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Amazonia: CPD Site SA5

MANAUS REGION
Brazil

Location:  In western Central Amazonia, a circle with 150-km radius around Manaus, between latitudes 1°30'-3°00'S and longitudes 58°15'-61°00'W.
Area: 
c. 70,700 km².
Altitude: 
.
16-130 m.
Vegetation: 
Several types of rain forests on non-flooded and in variously inundated areas, forested to open shrubby formations on white sandy soils.
Flora: 
High diversity, especially of tree species; high endemism.
Useful plants: 
Genetic reserves, e.g. for timber trees; edible fruits, medicinal plants.
Other values: 
Fauna refuge, watershed protection, research stations, tourism.
Threats: 
Road building; colonization; fire; conversion to pasture or plantation; removal of sand and fuelwood.
Conservation: 
Ecological Research Stations, Forest Reserves, Environmental Protection Zone (APA); nearby National Park, Amerindian reserve, Biological Reserve.

Map 41: CPD Site SA5
References

Geography

The Manaus region is in western Central Amazonia in Brazil's northern State of Amazonas, and includes the lower drainage area of the Rio Negro and its confluence with the Solimões River, which extends eastward as the Amazon River. Habitats have been selected within a circle with a radius of 150 km from the city of Manaus (3°S, 60°W), near the Rio Negro's mouth (Map 41) (G.T. Prance, E. Lleras and B.W. Nelson, pers. comm., Workshop 90, Manaus).

The mean annual temperature is 27.2°C, and the maximum is 37.8°C (Junk 1983; Leopoldo et al. 1987). The mean annual precipitation at Manaus is 2100 mm, with 73% received as heavy rains. About 26% of the water runs off, 20% becomes groundwater and the rest is evaporated and transpired by the vegetation, usually to recycle as rain in c. 5.5 days. The Lower Amazon Basin is somewhat drier from July to October (Daly and Prance 1989; Leopoldo et al. 1987).

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Vegetation

Three main types of vegetation occur in the vicinity of Manaus: (1) rain forests on "terra firme" (non-flooding areas), as well as small terra-firme savannas on compacted clay soil; (2) forests of inundated areas, as well as grassland and floating mats; and (3) formations on white-sand soils (Prance 1987b).

1. Rain forests on terra firme (uplands)
This vegetation type covers c. 50% of Amazonia (Prance 1987b); the Manaus forests are typical of these Central Amazonian forests that have developed on the dominant clayey latosolic sediments of Tertiary plateaux and slopes. The soils are well structured and well drained but of poor fertility (Brickmann 1989). Some differences in structure and composition occur according to topography: tall terra-firme rain forest on clay soil, Amazonian caatinga forest (campinarana) and open Amazonian caatinga scrub (campina) on small patches of hydromorphic podzols, and 130 km north of Manaus, campinas and campinaranas on sandstone.

Two main terra-firme subunits are recognized: (i) terra-firme lowland forest on rather flat areas, which presents the characteristic heterogenous climax; and (ii) terra-firme hill forest, above 250 m (e.g. north of the Manaus region). Locally, in addition to forest on rather flat areas, there are distinctive forests on slopes (Brickmann 1989; Guillaumet 1987; Prance 1987b).

The climax forest has large biomass, a high closed canopy, high species diversity with much local variation in composition, large lianas (of somewhat limited frequency) and a relatively sparse herbaceous ground cover. Four layers may be detected: the canopy at 30-35 m with a few emergents to 44 m, a stratum of 12-15 m tall trees, a stratum of small trees and shrubs 7-12 m tall and a stratum of low shrubs and saplings to 7 m (Guillaumet 1987).

Guillaumet (1987) emphasized the difficulty in comparing the few site inventories made in the region. He lists the 12 tallest tree species (35-44 m), those of largest dbh (c. 20 species over 85-200 cm) and the density of trees of various sizes. About 20% of the larger trees have buttresses. Characteristically, palms are abundant and diverse in several strata. At one site, in each of seven families more than 10 species occurred, with the most in Lecythidaceae (18 spp.), Moraceae (15) and Sapotaceae (14). Typically, no species of tree is common; among 235 species of trees in the area, the most frequent species (individuals per ha) at the site were Eschweilera coriacea (only 26) and Scleronema micranthum (only 9) (Prance, Rodrigues and Silva 1976). Lianas especially of Leguminosae were frequent at one site. There are abundant epiphytes, especially ferns, Bromeliaceae and Orchidaceae. The forest is particularly rich in hemi-epiphytes and pseudo-epiphytes such as species of Philodendron, Heteropsis and Anthurium (Araceae) and seven other families. Stranglers of Ficus spp. and Clusiaceae are abundant but not necessarily strangling their host trees. About 12 genera of semi-parasitic Loranthaceae are frequent in the canopy. Flowering plant saprophytes (Burmanniaceae, Gentianaceae, Triuridaceae) are also rich in species.

