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Central America: CPD Site MA20

DARIEN PROVINCE AND
DARIEN NATIONAL PARK

Panama
(Central)

Location: Eastern Panama, the park in south-east Darién Province between latitudes 7°12'-8°31'N and longitudes 77°09'-78°25'W.
Area:
Darién Province 16,671 km², Darién National Park 5790 km².
Altitude:
0-1875 m.
Vegetation:
Tropical lowland dry, moist and wet forests, perhaps 500-year-old secondary rain forest; tropical premontane moist (warm transition), wet and pluvial forests; lower montane pluvial forest. Marshes and swamps, tall non-flooding forests, cloud and elfin forests.
Flora:
Darién Province: 2440 species recorded; high endemism - e.g. Cerro Tacarcuna 23% endemism; threatened species.
Useful plants:
Timber species; medicinals.
Other values:
Wilderness; diverse fauna, including endemics and threatened species; watershed protection; Amerindian groups; intercontinental disease barrier; potential ecotourism.
Threats:
Colonization, logging, agriculture, grazing, road building, mining.
Conservation:
Darién National Park is a World Heritage Site and Biosphere Reserve, with a buffer zone and Punta Patiño Nature Reserve to east and Colombia's Los Katíos NP to west. Also, Canglón and Chepigana Forest Reserves; Comarca Emberá No. 1 (Cemaco District), Comarca Emberá No. 2 (Sambú District) and Kuna de Walá, Mortí y Nurrá Amerindian reserve.

Map 31: CPD Site MA20

References

Geography 

Located in the eastern portion of Panama which borders Colombia, Darién Province basically forms a long and flat lowland valley mostly fringed by mountains. Darién National Park is along 90% of the international border and extends from the Pacific coast to the continental divide, which is just 16 km from the Caribbean Sea.

The extensive central valley receives the drainages of the country's largest watershed (c. 13,371 km²), primarily from the south-east-flowing Chucunaque River and north-west-flowing Tuira River as well as the Balsas River, which join as the Tuira River to flow westward to the large Gulf of San Miguel on the Pacific Ocean (Map 31). Brackish to freshwater marshes and swamps extend well inland along the Tuira River's lower portion and the lower Chucunaque River. Twice a day strong Pacific tides (ranging 26 m) affect the Tuira and Chucunaque for many km inland.

The spine of Panama along the north-eastern border of the province is a fairly continuous mountain system, the Cordillera de San Blas and higher Serranía del Darién. The serranía emerges in the north with elevations of 300-600 m and in the south-east rises to 1875 m at Cerro Tacarcuna (8°10'N, 77°15'W), which is the highest mountain between the Andes and western Panama and only c. 32 km inland from Colombia's Caribbean coast at the Gulf of Urabá (which gives way to the vast lowland swamps and lagoons of the Atrato River).

The southern region of the province (Altos de Darién) is composed of a trident of somewhat discontinuous mountain systems extending northward. From east to west, they are: (1) Serranía de Pirre (with the flanking Serranía de Setetule), which extends into the central valley and contains the headwaters of the Tuira River; (2) Serranía de Jungurudó (with the Cordillera de Juradó to the south), extending northward to the Serranía de Bagre - which forms the western edge of the central valley; and (3) Serranía del Sapo (picture), bordering the Pacific Ocean. The somewhat convergent southern ends of several of these mountain systems and the Altos de Quía help to demarcate the international border. The principal heights (c. 1150-1550 m) in the southern area include Cerro Pirre, Alturas de Nique, Cerro Piña and Cerro Sapo.

Along the Pacific Ocean, the south-western edge of the province is a somewhat undulating coastline with rocky shores and sandy beaches. Farther to the north-west, the coast is broken by the Gulf of San Miguel, where the wide Lower Tuira River meets the sea. North of the gulf, the border of the province is the south-eastern end of the Serranía de Majé at the Serranía de Cañazas.

During the Tertiary's Middle Eocene an insular volcanic arc emerged, and the regional lowlands emerged from tectonic activity probably in the Late Pliocene. During the Pleistocene the sea-level fluctuated from 100 m lower to 50 m higher than at present. The southern mountains and the Caribbean slope of the north-eastern mountains are largely of volcanic origin, whereas the inland slope of the Serranía del Darién is of sedimentary origin. The lowland clay soils are generally derived from Late Miocene shale with layers of dolomite and calcareous sandstone (Golley et al. 1975; Hartshorn 1981; INRENARE and ANCON 1988).

