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LegendVegetation Map of the Chapada Diamantina National Park and Surrounding Areas

Roy Funch

Brazil is the fifth largest country in the world (more than 8,500,000 km²), and by far the largest tropical nation. It straddles the equator, from approximately 5° N to 34° S, and spans more than 4000 km , at its widest, from the Atlantic Ocean to the foot of the Andes mountains (approximately 33° to 70°W), encompassing a wide range of climatic, geographical and geomorphological conditions that help account for its enormous diversity of ecosystems/habitats and its tremendous biodiversity. There are, for example, more than 56,000 plant species (approx. 19 % of the world's flora) already identified from this country, including ~3,100 bryophytes, 1200-1300 species of pteridophytes, and more than 500 species of algae. Among the angiosperms, monocotyledons stand out with more than 8,000 species already described, of which 45% are endemic (MMA, 1998; Giulietti et al,. 2005).

While Brazil is most widely known for its vast tropical forests (more than 62% of the land area of the country, including the Amazon and Atlantic Coastal Forest biomes), there are also extensive areas occupied by savanna ( cerrado ) and semi-arid vegetation ( caatinga ), representing approximately 24% and 11% of its total area, respectively (IBGE, 2006). Caatinga vegetation is predominant in the NE sector of the country where the present work was carried out.

The north-eastern sector of Brazil (approximately 4° to 16° S and 35° to 46° W) has a complex climatic pattern due to its large land area (~1.5 million km²) and diverse geomorphology, as well as the fact that it lies at the junction of two principal weather systems (related to the north-east and south-east trade winds) that alternate in their influence over that region (Nimer, 1977; Harley, 1995). Additionally, the El Niño/Southern Oscillation (ENSO) phenomenon (Senado Federal, 1997) has profound effects on rainfall patterns in this region, with El Niño conditions being associated with unusually strong episodes of drought. In general, this NE region of Brazil experiences low monthly rainfall rates (usually less than 100 mm , although these can be extremely variable) and a long dry season, lasting from six to nine months (MME, 1981). The rainy season coincides with the astral summer months (roughly November through June), and bi-modal peaks of precipitation are usually observed, one occurring between November and January and the other from March to April. The climate at the study site is strongly influenced by altitudinal and rain-shadow factors, and will be discussed in more detail in the text.

Embedded within this semi-arid region of NE Brazil is the Espinhaço mountain chain (>60,000 km²), stretching from southern Minas Gerais State (~20° S latitude) to the northern border of Bahia State (~10° S) and parallel to the Atlantic Coast (Fig 1). This mountain chain is composed mostly of Precambrian sandstones and metamorphic rocks, and represents the eastern section of the extremely old and stable Brazilian Shield. The Espinhaço chain is effectively divided slightly north of the Minas Gerais/Bahia State border by a 300 km-wide corridor of lowlands (the Rio de Contas River basin). The northern sector of the range is known as the Chapada Diamantina, and is completely contained within the state of Bahia . While not very lofty (the highest peak is slightly over 2000 m ), large scale folding followed by erosion has resulted in a varied and highly dissected landscape (CPRM, 1994; Harley 1995). The Sincorá Range , focus of the present work, constitutes the eastern flank of the Chapada Diamantina. While the Sincorá Range does not have precisely defined limits, it is roughly bounded by the Sincorá River to the south and the town of Afrânio Peixoto to the north, where its easily recognizable scarps blend back into the high, dissected plateau that constitutes the bulk of the Chapada Diamantina. An almost continuous fault scarp approximately 250 m high and 80 km in extension sharply defines the western border of the Sincorá Range, while the eastern flank rises up quickly from the flat plain that stretches almost uninterrupted to the Atlantic Coast, about 300 km further east. This range has a strong N-S orientation, being approximately 160 km long, and from 20 to 30 km wide, and covers a total area of approximately 4000 km² (LS Funch, 2004).

The inland regions of north-eastern Brazil were originally occupied by nomadic Amerindians tribes, such as the Janduís and Paiacus in present day Bahia State . Colonization by the Portuguese (beginning in the early 16 th century) was originally limited to coastal areas where sugar cane was planted, while the semi-arid interior only began to be permanently occupied by Europeans after the introduction of cattle in 1550. The semi-arid climate favoured open-range grazing over crop production, with huge tracks of land eventually coming under the control of single individuals or a few large families. Population growth, poverty, the introduction of goats (with their more destructive grazing habits), the scarcity of wood products for construction and energy generation, reduced acreage suitable for planting combined with the irregular patterns of rainfall, resulted in a serious (and growing) degradation of the natural environment and the loss of biodiversity. This environmental crisis is reflected in the fact that less than 0.7% of the dry land caatinga vegetation in NE Brazil is protected within any sort of state or federal conservation area (MMA, 2006).

