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COLOMBIAN PACIFIC COAST REGION (CHOCO)
The Colombian Pacific Coast region (Chocó sensu lato) is the stretch of land mainly between the Pacific Ocean and Cordillera Occidental of the Andes, from west of the mouth of the Atrato River near Panama to the Mataje River bordering north-western Ecuador (Rangel-Ch. and Aguilar 1993). The Colombian departments represented are primarily Chocó, Valle del Cauca, Cauca and Nariño.
The region is composed of lowlands, with elevations rarely exceeding 600 m, and highlands, which are usually above 600 m. There are three major landforms: (1) plains of recent alluvium; (2) low hills formed by relatively recent stream dissection of Tertiary and Pleistocene sediments; and (3) complex mountainous areas of Mesozoic rocks (West 1957).
Five subregional zones may be distinguished (Map 74) (Rangel-Ch. et al. 1994):
1. Northern coast, with areas of the Serranía del Darién and Urabá;
2. Seaboard generally up to 500 m elevation, along the Pacific coast;
3. Central strip, which includes northern wet forest, central pluvial forest and the San Juan River area;
4. So-called highlands of El Carmen del Atrato, and the San José del Palmar area; and
5. Localities of Pacific slope from 500-1000 m in the departments of Valle del Cauca, Cauca and Nariño.
The region has a complex of vast river basins, the most important being the Atrato and San Juan, and in the south the (San Juan de) Micay, Patía and Mira. Alluvial plains extend along the valleys of the Atrato and the San Juan rivers, which are separated by a narrow watershed divide (c. 100 m in elevation) called the Isthmus of San Pablo, and empty respectively into the Caribbean Sea (Gulf of Urabá) and the Pacific Ocean. Since these two rivers parallel the Cordillera Occidental and receive numerous Andean subsidiaries, they are swollen torrents for much of the year. The waters of many of the rivers to the Pacific carve deep gorges through the mountain flanks, forming numerous spectacular rapids and waterfalls. The San Juan River discharges the most water to the Pacific of all the South American rivers (Barnes 1993), and the Atrato is the second largest river in South America in terms of the volume of water.
Swamps and shallow lakes ("ciénagas") cover the greater part of the lower half of the Atrato Basin. In contrast, along much of the San Juan River, swamps are absent or confined. Bordering the Atrato and San Juan rivers are wide belts of hills c. 100-300 m high (West 1957; Forero 1982). The widest hilly tract is to the west between the floodplains and the Serranía de Baudó which edges the Pacific. The hills present several landforms; terraces often occur, and are sites for settlement and agriculture (West 1957; Whitten 1974).
Mountainous areas include: in the east, the lower slopes of the Andean Cordillera Occidental, and large somewhat isolated massifs such as Cerro Torrá (2700 m) in the San José del Palmar area of the San Juan Basin; to the west, Serranía del Darién (highest at Cerro Tacarcuna, 1875 m, on the border with Panama) and Serranía de Baudó (highest at Alto del Buey, approximately 1400 m) (Forero and Gentry 1989; Sánchez-P. et al. 1990; Sota 1972).
The typical landscape of the Pacific platform is hills and mountains. Erosion processes are enhanced in sectors with more of an altitudinal gradient and particularly where drying is more intense due to the instability of the geologic materials, and on occasion due to scarcity of vegetation. The alluvial soils, which occupy extended areas along both banks of large rivers and smaller tributaries and marine flats, contain easily alterable minerals (feldspars and ferromagnesiums) in the sands and amorphous materials in the clays. Relatively high levels of nutrients are present in such areas, reflected in a greater capacity of cation exchange, high values of base saturation and low content of exchangeable aluminium; these soils are generally of open-clayey and clayey textures (Castiblanco 1990). In contrast, an ecological distinction is a tendency to highly leached nutrient-poor soils; the white-clay soil that occurs in some areas is one of the poorest in such essential elements as phosphorus as well as trace elements like boron and zinc (Faber-Langendoen and Gentry 1991).
The Chocó is probably the wettest sizeable region on Earth, with various locales receiving average yearly precipitation of 4000 mm to over 9000 mm; Tutunendó, central Chocó, has averaged 11,770 mm (Sota 1972). West (1957) assembled records of maximum precipitation for the region - Quibdó (capital of Chocó Department), with an average annual precipitation of 8558 mm, in 1939 reached 15,058 mm. The Chocó is one of few places in the neotropics where tropical pluvial forest occurs (tropical rain forest sensu Holdridge) (Gentry 1982).