2. Inundated forests, grassland and floating vegetation
There are substantial Amazonian areas of both permanently and periodically (e.g. annually) flooded forests. Species diversity is less than in terra-firme forests, with less regional endemism (Prance 1987b). The permanent types are: (i) permanent swamp forest of white water; and (ii) permanent "igapó" of black or clear water. The underlying soil is eutrophic humic gley; in some areas of dystrophic humic gley, palm swamps occur. Periodically flooded forests include: (iii) Seasonal "várzea", which is floodplain forest periodically flooded by fertile muddy-white waters, e.g. along the Solimões and Amazon floodplains. It is similar to the upland forest, with fewer species and more lianas. (iv) Seasonal igapó, which is floodplain forest periodically flooded by low-fertility black or clear water, e.g. along the Rio Negro. There are fewer, often different species than in seasonal várzea, and many of the trees have scleromorphic adaptations. Last, (v) sporadic floodplain forest occurs where there are quickly draining or flash floods at irregular times. This type is mainly in the upper portions of rivers and narrow streams and has greater species diversity, since many terra-firme species can survive such limited inundation.

The total flooded area of the Rio Negro and its tributaries probably exceeds 2000 km², and includes permanent and seasonal igapó, and sporadic floodplain forest (Goulding 1988). Annual fluctuation of the river-level is 9-12 m, with the inundation lasting 4-5 months along the upper and middle river and 7 months along the lower Rio Negro; the highest water is in June or July. Although data are limited, the annual variability appears large. Rarely forests remain flooded for 2-3 years, but this causes tree and shrub mortality; annual minimum and maximum fluctuations recorded at Manaus have been 5.5 m and 14 m (Goulding 1988; Irmler 1977). However, between November and February while the level of the Upper Rio Negro is falling with the dry season, the lower river is rising because it is dammed back by the rising Solimões-Amazon River (Goulding 1988), which results in somewhat intermediate vegetation from the influx of white water (Irmler 1977; Prance 1979).

Inundated forests have been described by Prance (1979, 1989; see also Junk 1983). Seasonal igapó has limited species diversity and generally lower biomass, since it is on poor sandy soils along riverbanks and the water lacks nutrients; the forest is usually not extensive and often is interspersed with open beaches. A few studies have been done on igapó vegetation near Manaus (e.g. Prance 1979, 1989; Keel and Prance 1979; Rodrigues 1961). The canopy is lower and the species tend to be different from those of várzea or terra firme. Only 54 species of trees and shrubs were found at one site. The most common trees are in Myrtaceae, such as Eugenia inundata; other common trees are Alchornea castaneifolia, Copaifera martii, Piranhea trifoliata and Triplaris surinamensis (Pires and Prance 1985; Prance 1989). Zonation of species occurs gradually along the moisture gradient away from the river from heavily to lightly flooded (Keel and Prance 1979).

Seasonal várzea is the most widespread type of inundated forest in Amazonia, often extending several km back from the riverbank (cf. Junk 1984, 1989). In Lower Amazonia it is associated with robust grass meadows (Pires and Prance 1985) and in the Manaus region with the Solimões River. Some characteristic trees of seasonal várzea around Manaus include Astrocaryum jauari, Calycophyllum spruceanum, Carapa guianensis, Ceiba pentandra, Hevea brasiliensis, Hura crepitans and Macrolobium acaciifolium. The herbaceous understorey is rich in individuals of Heliconia and Costus.

During the high-water season, vast mats of vegetation form on the extensive lakes and lake-like expanses where riverbanks often are completely inundated. Mats may cover several km² within a few months, yet with over 90% of the species reproducing and dying by the subsequent dry season. In white-water lakes and swamps that do not dry up, vegetational succession progresses from floating plants such as Eichhornia crassipes and Salvinia auriculata to a secondary community of grasses and sedges. Montrichardia arborescens, a tree-like Araceae to 6 m or more tall, then may colonize the mat, compacting it into a floating community from 20 cm above to partly 100 cm below the surface. Climbers such as Ipomoea spp. and even trees to 6-8 m (Bombax, Cassia, Cecropia, Ficus) may occur. Lago dos Patos (c. 100 km from Manaus) has a floating mat 200 m wide, 3 km long and more than 4 m thick. Usually, however, the mats sink at 1 m thickness or less (Junk 1983).