The central and northern sections of the Sincorá Range , as well as the non-montane areas adjacent to them, have experienced accelerated development in the past two decades due to the opening of new heavily mechanized agricultural areas, the settling of landless farmers, and the expansion of tourism. These activities have resulted in altered patterns of land and water use, increasing human population densities, land clearing and deforestation, and the intensive use of agro-chemicals in the headwaters of the regionally important Paraguaçu River Basin . The 1520 km² Chapada Diamantina National Park (CDNP), occupying approximately half of the entire Sincorá Range, has to some extent counterbalanced this regional growth by defining significant areas of mountain landscape as (legally) immune to anthropogenic alterations. Although it was founded more than 20 years ago (1985), the National Park still suffers from budget restrictions, limited staff and few planning tools. The publication of the first management plan for the park is being programmed, and its success as an operational document will depend on the availability of detailed and up-to-date information concerning the ecosystems that compose the reserve, their conservation status, the environmental threats they face, as well as the location and extent of human influences in the surrounding region. Additionally, this same information will be used to orient the National Park System in its acquisition of new areas contiguous to the existing reserve.

As such, vegetation maps are needed to indicate the types and distribution of regional vegetation assemblages in an easily interpretable manner and at an appropriate planning scale so as to be easily consulted by decision makers and other interested groups at all levels of conservation (and development) planning.

Description of the area

The study area included the entire legal limits of the Chapada Diamantina National Park (1,520 km²) as well as regions up to 10 km beyond its boundaries, in the northern sector of the Sincorá Range, Bahia State, NE Brazil (approximately 12° 11´- 13° 32´S x 40° 53´- 41° 42´W). For purposes of the present discussion, the mapping area has been divided into four geomorphological areas: eastern, central, north-western and south-western.

Almost the entire central section of the mapping region (the Sincorá Range and the Chapada Diamantina National Park ) is composed of extremely old (1.7+ b.y.) Precambrian sandstones, conglomerates, and quartzites, and is part of the ancient Brazilian Shield (CPRM, 1994) with elevations averaging about 900m above sea level and peaks up to 1700m. Soils are limited to sandy, usually thin, dystrophic neosols, with exposed rock surfaces covering a high percentage of the Sincorá mountain range. The regional climate is seasonally humid and mesothermic, with mild, wet summers and cool, dry winters. Average monthly temperatures range from about 18° to 22°, although these vary considerably with altitude. Freezing temperatures have never been reported in the region and summer temperatures rarely pass 35º. The period of heaviest rainfall is normally from November to March, and the dry season usually lasts from July to early November, although periods of prolonged drought are not uncommon. The average yearly rainfall in the Sincorá Range is 1192 mm , although total precipitation is extremely variable (e.g. 357 mm in 1993 and 1721 mm in 1989), as is its distribution throughout the year (Harley, 1995; Seixas, 2004). The principal vegetation types encountered within the Sincorá Range are: 1) sub-montane to montane semi-deciduous seasonal to evergreen forests on litholic neosols; 2) sub-montane to montane evergreen riparian forests along river courses; 3) campo rupestre , at higher altitudes with a mix of exposed rock surfaces and shallow soils; 4) sandy sedge meadows, at high altitudes on thin to deep sandy soils, and; 5) wetlands. Detailed descriptions of these vegetations are presented under the section “Vegetation types” that follows.

Soils

Pedological studies in Chapada Diamantina have revealed a mosaic of soil types reflecting the geological diversity of the region, climate, altitude, slope, exposure, hydrology, and vegetation (de Jesus et al., 1985; CPRM, 1994). A large percentage of the soils in the area mapped for vegetation were latosols, neosols (lithosols) and argisols (Rocha et al. , 2005), all of which are consistently dystrophic and highly acidic.

Soils were further categorized into very basic categories according to their colour and, to some extent, their composition (predominance of sand or clay). Colour was visually judged for fresh samples taken > 20 cm below the surface. Soil composition (sand or clay content) was subjectively tested using a simple technique of manipulating a small amount of moist soil.

Vegetation types

A diverse series of vegetation types were recognized, which are listed and briefly described below. Recognition was based partly on previous experience in the area, and further strengthened by observations made during the course of the project.

1. Campo rupestre (CR)

2. Sandy sedge meadows (M)

3. Latosolic forests (F1)

4. Montane lithosolic forests (F2)

5. Riparian forests (F3)

6. Wetlands (W)

7. Transition/Mosaic areas (T)

8. Alluvial sands (S)

9. Anthropogenic areas (A)

Campo rupestre ( CR ) – is an open, low vegetation form found on nearly continuous rock surfaces in the mountain (usually above 800 m a.s.l.) primarily composed of sclerophyllus and evergreen shrubs or sub-shrubs and small trees (Harley and Simmons, 1986) that are highly specialized and adapted to conditions of low soil pH and low nutrient content, high insolation, and diurnal variations in temperature and humidity, with dew often providing much of their water requirements during periods of drought. It is not a single physiognomic vegetation type, but varies according to the substrate in a continuous mosaic of habitats and species mixes, ranging from purely epilithic species (especially orchids, bromeliads) on exposed rock surfaces, through areas with blocks of loose stone or deep cracks that can collect slightly greater amounts of soil and humidity will harbour shrubs and small trees. The campo rupestre is notable for its high levels of plant endemism (Giulietti, et al., 1997) and the number of species per family and their frequency and cover will vary greatly from area to area within the same range, but with consistently significant presence of the families Velloziaceae, Asteraceae, Cyperaceae, Cactaceae, Orchidaceae, Bromeliaceae, Lamiaceae, Compositae, Ericaceae, Melastomataceae, Guttiferae, Amaryllidaceae, Leguminosae, Clusiaceae, Verbenaceae, Araceae, Asteraceae, Begoniaceae, Melastomataceae, Rubiaceae, Verbenaceae, and Aquifoliaceae, among others (Harley and Simmons, 1986; Harley, 1995; Conceição, 2003; Conceição et al. , 2005; Harley et al., 2005).