There is substantial precipitation in the north (where it is locally more varied), the greatest discharge of rain centrally, and the least in the south. At Riosucio rainfall averages 3700 mm, in the mid-region at Istmina 7560 mm and in the southern zone at La Florida 2250 mm. Dry seasons occur in the north toward Panama and the Caribbean Sea (about December-March), and in the south toward the Ecuadorian border. The precipitation is bimodal-tetraseasonal, with a greater concentration in the first half of the year. The rain tends to occur in the afternoon and at night. The climate is classified as B2rA´a´ (Thornthwaite system) with little deficiency of megathermal water (Rangel-Ch. and Aguilar 1993; Sota 1972).
Sunshine averages 110 hours per month through the year. The annual mean temperature is 23.6°C, the average maximum is 29.9°C and minimum 18.6°C; the monthly means rarely exceed 28°C (West 1957). At Quibdó, the maximum temperature is 41°C, the minimum 15°C (Forero 1982). Periods of high relative humidity (SeptemberJune) average 89%.
The Chocó phytogeographic region (Gentry 1982) may be broadly considered to include the border area of eastern Panama (cf. Darién Data Sheet, CPD Site MA20), and the coastal lowlands of western Colombia and north-western Ecuador mainly covered by wet forest and pluvial forest (cf. Ecuadorian Mesic Forests Data Sheet, CPD Site SA40). Based on the altitudinal gradient, the variation of water content in the soil and the influence of the sea, several vegetation types are recognized (Rangel-Ch. and Lowy 1993; West 1957; Cuatrecasas 1958; Zuluaga-R. 1987; Von Prahl, Guhl and Grogl 1979; Aguirre-C. and Rangel-Ch. 1990; Acosta-Solís 1970):
1. Mangroves ("manglares"): arboreal or shrubby associations that are amphibious in character, with evergreen foliage. They occupy large expanses along the coastal Pacific seaboard. The vegetation corresponds to the syntaxonomic Order Rhizophoretalia. These mangroves are the most diverse in species in the Western Hemisphere (Gentry 1982). The species that are characteristic and diagnostic of the vegetational syntaxa belong to Rhizophora (Rhizophoraceae), Pelliciera (Pellicieraceae), Avicennia (Verbenaceae) and Laguncularia and Conocarpus (Combretaceae).
Communities with the following dominants are present (Von Prahl, Guhl and Grogl 1979): (i) species of Rhizophora, which develop on the seaboard front and penetrate; (ii) species of Avicennia, established landward next to Rhizophora on more consolidated terrain; (iii) Laguncularia racemosa, which forms a strip next to the woods of Avicennia; and (iv) transitional communities - "natales" of Mora megistosperma (Leguminosae) and "cativales" of Prioria copaifera (Leguminosae).
2. Aquatic and marshy communities; in the sequential process, their point of origin is rivers and lentic environments. The following communities are present: (i) Eichhornia crassipes (Pontederiaceae) and Pistia stratiotes (Araceae); (ii) Marathrum haenkeanum (Podostemaceae) and Dicranopygium crinitum (Cyclanthaceae); and (iii) Polygonum acuminatum (Polygonaceae).
3. Riverbank and beach communities established on sandy terrain left by rivers: (i) "pajonales" with Panicum spp.; and (ii) canebrakes of Gynerium sagittatum in tall dense associations on riverbanks.
4. Firm-terrain rain-forest vegetation in the lowlands: in the northern zone, the vegetation is in the Alliance Brosimion utilis, which includes communities dominated by (i) Cavanillesia platanifolia (Bombacaceae); (ii) Anacardium excelsum (Anacardiaceae) and Castilla elastica (Moraceae); (iii) Anacardium excelsum and Pseudolmedia laevigata (Moraceae); and (iv) Chrysophyllum sp. (Sapotaceae) and Brosimum guianense (Moraceae). In the southern zone, the forest is pluri-stratified, with two arboreal strata, and climbing plants and epiphytes of vigourous growth.
The lowland pluvial forests are among the world's most dense and species-rich. Samples of 0.1 ha halfway between Quibdó and Tutunendó and at Bajo Calima averaged 510 plants 2.5 cm or more in dbh of 262 species, including 88 trees (and lianas) 10 cm or more in dbh (Gentry 1986). These wettest forests tend to have more but smaller trees (and lower biomass) than do equivalent forests elsewhere in the neotropics. They are relatively poor in free-climbing lianas, which are mostly replaced by woody hemi-epiphytes belonging to families like Guttiferae, Marcgraviaceae and Melastomataceae (Gentry 1986), all of which have world centres of diversity in this region.
