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(Tropical) Andes: CPD Site SA26


Location:  In Eastern Cordillera, Cocuy region c. 280 km north-east and Guantiva region c. 230 km north to north-east of Bogotá. Cocuy region about latitudes 6°10'-6°45'N and longitudes 72°00'-72°25'W; Guantiva region about latitudes 6°03'-6°20'N and longitudes 72°40'-73°00'W.
Cocuy region 3060 km², with 33 km² north-south line of 22 snow capped peaks; Guantiva region 1200 km².
Cocuy region c. 500-5493 m; Guantiva region from c. 2200 m on western slope to 4270 m.
Number of vegetation types and diversity highest of Eastern Cordillera: (1) on lower eastern slope (per)humid Andean rain forests, on lower western slope broad diversity of dry xerophytic scrub and low forest types of Chicamocha Valley; (2) on upper eastern slope Espeletia stem-rosette ­ bamboo páramo, on upper western slope Espeletia stem-rosette - bunchgrass páramo; (3) extensive superpáramo best developed of Eastern Cordillera.
Highest diversity - Cocuy páramo with over 220 genera of vascular plants and highest proportion of endemism in páramos (including superpáramos) of Eastern Cordillera; endemic species also prominent in uppermost forests; threatened species.
Useful plants: 
Medicinals - e.g. widespread in regional use are "lítamo real" (Draba litamo, D. cocuyensis), "granizo" (Hedyosmum spp.), "árnica" (Senecio spp.); spices; and for construction.
Other values: 
Endemic animals, threatened species; watershed protection and water resources for villages and settlements; indigenous people; spectacular mountain scenery.
Too much sheep-grazing, occasionally combined with burning and hunting; cutting in uppermost forests, especially on western slope; upslope shift to c. 4000 m of low-income agriculture; ecotourism, when not adequately managed.
El Cocuy Natural National Park (3060 km²), with 216 km² also an Amerindian reserve.

Map 62: CPD Site SA26


The Cocuy-Guantiva region comprises large areas of Andean forests and páramos in mountain ranges on both sides of the Chicamocha River Valley in Boyacá Department. The Sierra Nevada del Cocuy is east of this deep valley and the Páramo de Guantiva west of the valley (Map 62). Included approximately are the areas between Chita/Sácama and Güicán-Las Mercedes/Bócota, and between Belén and Onzaga.

El Cocuy Natural National Park is on the boundary of Boyacá, Casanare and Arauca departments; on the north-west, Santander Department borders the region. The Sierra Nevada del Cocuy (or Güicán or Chita), between c. 6°20'-6°35'N latitudes, is the highest mountain range of the Colombian Eastern Cordillera. It is the only range in the cordillera carrying a snowcap, which extends over some 33 km following the main divide (Notestein and King 1932). Among the snow peaks are Ritacuva Blanco (5330 m), Picacho (5030 m), Puntiagudo (5200 m), El Castillo (5100 m) and "picos sin nombre" (c. 5000 m). Glacial valleys contain lakes, among which stand out for beautiful scenery e.g. La Plaza, Laguna Grande de la Sierra, Laguna Grande de Los Verdes and the Lagunillas Valley lakes (La Pintada, La Cuadrada, La Parada, La Atravesada). In the northern section of the Cocuy range appears a parallel ridge extending north, separated by the upper Ratoncito Valley, with spectacular glacial lakes and cushion bogs. Also, the impressive El Castillo peak is separated to the east from the Cocuy main ridge. This branch, which forms the boundary between Boyacá and Arauca departments, continues eastward and some 18 km east of Ritacuva Blanco constitutes the isolated snowcapped Sirará summit (5200 m).

Among the rivers in the northern sector are the Tunebo, Ratoncito, Rudiván (or Cubugón), Orozco and Derrumbada. On the eastern slope the main rivers are the Cusay, Tame, Purare and Mortiñal/San Lope; on the south to south-eastern side is the upper Casanare River with the tributaries Quebrada Los Osos, Q. Maicillo and Q. El Playón. The main rivers on the western side are tributaries of the Nevado River, which flows into the Chicamocha River: Quebrada Rechiniga and the rivers Cardenillo, Corralitos and Lagunillas.