Sandy sedge meadows ( M ) - vegetation is a variation of campo rupestre and is often included in that category, but for the purposes of this work we consider it more useful to treat it as a separate vegetation type. It occurs on relatively flat landscapes in the mountains within the Sincorá Range on extremely thin, sandy, litholic neosols that are often saturated for fairly long periods of times during the rainy season. The same type of vegetation can also occur within high mountain valleys that have deep sandy soils but are subjected to frequent burning (which removes the forest cover). These very low and open sedge grasslands are dominated by Cyperaceae, Graminae, Xyridaceae, and Eriocaulaceae, and are often used for pasturing domesticated animals during the regional dry season.

Sub-montane to montane semi-deciduous seasonal forests on deep latosols (Latosolic forests) ( F1 ) – these are semi-deciduous forests subject to seasonal variations in rainfall (and, to a lesser extent, temperature) that occupy also the entire eastern border of the Chapada Diamantina at altitudes between approximately 400- 800 m (Funch et al. , 2006). The landscape is relatively flat to deeply dissected, and covered with deep clay latosols that range in colour from yellow, to orange, to dark red. These soils are generally very deep, well drained, dystrophic, highly acidic, with a high aluminium content and little organic matter (MME, 1981) (though there is usually a thin leaf litter layer).

Sub-montane to montane semi-deciduous seasonal to evergreen forests on litholic neosols (Montane lithosolic forests) ( F2 ) - cover the boulder-strewn and sometimes exceeding steep river valley slopes within the mountains of the Sincorá Range . Due to the near ubiquitous presence of sandstone rocks in the area of the Park, the soils of these valley sides are sandy, usually thin and stony, dystrophic, and become progressively drier at longer distances from the river (this effect can be modified depending on exposure, aspect, or the presence of natural seeps). As such, and although predominantly evergreen, the montane forests on the lower slopes demonstrate a certain degree of deciduousness in the dry season (Funch et al. , 2002).

At altitudes near 1,000 m and above, however, environmental conditions become increasingly more humid due to longer daytime cloud cover, more rainfall and dew, and the fact that these forests generally occupy more protected sites in the mountains such as narrow valleys and deep crevices in the sandstone rock (sheltered from both drying and wildfires). The soils are sandy, rocky, and not especially deep, but due to the high humidity and lack of fires there is a significant accumulation of damp organic material on the forest floor. These forests become increasingly more humid and evergreen as they occupy higher sites in the mountains and future studies will most likely suggest their treatment as a separate category.

Sub-montane to montane evergreen riparian forests (Riparian forests) ( F3 ) –are seasonal evergreen forests that follow river courses. In the Sincorá Range itself the rivers margins are strewn with boulders, but the sandy substrate that can collect between the rocks is continually humid (although dystrophic and having a very low organic material content). As these forests are only found adjacent to perennial water courses, they are usually of very restricted width (mostly less than 10 m ) and usually grade rapidly into sub-montane to montane semi-deciduous seasonal forests as the soil becomes drier and the inclination of valley sides becomes more pronounced. At higher altitudes, they become continuous with the montane evergreen forests.

Wetlands ( W ) – are permanently inundated or boggy areas. The largest wetlands are found at the foot of the eastern slopes of the Sincorá Range where the Santo Antônio and Utinga Rivers meander over low extensive plains. There are also many sites with poor drainage within the mountains themselves, although these tend to be very limited in area (usually less than one hectare).

Alluvial Sands ( S ) – are regionally extensive alluvial sand deposits that are mostly geologically very recent, resulting from diamond mining activities in the mountains or sometimes to road building. These soils are of essentially pure sand, dystrophic, deep, rapidly drained, and frequently re-worked during seasonal flooding, resulting in an extremely sparse vegetation cover.

Transition/Mosaic Areas ( T ) – As a result of many interacting ecological factors related to altitude, exposure, slope, local geology, soil composition, soil depth, microclimate, etc., the contact between two (or more) vegetation forms will result in a species-rich transition zone (often a mosaic), composed of elements from both communities.

Anthropogenic Zones ( A ) – These are areas that have been (or are presently) subjected to severe anthropogenic disruption of the natural vegetation cover.

In areas undergoing natural recovery, the vegetation encountered will depend on the previous vegetation cover, the duration of human use/occupation, time since abandonment, type of alteration/severity of the alteration to which the land was submitted, soil type, slope, water regime, etc.