5. Highlands belt; communities with the following species: (i) Inga sp. (Leguminosae), Billia columbiana (Hippocastanaceae), Brosimum sp.; (ii) Sorocea sp. (Moraceae), Jacaranda hesperia (Bignoniaceae), Pourouma chocoana (Urticaceae); (iii) Guatteria ferruginea (Annonaceae), Cecropia sp. (Cecropiaceae), Inga spp.; and (iv) Elaegia utilis (Rubiaceae), Brunellia sp. (Brunelliaceae), Pourouma cf. aspera, Inga spp.
The flora of the Chocó is poorly known mainly because of difficult accessibility and the very wet climate, which create arduous conditions for exploration and collection of specimens (Lellinger and Sota 1972). Although there were some collections in the 19th century, the first thorough botanical survey was in 1971 for pteridophytes (Sota 1972; Lellinger 1975). Current data suggest that the Chocó is floristically rich, with 8000-9000 vascular plant species (Forero 1982, 1985; Gentry 1982, 1993; Forero and Gentry 1989).
In the entire region, so far 4638 species of vascular plants have been recorded, belonging to 201 families and 1376 genera (cf. Murillo and Lozano 1989). The families with greatest generic diversity are Orchidaceae, Leguminosae, Asteraceae and Rubiaceae (Table 63) (Rangel-Ch. and Lowy 1993; Forero and Gentry 1989). Palm species and Bombacaceae (especially species of Quararibea and its segregate Phragmotheca) are notably prevalent.
According to the subdivisions established for the Pacific region (Rangel-Ch. et al. 1994), the central subregion (zone 3) has the greatest concentration of plant species: 1440 species in 639 genera of 131 families; next is the coastal lowlands subregion (zone 2), with 1389 species in 733 genera of 165 families. Within zone 3, the greatest diversity of species is in the central pluvial forest at the centre of Chocó Department, in the Atrato River Valley in the vicinity of Quibdó, which receives 7200-7600 mm of rain annually.
A unique feature of the Chocó pluvial forests is unusually large leaves (Gentry 1986). The largest, which may represent the largest simple leaves of any woody dicotyledon, are individual masses of tissue 1 m long and over 50 cm wide of Psittacanthus gigas (Loranthaceae). The Chocó has a dozen family or genus world records for leaf size, especially at Bajo Calima where both large size and extreme sclerophylly characterize the leaves of many unrelated species. Some of these large-leaved taxa range to eastern Panama or the Ecuadorian Chocó, but most seem to be narrow endemics in the unusual phosphorus-lacking white clay of the Bajo Calima area. Species of Bajo Calima with (putatively) the largest leaves in their families include Iryanthera megistophylla (Myristicaceae), Pleurothyrium new sp. (Lauraceae), Licania gentryi (Chrysobalanaceae), Psittacanthus gigas, Ilex new sp. (with leaves 15-25 cm × c. 10 cm) (Aquifoliaceae), Protium amplum (Burseraceae), Guarea cartaguenya (Meliaceae), Schlegelia dressleri (Bignoniaceae) and Pentagonia grandiflora (Rubiaceae).
Chocó forests are poor in wind-dispersal and rich in bird- and mammal-dispersal of fruits and seeds; well over 90% of the woody species are dispersed by these animals (Gentry 1986). Presumably the relatively large seeds (which dispersal by vertebrates made possible) have a strong selective advantage in the highly leached, nutrient-poor soils of the lowland Chocó. Several families and genera have larger seeds in the Chocó than elsewhere. Sacoglottis ovicarpa is the largest-fruited Humiriaceae. Orbignya cuatrecasana is exceeded in fruit size among palms only by coconut (Cocos) and double coconut (Lodoicea). Several endemic species of Myristicaceae have the largest fruits in their genera (Compsoneura, Iryanthera).
The region's flora has strong affinities with the flora of Central America (particularly Panama), which gradually decrease southward (Forero 1982; Gentry 1982; Lellinger and Sota 1978). In general the tree species belong to Amazonian genera, and sometimes the same species occur. The Amazonian affinities within the Chocó also decrease southward (Gentry 1982, 1989).
An interesting peculiarity of the Chocó lowlands is that families usually restricted to the Andean uplands occur here near sea-level (Gentry 1986). Typical montane taxa with lowland representatives are Podocarpus, Talauma, Hedyosmum, Meliosma, Brunellia, Panopsis and Ilex. The presence of these generally montane taxa is an indication of the cloud-forest-like nature of the wet Chocó forests.