3. Formations on white-sand soils
Interspersed within lowland terra-firme rain forest of the Rio Negro region are white-sand "campinas", which are islands of open shrub to low-forest formations on podzols of leached quartz-sand soils or regosols. Campinas occur from the headwaters of the Rio Negro to Manaus near the Solimões-Amazon River (Goulding 1985), and in large or small areas scattered over much of Amazonia (Prance 1987b). Campinas often gradually grade (physiognomically and floristically) into the surrounding forest. These soils are not only extremely poor in nutrients, but too porous to retain water. The extreme ecological restrictions have resulted in a continuum of highly distinctive formations, generally characterized by scleromorphic leaves, gnarled trunks and unusual floristic composition (Macedo and Prance 1978). The formations may cause the black water, because they are unable to filter out humic acids from organic decomposition (Anderson 1981; Anderson, Prance and Albuquerque 1975; Goulding, Carvalho and Ferreira 1988).

There are no accurate estimates of the total area of Amazonia or the Rio Negro Basin occupied by these formations, but it is considerable - thousands of km². There are four phases to the vegetation: savanna; scrub; woodland with a patchy canopy 5-15 m high (with emergents to 20 m tall); and forest with a generally continuous canopy 20 m to almost 30 m high. The campina-forest areas around Manaus are lower (up to 15 m) and poorer in species than the immense relatively rich areas (especially in understorey and ground-cover species) found in the Upper Rio Negro region (CPD Site SA6) (Anderson 1981; Prance 1987b). Around Manaus the campina forest is dominated by tree species such as Aldina heterophylla, Glycoxylon inophyllum and Humiria balsamifera. The twisted, much-branched trees are loaded with many epiphytic orchids, bromeliads, Araceae, Gesneriaceae (e.g. Codonanthe) and pteridophytes. The many open areas of sand are covered by the blue-green alga Stigonema tomentosum. Lichens and mosses are abundant on branches and the soil surface. Sometimes the ground is covered with a spongy mat of Cladonia; Sphagnum is less frequent (Pires and Prance 1985; Prance 1987b, 1989).

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Flora

Being at the confluence of the Negro and Amazon rivers, the Manaus region has extensive periodically flooded habitats (igapó and várzea), each with a mosaic of communities determined by the duration of flooding, water chemistry and sedimentation characteristics. The modern-age dry belt east of Manaus cutting north-west to south-east across the Amazon River (CPD Site SA4) may impede the eastward migration of species, causing decreased diversity near Belém (in the State of Pará) relative to Manaus and westward from Manaus. Finally, the first direct dated evidence for a dry Pleistocene Amazon, in Acre (Kronberg, Benchimol and Bird 1991) and at Carajás (Absy et al. 1989), as well as indirect evidence for relative aridity on the Guayanan and Brazilian shields (Veiga 1991), lend new weight to the Pleistocene refugium hypothesis (see Whitmore and Prance 1987) as an explanation for high a diversity, if Manaus was spared this dryness.

Each of the vegetation types is more or less floristically distinct, so there is high [beta] diversity within this small circle. Total floristic diversity is unknown, but a diversity in terra-firme tree plots is high at c. 200-230 spp./ha (of trees with dbh 10 cm or more) and the total tree flora is well over 1000 species. The terra-firme list (represented by the Ducke Forest Reserve) is dominated by the speciose tree genera Licania, Inga, Protium, Eschweilera, Swartzia, Aniba, Miconia, Ocotea, Casearia and Couepia. Apparently there is high non-edaphic endemism in several families (e.g. Lecythidaceae, Chrysobalanaceae, Bignoniaceae, possibly Sapotaceae), and one endemic family - Duckeodendraceae.

The Manaus region is probably a centre of high diversity and possibly also local endemism. Species from western Amazonia, the Guianas and eastern Amazonia overlap, making this a priority area for conservation. There is a unique concentration of diversity in tree taxa, and families not speciose elsewhere: Chrysobalanaceae, Burseraceae, Lecythidaceae and Vochysiaceae.