Three additional vegetation types were encountered, but are not included in the numbered series above, either because they did not fall within the area of the National Park, or in the case of the last (capitinga), occupied too small an area to significantly contribute to the overall picture:

10. Cerrado (C)

11. Caatinga (Caat)

12. Capitinga (CAP)

Cerrado (Brazilian savannas) ( C ) - vegetation is characterized, in general, by the presence of both a well defined ground and an arboreal layer. The ground layer is continuous in all but the pure forest physiognomy (called cerradão ), and is composed principally of grasses, sedges, and numerous subshrubs (which can appear to be herbaceous but often possess a well developed root systems with xylopods), acaulescent palms, and very few annual species. The arboreal layer is generally discontinuous and up to 10 m tall, with twisted branches, thick bark, and leaves that are rarely absent, large, and thick.

Cerrados demonstrate wide and continuous variations in their physiognomy, ranging from open savannas, with predominantly ground plants, to forest formations. They have been traditionally classified into five basic classes: “ campo limpo ” ( C1 ), open savannas with only occasional ground plants; “ campo sujo ” ( C2 ), open savannas with predominantly ground plants with low ( £ 3m), well-spaced shrubs; “ campo cerrado ” ( C3 ), with taller ( £ 5m), more closely spaced shrubs; “ cerrado sensu stricto ” ( C4 ) has taller shrubs and small trees that are more densely spaced, but still retains an open canopy layer; and “ cerradão ” ( C5 ), with a forest physiognomy of an essentially closed canopy, and a reduced or absent ground layer (Coutinho, 1978).

Cerrado vegetation typically grows on deep, well drained, dystrophic soils, with low pH and consequently very high levels of exchangeable aluminium. They are subject to a markedly seasonal climate of heavy (~1500 mm yr -1 ) summer rains (October to March) followed by a long drought period during the winter months. As a consequence of the prolonged dry period, fires are extremely common in this biome and the plants have many adaptations to this fire regime, including thick bark, bud protection, xylopods that aid in root-sprouting, while some plants have fire-responses that synchronize flowering and induce fruit dehiscence.

Botanical studies have generated an estimate of some 7,700 different species in the widespread cerrado biome (Mendonça et al. , 1998). Ratter and Dargie (1992), for example, undertook plant surveys in 26 cerrado areas (principally in central Brazil ) and identified 485 shrub and arboreal species. Of these 485 species, however, only 27 occurred in 15 or more of the 26 localities, demonstrating a great floristic heterogeneity within this vegetation.

Caatinga (Caat) – vegetation occupies the extensive semi-arid region of NE Brazil (between 700,000 and 800,000 km²), corresponding to 10% of the entire country. Rainfall levels there are low (usually between 400 – 700 mm yr -1 ) and extremely variable from year to year, with an extended dry period of up to 9 months. The vegetation is highly xeromorphic with a predominance of profusely branched low trees and shrubs. The plants generally have small leaves, spines, thick bark, and a well-developed root system often with tubers. With few exceptions (such as palms and a few dicots such as Zizyphus joazeiro Mart.), the plants are deciduous. Cacti are conspicuous, and there are many terrestrial bromeliads (Harley et al. , 2005). Annual herbaceous plants flourish during the short rainy season, and in many regions vernal pools appear although these are rarely seen in the present study area, due to the very irregular topography near the Sincorá Range and the karst topography slightly further west. Caatinga also occurs at low altitudes (below 500 m ) throughout most of its range, but flourishes at elevations between 700-1000m in the study area, in the rain shadow of the Sincorá Range .

Capitinga ( CAP ) – Small, scattered, and isolated areas of capitinga vegetation (up to a few hectares) exist throughout the region around the Sincorá Range growing on deep, extremely well-drained, fine sandy soils. Different from alluvial sand deposits, these sandy areas seem to be associated with the direct on-site weathering of friable sandstone outcrops (personal observation) and they are usually completely surrounded by latosol landscapes. The vegetation in these areas of capitinga is low and open, with many shrubs, acaulescent palms, scattered and infrequent small trees and cacti, annual plants are rare and there is a high percentage of bare ground. While similar in physiognomy to the coastal restinga vegetation and the dune areas bordering the São Francisco River (Rocha et al. , 2004), caatinga is distinguishable from these other sites (LS Funch, UEFS, Bahia, Brazil, unpubl. res.); the species composition quite different, there is no saline component to the environment, and they are not derived from wind or water transport and subsequent re-deposition. For these reasons, the local denomination for these sites has been retained

Disturbance categories of forested areas:

Six disturbance categories were employed to describe the conservation status of the diverse forest formations:

Good ( Gd ) – good state of conservation; no signs of harvesting or fire damage; diverse mix of tree diameters and ample spacing of individuals; light underbrush.

Medium ( M ) – medium state of conservation; signs of selective harvesting; few large trees and close spacing of small trees; heavy underbrush.

Poor ( P ) – damaged forests; visible signs of fire damage or harvesting; very dense spacing of small trees; heavy underbrush.

Fire Damage ( FD ) – mature, primary forest that has visibly suffered recent heavy fire damage; many dead trees still standing; many live trees showing canopy damage; heavy underbrush.

