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PETEN REGION AND
The Department of Petén, which comprises the northern third of Guatemala, is one of the last remaining large wildland areas in Central America. Northern Petén is a plateau at an elevation of 200-400 m and forms the beginning of the Yucatán Peninsula of Mexico (Ferrusquía-Villafranca 1993). It is bounded to the south by a transverse chain of lakes extending eastward from near the Sierra del Lacandon (to 600 m elevation) and beyond Laguna Perdida to lakes Yaxhá and Sacnab. Central Petén, the low area between Lake Petén Itzá and the Subín River, encompasses only a small portion of the region. It is made up of Cretaceous limestone beds overlain with broad, deep clay. The area has many limestone hills and scattered sinkholes. Southern Petén for the most part is a broad and higher slightly undulating basin, except in the south-east where the Maya Mountains extend westward, descending from 1120 m in Belize - they are significant since the average elevation in the department is 200 m. The Petén is dominated by often thin soils of red clay ("tierra rosa") and black or dark brown clay ("rendzina"); humic gleys, grumusols and red-yellow podzols are interspersed throughout.
The Petén's main rivers - the San Pedro, Machaquilá (Santa Amelia) and La Pasión - flow westward through the department and empty into the Usumacinta River, which forms the western border with Mexico (see Lacandon Data Sheet, CPD Site MA1). The mean annual temperature is 26.5°C; the extremes are 12°C and 40°C. The mean annual precipitation ranges from 900-3500 mm, increasing from north-east to south. There is a pronounced dry season in January through April-May (Leyden 1984; Heinzman and Reining 1990; Schwartz 1990). Winds are stronger from February to June, and there are sporadic hurricanes.
The Maya Biosphere Reserve (15,000-16,000 km²) occupies the northern 40% of the Petén, encompassing nearly 10% of Guatemala's land area. It includes part of the municipalities La Libertad, San Andrés, San José, Flores and Melchor de Mencos. The landscape varies from gently undulating plains to karst topography with rounded to steep hills and narrow valleys. It is underlain by Early Tertiary limestone. Soils in the seasonally inundated lowlands are deep, poorly drained clay substrates, whereas in the uplands are shallow clay soils - neither are suitable for sustained low-input agricultural production. The precipitation averages 1200-1500 mm annually. The warmest period is April to September, with an average temperature of 32°C, and the coolest is November-January, with an average minimum of 20°C.
About 85% (30,000 km²) of the Petén was covered with semi-deciduous (seasonal) subtropical moist forest - the majority of the closed tropical broadleaved forest in Guatemala (cf. Nations and Komer 1984); less than 50% remains (Schwartz 1990). The northern Petén's vegetation (including the Maya BR) has much of the same flora as in the Yucatán Peninsula of Mexico and northern Belize. The Maya BR with the contiguous forests of Mexico and Belize (Map 24) is now Mesoamerica's last large lowland forest, c. 20,000 km². The canopy is 10-25 m high, being lower in seasonally flooded forest. Lundell (1937) defines three major tree associations within upland climax forest: (1) "ramonal" - groves of Brosimum alicastrum, especially found at sites of Maya ruins; (2) "caobal" - with Swietenia macrophylla (mahogany); and (3) "zapotal" - with Manilkara zapota, which is characteristic of dry rather than mesic upland forest. Epiphytes (e.g. orchids, aroids, bromeliads, cacti), ferns, bamboo and lianas are very abundant. In low-lying basins as at Laguna del Tigre, Laguna Perdida and lakes Petén Itzá and Yaxhá are swamps or marshes, which may be fringed by a "botanal" - with Sabal sp. Dense communities are found along rivers and the edges of lakes. Floating fern and sedge bogs occur in some lakes, and water-lilies in shallow open waters - the dominants are Nymphaea ampla and Nymphoides humboldtianum.
The central Petén has savanna with forested hills. Sinkholes are covered by herbaceous and subclimax forest (Lundell 1937). The savanna vegetation may have been created during the ancient Maya times of 3000-1700 BP (Leyden 1984), when they burned the forest for cultivation; continued burning prevents the forest from re-establishing. The savanna supports a diverse and complex herbaceous flora, most of which is fire resistant. The grasslands are surrounded by a barrier of scrub that acts as a buffer, protecting the mesophytic forest from fire.