Collections in the central Chocó from Valle del Cauca Department are rich in endemic species (Gentry 1982). The southern Chocó probably has the entire continent's highest levels of biological endemism (Gentry 1993, pers. comm.). About 20% of the Chocó species are regionally endemic; also, high very local endemism within Chocó areas may be characteristic (Gentry 1989, 1992). This high endemism is the result of active speciation among epiphytes (e.g. Araceae, Orchidaceae, Bromeliaceae), palmettos (e.g. Heliconia, Renealmia, Costus) and shrubs (e.g. Melastomataceae, Psychotria). At the generic level the Chocó also is a significant centre of endemism - genera endemic or strongly centred here include Otoba (Myristicaceae), Trianaeopiper (Piperaceae), Schlegelia (Scrophulariaceae) and Cremosperma (Gesneriaceae). The high endemism is strong evidence suggesting there was a Pleistocene Chocó refugium (Forero 1982; Gentry 1982).
Some of the unusually large fruits and seeds of Chocó species are also edible for humans. The large mammal-dispersed fruit of "borojó" (Borojoa patinoi) has resulted in the recent advent through Colombia of a delicious "refresco". Extractive reserves based on extra-large-fruited species like borojó, that are adapted to flourish on ultra-poor soils under high rainfall, might be a better developmental option than typical practices.
The vegetational resources of this extensive region are used for many purposes, such as food, folk medicine, lumber, fibre, dyes and tannin, and rubber and gums. The following is just a sketch:
Social and environmental values
The inhabitants of the Pacific coast represent 2.7% of Colombia's population of 33 million. The population generally is sparse, and is 90% blacks (Whitten 1974) with 5% mestizos and 5% indigenous peoples. Most of the inhabitants depend for their livelihoods on subsistence agriculture, hunting, fishing, forestry and mining (Barnes 1993; West 1957). The most important urban areas are Buenaventura, Tumaco and Quibdó (Castiblanco 1990).
The vast Pacific region is rich in mineral and natural resources, which have attracted outsiders since the Spanish conquest. This region is the country's main producer of platinum and second largest producer of gold, which have been exploited especially since the 19th century. There are considerable deposits of bauxite, manganese, tin, zinc, nickel, tungsten, copper and chromium, as well as possible petroleum reserves (Barnes 1993). The forest resources have been over-exploited; the Amerindian groups (Emberá, Waunana, Eperara), blacks and mestizos have not reaped benefit from these exploitations. The great biodiversity is probably what contributes most to the richness of the region.
Birds recorded total 577 species in 353 genera of 67 families; Tyrannidae is the most diverse family with 28 genera and 60 species, and the most speciose genus is Tangara (16 tanager spp.) (Roda and Styles 1993). The Chocó region is one of the richest areas of avian endemism in the neotropics; there are two Endemic Bird Areas (EBAs). No fewer than 62 species of restricted range occur within the Chocó and Pacific slope of the Andes EBA (B14), which embraces the lowlands and Andes south of the Serranía de Baudó. A further 14 such species occur in the lowlands of eastern Panama and northern Colombia (north of the Serranía de Baudó) in the Darién and Urabá lowlands EBA (A19), which embraces the entire Atrato drainage; six are restricted to the lowland and foothill forests bordering the serranía. At least 28 of the restricted-range species are confined to lowlands below 800-900m. Generally widespread forest destruction has caused 18 species to be threatened; three of the endemics are considered threatened but primarily because of poor observer coverage: Chocó tinamou (Crypturellus kerriae), speckled antshrike (Xenornis setifrons) and Baudó oropendola (Psarocolius cassini) - none of them has been recorded in Colombia since the 1940s.
There have been 97 species of reptiles recorded: the suborder Serpentes has 52 - Colubridae is the most diverse family with 25 genera and 35 species, and the most speciose genus is Micrurus (7 spp.); the suborder Sauria has 45 species representing 17 genera in six families - Iguanidae is the most diverse family with six genera and 26 species, and the most speciose genus is Anolis (20 spp.) (Sánchez and Castaño 1994). There are 127 recorded species of amphibians in 26 genera of seven families; Leptodactylidae is the most diverse family with three genera and 45 species, and the most speciose genus is Eleutherodactylus (40 spp.) (Roa and Ruiz 1993).
The principal threats are indiscriminate felling of trees; expanding permanent agriculture with plantations especially of African oil palm (Elaeis guineensis); raising cattle; dredging of rivers; shrimp-farming that deforests the mangroves; excessive direct exploitation of mangroves and palm groves; and road building (cf. Budowski 1990). The current rate of deforestation is 600 km² annually.