The Cocuy range consists mainly of folded sedimentary Cretaceous (Albian-Aptian) quartzitic and sandstone rocks, shales and occasional limestone inclusions (van der Hammen et al. 1981; INDERENA 1984). The western dip slope of Cocuy has quartzite. In the westernmost section appears coal of the Guaduas (Maestrichtien) and Socha (Palaeocene) formations, while north to north-west of the main range are Palaeozoic (Mid-Devonian to Permian) sedimentary rocks. On the eastern slope, siltstones and calcareous layers near Patio Bolos contain numerous plant fossils. On the eastern slope between 5000-1000 m sedimentary rock is present, consisting of conglomerates and clayey-sandy layers related to Tertiary (Oligocene to Pliocene) river deposition. At lower elevations appear alluvial terraces and cones.

Quaternary deposits as a consequence of glacial action are common in the highest parts of the Cocuy range (Kraus and van der Hammen 1960; González, van der Hammen and Flint 1965; van der Hammen et al. 1981). In the southern section of the range there are at least five (possibly six) glacial drift bodies. Drifts numbered 2-5 are of Last Glacial age. Drift 6 represents the historic Neoglacial and its terminal moraines mark the grass-páramo ­ superpáramo border, e.g. at páramo Cóncavo. Probably in the period between 45,000-25,000 BP the Cocuy glaciers expanded the most, under a relatively wet climate - down to elevations of at least 2700 m, where they were in contact with forest. At that time the narrow páramo belt contained abundant Polylepis forests. Today the Cocuy icecap and glaciers are still shrinking [see van der Hammen et al. (1981) Figs. 9-11 and the 1:80,000 scale glacio-morphological map].

The climate is very wet on the eastern slope of the Cocuy range, with yearly precipitation up to 4000-5000 mm at around 1000 m; the western slope is dry, with a yearly average (over 10 years) of c. 901 mm at El Cocuy (2749 m) and 959 mm (over 5 years) at Chita (3005 m). Permanently high atmospheric humidity and generally just one dry period (December to February) are present (van der Hammen et al. 1981). Annual average temperatures in El Cocuy NNP are in a wide spectrum, between 23.6°C and -3°C, with an average of 0°C at c. 4800 m.

The Páramo de Guantiva region (roughly 40 km × 30 km) according to local inhabitants largely encloses the land on the eastern border between Soatá, Susacón and Sátivasur, to the mountains west of Belén at the south-western edge, to the Cuchilla de San José and Onzaga at the northern border. The Páramo de Guantiva itself is some 5 km west to south-west of Susacón and reaches c. 3900 m. The mountain range north-west of Belén has a number of glacial lakes (e.g. Grande, El Alcohol, Redonda, Negra Cazadero) in deep U-shaped valleys with moraines. This section contains the highest summit, Pan de Azúcar (c. 4270 m). The mountain range runs in a north to north-east direction and is split into the south-north valleys of the Susa, Chaguasca and Chicamocha rivers.

With respect to the geology of the Belén cordillera area, Cleef (1981) referred to reddish sandstones, siltstones and conglomerates of the Girón Formation. The summit of Pan de Azúcar consists of Palaeozoic (Devonian to Permian) gneiss and granites. West and north-west of this peak are intrusive and extrusive rocks of Triassic to Jurassic age, and to the north is a narrow zone of Cretaceous sandstone.

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Sierra Nevada del Cocuy

The vegetation of the Sierra Nevada del Cocuy includes montane vegetation - mainly forest types, and tropical alpine páramo. The treeline is c. 3500-3700 m on the dry western slope, but close to 3000 m on the wet eastern slope (Cleef 1981; van der Hammen and Cleef 1986). Probably there is a strong relationship with local climate. The more upslope treeline on the western slope may be related to the ascending warm dry air masses from the deep Chicamocha Valley; the much lower treeline of the eastern slope is related to the superwet environmental conditions prevailing there. The great height of the Sierra Nevada del Cocuy apparently largely prevents exchange from the dry and wet climates to their opposite sides.