The southern Petén, which is the least explored botanically, has much of the same vegetation as the northern Petén but some species are less common, such as Manilkara zapota, perhaps replaced by M. chicle; Swietenia macrophylla has been logged extensively (Lundell 1937; Nations and Komer 1984). Species of Sapotaceae are most characteristic of this wetter area. The vegetation is allied more with southern Guatemala and southern Belize. Extensive aquatic and riparian vegetation occurs.
The tropical rain forests of Guatemala are of special value because their presence in northern Central America and southern Mexico is particularly complex. Guatemala has some rain forests no older than 10,000-11,000 years, and some in the Petén are considerably younger (1000 years old) due to former Mayan disturbance (Binford et al. 1987). Late Glacial vegetation consisted of marsh, savanna and juniper scrub. The Petén thus was not a Pleistocene refugium for mesophytic taxa (Leyden 1984), but the adjacent southern Izabal area of Guatemala and Belize may have been (Wendt 1993).
The Petén floristic associations are continuous with associations in the Mexican Lacandon region (CPD Site MA1). Since much of the Petén is not well known botanically, it is difficult to estimate the number of vascular plants or endemics. The floristic diversity of the Maya BR is considered exceptional, with over 3000 plant species (CONAP 1992). The overall region's flora is considered distinctive; e.g. many of the Petén's regionally endemic taxa are shared to varying extents with northern Belize and Mexico's majority of the Yucatán Peninsula to eastern Tabasco and the eastern highlands of Chiapas (Breedlove 1981; Wendt 1993). Most local Petén endemics have been found in areas that allow little human intervention, such as steep hills and swamps (Lundell 1937).
Information on the distribution of plants in the Petén, for example to determine which are the rare species, is still very limited and incomplete. Subtropical forests contain relatively low tree-species diversity and a higher number of individuals per species (Salafsky, Dugelby and Terborgh 1993). The high densities of species such as Manilkara zapota, Chamaedorea spp. and Pimenta dioica facilitate extractive industry.
The Petén region is rich in serviceable plants, for example thatching palms, construction materials, fuelwood, fibres - e.g. Desmoncus sp. ("bayal") and Philodendron sp. ("mimbre") for basketry and furniture, forest fruits, medicinal plants and species marketable from upland forests, such as Manilkara zapota ("chicozapote"), Chamaedorea spp. (mostly two understorey palms) and Pimenta dioica. A few studies have analysed the economic benefits of a conserving, sustainable use of Guatemala's tropical forests and renewable resources (Nations et al. 1988; Reining and Heinzman 1992; Salafsky, Dugelby and Terborgh 1993). An estimated 80% of the hardwoods in Guatemala occur in the Petén, such as Swietenia macrophylla, Cedrela odorata, Calophyllum brasiliense var. rekoi, Pouteria spp., Bursera simaruba, Spondias and Ficus (Leyden 1984). The Maya BR contains more than 300 species of useful trees (CONAP 1990).
A potentially important forest resource is Brosimum alicastrum ("ramón") - a common tree occasionally up to 30 m tall, which may have been nurtured by the Maya (Leyden 1984). The seeds were an important food source in pre-Columbian times, but present human consumption is quite low (Heinzman and Reining 1990); the fruits, foliage and bark are gathered as forage for mules and horses. These parts are rich in protein and other essential nutrients.
Economic assessment of non-timber forest products
Heinzman and Reining (1990) analysed some potentially sustainable rural extraction practices in the northern Petén. Collecting several products more or less sustainably for export represents a wage resource for over 6000 people who otherwise subsist mainly on slash-and-burn ("milpa") agriculture (see Lacandon Data Sheet, CPD Site MA1), but only 13% of the Petén soils are deep and well drained. The total economic return from these non-timber forest products is greater than if the forest were converted to pasture (Nations et al. 1988; Heinzman and Reining 1990).