Fire Damage/Poor Forest ( FD/P ) – already damaged forests (P) that have suffered additional burning; signs of recent fire; no large trees, very dense spacing of small trees; heavy underbrush.

Heavily Disturbed ( HD ) – by non-fire factors (mechanical disturbances)

Overview

The eastern sector of the mapping area is a generally flat but deeply dissected plain almost completely covered by sub-montane to montane semi-deciduous seasonal forests on deep latosols (Latosolic forests) ( F1 ). This was formerly a solid forest block with tall trees (20+ m) and many economically valuable species such as pau d'árco, ipê rosa ( Tabebuia spp.) and jacarandá ( Dalbergia sp.). Recently, however, most of the area has been subject to intensive exploitation, including clearing for pasture formation, selective cutting, or has suffered from extensive burning. The most level terrain is most heavily developed, probably because of its facility of access for men and machines, and is occupied by a number of large farms (> 15,000 ha .), government land-grant communities for (previously) landless farmers, as well as hundreds of smaller individual property holdings. These lands are principally used for cattle raising, but some crop production is encountered (the latter activity being restricted to a great extent to drainage areas). Areas not directly cleared have all been selectively logged for commercially valuable lumber species, fence posts, rough hewn housing timbers, etc.

Latosolic forests areas situated nearest the Sincorá Range have a more irregular topography, are less suitable for agricultural ends, and have experienced the least mechanical impact of human intervention. Only small tracts in this region can be considered well-preserved, however, because serious and repeated forest fires have affected most of the region. The frequent presence of fires in these less populated areas reflects a number of underlying factors: these marginal sites are often government lands or tracts owned by traditional families who have no control over, or interest in, these distant sites. As such, they are often occupied by otherwise landless citizens who will only have (possible future) ownership rights over lands that they can demonstrate to have cleared and occupied. With no tools beyond a hoe, an axe, and a box of matches, and nothing to lose in a wide-spread conflagration, forest fires usually follow occupation. Wildfires can also spread from the mountain vegetation of the Sincorá Range to adjacent forest sites and, somewhat ironically, the lack of cleared areas (pastures, fields) facilitates the propagation of the flames to contiguous areas. There is almost no fire-fighting capacity in the region.

The north-western sector of the mapping area presents an irregular topography of low hills composed of sandstone, quartzite, and conglomerate rocks (rising approx. 100 – 150 m above the local landscape) that are covered by thin, generally rocky, neosols. This region falls within the rain-shadow of the Sincorá Range immediately to the east, which is reflected by the predominant caatinga dry land vegetation encountered there. In spite of the locally poor soils and the lack of rainfall, this area, like most of the caatinga region, has been extensively (though not densely) occupied and altered, principally for cattle raising. Most of the land holdings are relatively small (tens to hundreds of hectares), and crop production is limited to drainage areas with permanently flowing streams. To the south and to the northeast of this area the soils become deeper and have a higher clay content, which favors a transition to cerrado vegetation. Along the eastern edge of this sector, along its contact with the Sincorá Range and in apparent response to an increasing rainfall gradient, the caatinga vegetation becomes progressively more arboreal, grading into the forest formations that occupy the western slopes of the mountains or into arboreal cerradão (slightly more to the north).

The south-western sector of the mapping area is a large, relatively flat plain (although more dissected in the west-central zone) at approximately 900- 1000 m , covered principally by dystrophic argisols (moderately deep, well-drained soils) and latosols and occupied principally by cerrado vegetation, although significant areas of forest are encountered below the western flank of the Sincorá Range. Until the beginning of the 1980's there was very little development in the region. Open-range cattle were present to a fairly limited extent on the extensive plains, but human occupation (and farming) was generally restricted to the western flanks of the Sincorá Range and the lowland areas at its more dissected northern end, and the forested areas to the extreme S and SE of this sector were also mostly intact. In the intervening decades, however, the region has become more densely occupied, and its landscape and land cover profoundly altered by crop production, cattle ranching, and dam construction. Satellite imagery reveals the presence of more than 100 circular areas cultivated (currently or in the recent past) using central-pivot irrigation, while countless rectangular to irregular non-mechanized agricultural/pasture plots dominate the landscape, especially in the more dissected northern zone, leaving only ragged remains of (modified) native vegetation. Coffee plantations have likewise decimated the forest cover in the extreme S and SE of this sector.

The central section of the mapping area represents the axis of the Sincorá Range itself, including the Chapada Diamantina National Park (outlined) that occupies the bulk of this central sector. The Sincorá Range has elevations averaging about 900m above sea level and peaks up to 1700m. Soils are generally limited to sandy, usually thin, dystrophic neosols, while exposed rock surfaces cover a high percentage of the mountain range. The principal vegetation forms there are campo rupestre , sub-montane to montane semi-deciduous seasonal to evergreen forests on litholic neosols (montane lithosolic forests), and sandy sedge meadows. Rain fall is usually very high (average ~1200 mm yr -1 ) and is supplemented by dew and cloud-mist at higher elevations. While only two roads cross the mountains within the mapping zone, the Sincorá Range has experienced extreme anthropogenic pressure from diamond mining activities, hunting, cattle grazing, and subsistence farming (the former three associated with extensive burning). With the decadence of mining in the late 1800's and the migration of huge contingents of the population to urban centres in the mid 1900's, a gradual (but partial) natural recovery of the native vegetation on previously mined areas was possible, although burning continued unabated, with its especially detriment effects on forest formations (this topic will be discussed in detail in the section on vegetation that follows).