The forests around Manaus and Belém are the most heavily collected sites in Amazonia (Daly and Prance 1989). Species lists have been compiled for some sites around Manaus and some detailed information has been gathered on composition and distribution (e.g. Prance, Rodrigues and Silva 1976; Anderson, Prance and Albuquerque 1975; Rodrigues 1961, 1967). The most important feature of the tropical rain forest on terra firme is its outstanding species diversity. Many studies have shown this richness throughout the region, with variation according to local rainfall. For example, Prance, Rodrigues and Silva (1976) in 1 ha of forest near Manaus recorded 179 tree species 15 cm or more in dbh; the 235 tree species in the general area were in 43 families. Fittkau and Klinge (1973) reported 502 species of trees and shrubs per ha in Manaus locations. Guillaumet (1987) provides summaries and analysis of this variable, complex ecosystem.

Because of physiological stress caused by the immersion, as well as the likelihood of species with adaptations to water dispersal being widely distributed, inundated forests have less species diversity than terra-firme forests and also less regional endemism (Prance 1987b). However, a significant number of species of trees and shrubs that occur in seasonal igapó are endemic to this habitat, and a few species are endemic to both seasonal igapó and várzea.

Plant diversity in campinas is reduced in comparison to contiguous rain forest. In the Lower Rio Negro region where the campinas are small and scattered, species richness is low in contrast to the extensive, older yet often isolated areas of the Upper Rio Negro region. The campinas are in effect ecological islands. Macedo and Prance (1978) showed that in the Manaus area, long-distance dispersal plays a very important role in distributing the species - 60% were dispersed by birds. Many species of plants are endemic to campinas (c. 55%) and occur in most of them; a few rare species are endemic to only one or two campinas (e.g. Erythroxylum campinense).

The Manaus region is the most controversial natural refugium of Amazonia. Some botanists regard it as an artefact of the large amount of collecting conducted there (Daly and Prance 1989). However, Prance (1987a) estimated that although the amount of endemism seemingly present could be greatly reduced by intensive collecting elsewhere, endemism would never drop to a level such that the Manaus region would no longer be considered an important centre.

Gentry (1990) compared a checklist (derived from Brazil's Programa Flora database) of 825 vascular plant species at the Adolpho Ducke Forest Reserve (100 km²) near Manaus (Prance 1990) with checklists from Central American wet and seasonal sites and a Peruvian Amazon seasonal site. The Central Amazonian flora represented by the Ducke reserve stands out as having a unique set of speciose families. The most speciose genera were almost exclusively trees, rather than herb, shrub or epiphyte genera which were most diversified at the three other sites. The peculiarity of the Manaus flora carries over into the family level: Manaus has the most unusual suite of families among the 20 most speciose families at each of the four sites. Chrysobalanaceae, Burseraceae, Lecythidaceae and Vochysiaceae are speciose at Manaus but not in the Peruvian Amazon or Central America, whereas Leguminosae, Moraceae, Lauraceae and Sapotaceae are speciose at Manaus and one or more of the other sites. The Manaus pattern of dominance by trees (in the listing of species) would probably hold as well for the less diversified flora of eastern Amazonia, but its checklists have been based solely on tree plots, so comparison of diversity by habit is not possible.

However, the comparison by simple presence or absence of families and genera for the Manaus flora with the flora of other sites is misleading since it is based on the single checklist (825 spp.) from the Ducke reserve. About 30% of the Instituto Nacional de Pesquisas da Amazônia (INPA) herbarium's 115,000 Brazilian Amazon specimens (as of 1989) are from the few degree-squares around Manaus (and a large percentage are not duplicated in other herbaria), and additionally there are more than 50,000 vouchers from the Minimum Critical Size of Ecosystems Project (MCSE) 70-90 km north of Manaus. Most of the families conspicuously absent from the Ducke reserve checklist (Gentry 1990) do occur regionally outside this reserve.

The monotypic Duckeodendraceae can be considered regionally endemic to Manaus - Duckeodendron cestroides penetrates at least as far as the lower Madeira River. Many endemic species are recorded for Manaus (Prance 1987a, 1990), but the very low density of scientific collecting for 1500 km west and south-west from Manaus does not allow a reliable demarcation of endemic limits in those directions (Nelson et al. 1990).