In 1990 Guatemala passed a law (Decree 5-90) for a Maya Biosphere Reserve, designating 7500 km² of the BR for extractive industry based on non-timber forest products. During the past 30-100 years, three such major products have been harvested: (1) Millions of "xate" palm leaves (Chamaedorea elegans, C. oblongata) are exported through the year for greenery in floral arrangements. Xate produces US$4-6 million annually (Morell 1990). (2) Chicle, the latex of Manilkara zapota (a tree to more than 30 m tall), is extracted for the manufacture of chewing gum. The largest concentration of high-grade chicle is found in the Maya BR and sold primarily to the Japanese. Small quantities of latex from Ficus lundellii, Bumelia mayana and Stemmadenia donnell-smithii may be used as enhancing supplements. In 1990-1991 the high quality latex sold for US$3.75 per kg (Reining and Heinzman 1992). (3) Another important annual product harvested on a rather sustainable basis is allspice ("pimienta gorda", Pimenta dioica), a common tree rarely over 20 m tall. Currently Guatemala (and Mexico) supply almost 30% of the international market.
An important aspect of harvesting these renewable resources is that they promote conservation of the forest as well as providing income locally. A family can earn three times more as an average daily wage from the forest products than from clearing forest, planting maize and raising cattle. Heinzman and Reining (1990) recommend development of institutions to ensure sustainable use of these common resources, and to further diversify in non-timber forest products that are subject to sustainable exploitation.
Social and environmental values
The Petén has a rich fauna which is poorly known. The country's list so far includes 1453 vertebrate species (not including saltwater fishes); at least 333 bird species occur in the Petén. The Petén wetlands provide significant wintering grounds for many North American migratory bird species. The Laguna del Tigre complex of diverse wetlands represents one of the most extensive freshwater wetland areas in Central America, of which 484 km² are recognized under the world's RAMSAR convention.
The Petén is an important refuge for many species, such as howler monkey, ocelot, margay cat, jaguar, puma, Baird's tapir, harpy eagle, macaws, Moreletti's and American crocodiles, iguana, beaded lizard and boa constrictor. About 133 of the animal species are considered threatened; some species are listed in CITES appendices as at risk from international trade (URL 1984). No globally threatened bird species or Endemic Bird Areas are in this rain forest. Nevertheless, the area is of national importance for a number of species of birds of prey, including the near-threatened orange-breasted falcon (Falco deiroleucus).
Other economic benefits of conserving the biological diversity of the Petén include stabilization of hydrological functions, soil protection, tourism and the opportunities to create jobs. Tourism is a large and growing industry vital to the economy of Guatemala - in 1990 the country earned US$185 million. Tours in the Petén for nature and archaeological interest show that there is strong potential. In 1970 an all-weather road opened the central Petén to southern Guatemala. Tikal National Park (picture) draws 15% of the tourists who visit the country; it is estimated that 2500 archaeological sites occur in the Petén, perhaps half in the Maya BR. Several Maya peoples live in the Petén, e.g. the Lacandon, Itzá and Mopán.
The combination of the tropical forest environment and Maya history and culture is most appealing and should be conserved and promoted. A Mundo Maya (Maya World) or Ruta Maya (Maya Trail) is being organized, similar to the Inca Trail in South America. The Mundo Maya will connect the major archaeological sites and National Parks of Mexico, Guatemala, Belize, Honduras and El Salvador (Hagman 1989).
There are many threats to the vegetation and flora of the Petén, including colonization, road building, logging, fire, ranching cattle and pigs, oil exploration and over-collecting (D'Arcy 1977; Schwartz 1990; Stuart 1992). The population, which for decades was 15,000 persons or less in Petén Department (c. 36,000 km²), rose to 65,000 in 1973 and presently has reached over 300,000 people, increasing at the annual rate of 5.5% (compared to 2.9% for the rest of the nation) (Southgate and Basterrechea 1992). Colonists have included "ladinos" (mestizos) from elsewhere in Guatemala, refugees from El Salvador and Kekchí Amerindians from Alta Verapaz Department. The region is becoming more accessible with construction of a major road linking the Petén with Guatemala City and Belize (Prensa Libre 1991).
Agricultural expansion and logging are the major reasons for forest loss. The rate of deforestation in the Petén is very high in the last 10-15 years, 200-400 km² have been converted annually. Colonization is occurring in the western Petén and the Franja Transversal del Norte, which stretches along the southern Petén border. Two government agencies are granting parcels of land between 20 ha and 100 ha in the department for colonization. Milpa cultivation is destroying forest because the increasing human populations are not allowing adequate fallow periods for forest regeneration (cf. Lundell 1937). Severe erosion is evident in some places, e.g. the south-eastern Petén (Schwartz 1987, 1990).