Montane forest zone
On the Cocuy western slope the following observations have been made (van der Hammen et al. 1981; Cleef, unpublished):

  • In the lower northern part near Capitanejo is dry xerophytic vegetation. From the Chicamocha Valley floor to 2800 m, most natural forest has been cleared, except for small stands on very steep slopes. Between c. 2800-3500 m Weinmannia fagaroides seems dominant in the patches of Andean forest (c. 15 m high), associated with species of Ilex, Styrax and Prunus, as well as Clusia, Clethra, Myrsine, Rhamnus, Viburnum, Vallea, Psychotria, Xylosma, Cestrum, Oreopanax and Lauraceae and Melastomataceae. Low (6-8 m) high-Andean forest of Hesperomeles lanuginosa occurs to c. 3750 m, interchanged with Polylepis quadrijuga forest to over 4000 m.
  • In addition Cleef (unpublished) noted above the town of El Cocuy (2760 m): Alnus acuminata common (3080 m) along streams; residual forests of Gynoxys, Oreopanax and Vallea stipularis (c. 3250-3350 m); solitary Polylepis quadrijuga from 3550 m upslope. At 3800 m near Alto de la Cueva, are frequent Espeletiopsis colombiana patches (from 3600 m) and a shrubby species of Lupinus. Locally in the Andean forest belt (e.g. c. 2800 m at El Claval), there are small Ludwigia peruviana-Carex acutata swamps.

The Andean rain-forest zonation on the Cocuy eastern slope according to van der Hammen et al. (1981) is as follows:

1. From 2100-2550 m, very wet sub-Andean rain forest with Weinmannia cf. pinnata as the dominant tree, associated with cyatheaceous tree ferns and species of Cecropia and Heliocarpus (to 2200 m) and Alchornea and Acalypha (to 2400 m). In addition species of Brunellia, Stylogyne, Myrsine, Eugenia, Hedyosmum, Ocotea, Ternstroemia, Guarea, Billia, Piper, Saurauia, Freziera, Psychotria and Sapium are recorded from this forest (to 40 m high) by T. van der Hammen and R. Jaramillo-Mejía (in prep.) and van der Hammen et al. (1981).

2. From c. 2550-3100 (3300) m, Weinmannia rollottii dominates the Andean rain forest (20-35 m high), associated with W. cf. pinnata and Cyatheaceae (up to 2950 m) - replaced at higher altitudes by Blechnum sp. The entire-leaved Geonoma weberbaueri and Chusquea bamboos are very conspicuous. Additional associates are species of Miconia, Clusia, Clethra, Brunellia, Myrsine, Drimys, Ternstroemia, Freziera, Geissanthus, Ocotea, Hedyosmum, Piper, Monnina and Oreopanax. In the uppermost wet forest, Escallonia myrtilloides is common.

Páramo vegetation
The zonal páramo contains a number of vegetation types, grouped into subpáramo, grass páramo and superpáramo; an asterisk (*) means the community has not been reported from elsewhere.

1. Subpáramo communities include (i) on the western slope, Myrica parvifolia shrubs on lowermost páramo Cóncavo (3700 m); *Pentacalia vaccinioides shrub-páramo with Espletiopsis jimenez-quesadae; between c. 3750-3900 m, dense *Arcytophyllum nitidum dwarf shrubs with Masdevallia coriacea and Scaphosepalum sp.; and (ii) on the eastern slope, Hypericum magniflorum (3000-3350 m) and H. lycopodioides (3250-3700 m) shrubs with Pentacalia cacaosensis; and Chusquea-Ageratina tinifolia shrubs.