Rainfall is abundant (3000-4000 mm) on the mountains next to the Caribbean coast, still more abundant (4000-5000 mm) on the inland mountains, and diminishes (2800-1700 mm) in the central valley and gulf areas. There is a marked dry season (weaker near Colombia) with less than 100 mm monthly from January through March-April. The temperature generally varies between 16°-35°C, with an annual mean of c. 25°-27°C (OEA 1978; INRENARE and ANCON 1988).

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Vegetation 

The Darién is one of the most diverse and species-rich regions in Central America. It includes almost ten major vegetation types (see below), from the littoral strand and coastal dry forests to mangroves, brackish and freshwater swamps, and various lowland and premontane to lower montane rain- forest life zones, including cloud forests and elfin forests at rather low elevations because of the isthmian effect (see maps in Duke and Porter 1970; INRENARE and ANCON 1988). Five Holdridge life zones are represented in Darién National Park.

There is speculation whether the central Darién lowland forests are primary or much is secondary growth from a savanna created by the Amerindians and abandoned after the coming of the Spanish c. 500 years ago. Historical accounts, such as those of Balboa in 1513 and Wafer in 1681 are quoted in Sauer (1966) and Prebble (1968); also see Jaén-Suárez (1985). Some dominant trees may be secondary species that are not reproducing but have attained great size (Golley et al. 1975; INRENARE and ANCON 1988), although Hartshorn (1981) suggested that studies of forest dynamics and the regeneration potential of Cavanillesia platanifolia ("cuipo") (picture) did not support a secondary status.

Bordering the Gulf of San Miguel's southern embayments (from about Punta Garachiné eastward to Punta Alegre) is an isolated strip of tropical dry forests (thorn and seasonally strongly deciduous to semi-deciduous tropical dry forests). Characteristic trees include Albizia caribaea, Bombacopsis quinata, Cochlospermum vitifolium, Prosopis juliflora and Sabal allenii.

Mangrove forests occur along the Pacific and Atlantic coasts; their greatest representation is in the Gulf of San Miguel area, which has c. 46% of the mangrove forest of the entire country (G. Palacios 1992, pers. comm.). The major Pacific brackish water community is nearly pure stands of Rhizophora brevistyla to 41 m, which ranks them among the world's tallest mangroves. Pelliciera rhizophorae is an occasional associate. Minor brackish water communities develop in estuaries and rivers changing with decreasing salinity from Avicennia germinans to Mora oleifera and then Montrichardia arborescens. Caribbean coastal islands, inlets and bays support narrow low bands of Rhizophora mangle and the less abundant and taller (to 5 m) Laguncularia racemosa (Golley et al. 1975; Hartshorn 1981).

In the Gulf of San Miguel area, saltmarsh vegetation gives way inland to freshwater marshes and swamp forests, often with palms (e.g. Manicaria, Jessenia, Euterpe) or stands of Copaifera aromatica, C. panamensis, Pachira aquatica, Pterocarpus rohrii and various canopy dominants Prioria copaifera, Pterocarpus officinalis, Tabebuia rosea and Swartzia panamensis (Porter 1973; Nations and Komer 1983). Along the Tuira, Chucunaque and Balsas rivers is broad evergreen riparian forest with many understorey palms and some pure stands of Prioria copaifera ("cativo"), which can reach 55 m tall (Hartshorn 1981); details on the cativo swamps are in Duke and Porter (1970).

The most widespread vegetation type in the Darién is seasonally semi-deciduous tropical moist forest, which occupies non-flooded lowlands of the Chucunaque-Tuira Basin and western areas; c. 10% of the park is in this life zone. The most dominant, emergent trees are especially the deciduous Cavanillesia platanifolia (to 40 m and 2 m in dbh) and Ceiba pentandra, as well as Anacardium excelsum. Frequent canopy trees include Bombacopsis quinata, B. sessilis, Enterolobium cyclocarpum, Licania hypoleuca, Platypodium elegans, Pseudobombax septenatum, Sterculia apetala, Terminalia amazonia, Tetragastris panamensis and Vitex cymosa. In the subcanopy Mouriri parviflora is dominant and palms (e.g. Sabal allenii) are frequent. The dominant shrubs are Faramea luteovirens, Mabea occidentalis and Piper pinoganense. Lianas do very well in this formation (in the canopy and also the subcanopy), whereas epiphytes and ferns are relatively scarce (Golley et al. 1975; Hartshorn 1981).