Although logging permits within the Maya BR were revoked in March 1991, some residents are illegally cutting mahogany (Swietenia macrophylla) for construction or exporting the wood to Belize and Mexico for income; this valuable species is becoming scarce (Moser et al. 1975; Stuart 1992). Chamaedorea (xate) densities are being reduced (especially C. elegans) by over-collecting, and Manilkara zapota also probably has declined during the past 100 years. Inadequate transportation in the Petén still prevents large-scale cattle production, but there is some cattle-raising near Tikal and in the south-eastern and west-central portions of the department. Most of the cattle are shipped to Belize to be transported and sold in Mexico. Oil and mineral explorations have destroyed some forest with construction of roads and test sites; these threats are not intense, as the anticipated amounts of oil and mineral strikes have not been found.
Mexico and Guatemala have considered constructing a hydroelectric dam on the Usumacinta River. The proposed dam would have flooded more than 1000 km² of the Petén forest. The likely sediment load in the river would have required check dams farther upstream to keep the main dam from silting up (Nations and Komer 1984; Wilkerson 1985). The project was halted in 1992 by Mexico's president (see the Lacandon Data Sheet, CPD Site MA1).
Between 1955 and 1988 Guatemala declared 52 conservation areas, although the majority did not meet international criteria for protected areas. Areas were reserved for scenic landscapes, pre-Columbian Maya ruins and particular plant or animal species (biotopes) (Godoy 1988). For the last three decades, Guatemalan conservationists have been aware that much more of the country's biological diversity needed to be conserved and managed, and they recommended the creation of other reserves (Godoy 1987a, 1987b). A regional network of protected areas has been recommended for Petén Department - the Integrated System of Protected Areas of the Petén (SIAP). SIAP would comprise National Parks, Forest Reserves, Wildlife Refuges and other types of protected areas. These areas have been put in priority order for the instigation and development of conservation measures (Godoy and Castro 1990).
In 1989 Guatemala passed a Forestry Law (Decree No. 70-89) due to the increasing degradation of forests, and stated the importance of protecting and renovating forest resources. A new agency was created to administer and manage forest resources, wild life and some National Parks - the Dirección General de Bosques y Vida Silvestre (DIGEBOS), incorporating the former Instituto Nacional Forestal (INAFOR). Also in 1989, a significant step was made towards increasing the number and effectiveness of conservation units with passage of a Protected Areas Law (Decree No. 4-89). An extensive national system of conservation units was created as the Guatemalan System of Protected Areas (SIGAP) and a new organization was established to manage them - the Consejo Nacional de Areas Protegidas (CONAP) (cf. Nations et al. 1988). This law established boundaries for 6 existing reserves and created 44 new Special Protection Areas (TNC 1989). About 44.4% of these areas are in the Petén (WCMC 1992) c. 14,000 km² of the area north of 17°10'N and the Sierra del Lacandon in the western Petén.
In 1990, the Maya Biosphere Reserve was established (see Map 24). In three grouped areas, it has five National Parks and three reserves (biotopes): (1) Laguna del Tigre NP and Laguna del Tigre-Río Escondido Biotope (3508 km²), with Sierra del Lacandon NP nearby (1950 km²); (2) El Mirador NP, Dos Lagunas Biotope and Río Azul NP (1470 km²); and (3) Tikal NP (573 km²) and the adjacent San Miguel La Palotada (El Zotz) Biotope (435 km²). A multiple-use area (7500 km²) adjoins these protected areas for their protection and for the management of renewable forest resources, and next to the BR's southern boundary is a 10-15 km² wide buffer zone (250 km²).
The Maya BR is cooperatively administered: the lead agency is CONAP; other key participants include the Centro de Estudios Conservacionistas (CECON) the academic unit of the Universidad de San Carlos de Guatemala that is responsible for promoting field research and conservation of renewable resources; and the Instituto de Antropología e Historia (IDAEH) the government institution responsible for administering the marvellous cultural heritage. The Maya BR is one of 11 areas given priority under the region's 1992 Convenio para la Conservación de la Biodiversidad y Protección de Areas Silvestres Prioritarias en América Central.