No fully vouchered inventory of plot diversity of trees has been published using a 10-cm dbh cutoff for 1 ha of terra-firme forest near Manaus. Using a 15-cm dbh cutoff, Prance, Rodrigues and Silva (1976) found 179 tree species in 1 ha. Milliken et al. (1992) in 1 ha c. 200 km north-west of Manaus found 201 species of trees [>=] 10 cm dbh. At the Ducke reserve, Alencar (1986) found 215 trees of [>=] 10 cm dbh or more in 1 ha and almost 400 different tree species in 2.5 ha. Gentry (in prep.), using a 2.5-cm dbh cutoff in a 0.1-ha plot at the Ducke reserve, found c. 233 tree, liana and acaulescent palm species among 365 individuals about as high as the richest sites near Iquitos, Peru (CPD Site SA9).

These few data suggest that single hectares of well-drained terra-firme clay near Manaus have c. 200-230 tree and liana species 10 cm or more in dbh/ha, considerably fewer than the almost 300 species on 1 ha near Iquitos (Gentry 1988). On the other hand, Central Amazonia has very much higher plot diversity than Central America or eastern Amazonia. Fifty contiguous 1-ha plots on Barro Colorado Island (BCI) in Panama averaged just 93 species [>=] 10 cm dbh per ha (Foster and Hubbell 1990), and seven 1-ha tree plots in eastern Amazonia near Marabá and Carajás had 119-130 spp. per ha (Silva et al. 1988).

Of all the 0.1-ha inventories done by Gentry in neotropical forests, that at the Ducke reserve is the most above the predicted diversity based on an otherwise consistent relationship between increasing rainfall and diversity (cf. Gentry 1988). Again, this remarkably high diversity actually may be the norm for a great distance from Manaus - published plot-diversity data are lacking for terra firme between Manaus and Iquitos, but two fully vouchered discontinuous 1-ha plots of várzea floodplain forest at the lower Japurá River have been inventoried (Ayres 1986). For trees [>=] 10 cm dbh, they held 135 and 109 species separately and 176 species in total, similar to the diversity of terra-firme plots in eastern Amazonia.

For larger plot sizes the Manaus tree flora also ranks very high in diversity. On a family by family basis, Rankin-de-Mérona et al. (1992) so far have identified 698 tree species [>=] 10 cm dbh in 53 families for 70 fully vouchered non-contiguous hectares on terra-firme clay and sand soils at the MCSE Project; mean density is 636 individuals per ha. They estimate that the 70 ha eventually will yield 1000 tree species [>=] 10 cm dbh. This major study is the best indication of the total tree diversity of Manaus. Much lower numbers are known from other large inventories: 320 tree and liana species [>=] 10 cm dbh in 10 ha spread over a large area east and south of Belém (Salomão et al., in press) and 306 species (trees and shrubs [>=] 2 cm dbh) for 50 ha of seasonal forest on Barro Colorado Island (Foster and Hubbell 1990). The entire known BCI tree and shrub flora is 409 species, which is probably exceeded by just the tree species with dbh 10 cm or more on 10 ha near Manaus.

Species × area data for the MCSE hectares along a 40-km strip are also indicative of very high diversity. Myristicaceae are evenly distributed - reaching 90% of their 70-ha species total after tallying just 7 ha, whereas Annonaceae require more than 36 ha to attain 90% of their species total in the 70 ha. Most remarkable is the linear ascending species area relationship for Sapotaceae in the 35 ha analyzed so far. Most of the species in five major families (Burseraceae, Annonaceae, Apocynaceae, Melastomataceae, Myristicaceae) occur at an average density of less than one tree per ha. The most abundant species are often clumped and therefore absent from many of the hectares. The data are not yet available for many families, so they cannot yet be ranked by species diversity at the MCSE sites.

Several observations may explain the exceptionally high diversity of Central Amazonian tree plots. Part of this diversity near Manaus is due to its central location a spillover effect providing species typical of the western Amazon, the Guianas and the eastern Amazon (Prance 1978, 1990), all added to the widespread Amazonian species. A heterogeneity of substrates near Manaus also contributes [beta] diversity and then [alpha] diversity by smaller scale overlap. For example, the Guayana Shield lies only 140 km north of the city, apparently permitting some Guayanan elements to occur at least as far as the MCSE sites (Prance 1990). Off the southern edge of the Guayana Shield are patches of open xeromorphic flora on Palaeozoic sandstone, with some new records for Brazil of typically Venezuelan species (O. Huber, pers. comm.) as well as new species. On the opposite (southern) bank of the Rio Negro are two small clay-soil savannas which contribute typical widespread savanna elements such as Physocalymma scaberrimum, Curatella americana and Curculigo scorzoneraefolia, whereas podzol campinas and sandy igapó have provided niches for penetration of families and genera from the sandy lowlands and highlands of the Guayana sandstone floristic province, e.g. Humiriaceae, Theaceae, Rapateaceae, Micrandra, Eperua, Pagamea, Glycoxylon and Neoythece (Kubitzki 1989).