In slightly higher (to c. 200-250 m) and/or wetter areas in the Darién occurs premontane wet forest - warm transition, and then to c. 500-600 m there is tropical wet forest (seasonal yet evergreen tropical rain forest), which extends broadly into the northern Chocó of Colombia (in CPD Site SA39). About 60% of the park is in these two life zones. Anacardium excelsum is dominant in the canopy and the following trees are frequent: the above Bombacopsis spp., Brosimum guianense, Ceiba pentandra, Cochlospermum williamsii, Dipteryx panamensis and Myroxylon balsamum; the main subcanopy tree is Oenocarpus panamanus. The dominant understorey shrub is Mabea occidentalis and frequent shrubs include Clidemia spp., Conostegia spp. and Miconia spp. Epiphytes are abundant and canopy lianas common.

At higher elevations occurs premontane pluvial forest and on the highest peaks and ridges lower montane pluvial forest; these life zones cover c. 30% of the park. Cloud forest can commence at c. 750 m and elfin forest may be found on the highest exposed elevations. The premontane forest is low (30 m) and dense, with many subcanopy palms; lianas are common. The same Brosimum and Dipteryx species are frequent in the canopy and Cephaelis elata is a dominant shrub; ferns are most common in this zone. A distinctive montane oak forest (Quercus humboldtii) is present above 1400 m on Cerro Malí and 1500 m on nearby Cerro Tacarcuna. The dominant trees in cloud forest are Oenocarpus panamanus, in elfin forest Clusia spp. (Porter 1973; Golley et al. 1975; Gentry 1977, 1985).

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Flora 

Although the Flora of Panama has been finished, the coverage is very incomplete; a revised checklist being maintained by the Missouri Botanical Garden has 2440 species recorded for the Darién (ANCON 1993, pers. comm.). Since 1970, collecting in the wetter forests of Panama has revealed many new species (e.g. D'Arcy 1977; Gentry 1985, 1986). Limited collecting in the region of the park and the lack of synoptic studies preclude determining the total number of species or endemics (only 700 species have been recorded).

The diversity of the Pacific mangrove forest is pronounced, with c. 15 species compared to c. 6 on the Caribbean side. Duke (1975) provided a checklist for the half of collections identified for an ecological study of several Darién forest types; c. 400 species - including over 200 canopy and subcanopy trees had been identified in tropical moist forest, and even riverine forest had c. 220 identified species - including 66 tree species. The samples suggest that these forests are as rich as forests in Amazonia.

Due to the mountainous conditions that reflect the Darién's past geological insularity and isolation, and as well its relatively recently formed lowlands, the Darién has its own characteristic flora. The flora has complex ancient and more recent affinities, which reflect the varying land's mixed history of location, isolation and connection (Gentry 1985). Upland elements show northern affinities with Central America (e.g. Costa Rica) that suggest floristic migration during cooler palaeohistoric periods. With Pleistocene climatic shifts, there may have been a refugium in this region. Lowland trees and lianas have Amazonian affinities, whereas the epiphytes and understorey shrubs have northern Andean relationships. The flora has particular affinities with the flora of the Chocó region of Colombia (in CPD Site SA39), as well as the ancient Guayana region (see CPD Site SA2).

Some studies have been conducted on and near Cerro Pirre and Cerro Tacarcuna, where many endemics occur (Lewis 1971; Gentry 1985). The limited collections (90% identified) from Cerro Tacarcuna show 23% endemism and c. 25% new species, including the new genus Tacarcuna (Euphorbiaceae) and such remarkable new species as Freziera forerorum (Theaceae), a tree which may have the most asymmetric leaf base among flowering plants (Gentry 1978). Other appropriate areas are presumed to have high angiosperm endemism, especially isolated cloud forests.

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Useful plants 

The Darién forests contain important reserves of timber, such as Prioria copaifera ("cativo"). This region has contributed 75% of the logs to the national market, with cativo comprising half of the total. In 1976, of the five species supplying 94% of all the logs Panama marketed (cativo, Anacardium excelsum, Bombacopsis quinata, Vatairea sp., Hyeronima oblonga), the first four were species of the Darién (Hartshorn 1981). A forest inventory of the province found adequate quantities of excellent timber trees and some that also can be used for plywood exports and paper and pulp products e.g. Anacardium excelsum, Capara guianensis, Cedrela odorata, Cordia alliodora, Dialium guianense, Myroxylon balsamum, Swietenia macrophylla, Tabebuia guayacan, T. rosea and Terminalia amazonia (Lamb 1953; OEA 1978). About 35.5% of the province's land is in class VII, which is marginal for agriculture but suitable for forestry.