2. Grass-páramo communities include Acaena cylindristachya-Plantago sericea subsp. argyrophylla herb fields (c. 4000-4100 m); only on the eastern slope, Espeletia annemariana/E. cleefii-Chusquea tessellata bamboo páramo (c. 3400-3900 m); upper páramo of *Espeletiopsis colombiana/Espeletia cleefiiCalamagrostis effusa [3900-4250 m according to Sturm and Rangel (1985), locally to 4500 m]; Stipa hans-meyeri patches on western slope, c. 4350 m on páramo Cóncavo this vegetation type had only been reported from Sabana de los Leones, Chirripó páramo, Costa Rica and from Ecuador; on the western slope (4300-4400 m), *Diplostephium rhomboidale-Pentacalia vaccinioides shrubs on terminal moraines.

3. Superpáramo communities include low shrubs of Loricaria complanata (c. 4200-4400 m) and of *Niphogeton josei (4270-4340 m); on the eastern slope at c. 4250-4300 m, *Espeletiopsis colombiana/Espeletia cleefii lower superpáramo stands with Geranium sibbaldioides; Agrostis breviculmis-Acaulimalva spp., lower superpáramo herb field; on eastern slope at c. 4350 m, Racomitrium crispulum-Valeriana plantaginea lower superpáramo vegetation; low scrub of *Pentacalia guicanensis in small patches from 4200 m upslope; Senecio niveo-aureus communities (4250 m up to snowcap); Luzula racemosa-Pernettya prostrata superpáramo moraine vegetation (from c. 4250 m up to icecap).

Noteworthy among many azonal páramo vegetation types are:

4. Aquatic and wetland communities

  • Glacial-lake bottom vegetation dominated by respectively Isoetes karstenii, I. glacialis and I. socia, among others.
  • Marsh vegetation with Carex peucophila and Werneria crassa subsp. orientalis (c. 4000-4425 m); occasionally peaty facies with the rare monotypic Floscaldasia hypsophila (shared only with Los Nevados NNP of the Central Cordillera) and/or with dense stands of stem-rosettes of Espeletia lopezii (Cleef 1981; Sturm and Rangel 1985).
  • Cushion bogs of the juncaceous Distichia muscoides (altitudinally the highest cushion bog), Plantago rigida, or the cyperaceous Oreobolus cleefii (of limited occurrence). On the western slope, reported only from páramo Cóncavo (3550-3770m), Puya aristiguietae-Sphagnum bog. In the eastern-slope bamboo páramo, Chusquea tessellata-Sphagnum bogs with Espeletia lopezii.

5. Dwarf forests

  • About 2700-3700 m, Escallonia myrtilloides dwarf forest; from 3450-4250 m, Gynoxys albivestita dwarf forest with an unknown Echeveria sp. (Crassulaceae); at 4000 m and upslope, Polylepis quadrijuga dwarf forest; from 4100-4300 m, gnarled dwarfed tree patches of Pentacalia flos-fragrans.
  • Only on western side between c. 3700-4050 m, apparently caused by frequent grazing and trampling, Aciachne acicularis cushion meadows as subseral vegetation. On the western slope up to 3800 m, Hesperomeles lanuginosa dwarf forest; from 3800-4400 m, *Diplostephium rhomboidale dwarf forest patches. Mainly on the eastern slope, Sphagnum-Diplostephium revolutum dwarf forest.

Páramo de Guantiva

The Andean forest vegetation in the Guantiva Páramo region also has been studied by van der Hammen and Jaramillo-Mejía (in prep.). Grabandt (1980) provided a summary, noting that oak (Quercus) forests are directly in contact with the páramo in the Onzaga area (van der Hammen and González 1965; van der Hammen 1962).

The páramo belt includes subpáramo thickets, dwarf shrubs and grass páramo; on the western and north-western Magdalena Valley slopes, Chusquea tessellata bamboo páramo; and on drier eastern slopes, Calamagrostis bunchgrass páramo. The summit area of Pan de Azúcar at c. 4270 m just reaches into the lowermost superpáramo belt.

