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The Tambopata region (picture) of the south-western Amazon Basin and nearby Andes comprises the lowlands of the area south of the Madre de Dios River and east of the town of Puerto Maldonado in Madre de Dios Department, and the montane headwaters of the Tambopata River in Puno Department (Map 46). The region could be expanded phytogeographically to include parts of adjacent Acre in western Brazil and Pando and La Paz departments in northern Bolivia.
The montane portion of this region (500-3000 m) belongs to the Madre de Dios physiographic province (Young 1992). Geologically, it is characterized by very steep slopes and Early Palaeozoic and Early Tertiary sedimentary bedrock (Peñaherrera 1989). The information on the vegetation, the flora and conservation of this upland portion of the region are covered in the Data Sheet for the Eastern Slopes of the Peruvian Andes (CPD Site SA37).
The lowland portion of the Tambopata region, below 500 m, is characterized by gentle relief and Quaternary sediments. Rainfall at Puerto Maldonado is near 2000 mm annually and the 5-month dry season has 3 months that average less than 60 mm (ONERN 1972). The median annual temperature is 12°-13°C, so the Tambopata region is considered subtropical in the Holdridge life-zone system (ONERN 1976). During the austral winter, strong cold fronts ("friajes") are common from May to July; temperatures can drop to 7°C, which is a potential limiting factor for many tropical plants.
Another geographical distinction of this region is the absence of many of the habitats that characterize northern Upper Amazonia. The Tambopata region does not have white-sand soils and the various seasonally inundated types of forest called "tahuampa" in Peru and "várzea" and "igapó" in Brazil. Instead, the habitat mosaic is related to riverine dynamics and associated successional changes. Riverine meander loops may grow 25 m every year, implying that the entire meander belt is swept out and replaced by younger deposits every 500-1000 years (Terborgh 1990). There is evidence of these riverine processes in 26% of lowland Peruvian Amazonia (Salo et al. 1986).
In addition to the younger succession, there are more subtle vegetational differences, which are associated with relatively rich alluvial soils, older less fertile and leached clay soils, or presumably old and less fertile sandy-clay soils, each supporting distinctive floristic ensembles (Gentry 1988). Part of the habitat mosaic is also made up of a complex suite of sites with impeded drainage, ranging from oxbow lakes to swamp forests.
Open savannas of the Pampas del Heath occupy shallow basins along the Heath River on the Bolivian border. A variety of inceptisols constitute the soils, which are mostly acidic and with organic content ranging from 8.1% to 1.6% (Denevan 1980).
Above 500 m three upland subtropical forests occur: upper montane, lower montane and premontane (see Data Sheet for the Eastern Slopes of the Peruvian Andes, CPD Site SA37). The upper and lower montane forests occupy a narrow belt on steep slopes.
The dominant vegetation of the lowland portion of the Tambopata region is tropical moist forest. Since the area is at the south-western periphery of Amazonia, there is a strong dry season when many of the trees are deciduous. Nevertheless, the mature forest is physiognomically typical rain forest with a canopy 30 m high, frequent emergents (the most frequent being Bertholletia excelsa, the Brazil nut), and numerous palms, lianas and epiphytes.
Natural successional processes associated with changes in river channels are prevalent in much of Amazonia (Salo et al. 1986; Kalliola, Puhakka and Danjoy 1993). Similar to other sites in southern Amazonian Peru, the lowland portion of the Tambopata region is especially notorious for the conspicuous role of bamboo (mostly Guadua weberbaueri), which covers large patches (e.g. Terborgh 1985). In addition, where succession is further complicated by cyclical post-flowering die-offs, there are various younger successional stages along the meander belts of larger rivers. Successional growth in these areas typically consists of riverside strips dominated by Tessaria and Gynerium, followed by dominance of Cecropia membranacea, and then forests dominated by Ficus insipida and/or Cedrela odorata (e.g. Foster, Arce-B. and Wachter 1986; Salo et al. 1986; Foster 1990).
While part of the conspicuous habitat mosaic that characterizes north-western Amazonia is absent in Madre de Dios, there remain a number of conspicuously distinct types of vegetation other than the successional areas. At Tambopata, there are at least four main types of mature forest, associated respectively with alluvial soil, clayey soil, sandy-clay soil and poorly drained swamps (Erwin 1985). One-ha plots of trees and lianas 10 cm or more in dbh document the striking floristic differences. The overlap in species between pairs of plots on similar substrates is over 30%, but on different substrates less than 10% (Gentry 1988). About 600 such species occur in each plot - which is impressive, but significantly lower than in similar samples from the Iquitos region (CPD Site SA9). The overall number of species at Tambopata is high due to the many distinctive habitats.