The rain forests of the Chocó have been a source of lumber since the beginning of the 20th century; West (1957) found that hardwoods scattered through the region and more concentrated softer woods of swamp forests had already been logged out from most accessible areas; see also Lellinger and Sota (1972). Deforestation has intensified since 1976. Gentry (1989) reported that most slopes in the Darién Gap on the border with Panama have been cleared. In the Urabá area, only small patches of forest are left after expansion of the production of banana and star-fruit (Averrhoa carambola). North of Buenaventura in the Bajo Calima area, commercial paper-pulp operations have clear-cut extensive areas, and considerable land has been deforested in the Tumaco area of Nariño Department. Installation of commercial shrimp farms particularly in the south has caused destruction of mangroves (Barnes 1993). By 1984, over 15,000 km² of coastal forest had been destroyed for domestic use for wood and paper or to make way for agro-industrial production of the African oil palm.
Uncontrolled gold mining has produced massive sedimentation in rivers and contaminated them with mercury. Motorized equipment is eroding riverbanks, causing the riverbeds to drop, and threatening fish stocks and the ability to travel and transport goods between communities (Barnes 1993).
As part of Colombia's development strategy, the Chocó region is coming under increasing pressure (WWF 1993; Barnes 1993). The overall region is managed by regional autonomous corporations: CORPURABA, CODECHOCO, CVC (for the Dept. Valle del Cauca), CRC (for Cauca) and CORPONARIÑO. Forestry research is assisted by the Corporación Nacional de Investigaciones Forestales (CONIF) (Budowski 1990). Development is underway, with new roads promoting an influx of settlers, conversion of primary forests into pasturage and agricultural fields, mining, and new exploitation by timber companies.
Although not being advanced presently, the government in 1984 approved a plan to construct an interoceanic canal (similar to the Panama Canal) by dredging the Atrato and Truandó rivers (Palacios-M. 1985; Ramírez 1967). With this project would come major changes, such as hydroelectric plants and intensification of road building and land conversion (Convers-P. 1985).
The four Natural National Parks Los Katíos (720 km²), Utría (543 km², terrestrial and marine), Isla de Gorgona (including Gorgonilla Islet) (492 km², 16 km² terrestrial) and Sanquianga (800 km²) protect 2013 km² of the region's terrain, which also includes the 200-1000 m portion of Farallones de Cali NNP (totalling 1500 km²) and 500-1000 m portion of Munchique NNP (440 km²) (INDERENA 1984; Sánchez-P. et al. 1990). The relatively new (1987) Utría Natural National Park (Chávez 1988) is receiving fundamental help through the Parks in Peril Program of The Nature Conservancy (U.S.) and Fundación Natura Colombia working with INDERENA (the Instituto Nacional de los Recursos Naturales Renovables y del Medio Ambiente) and the local communities (FNC 1993).
In a 1985 meeting in Bogotá (organized by WWF-U.S.), national and international experts assessed conservation priorities for the Chocó region, evaluating established reserves and recommending new ones. The recommendations and priorities were formulated to be used in the region's future development and management (García-Kirkbride 1986a, 1986b; Hernández-Camacho 1984; Gentry 1989). The government in 1993 initiated a 6-year Proyecto Biopacífico to fuse a strategy for conservation and sustainable management of the region's biodiversity (INDERENA 1993).
Additional areas need to be delineated and their protection promoted (cf. INDERENA 1993), e.g. (1) in the northernmost zone of the Chocó beyond the banks of the Atrato River; (2) an area along the Atrato riverbanks that includes the plant communities "manglares, natales, sajales"; (3) between the Atrato, Baudó and San Juan rivers; (4) in Valle del Cauca Department between the Calima River and Buenaventura (cf. Faber-Langendoen and Gentry 1991); and (5) Isla de Gallo in Tumaco Cove. The central Chocó, except where accessible from the Atrato and San Juan rivers or the expanding road system around Quibdó, currently remains covered by essentially intact forest. Nariño Department needs to receive special consideration to preserve representative areas of its biotic conditions.
Map 74. Colombian Pacific Coast Region (Chocó) (CPD Site SA39), showing subregional zones
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This Data Sheet was written by Olga Herrera-MacBryde (Smithsonian Institution, SI/MAB
Program, S. Dillon Ripley Center, Suite 3123, Washington, DC 20560-0705, U.S.A.), Orlando
Rangel-Ch., Mauricio Aguilar-P., Hernán Sánchez-C., Petter Lowy-C., David Cuartas-Ch.
and A. Garzón-C. (Universidad Nacional de Colombia, Instituto de Ciencias Naturales,
Museo de Historia Natural, Apartado Aéreo 7495, Santafé de Bogotá, Colombia).
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