Considerations on the Vegetation within the National Park and the full mapping region

The Chapada Diamantina National Park (1524 km²) comprises 9 basic vegetation types/landscapes: 1) campo rupestre (63.4 % of the Park area); 2) sandy sedge meadows (13.4%); 3) sub-montane to montane semi-deciduous seasonal to evergreen forests on litholic neosols (montane lithosolic forest), and; 4) sub-montane to montane evergreen riparian forests (3 & 4 together, 7.1%); 5) sub-montane to montane semi-deciduous seasonal forests on deep latosols (latosolic forests) (6%); 6) transition zones (7.3%); 7) wetlands (0.7%); 8) alluvial sands (0.5%), and; 9) anthropogenic landscapes (1.6%).

1) The Campo Rupestre (CR) vegetation throughout the Espinhaço mountain chain (which includes the Chapada Diamantina and the Sincorá Range) is distributed in the form of an archipelago of high elevation areas separated by lower lands with distinct environments and vegetation types (forests, caatinga , cerrado , etc.) (Harley, 1995). This isolation contributes to its high degree of endemism and the generally high b diversity observed between adjacent sites (Conceição et al. , 2005). The floristic composition includes elements with wide distribution as well as components common to the mountain vegetation of the tepuis of northern South America , the Brazilian Central Plains, and the coastal sandy restinga areas, all of which have dystrophic, sandy soils (Giulietti and Pirani, 1988; Harley, 1995; Giulietti et al. , 1997).

Campo rupestre is the dominant vegetation form within the Chapada Diamantina National Park (63.4 %) and, together with c aatinga (dry land) vegetation and the cerrados , one of the most characteristic of the Chapada Diamantina mountain range as a whole. Campo rupestre vegetation has an open herbaceous-shrub physiognomy and occupies rocky outcrops and sandy areas in the mountains above ca. 900 m in the central portion of the present study area. (Harley, 1995; Conceição et al. , 2005).

In spite of the fact that the CR vegetation experiences severe and extremely frequent burning cycles within the Sincorá Range (less than a decade in more protected areas, and almost annually in others) this vegetation form (as well as the related sedge meadows) has been able greatly to expand in this area as a result of anthropogenic disturbances. Mining allied with burning have eliminated the soil and the forest vegetation from enormous swathes of the Sincorá Range (on the order of 200 km²), and much of the landscape above 800m has been occupied by CR vegetation (at lower altitudes, a transition vegetation with representatives of CR, cerrado , and invasive species).

2) There are basically two forms of sandy sedge meadows (M) found within the Sincorá Range (and the CDNP), but the important difference between them do not lies in their physiognomy or species composition, but rather in their origin.

The more original form of sandy sedge meadows grew on relatively flat landscapes in the mountains within the Sincorá Range on extremely thin, sandy, litholic neosols that are often saturated for fairly long periods of times during the rainy season. These sites have become more open and exposed, with less shrubs and small trees, due to extensive burning, but are otherwise truly natural sites.

The anthropogenic form of sandy sedge meadows now occupy the central axis of the Sincorá Range in an area of the CDNP dominated by a narrow anticlinal valley that stretches more than 53 km and averages over 900m in altitude. The abundant rainfall and mist in the high mountains and the deep sandy sediments filling the valley floor favored dense forest formations in the past. However, constant and frequent anthropogenic fires, related principally to mining, agriculture, hunting, and native pasture management, essentially eliminated the forest formations in this series of valleys, and the now open habitat was occupied principally by the fire-tolerant low-growing herbs and sub-shrubs of the original sedge meadow vegetation.

3) Sub-montane to montane semi-deciduous season to evergreen forests on neosols (montane lithosolic forests) (F2) can be found throughout the Sincorá Range . As the sandstone and conglomerate rocks that compose the mountains erode into loose sand and coarse gravel, they form infertile, rapidly drained, and (usually) thin soils (neosols) that will support forest formations under favorable combinations of soil depth and humidity. Although predominantly evergreen, these montane lithosolic forests demonstrate a degree of deciduousness in the dry season (Funch and Barroso, 2002) due to the relative abundance of a group of deciduous or semi-deciduous trees such as Simaruba amara Aubl. , Diospyros sericea A.DC. , Bowdichia virgilioides Kunth. , Maprounea guianensis Aubl. , Zanthoxylum rhoifolium Aubl. , Emmotum nitens Miers, and Aspidosperma discolour A.DC., which contribute to the formation of moderate peaks of leaf fall during the dry season (Funch et al. , 2006)

Sub-montane semi-deciduous seasonal forests occupy the lower eastern facing slopes (starting at about 400 m a.s.l.) in exposed areas with sufficiently deep soils, as well as the sides of deep river valleys among boulders and rock outcrops. These forests, however, have been eliminated from huge areas on the slopes and within the valleys of the Serra do Sincorá due to both mining activities (which removes the soil) and fire, and these sites have been occupied by: 1) campo rupestre vegetation (following soil loss on mountains slopes above 800 m ); 2) transition vegetation composed of forest/shrubs and some elements of campo rupestre (following soil loss on the slopes below 800m); 3) sedge meadows, in high mountain valleys on deep sandy soils (due to the very high frequency of fires), or; 4) damaged forest (maintained in the earliest stages of growth and succession by repeated burning).