The Maya BR is among the key sites in The Nature Conservancy (TNC) Parks in Peril campaign, to build a conservation infrastructure and secure long-term funding to sustain local management of the protected areas and integrate them into local economies (Houseal 1990). The USAID Maya Resource Management project (MAYAREMA) is offering financial and technical assistance to CONAP to manage the resources of the BR more sustainably.
The Maya BR abuts two neighbouring protected areas: in Mexico, Calakmul Biosphere Reserve (8250 km²); in Belize, Río Bravo Conservation Area (610 km²). The U.S. Man and the Biosphere Program in 1992 approved a proposal to develop a regional approach for sustainable development and conservation of natural resources in the Maya tri-national region of Mexico, Guatemala and Belize. Paseo Pantera, a consortium of U.S. and Central American non-governmental and governmental organizations and institutions, helps regionally to enhance wildlands management and preserve natural diversity by means of a biological corridor from Guatemala through Panama, and Tikal NP is one of the five key areas in as many countries for its pilot work (Jukofsky 1992; Marynowski 1993).
Other international conservation organizations are supporting the Guatemalan efforts, including the Center for International Development and Environment/World Resources Institute (Nations et al. 1988), WWF - U.S. (1989; Cohn 1989) and Conservation International - which is aiding Guatemalan decision-makers and local communities by promoting low-impact tourism and developing markets for sustainably harvested products (CI 1991). The Nature Conservancy is preparing technical studies on 14 protection areas in the Petén to secure their status as National Parks and has established a Conservation Data Centre in cooperation with the Universidad de San Carlos (TNC 1989).
Map 24. Petén Region and Maya Biosphere Reserve, Guatemala (CPD Site MA13) and adjacent areas
Binford, M.W., Brenner, M., Whitmore, T.J., Higuera-Gundy, A., Deevey, E.S. and Leyden, B.W. (1987). Ecosystems, paleoecology and human disturbance in subtropical and tropical America. Quat. Sci. Rev. 6: 115-128.
Breedlove, D.E. (1981). Introduction to the Flora of Chiapas. In Breedlove, D.E. (ed.), Flora of Chiapas. Part 1. California Academy of Sciences, San Francisco. 35 pp.
CI (1991). CI launches program in the Maya heartland. Tropicus 5(3): 3.
Cohn, J.P. (1989). A will to protect. Américas 41(2): 46-53.
CONAP (1990). Reserva de la Biósfera Maya. Consejo Nacional de Areas Protegidas (CONAP), Guatemala City. 8 pp.
CONAP (1992). Plan maestro de la Reserva de la Biósfera Maya RBM. CONAP, Guatemala City. 25 pp.
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.
Ferrusquía-Villafranca, I. (1993). Geology of Mexico: a synopsis. In Ramamoorthy, T.P., Bye, R., Lot, A. and Fa, J.E. (eds), Biological diversity of Mexico: origins and distribution. Oxford University Press, New York. Pp. 3-107.
Godoy, J.C. (1987a). Areas silvestres protegidas potenciales de El Petén, Guatemala. Perspectiva 8(1): 166-178.
Godoy, J.C. (1987b). Memorias minitaller de áreas silvestres protegidas. Centro de Estudios Conservacionistas (CECON), Centro Agronómico Tropical de Investigación y Enseñanza (CATIE) and WWF U.S. Guatemala City. 21 pp.
Godoy, J.C. (1988). Algunas consideraciones sobre áreas silvestres protegidas fronterizas. CECON and CONAMA (Comisión Nacional del Medio Ambiente), Guatemala City. 3 pp.
Godoy, J.C. and Castro, F. (1990). Plan estratégico del Sistema de Areas Protegidas de El Petén, Guatemala (SIAP). Proyecto de Conservación para el Desarrollo Sostenido en América Central. CATIE y el Unión Internacional para la Conservación de la Naturaleza y los Recursos Naturales (UICN). Turrialba, Costa Rica. 105 pp.
Hagman, H. (1989). Maya plan for tourists is, at heart, for ecology. Washington Times (9 October): E1, E5.