1. In the subpáramo, among other types appear shrub-páramo with Espeletia muiska, dwarf shrubs of Arcytophyllum nitidum and Sporobolus lasiophyllus.

2. From the grass páramo have been reported different types of Chusquea tessellata bamboo páramo, Espeletiopsis muiska/Espeletia boyacensis-Calamagrostis effusa bunchgrass páramo with Oreobolus sp. and Castratella piloselloides, Acaena cylindristachya-Plantago sericea herb field and Espeletiopsis guacharaco/Espeletia azucarina-Calamagrostis effusa bunchgrass upper páramo.

3. The lower superpáramo at Pan de Azúcar consists of fragments of Loricaria complanata shrubs and scree vegetation of endemic Poa and Halenia sp., species of Arenaria and Cerastium, Montia meridensis, Rhacocarpus purpurascens and Racomitrium crispulum, giving evidence of the upper condensation zone, and small valley floors covered by Senecio canescens. On the opposite Morro Verde probably appear stands of Lupinus alopecuroides (Guillermo Merchán, pers. comm.).

Extrazonal and azonal vegetation types

Important are isolated patches of Polylepis quadrijuga dwarf forest in the páramo, which are common north-west of Belén. In the same area are a number of crystalline lakes and Plantago rigida cushion bogs on valley floors. Peaty valley floors near the treeline contain extensive Sphagnum peat-bog swamps with endemic species such as Espeletia nemenkenii or Espeletia arbelaeziana.

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Sierra Nevada del Cocuy

In the páramo flora of the Sierra Nevada del Cocuy have been reported (thus far) c. 220 vascular plant genera (cf. Cleef 1983). In the superpáramo have been recorded 110 vascular plant species (in c. 55 genera), c. 20 of which are endemic, including the genus Floscaldasia (Compositae) - whereas in the superpáramo of Sumapaz (southward) are three endemic species and in Almorzadero (to the north-west) just two endemics. About two-thirds of the superpáramo species are shared with the Sumapaz superpáramo (see CPD Site SA27). The first sample collections of the region were made in 1939 in the upper Lagunillas Valley.

Endemic species of the Cocuy mountain range (cf. Cuatrecasas and Cleef 1978; Al-Shehbaz 1989; Rangel-Ch. and Santana-C. 1989) include Aragoa hammenii, Draba cocuyensis, D. hammenii, D. litamo, Diplostephium rhomboidale, Hypericum lycopodioides, H. papillosum, Niphogeton josei, Salvia nubigena, Oritrophium cocuyense, Paepalanthus lodiculoides var. floccosus, Pentacalia guicanensis, P. cleefii, Senecio cocuyanus, S. adglacialis, S. pasqui-andinus, S. tergolanatus, S. virido-albus (also at Guantiva), Espeletia cleefii, Espeletiopsis jimenez-quesadae, Puya cleefii and Acaulimalva species.

From the headwaters of the Casanare River on the southern slope of the Cocuy have been reported endemics such as Aragoa dugandii, Castratella rosea, Pentacalia cacaosensis and Espeletia curialensis.

The Sierra Nevada del Cocuy is the southernmost outpost known (so far) of e.g. Carex peucophila, Ilex tamana and Libanothamnus tamanus. Highly interesting was the 1977 find of the aquatic liverwort Herbertus oblongifolius predominant in a small lake at 4060 m in the headwaters of Quebrada El Amarillal. This rare species was known only from the Itatiaia massif near Rio de Janeiro, Brazil.

Páramo de Guantiva

Endemic species of the Guantiva Páramo include Hypericum sabiniforme, Niphogeton fruticosa, Espeletia arbelaezii, E. azucarina, E. brachyaxiantha, E. discoidea, E. nemenkenii and Espeletiopsis muiska. Also different oak (Quercus) species have been described from the Onzaga forests.

