated mountain range in northern Colombia, separate from the Andes range that runs through the north of the country. Reaching an elevation of ...
,
Tayrona National Natural Park The Tayrona National Natural Park ( es, Parque Nacional Natural Tayrona) is a protected area in the Colombian northern Caribbean region and within the jurisdiction of the city of Santa Marta, from the city centre. The park presents a biodi ...
and Sierra Nevada de Santa Marta Biosphere Reserve, but these areas were still being cleared in 1998. The indigenous reserves of the
Cogui The Kogi ( ), or Cogui, or Kágaba, meaning "jaguar" in the Kogi language, are an indigenous group that resides in the Sierra Nevada de Santa Marta mountains in northern Colombia. Their culture has continued since the Pre-Columbian era. Langua ...
, Arsario and Arhuaco people overlap the Sierra Nevada de Santa Marta National Park.


Notes


Sources

* * * * * * {{authority control Ecoregions of Colombia Sierra Nevada de Santa Marta * Trees of Colombia Neotropical tropical and subtropical moist broadleaf forests Andean forests Montane forests