The indigenous peoples use many species, for many purposes (Duke 1968, 1970; Torres de Araúz 1985). Cooking oil extracted from the palm Jessenia bataua ("trupa") is particularly valued; the palm Phytelephas seemannii is used to make vegetable-ivory carvings.

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Social and environmental values 

The Darién includes three indigenous groups, Emberá and Wounaan (from the Colombian Chocó) and Kuna (Houseal et al. 1985), and early historic Negroes. Although just 400-500 Kuna are still in the Darién, 40,000-50,000 inhabit the nearby Comarca de Kuna Yala (San Blas). Within the park there were c. 2500 people in 1987: 250 Kuna, 1700 Emberá, few Wounaan, 400 Negroes and some colonists. The Amerindians have extensive knowledge of the forest, and despite long contact with outside societies have managed to keep some of their traditional forest cultures. The Kuna have been studied extensively (see D'Arcy and Correa-A. 1985); their language and behaviour are in part directed by their relationships with wild animals and plants and the symbolic and magical features they represent (Chapin 1991).

The lowland wet forest is of special importance because it may be a secondary forest almost 500 years old. If further research reveals that this forest is secondary, it may be possible to learn how to regenerate such tropical forest with greater success, more regenerative capacity and biodiversity.

The region has a rich fauna which is still poorly known. There are c. 770 vertebrate species in Darién Province, with considerable endemism; nearly all of them have been recorded in the park. Over 60% of Panama's mammal species occur in Darién Province, including six of the eight species of primates - three-banded douroucouli, Geoffroy's spider monkey, brown-headed spider monkey, crested bare-faced tamarin, mantled howler monkey and white-faced capuchin; and five or six cat species - puma, jaguar, ocelot, margay, jaguarundi and probably oncilla (Felis tigrina). Endemic reptile and amphibian species include two snakes, a lizard, a salamander and five frogs.

There are also many bird species in the region that are threatened elsewhere, such as crested guan, marbled wood-quail, harpy eagle, crested eagle, crested owl, brown violetear, green-crowned brilliant, great curassow, macaws and black-cheeked woodpecker. With its extensive lowland forests bounded by isolated mountains, the region is at the centre of two Endemic Bird Areas (EBAs): the Darién and Urabá Lowlands EBA (A19) below c. 900m, and higher up the East Panama and Darién Highlands EBA (A20). Thirty restricted-range species occur in the two EBAs; some are confined to single mountain areas such as Cerro Tacarcuna.

The park protects one-third of the watersheds in Darién Province. Their conservation is highly important for its social and economic development. The region has potential for expanded ecotourism and for ecologically broad scientific research.

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Threats 

The eastern Darién and adjacent Colombia have the only gap (106 km) in the Pan American highway, which extends from Alaska to Tierra del Fuego. The highway penetrated the Darién in 1973, reaching Yaviza in 1984 - 24 km from the park. Secondary roads have opened some of the Darién interior, especially southward. The plans to complete the highway have been slowed more by limited funds than environmental concerns (Herlihy 1989). The almost inevitable opening of the Darién by a highway through the Tuira River Valley will bisect the park with a 22-km roadway and introduce numerous hazards and pressures on this fragile environment (D'Arcy 1977; Gentry 1977; INRENARE and ANCON 1988; Polsky 1992).

The Darién natural environment is already under continued and increasing pressure (Holz 1980; Herlihy 1989). On Panama's Azuero Peninsula west of Panama City, most forests have been cut down and burned; farmers are leaving and settling in other regions of the country and the Darién is their first choice. In 1978, only 1.6% of Panama's population lived within Darién Province. The government has made the province a priority for major development of natural resources. Programmes have been started to make use of these resources as part of Panama's solution to alleviate immediate economic problems (OEA 1978; RARE 1983; INRENARE and ANCON 1988). Mining concessions have been granted in the park. Under the national colonization plan, settlers are steadily coming into the Darién and opening the frontier (Torres de Araúz 1970; D'Arcy 1977; Herlihy 1989). By 1986 the population had increased to c. 34,000, a third larger than in 1970, primarily by migration of colonists using access by road.