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Useful plants

Cavalcante (1976, 1979) includes 171 species in his survey of edible fruits of the Amazon; 84 occur in the extensive terra-firme primary forests or along their small streams and 34 are in a preliminary checklist of vascular plants from a reserve near Manaus (Prance 1990). Nonetheless, very few native fruits (or medicinals) are sold in local Manaus markets, for cultural reasons and because native fruit species on the poor soils north of Manaus produce little edible biomass (M. van Roosmalen, pers. comm.). Three edible palms occur in high densities near Manaus, and could be exploited to a much greater degree: Oenocarpus (Jessenia) bataua (fruit, oil), Oenocarpus bacaba (fruit) and Mauritia flexuosa (fruit). Important species exploited by a limited number of extractivists near Manaus are Minquartia guianensis for durable posts; Mezilaurus itauba for boat construction; other timber species for local use; and hardwoods in general for fuelwood consumed by city bakeries.

Within 150 km of Manaus the only major extractive species exploited are Aniba rosaeodora and A. duckei for an essential oil. In total from the general region, rosewood oil valued at US$1.4 million FOB (free on board) was exported in 1984 through Manaus, according to the Association of Exporters of the Manaus Free Zone.

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Economic assessment

Central Amazonia has no major concentrations of prime export hardwoods. Várzea-forest peeler-log species support three large plywood factories for export, but they are depleted near Manaus. The logs are extracted by floating them out of flood forest, causing little ecological damage. The State of Amazonas produces c. 20% of the Brazilian Amazon's annual 285,000 m³ of laminates and plywoods (Silva et al. 1991). These faster growing (softwood) species could be planted on floodplains, but presently there is no significant planting of native timber or plywood species anywhere in Amazonia.

In contrast with Acre, Iquitos and Belém, minor forest products are of little economic importance in Central Amazonia (Vantomme 1990). Nevertheless, extractive products and flooded-forest fish consumed locally and generally not tallied in statistics are important for the survival of hundreds of thousands of people. Floodplain forests are necessary for survival of the fruit-eating "tambaqui" (Colossoma macropomum), which leads the list in tonnage for food-fish species sold in Manaus markets. About 90% of the fish sold are taken from várzea lakes, forests and rivers (Bittencourt and Cox-Fernandes 1990); many fishes depend on food chains in the flooded forest for part of the year (Goulding 1980).

Green tourism attracts many foreign visitors, who leave a large share of the annual US$200 million (1989) of tourism receipts in Manaus. Jungle lodges and riverboat tours are growing in number to attend this market, though many are poorly organized and marketed. Although receipts in hard currency are grossly under-reported, thousands of people are directly employed in hotels, lodges, restaurants and transportation.

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Threats

Daly and Prance (1989) provide a basic overview of the conversion of Amazonia, including construction of major roads and plans for hydroelectric, agricultural and agro-forestry development. Lovejoy (1985) gives an extensive review, and Hecht (1981) gives a detailed analysis especially of natural-resource uses and their effects on the Amazonian soils.

Road building stimulates subsistence colonization, hunting and invasion of reserves. Fragile dry sandstone flora and unique clear-water stream ecosystems on sandstone are threatened 130 km north of Manaus in Presidente Figueiredo by deforestation, erosion and sedimentation. Large-scale deforestation for subsidized pastures had been generally diminishing, but it is being renewed north of Manaus for establishment of oil-palm plantations. Várzea forests have had their economic species heavily exploited.

Urban expansion/invasion occurred into one INPA reserve near Manaus. Campina is the most threatened type of vegetation in the Manaus vicinity. Campinas have been used as a source of sand for road building and construction projects throughout Amazonia, resulting in complete loss of many areas. Many campinas have been burned, resulting in a considerable loss of habitat as well as the extirpation or extinction of many endemic species, and their replacement by secondary growth "capoeira". The degree, intensity and frequency of widespread burning are factors affecting regeneration potential. Recuperation may take many centuries. The long-term effects of these factors can be seen at a number of campina sites where radiocarbon-dated pottery shard remains and charcoal indicate former human occupation and cultivation practices 800-1100 years ago (Anderson 1981).