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Useful plants

A number of plants are gathered - e.g. for medicinals, spices, construction - and frequently sold by inhabitants of the region, as well as by indigenous tribes (e.g. Tunebo) on the lower Llanos side of the eastern slope of the Cocuy range. "Lítamo real" (Draba litamo) is the medicinal plant most known from the region; also well known are "chichoria" (Hypochoeris sessiliflora), "verdolaga" (Peperomia sp.), "granizo" (Hedyosmum spp.) and "árnica" (Senecio spp.). "Palo colorado" (Polylepis quadrijuga) is not only used for fuelwood and fences but also construction of walls or dwellings, together with Espeletia lopezii stem-rosettes (trunks).

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Social and environmental values

El Cocuy NNP harbours a great diversity of large mammals, such as four primate species, all the cats known from Colombia, the tapir Tapirus terrestris and two species of deer. There also are many species of birds in the region (Borrero 1955; Olivares 1973; INDERENA 1984).

The Sierra Nevada del Cocuy region was originally inhabited by indigenous groups of Lache (now extinct) and Tunebo. People living at these high altitudes survive in marginal conditions, because of low income and the harsh environment. Also the Tunebo Amerindians who cross the cordillera are very poor and in poor health. Development of the Cocuy NNP area must be in accordance with the needs, and enhance possibilities and the quality of life, of the local peasants. Presently the conditions in the Cocuy NNP area are unsafe and do not allow studies towards this goal. Peasants living in the Guantiva region are also poor, but probably in better conditions than those living in the Cocuy.

Economic assessment

The Sierra Nevada del Cocuy apparently has the largest sheep population of all the páramos of the Colombian Eastern Cordillera, where cattle-ranging usually prevails. Herds of sheep are mostly grazed on the dry western slope of the Cocuy, and their influence on the páramo vegetation is considerable.

The same region seems also to be the only place in the Eastern Cordillera where potato agriculture with some onions and other Andean tubers is possible up to 4000 m. The potato production for the outside market (Cúcuta near Venezuela) is important.

However, most important for the region may be the permanent water supply from the high snowcapped Cocuy mountains, not only for the eastern slope Orinoco drainage, but especially for the dry and almost deforested lands on the western slope towards the deep Chicamocha Valley. Coal and salt (near Salinas, headwaters of the Casanare River) are of minor importance.

The Guantiva Páramo is also an important regional watershed area, which is used as well for extensive ranging of cattle, its wood supply for construction, fences and fuel, for the gathering of medicinal and spice plants, and for hunting.

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The relatively dense human population on the western side has had a very strong impact on the residual montane forests, whereas the eastern slope rain forests have so far remained largely intact. The shift upslope to c. 4000 m of low-income agriculture has converted habitats. Ecotourism, when not adequately managed, can be destructive.

Major threats are uncontrolled activities such as burning, grazing, hunting and especially tree cutting in the uppermost forest belt and forest patches, since the trees grow very slowly because of prevailing low temperatures.

Espeletia nemenkenii is a highly endangered species endemic in the Belén páramos; its habitat of treeline Sphagnum-Blechnum bogs has almost been drained and disappeared. The same situation may apply for the endangered Espeletia arbelaeziana.

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El Cocuy Natural National Park (3060 km²) was established in 1977; in 1974/1979, 216 km² were set aside in the Tunebos Indigenous Reserve (INDERENA 1984). The NNP could use a conservation and management plan, taking into special account the needs and priorities of the local poor inhabitants.

The Guantiva Páramo region does not so far have conservation status, through which a conservation and management plan could establish wise-use practices and conservation of representative areas of páramo and Andean forest. It is strongly recommended that the páramo-forest border be preserved along the western slope of the Guantiva region down to at least 2200-2500 m, in order to allow for the exchange of biotic elements.

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Map 62. Sierra Nevada del Cocuy-Guantiva, Colombia (CPD Site SA26)


Al-Shehbaz, I.A. (1989). New or noteworthy Draba (Brassicaceae) from South America. J. Arnold Arboretum 70: 427-437.