The areas predominantly under pressure from colonists and timber companies are wetlands and lowland forests (Mayo-Meléndez 1965; Duke 1968, 1975). The wetlands around the Tuira River were first settled, e.g. in the district of Chepigana. They are mostly cultivated with rice and maize (Duke 1968). Rice producers prefer creating new fields because of higher yields and lower weed control, rather than using the same fields year after year (Torres de Araúz 1970). New settlers have taught local inhabitants that clearing the land for pasture to raise cattle is as profitable if not more so than producing field crops. The environment of the region is undergoing enormous change; probably over 2000 km² are already farmlands and pasturage (Herlihy 1989), which is over 36% of the province's land suitable for such uses (i.e. classes II-VI - OEA 1978).

Darién lumber is an important factor in Panama's exports; it is mainly used for construction. The wetland areas are the prime logging areas (Mayo-Meléndez 1965). Mangroves also are used to make charcoal for cooking or to sell. The bark of Rhizophora brevistyla (red mangrove) is exported for use in the tannin industry. As the population of settlers increases, the mangroves are likely to be severely degraded, disrupting this fragile ecosystem - even though the mangrove forest is a breeding ground for white shrimp, the keystone of Panama's shrimp industry, which in 1984 resulted in exports of US$30.7 million.

From neighbouring Colombia potential threats could be realized with establishment of convenient access into the Darién. Most of the usable slopes on the Colombian border have been cleared (except in Los Katíos Natural National Park), but due to the natural barrier of mountains and swamps and no direct access, the Panamanian slopes have been relatively untouched. Disease is another problem that will confront Panama and northern America with completion of the Pan American highway. An example is the dreaded hoof-and-mouth disease ("aftosa") which affects cattle and pigs, and also over 50 plant diseases that may strike native species and crops.

Already Darién NP is facing encroachment by settlers. In the park's buffer zone there are c. 40 communities with more than 8000 people (INRENARE and ANCON 1988). Many inhabitants for example of the towns El Real de Santa María and Boca de Cupe have farms near the park boundaries. However, 78% of the park lands are in class VIII (48%), which is unsuitable for agriculture and forestry, or VII-VIII (30%), which is marginal for forestry (OEA 1978). The number of Emberá in the park has increased to 3000 (Polsky 1992).

The increasing population of colonists from western Panama disrupts cultural values of the indigenous peoples who have immigrated to or have been in the region (Herlihy 1989). The Chocó and the remaining Kuna have small agricultural plots predominantly along rivers and streams and do a limited amount of hunting. In the nearby territory Comarca de Kuna Yala (San Blas Province), the Kuna initiated a series of projects to secure their traditions and protect their forest from illegal encroachment. The projects include a wild-life reserve, a forest park and a botanical park (Nations and Komer 1983; Chapin 1984). Their endeavours have potential to be a model for protection of tropical rain forest by indigenous peoples, although the Kuna are being affected by the strong pressures of cultural change (cf. Chapin 1991).

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Conservation 

The heavy influx of settlers has prompted the Panamanian government to recognize the need for more management and conservation. Darién National Park was established in 1980, partly having been preceded in 1972 by a Protection Forest. In 1973 Colombia established the contiguous Los Katíos Natural National Park (520 km²) for the southern end of the Serranía del Darién. The Panamanian region was accepted as a World Heritage Site in 1981 and a Biosphere Reserve in 1983. Definitive boundaries apparently still remain to be legalized by decree. Darién NP encompasses c. 5790 km², making it the largest Panamanian park and one of the largest protected areas in Central America, which conserves the largest tropical rain forest on the Pacific slope.

Other protected areas in Darién Province are: Kuna de Walá, Mortí y Nurrá Indigenous Reserve (in the headwaters area of the Chucunaque River, the boundaries being established); the Chocó homelands Comarca Emberá No. 1 (Distrito de Cemaco) (1826 km² in the Chucunaque-Tuira Basin) and Comarca Emberá No. 2 (Distrito de Sambú) (931 km² in the Sambú Basin); Canglón Forest Reserve (316 km²); Chepigana Forest Reserve (1460 km² in 1960, but the lands were reclassified and the smaller area is being redetermined); and Punta Patiño Nature Reserve (263 km²) (ANCON 1993, pers. comm.). Over half the province thus is under some management category involving environmental protection. The Darién Region is one of eleven areas given priority under the 1992 Convenio para la Conservación de la Biodiversidad y Protección de Areas Silvestres Prioritarias en América Central. Standards have been established for entrance and use of the buffer zone (Map 31) and the Darién National Park, to develop in an orderly manner ecotourism with conservation, which will benefit both the park and the communities in and near the park. Darién NP has been given top priority for preservation of representative species of terrestrial and aquatic ecosystems (LaBastille 1978; INRENARE and ANCON 1988).