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Conservation

The economy of the Manaus Free Zone has a conservation effect: it permits cheaper transport of foodstuffs from distant southern Brazil, and diverts capital from deforestation and agriculture. Brazilian institutions (e.g. INPA) with international collaboration are undertaking major ecological studies to understand the dynamics of the forests with a view to their long-term use (Schubart and Walker 1987). Just 1.48% of the State of Amazonas had been deforested as of August 1991 (INPE 1992), so most conservation areas are simply lines drawn through a continuous forest cover.

Four governmental research stations are established in the Manaus region and larger areas are being considered. Over 320 km² of INPA reserves include terra firme, campinarana and campina, and a 1150-km² military base with predominantly terra-firme forest is well protected (Table 56). The Anavilhanas Ecological Station has large protected tracts of terra firme and igapó. No várzea ecosystems near Manaus are protected, but westward on the Lower Japurá River, Mamirauá Ecological Station is on várzea.

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Map 41. Manaus Region, Brazil (CPD Site SA5), showing selected conservation areas

References

Absy, M.L., van der Hammen, T., Soubies, F., Suguio, K., Martin, L., Fournier, M. and Turcq, B. (1989). Data on the history of vegetation and climate in Carajás, eastern Amazonia. In International Symposium on Global Changes in South America During the Quaternary, São Paulo, 1989. ABEQUA/INQUA, Special Publication No. 1, São Paulo. Pp. 129-131.

Alencar, J.C. (1986). Análise de associação e estrutura de uma comunidade de floresta tropical úmida onde ocorre Aniba rosaeodora Ducke (Leguminosae). Ph.D. thesis. INPA/Universidade do Amazonas, Manaus. 332 pp.

Anderson, A.B. (1981). White-sand vegetation of Brazilian Amazonia. Biotropica 13: 199-210.

Anderson, A.B., Prance, G.T. and Albuquerque, B.W.P. de (1975). Estudos sobre a vegetação das campinas Amazonicas III. A vegetação lenhosa da campina da Reserva Biológica INPA-SUFRAMA. Acta Amazonica 5: 225-246.

Ayres, J.M. (1986). Uakaris and Amazonian flooded forest. Ph.D. dissertation. Cambridge University, Sidney Sussex College, Cambridge, U.K. 337 pp.

Bittencourt, M.M. and Cox-Fernandes, C. (1990). Pesca comercial na Amazônia Central: uma atividade sustentada por peixes migradores. Ciência Hoje 11(64): 20-24.

Brickmann, W.L.F. (1989). System propulsion of an Amazonian lowland forest: an outline. GeoJournal 19: 369-380.

Cavalcante, P.B. (1976). Frutos comestíveis da Amazônia, Vols. 1 and 2. INPA, Manaus. 166 pp.

Cavalcante, P.B. (1979). Frutos comestíveis da Amazônia, Vol. 3. Public. Avulsas No. 33, Museu Paraense Emílio Goeldi, Belém. 61 pp.

Daly, D.C. and Prance, G.T. (1989). Brazilian Amazon. In Campbell, D.G. and Hammond, H.D. (eds), Floristic inventory of tropical countries: the status of plant systematics, collections, and vegetation, plus recommendations for the future. New York Botanical Garden, Bronx. Pp. 401-426.

Fittkau, E.J. and Klinge, H. (1973). On biomass and trophic structure of the Central Amazonia rain forest ecosystem. Biotropica 5: 2-14.

Foster, R.B. and Hubbell, S.P. (1990). The floristic composition of the Barro Colorado Island forest. In Gentry, A.H. (ed.), Four neotropical rainforests. Yale University Press, New Haven. Pp. 85-111.

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Authors

This Data Sheet was written by Dr Bruce W. Nelson [Instituto Nacional de Pesquisas da Amazônia (INPA), Caixa Postal 478, 69000 Manaus, AM, Brazil] and Olga Herrera-MacBryde (Smithsonian Institution, SI/MAB Program, S. Dillon Ripley Center, Suite 3123, Washington, DC 20560-0705, U.S.A.).

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