Borrero, J.I. (1955). Avifauna de la región de Soatá, departamento de Boyacá, Colombia. Caldasia 7: 52-86.

Cleef, A.M. (1978). Characteristics of neotropical páramo vegetation and its subantarctic relations between the southern temperate zone and tropical mountains. Erdwissenschaftliche Forschung (Wiesbaden) 11: 365-390.

Cleef, A.M. (1981). The vegetation of the páramos of the Colombian Cordillera Oriental. Dissert. Bot. 61. J. Cramer, Vaduz. 321 pp.

Cleef, A.M. (1983). Fitogeografía y composición de la flora vascular de los páramos de la Cordillera Oriental colombiana (Estudio comparativo con otras altas montañas del trópico). Rev. Acad. Colomb. Cienc. Exactas Fís. Nat. 15: 23- 29.

Cuatrecasas, J. and Cleef, A.M. (1978). Una nueva Crucifera de la Sierra Nevada del Cocuy (Colombia). Caldasia 12: 145-158.

Gonzáles, E., van der Hammen, T. and Flint, R.F. (1965). Late Quaternary glacial and vegetational sequence in Valle de Lagunillas, Sierra Nevada del Cocuy, Colombia. Leidse Geologische Mededelingen 32: 157-182.

Grabandt, R.A.J. (1980). Pollen rain in relation to arboreal vegetation in the Cordilleria Oriental. Rev. Palaeobot. Palynol. 29: 65-147.

INDERENA (1984). Colombia parques nacionales. Instituto Nacional de los Recursos Naturales Renovables y del Medio Ambiente (INDERENA), Bogotá. 263 pp.

Kraus, E. and van der Hammen, T. (1960). Las expediciones de glaciología del A.G.I. a las Serras Nevadas de Santa Marta y El Cocuy. Comite Nac. del Año Geofísico, Instituto Geográfico Agustín Codazzi, Colombia. 9 pp.

Notestein, F.B. and King, R.E. (1932). The Sierra Nevada del Cocuy. Geogr. Review 22: 423-430.

Olivares, A. (1973). Aves de la Sierra Nevada del Cocuy, Colombia. Rev. Acad. Colomb. Cienc. Exactas Fís. Nat. 14: 39- 48.

Rangel-Ch., J.O. and Santana-C., E. (1989). Estudios en Draba (Cruciferae) de Colombia. I. Cuatro especies nuevas de la Cordillera Oriental. Rev. Acad. Colomb. Cienc. Exactas Fís. Nat. 17: 347-355.

Sturm, H. and Rangel-Ch., J.O. (1985). Ecología de los páramos andinos: una visión preliminar integrada. Biblioteca J.J. Triana No. 9. Instituto de Ciencias Naturales, Museo de Historia Natural, Universidad Nacional de Colombia, Bogotá. 292 pp.

van der Hammen, T. (1962). Palinología de la región de Laguna de los Bobos. Historia de su clima, vegetación y agricultura durante los ultimos 5000 años. Rev. Acad. Colomb. Cienc. Exactas Fís. Nat. 11: 359-361.

van der Hammen, T. and Cleef, A.M. (1986). Development of the high Andean páramo flora and vegetation. In Vuilleumier, F. and Monasterio, M. (eds), High altitude tropical biogeography. Oxford University Press, New York. Pp. 153-201.

van der Hammen, T. and González, E. (1965). A Late-Glacial and Holocene pollen diagram from Ciénaga del Visitador (Dept. Boyacá, Colombia). Leidse Geologische Mededelingen 32: 193-201.

van der Hammen, T., Barelds, J., de Jong, H. and de Veer, A.A. (1981). Glacial sequence and environmental history in the Sierra Nevada del Cocuy (Colombia). Palaeogeography, Palaeoclimatogy, Palaeoecology 32: 247-340.


This Data Sheet was written by Dr Antoine M. Cleef (University of Amsterdam, Hugo de Vries-Laboratorium, Kruislaan 318, 1098 SM Amsterdam, The Netherlands).

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