The park has an administrative office in El Real de Santa María and three ranger stations (Balsas, Cruce de Mono, Pirre) for protection, conservation and management. From 1989 INRENARE and ANCON have delimited and labelled 100 km of trails between the three ranger stations. The park rangers are provided with training and basic equipment for effective park protection and enforcement. There are monthly visits to the park's most critical areas to evaluate the effects of settlers, hunters, etc.

Environmental education is undertaken in every community in or neighbouring the park, including collaboration (by Panama and U.S.A.) in the Sistema Nacional para la Erradicación de la Malaria (SNEM) and the Comité para la Prevención de la Fiebre Aftosa (COPFA). Precautions have been taken (including regulations) to avoid and minimize impacts from the planned Pan American highway. The ecological dangers of opening roads and the highway without necessary controls are explained to the public.

In the buffer zone, ANCON (the Asociación Nacional para la Conservación de la Naturaleza) has established Peresénico Agroforestry Farm (near Pirre Ranger Station), a project of Finca Agroforestal. This provides opportunities for the neighbouring communities to benefit from a learning centre for agroforestry techniques and park information, and cheaper access tickets, etc., with a goal of reducing the pressure these communities exert on the protected zone. ANCON has an environmental education centre at El Real which works cooperatively with the park personnel and has research and development centres for ecotourism at Santa Cruz de Cana Mining Centre and Punta Patiño Nature Reserve.

Punta Patiño Nature Reserve, which is located in the park's buffer zone, has extensive mangrove forest and semi-deciduous dry forest. The reserve protects many mammal and bird species threatened with extinction (43 such species live in the area). ANCON is establishing an agroforestry programme in the area for habitat restoration using native plants and traditional crops. Notwithstanding the scientific character of this reserve, there is potential for developing some tourism without altering the natural ecosystems.

Conservation efforts in the Darién developed by INRENARE and ANCON are possible thanks to financial support of several international organizations such as The Nature Conservancy (partly through the Parks in Peril and the Last Great Places campaigns), U.S. Agency for International Development, Oro Verde, World Wildlife Fund - U.S. and World Wildlife Fund - U.K.

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Map 31. Darién Province and Darién National Park, Panama (CPD Site MA20)

References

Chapin, M. (1984). Kuna Indians: initiate rain forest reserve in Panama. Focus (WWF-US): Pg. 6.

Chapin, M. (1991). Losing the way of the Great Father. New Scientist 131 (1781): 40-44.

D'Arcy, W.G. (1977). Endangered landscapes in Panama and Central America: the threat to plant species. In Prance, G.T. and Elias, T.S. (eds), Extinction is forever. New York Botanical Garden, Bronx. Pp. 89-104.

D'Arcy, W.G. and Correa-A., M.D. (eds) (1985). The botany and natural history of Panama: la botánica e historia natural de Panamá. Monogr. Syst. Bot. No. 10, Missouri Botanical Garden, St. Louis. 455 pp.

Duke, J.A. (1968). Darien ethnobotanical dictionary. Battelle Memorial Institute, Columbus, Ohio. 131 pp.

Duke, J.A. (1970). Ethnobotanical observations on the Choco Indians. Econ. Bot. 24: 344-366.

Duke, J.A. (1975). Plant species in the forest of Darien, Panama. In Golley, F.B., et al., Mineral cycling in a tropical moist forest ecosystem. University Georgia Press, Athens. Pp. 189-221.

Duke, J.A. and Porter, D.M. (197O). Darien phytosociological dictionary. Battelle Memorial Institute, Columbus, Ohio. 70 pp.

Gentry, A.H. (1977). Endangered plant species and habitats of Ecuador and Amazonian Peru. In Prance, G.T. and Elias, T.S. (eds), Extinction is forever. New York Botanical Garden, Bronx. Pp. 136- 149.