Heinzman, R.M. and Reining, C.C.S. (1990). Sustained rural development: extractive forest reserves in the northern Petén of Guatemala. Trop. Resources Inst. Working Paper No. 37, Yale School of Forestry and Environmental Studies, New Haven.
Houseal, B. (1990). Maya riches. Nature Conservancy Mag. 40(3): 16-21.
Jukofsky, D. (1992). Path of the Panther. Wildlife Conservation 95(5): 18-24.
Leyden, B.W. (1984). Guatemalan forest synthesis after Pleistocene aridity. Proc. Natl. Acad. Sci. 81: 4856-4859.
Lundell, C.L. (1937). The vegetation of Petén. Carnegie Institution of Washington Public. No. 478, Washington, D.C. 244 pp.
Marynowski, S. (1993). Paseo Pantera: the great American biotic interchange. Wild Earth (Special Issue): 71-74.
Morell, V. (1990). Bringing home a piece of the jungle. Int. Wildlife 20(5): 12-15.
Moser, D. and the editors of Time-Life Books (1975). Central American jungles. Time-Life Books, New York. 184 pp.
Nations, J.D and Komer, D.I. (1984). Conservation in Guatemala. Center for Human Ecology, Austin, Texas. 17O pp.
Nations, J.D., Houseal, B., Ponciano, I., Billy, S., Godoy, J.C., Castro, F., Miller, G., Rose, D., Rosa, M.R. and Azurdia, C. (1988). Biodiversity in Guatemala: biological diversity and tropical forest assessment. Center for International Development and Environment/World Resources Institute, Washington, D.C. 185 pp.
Prensa Libre (13/9/1991). Carretera a Petén será una realidad.
Reining, C.C.S. and Heinzman, R.M. (1992). Nontimber forest products in the Petén, Guatemala: why extractive reserves are critical for both conservation and development. In Plotkin, M. and Famolare, L. (eds), Sustainable harvest and marketing of rain forest products. Island Press, Washington, D.C. Pp. 110-117.
Salafsky, N., Dugelby, B. and Terborgh, J. (1993). Can extractive reserves save the rain forest? An ecological and socioeconomic comparison of nontimber forest product extraction systems in Petén, Guatemala, and West Kalimantan, Indonesia. Conserv. Biol. 7(1): 39-52.
Schwartz, N.B. (1987). Colonization of northern Guatemala: the Petén. J. Anthr. Res. 43: 164-183.
Schwartz, N.B. (1990). Forest society: a social history of Petén, Guatemala. University Pennsylvania Press, Philadelphia. 367 pp.
Southgate, D. and Basterrecha, M. (1992). Population growth, public policy and resource degradation: the case of Guatemala. Ambio 21: 460-464.
Stuart, G.E. (1992). Maya heartland under siege. Natl. Geog. 182(5): 95-106.
TNC (The Nature Conservancy) (1989). Guatemala enacts landmark conservation law. Nature Conservancy Mag. 39(3): 36.
URL (1984). Perfil ambiental de la República de Guatemala, Tomo II. Universidad Rafael Landívar (URL), Instituto de Ciencias Ambientales y Tecnología Agrícola (ICATA). URL/USAID-Guatemala-ROCAP Contract No. 596-0000-C-00-3060-00, Guatemala City. 249 pp.
WCMC (World Conservation Monitoring Centre) (1992). Protected areas of the world. A review of national systems. Vol. 4: Nearctic and Neotropical. IUCN, Gland, Switzerland and Cambridge, U.K. 459 pp.
Wendt, T. (1993). Composition, floristic affinities, and origins of the canopy tree flora of the Mexican Atlantic slope rain forests. In Ramamoorthy, T.P., Bye, R., Lot, A. and Fa, J.E. (eds), Biological diversity of Mexico: origins and distribution. Oxford University Press, London. Pp. 595-680.
Wilkerson, J.K. (1985). The Usumacinta River: troubles on a wild frontier. Natl. Geog. 168(4): 514-543.
WWF-US (1989). Latin America and the Caribbean new and ongoing projects. In WWF-US 1989/90 Program. Washington, D.C. Pp. 267-275.
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 Jane Villa-Lobos (Smithsonian Institution, Department of Botany, NHB- 166, Washington, DC 20560, U.S.A.).
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