Another important type of vegetation in the lowlands is the open savanna that occurs near the border with Bolivia, east of Palma Real River. This savanna is surrounded by or mixed with three other vegetation types: palm swamp forest (with Euterpe precatoria and Jessenia sp.), open scrubland dominated by Curatella americana and seasonal forest with Calophyllum (Denevan 1980).
The Tambopata region is botanically almost unexplored. The upland portion may comprise c. 140 families of vascular plants (León, Young and Brako 1992); notable are Alzateaceae, Arecaceae, Lauraceae, Moraceae and Rubiaceae (see CPD Site SA37, on Eastern Slopes of the Peruvian Andes).
The flora of the lowland forests includes nearly 2000 species. Records from the Tambopata Explorer's Inn Reserve (5.5 km²) show almost 1400 species of vascular plants (Reynel and Gentry, in prep.). At this locality, the two largest families are Leguminosae (106 spp.) and Rubiaceae (101 spp.), as is typical in much of Amazonia. Bignoniaceae, the predominant liana family, is third with 58 species, followed closely by Moraceae and ferns and their allies with 55 and 54 species. While these are always among the most speciose families and groups in Amazonia, the preeminence of Bignoniaceae is greater than at any other site for which comparable data are available. Euphorbiaceae, Melastomataceae and Piperaceae each have 44 species in the Florula list, whereas Araceae, palms, Solanaceae, Gramineae, orchids, Annonaceae, Lauraceae and Sapindaceae each have 28-33 species. Families with c. 20 species each include Meliaceae, Flacourtiaceae, Cyperaceae, Acanthaceae and Apocynaceae. Thus the familial composition of this Tambopata Reserve is typically Upper Amazonian (cf. Gentry 1990; Gentry and Ortiz-S. 1993).
At the generic level the flora of the Tambopata Explorer's Inn Reserve is also neotropically typical. The largest genera are Piper with 34 species and Psychotria and Inga with 32. These are the same genera that are most speciose on Barro Colorado Island, Panama and among the most speciose genera at all sites for which comparable data are available (Gentry 1990). Other especially speciose genera at Tambopata are Miconia, Solanum, Arrabidaea, Paullinia, Ficus and Philodendron. Of these, the 14 species of Arrabidaea are most distinctive, since it is usually the largest Bignoniaceae genus present but not among the most speciose genera at any other site known. While the data are very incomplete and not fully tabulated, c. 12% of the Tambopata Florula species may be endemic to the south-western part of Upper Amazonia (i.e. Madre de Dios in Peru, Acre in Brazil and parts of Pando and La Paz in Bolivia). Thus, even though its flora is typically Amazonian at the familial and generic levels, south-western Amazonia clearly warrants separate conservation focus.
Moreover, there are some striking floristic differences between Tambopata and Manu National Park (CPD Site SA11), which is also in Madre de Dios Department but farther north-west and nearer the base of the Andes. For example, Bertholletia excelsa is common at Tambopata but very rare at Manu; locally dominant species such as Lueheopsis hoehnei and Tabebuia insignis are unknown in Manu NP.
There are two additional floristically distinct habitats. The open savannas of the Pampas del Heath have a completely different flora. The majority of species of the Pampas del Heath are not represented in other areas of the Tambopata region and are otherwise unknown from Peru, although they are known from similar poorly drained habitats to the south. The foothill ridges along the Tavara River also have a flora that is quite different from that at the Tambopata Reserve (Conservation International Rapid Assessment Program team, unpublished data).
The upland portion of the Tambopata region was explored early by extractors of timber and quinine bark (Cinchona spp.); today only the former activity continues. Huge Podocarpus trees, perhaps the most valuable Andean timber, occur on the foothill ridges. A large number of species are commercialized for their timber, and saw-wood from at least 150 species mainly from the lowlands is sold in Puerto Maldonado. There are several wild conspecifics of cultivated fruit trees - including Theobroma cacao, Bactris aff. gasipaes, Pourouma cecropiifolia, Inga edulis, Pouteria macrophylla - mostly concentrated in alluvial-soil forests (Phillips et al. 1994). A wild pineapple (Ananas sp.) that might be the ancestor of the cultivated pineapple is common in the Pampas del Heath.
Harvesting of Brazil nuts constitutes a major local industry, the species occurring at low density (c. 1 tree per ha) in most of the lowland forest types. Rubber tapping was formerly important in the region and continues along the Brazilian border; the only rubber species in the reserve is probably Hevea guianensis (which produces inferior rubber to H. brasiliensis). Some local species provide useful fruits. Ethnobotanical studies indicate that some of the local "mestizos" know uses for up to 90% of the forest trees, mostly for subsistence construction, food and medicinal purposes (Phillips 1993; Phillips and Gentry 1993a, 1993b). The most useful wild species for local mestizo and indigenous peoples are palms, especially Iriartea deltoidea and Euterpe precatoria.