Gentry, A.H. (1978). A new Freziera (Theaceae) from the Panama/Colombia border. Ann. Missouri Bot. Gard. 65: 773-774.

Gentry, A.H. (1985). Contrasting phytogeographic patterns of upland and lowland Panamanian plants. In D'Arcy, W.G. and Correa-A., M.D. (eds), The botany and natural history of Panama: La botánica e historia natural de Panamá. Monogr. Syst. Bot. 10, Missouri Botanical Garden, St. Louis. Pp. 147-160.

Gentry, A.H. (1986). Endemism in tropical versus temperate plant communities. In Soulé, M.E. (ed.), Conservation biology: the science of scarcity and diversity. Sinauer Associates, Sunderland, Massachusetts. Pp. 153-181.

Golley, F.B., McGinnis, J.T., Clements, R.G., Child, G.I. and Duever, M.J. (1975). Mineral cycling in a tropical moist forest ecosystem. University Georgia Press, Athens. 248 pp.

Hartshorn, G.S. (1981). Forests and forestry in Panama. Institute of Current World Affairs, GSH-14. Hanover, New Hampshire. 17 pp.

Herlihy, P.H. (1989). Opening Panama's Darién gap. J. Cultural Geogr. 9(2): 41-59.

Holz, R.K. (1980). The Darién of Panama: the twilight of a unique environment. Explorers J. 58(4): 158- 164.

Houseal, B.L., MacFarland, C., Archibold, G. and Chiari, A. (1985). Indigenous cultures and protected areas in Central America. Cultural Survival Quarterly 9(1): 10-20.

INRENARE and ANCON (1988). [Plan de manejo y desarrollo integrado] Reserva de la Biósfera Darién, basado en la labor de R.E. Weber. Instituto Nacional de Recursos Naturales Renovables (INRENARE) and Asociación Nacional para la Conservación de la Naturaleza (ANCON), Panama City. 176 pp.

Jaén-Suárez, O. (1985). Nuevos hombres y ganados y su impacto en el paisaje geográfico panameño entre 1500 y 1980. In D'Arcy, W.G. and Correa-A., M.D. (eds), The botany and natural history of Panama. Monogr. Syst. Bot. Missouri Bot. Gard. 10. Pp. 379-392.

LaBastille, A. (1978). Facets of wildland conservation in Middle America. Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica. 37 pp.

Lamb, F.B. (1953). The forests of Darién, Panama. Caribbean Forester 14: 128-135.

Lewis, W.H. (1971). High floristic endemism in low cloud forests of Panama. Biotropica 3: 78-80.

Mayo-Meléndez, E. (1965). Algunas características ecológicas de los bosques inundables de Darién, Panamá, con miras a su posible utilización. Turrialba 15: 336-347.

Nations, J.D. and Komer, D.I. (1983). Central America's tropical rain forests: positive steps for survival. Ambio 12: 232-238.

OEA (1978). Proyecto de desarrollo integrado de la región oriental de Panamá - Darién. Secretaría General de la Organización de los Estados Americanos (OEA), Washington, D.C. 308 pp.

Polsky, C. (1992). Crossroads of the continents. Nat. Conservancy (M/A): 14-21.

Porter, D.M. (1973). The vegetation of Panama: a review. In Graham, A. (ed.), Vegetation and vegetational history of northern Latin America. Elsevier, New York. Pp. 167-201.

Prebble, J. (1968). The Darien disaster. Penguin Books, Middlesex, England, U.K. 384 pp.

RARE (1983). Draft plan for the development of a private sector initiative in natural resource and environmental programs in the Republic of Panama. RARE Report, Panama City. 55 pp.

Sauer, C.O. (1966). The early Spanish Main. University California Press, Berkeley and Los Angeles. 306 pp.

Torres de Araúz, R. (1970). Human ecology of route 17 (Sasardí- Mortí) region, Darién, Panama. Battelle Memorial Institute, Columbus, Ohio. 200 pp.

Torres de Araúz, R. (1985). Etnobotánica Cuna. In D'Arcy, W.G. and Correa-A., M.D. (eds), The botany and natural history of Panama. Monogr. Syst. Bot. Missouri Bot. Gard. 10. Pp. 291-298.

Authors

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.) and the Asociación Nacional para la Conservación de la Naturaleza (ANCON) (Apartado 1387, Panama City 1, Panama).

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