As the sub-montane lithosolic forests gain in altitude within the mountains, there is a gradual alteration of the species composition, and they grade into a more evergreen physiognomy as environmental conditions become increasingly more humid. There are also more protected sites available in the mountains in deep valleys and crevices in the sandstone rock where very little mining activity was ever developed and the vegetation is sheltered from both drying and wildfires. These sites have a significant accumulation of damp organic material on the forest floor, and are populated by species such as Podocarpus transiens (Plig.)deLaub. , Hedyosmum brasiliensis Mart. , Weinmannia paulliniaefolia Pohl ex Ser. , and Drimys brasiliensis Miers.

Large open valleys in the central portion of this range ( 900 m and above) that were once densely forested have been extensively burned or cleared for farming (see section on sandy sedge meadows above), converting approx. 130 Km² of the montane lithosolic forest into open sedge meadows.

4) The river courses in the mountain canyons are bordered by a narrow strip of sub-montane to montane evergreen riparian forests (F3), with its own unique species composition (see LS Funch, 1998) that includes Vochysia pyramidalis Mart., Bonnetia stricta Mart. (although these species can also be associated with water seeps on otherwise drier slopes), Clusia nemorosa G.Mey., Couma rigida Müll.Arg., Balizia pedicellaris (DC.)Barneby&J.W.Grimes, and Anaxagorea dolichocarpa Spraque&Sandwith (LS Funch, 1997; Ribeiro Filho, 2002; Stradman, 1997, 2000). The riparian forests usually grade rapidly into sub-montane to montane semi-deciduous seasonal forests at very short distances from the river courses as the soil becomes drier and the inclination of valley sides become more pronounced. At higher altitudes, they become continuous with the montane evergreen forests.

Riparian forests have been severely damaged and reduced within the entire Sincorá Range by mining, farming, and fire, and may be restricted to the width of just a few meters, although their importance as corridors for dispersal, especially for animals (Machtans et al. , 1996; Spackman and Hughes, 1995) can be disproportional to their actual size. An important refuge from the damage being inflicted on this forest type is found in the deep, narrow, and more protected valleys on the mid-slopes on the eastern side of the mountain range where no significant mining activity was undertaken and the humidity in the narrow canyons protects the vegetation from wild fires.

5) Sub-montane semi-deciduous seasonal forests on deep latosols (latosolic forests) (F1) are found east of the Sincorá Range growing on deep latosols at altitudes from 400- 600 m a.s.l. (Fig. 5 & 13). The landscape is generally that of a slightly undulating plain occasionally dissected by relatively narrow but often fairly deep “V” shaped river valleys. Rainfall is high at the foot of the Sincorá Range (approx. 1200 mm yr -1 ) but the region becomes progressively drier to the east and the forests there more deciduous.

Previous studies in the region of the CDNP (LS Funch et al. , 2005; LS Funch et al. , 2006) have demonstrated that the canopy layer of these forests attains 10-20m, and is dominated by species such as Copaifera langsdorffii Desf. and Aspidosperma discolor A.DC. (often emergents) as well as Pogonophora schomburgkiana Miers ex Benth. , Protium heptaphyllum March., Pouteria ramiflora Radlk., Terminalia brasiliensis Eichl., and Hymenolobium janeirense var. s tipulatum (N.Mattos)H.C.de Lima. With the possible exception of Pouteria ramiflora (known locally as massaranduba), these remaining species have little commercial value in terms of their lumber. Species such as Tapirira guianensis Aubl. (pau-pombo ), Chaetocarpus echinocarpus (Baill.)Ducke (pau-de-colher), and Simaouba amara (paraíba) are early successional species, and usually associated with more recently disturbed forest areas.

The sub-canopy of these same forests attains from 6 to 8 m , and is composed of species such as Micropholis gardneriana Micropholis gardneriana Pierre , Schoepfia obliquifolia Turcz ., Myrcia detergens Miq., and Pouteria torta Pouteria torta Radlk . The understory is rich in species of Rubiaceae and Polygalaceae as well as immature individuals of the canopy and sub-canopy species. Vines are also present, including Coccoloba confuse R.A.Howard , Bauhinia sp., and Phryganocydia corymbosa Bureau ex. K.Schum., while epiphytes are usually rare, but include Vanilla sp.