Social and environmental values
The human population in the lowlands of the Tambopata region consists of several indigenous groups (Arasaire, Ese-Eje, Huarayo), and an ethnically and culturally mixed group of recent migrants and longtime residents, mostly from the neighbouring highlands of Cuzco and Puno. The savanna in Pampas del Heath was inhabited by the Tiatinagua, a group considered to have gone extinct (Denevan 1980). Local settlers, especially the now largely acculturated Ese-Eje Amerindians, continue to harvest many products from the nearby forests, which are important to them not only economically but as part of their cultural heritage.
Living conditions in the region are at the medium poverty level. The agricultural products are maize, cassava, rice and plantains, which are sold in the local market or exported to Cuzco. All the major economic activities are extractive - gold mining and timber exploitation are the principal endeavours.
An important environmental value of the Tambopata-Candamo Reserved Zone is that it protects the entire drainage of a significant river system - one of the few places in the world where this has been established. Thus the now well-known values of natural forest in controlling flooding and erosion may be especially important here.
The South-east Peruvian lowlands Endemic Bird Area (EBA B30), which embraces the continuous lowlands from Manu south-eastward to the Tambopata region, is home to 15 restricted-range bird species, none of which occurs elsewhere. They are primarily confined to humid riverine and floodplain forests, with associated successional habitats also utilized. Two of these endemics are considered threatened, and the integrity of this reserve and the Manu region is essential for their long-term survival. The adjacent slopes of the Andes are home to a further 43 restricted-range species occurring in two EBAs.
Several tourist camps in operation along the Tambopata River and several on the Madre de Dios River near Puerto Maldonado call attention to the biotic richness of the region. The Tambopata Reserve is internationally famous as the site of the world's greatest lowland records of birds (554 spp., Parker 1991) and butterflies (1217 spp., Lamas, Robbins and Harvey 1991 and pers. comm.), and many of the tourists are drawn by this biological diversity. Puerto Maldonado proudly features signs proclaiming it the world's centre of biodiversity. The overall impact of tourism in the local economy is unknown.
Due to the importance of Brazil-nut harvesting to the local economy, this is one of the parts of Amazonia where extractive reserves might be most economically feasible. Unfortunately, political unrest, low market prices and transportation problems (the harvest is flown out) have all but destroyed this industry. Only one of the several Brazil-nut processing companies that once operated in Puerto Maldonado is still functioning.
While a recent study of the value of fruits and other natural products produced by the forest at Tambopata was disappointing as compared to some estimated potential values of Amazonian minor forest products, it did show that there are many valuable forest resources in the region and that harvest values could be substantially increased with appropriate technology and market development (Phillips 1993). The region's many wild congeners of cultivated crops are also valuable as sources of germplasm, although their actual economic worth is difficult to quantify. This region is a part of one of the major Vavilov centres of plant domestication, suggesting that it has special significance.
Subsistence agriculture and cattle-ranching have expanded in the region in recent years with a population explosion, largely due to immigration from the adjacent highlands. It is especially unfortunate that the alluvial-soil forests along the Tambopata River have been the most impacted by subsistence agriculture, as they have the most potential as subsistence extractive reserves. Gold mining has been the main attraction for migrants, and increasingly pollutes the Madre de Dios River and its major tributaries just west of the Tambopata region with mercury used for extraction.
Several valuable timber species are locally threatened due to over-exploitation, including Cedrela odorata, C. fissilis and Cedrelinga cateniformis. There are few incentives for the local people to conserve valuable species, in part because the loggers receive only a small fraction of market prices and in part due to the difficulty of obtaining secure land tenure. Even the original Tambopata Reserve (5.5 km²) adjacent to the Explorer's Inn Tourist Camp has been subjected to poaching of wild animals and timber trees.
The very interesting Pampas del Heath has been formally designated as a national sanctuary encompassing 1021 km². The Tambopata-Candamo Reserved Zone (picture) (14,000 km²) was formally declared in 1990; its status has yet to be clarified by the Peruvian Government. Many settlers live within the boundaries of this zone but there is little or no effective control of their activities. There is hope that a significant portion of the region will be established as a National Park.
Map 46. Tambopata Region, Peru (CPD Site SA10)
Denevan, W.M. (1980). The Rio Heath savannas of southeastern Peru. Geoscience and Man 21: 157-163.
Erwin, T.L. (1985). Tambopata Reserved Zone, Madre de Dios, Peru: history and description of the reserve. Rev. Peruana Entomología 27: 1-8.
Foster, R.B. (1990). Long-term change in the successional forest community of the Rio Manu floodplain. In Gentry, A.H. (ed.), Four neotropical rainforests. Yale University Press, New Haven. Pp. 565-572.