A majority of the arboreal species registered in the area are widely distributed both within the Chapada Diamantina and throughout the diverse forest formations from northern South America (sometimes Central America ) to southern Brazil and northern Argentina , such as Aspidosperma discolor , Tapirira guianensis , Protium heptaphyllum, and Copaifera langsdorffii (LS Funch, 1997). The species restricted to the regional tableland forests include Eschweilera tetrapetala S.A.Mori (LS Funch et al. , 2006), Dalbergia cf. nigra Allem. Ex. Benth., Tabebuia cf. serratifolia Rolfe, and T . cf. avellanedae Lorentz ex Griseb.

More towards the west, near the Sincorá Range itself, the landscape becomes irregular and hilly, and the latosols terminate rather abruptly at the very foot of the mountains, except for some isolated “islands” of red clay soil almost invariably located on the rounded crests (usually between 600 and 800m) of the otherwise rocky and thin-soiled mountain slopes of the eastern flank of this range. The only intact forests growing on latosol islands in the Sincorá Range are limited now to the areas on either side of the principal diamond mining zone (that is, north of the Mandassaia River , and south of the Piaba River ). This central gap resulted from intensive mining activities in the late 19 th and early 20 th century that left behind only scattered remnant scraps of intact soil (and forest). But even the forest islands outside of the mining zone are at least as heavily damaged as the lowland tableland forests, for they have been occupied by landless farmers and are subject to more fire damage (they are burned through agricultural practices but are also surrounded by the drier and more frequently burned mountain vegetation).

As such, the once continuous forest that blanketed all of the area east of the Sincorá Range, as well as portions of its flanks, has now been replaced by a tattered and discontinuous fabric of pillaged patches as a consequence of selective logging, burning, and clearing for farming and pasture formation (usually involving the production of charcoal after the more commercially valuable wood has been cut for lumber). Even what appears on satellite images to be a more compact forest block to the NE of the National Park is, for the most part, a heavily fire-damaged early sucessional forest.

While spontaneous (natural) fires almost certainly can occur in the Chapada Diamantina, an overwhelming percentage of wildfires are anthropogenic and related to escaped burns intended to clear lands for pasture/agricultural transformation (or renovation), to scorch native savannas and grass lands to improve grazing conditions (these uncontrolled fires can then spread to other vegetation formations), or to hunting (either to drive game, improve their “pasture”, or remove cover). Criminal or accidental burning are also very common on otherwise unoccupied lands.

Nonetheless, the best preserved sections of sub-montane semi-deciduous seasonal forests on deep latosols are nestled right against the slopes of the Sincorá Range where both rainfall and humidity tend to be greater and the steep terrain presents less favourable conditions for agricultural/pastoral invasion, harvesting, mining, and wildfires.

6) Transition/Mosaic vegetation (T) forms are extremely common in the mapping region, as would be expected in an area with a mountainous landscape. The abrupt altitudinal variations, which generate changes in rainfall, temperature, solar radiation, wind velocity, dew point, etc. combined with landscape factors such as local geological and edaphic conditions, aspect, and slope, generate an extremely wide range of habitats and niches that is in turn reflected in the composition of the local vegetation.

A number of vegetation types present in the Sincorá Range are not present within the CDNP (or at best very poorly represented in terms of their area), and include the xerophilic caatinga and cerrado vegetations on the western side of the range.

Cerrado (C) vegetation dominates a rather distinct sector in the west-central to south-western region of the study area (approx. 1600 km²). It can be readily seen in the field, as well as on the satellite images, that the cerrado zone in the mapping area is divided into two principal geomorphological areas. In the smaller, very northern sector the landscape is deeply dissected and more heavily occupied for traditional (manual) agricultural uses, due to the greater density of permanent water courses in these deep valleys and their generally more humid conditions. The bulk of the cerrado zone, however, has a generally gently rolling landscape of extremely deep soils that range from sandy argisols to white, yellow, or (more rarely) red clays (latosols), and it is dominated by more open forms of cerrado ( campo sujo, campo cerrado ), with more arboreal forms ( cerradão ) occupying only the deeper depressions/drainage courses. Some of the typical cerrado species in this area include Anacardium humile Mart., Annona coriacea Mart., Duguetia furfuracea R.E.Fr., Croton campestris A.St.-Hil., Axonopus sp., and Hancornia speciosa Gomes. Intensive mechanized agricultural use of these lands initiated in the late 1970's, and significant areas have cleared and are currently under cultivation (especially using central pivot irrigation techniques). Nonetheless, this region has the highest percentage (approx. 35%) of intact landscapes within the study area (lands that have not been mechanically disturbed, but only used for grazing).

Caatinga (caat) vegetation is encountered only in the north-western mapping sector of this study although it is by far the dominant vegetation type in the entire region to the west of the study area. Almost all of this caatinga vegetation has been severely modified by anthropogenic processes – principally intensive cattle grazing. The seasonal droughts force the cattle to scavenge widely and intensively for forage, and the generally flat to rolling landscape facilitates their movement. As a result, very few areas have escaped this intensive use. Interestingly, wildfires do not seem to be a significant factor in determining the conservation status of the caatinga vegetation, in spite of the extreme dryness of this region. Whether this is due to a more careful control of fire by local residents or some natural tendency of the caatinga vegetation to propagate combustion poorly is not known.

Vegetation Map – Legend

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