Foster, R.B., Arce-B., J. and Wachter, T.S. (1986). Dispersal and sequential plant communities in Amazonian Peru floodplain. In Estrada, A. and Fleming, T.H. (eds), Frugivores and seed dispersal. W. Junk, Dordrecht, The Netherlands. Pp. 357-370.
Gentry, A.H. (1988). Patterns of plant community diversity and floristic composition on environmental and geographical gradients. Ann. Missouri Bot. Gard. 75: 1-34.
Gentry, A.H. (1990). Floristic similarities and differences between southern Central America and Upper and Central Amazonia. In Gentry, A.H. (ed.), Four neotropical rainforests. Yale University Press, New Haven. Pp. 141-157.
Gentry, A.H. and Ortiz-S., R. (1993). Patrones de composición florística en la Amazonia peruana. In Kalliola, R., Puhakka, M. and Danjoy, W. (eds), Amazonia peruana vegetación húmeda tropical en el llano subandino. PAUT and ONERN. Jyväskylä, Finland. Pp. 155-166.
Kalliola, R., Puhakka, M. and Danjoy, W. (eds) (1993). Amazonia peruana vegetación húmeda tropical en el llano subandino. Proyecto Amazonia Universidad de Turku (PAUT) and Oficina Nacional de Evaluación de Recursos Naturales (ONERN). Jyväskylä, Finland. 265 pp.
Lamas, G., Robbins, R. and Harvey, D. (1991). A preliminary survey of the butterfly fauna of Pakitza, Parque Nacional Manu, Perú, with an estimate of its species richness. Publ. Museo Hist. Nat., Ser. A. Zool. 40: 1-19.
León, B., Young, K.R. and Brako, L. (1992). Análisis de la composición florística del bosque montano oriental del Perú. In Young, K.R. and Valencia, N. (eds), Biogeografía, ecología y conservación del bosque montano en el Perú. Mem. Museo Hist. Nat., Universidad Nacional Mayor de San Marcos (UNMSM), Lima. Vol. 21. Pp. 141-154.
ONERN (1972). Inventario, evaluación e integración de los recursos naturales de la zona de los ríos Inambari y Madre de Dios. Oficina Nacional de Evaluación de Recursos Naturales (ONERN), Lima.
ONERN (1976). Mapa ecológico del Perú. Guía explicativa. ONERN, Lima. 147 pp.
Parker, T.A., III (1991). Birds of Alto Madidi. In Parker, T.A., III and Bailey, B. (eds), A biological assessment of the Alto Madidi region and adjacent areas of Northwest Bolivia, May 18-June 15, 1990. RAP (Rapid Assessment Program) Working Papers 1. Conservation International, Washington, D.C. Pp. 21-23.
Peñaherrera, C. (1989). Atlas del Perú. Instituto Geográfico Nacional, Lima. 399 pp.
Phillips, O. (1993). The potential for harvesting fruit in tropical rainforests: new data from Amazonian Peru. Biodiversity and Conservation 2: 18-38
Phillips, O. and Gentry, A.H. (1993a). The useful plants of Tambopata, Peru I: statistical hypothesis tests with a new quantitative technique. Econ. Bot. 47: 15-32.
Phillips, O. and Gentry, A.H. (1993b). The useful plants of Tambopata, Peru II: additional hypothesis testing in quantitative ethnobotany. Econ. Bot. 47: 33-43.
Phillips, O., Gentry, A.H., Reynel, C., Wilken, P. and Gálvez-D., C. (1994). Quantitative ethnobotany and Amazonian conservation. Conserv. Biol. 8: 225-248.
Salo, J., Kalliola, R., Häkkinen, I., Mäkinen, Y., Niemelä, P., Puhakka, M. and Coley, P.D. (1986). River dynamics and the diversity of Amazon lowland forest. Nature 322: 254-258.
Terborgh, J. (1985). Habitat selection in Amazonian birds. In Cody, M.L. (ed.), Habitat selection in birds. Academic Press, New York. Pp. 311-338.
Terborgh, J. (1990). An overview of research at Cocha Cashu Biological Station. In Gentry, A.H. (ed.), Four neotropical rainforests. Yale University Press, New Haven. Pp. 48-59.
Young, K.R. (1992). Biogeography of the montane forest zone of the eastern slopes of Peru. In Young, K.R. and Valencia, N. (eds), Biogeografía, ecología y conservación del bosque montano en el Perú. Mem. Museo Hist. Nat., UNMSM, Lima. Vol. 21. Pp. 119-140.
This Data Sheet was written by the late Dr Alwyn H. Gentry and
Dra. Blanca León (Museo de Historia Natural, Apartado 14-0434, Lima- 14, Peru).
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