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Interior Dry and Mesic Forests: CPD Site SA20

ESPINHAÇO RANGE REGION
Eastern Brazil

Location:  Phytogeographic province of Espinhaço mountains in Bahia and Minas Gerais states, between latitudes 10°-20°35'S and longitudes 40°10'-44°30'W.
Area: 
6000-7000 km².
Altitude: 
.
Especially 1000-2107 m.
Vegetation: 
Mosaic of formations with extensive ecotonal areas. Largely "campos rupestres": open-rock pioneer vegetation and rock-dwelling plants; oligotrophic marshes; gallery forests; (savanna-like) "cerrado"; montane (cloud) forests; semi-deciduous to deciduous forests.
Flora: 
Very high diversity, probably more than 4000 species of vascular plants. Campos rupestres with very high generic and specific endemism, and large number of disjunct taxa of coastal "restingas", other mountains of central Brazil (mainly Goiás) or Guayana Highlands.
Useful plants: 
Ornamentals, especially everlasting plants and Velloziaceae, and other tepaloid monocotyledons e.g. orchids, bromeliads; medicinals; potentially, other genetic resources.
Other values: 
Watershed protection for drinking water, hydroelectric energy; tourist attraction.
Threats: 
Fire; cattle-grazing; mining; timber extraction for charcoal, building, fences, etc.; over-collecting of horticultural species; increasing tourist pressure; inappropriate hydroelectric schemes and road construction.
Conservation: 
Serra do Cipó National Park (Minas Gerais), Chapada Diamantina NP decreed (Bahia). An Ecological Station, State Parks, Environmental Protection Zones (APAs).

Map 56: CPD Site SA20
References

Geography

The Cadeia do Espinhaço (Espinhaço Range) forms the watershed between the Atlantic Ocean and the basin of the São Francisco River in eastern Brazil (Map 56). From 50 km to 100 km wide, it is formed of numerous low mountains or "serras" (900-1500 m high, peaking at over 2000 m), interrupted by river valleys that can be extensive and deep. The region naturally falls into a number of mountainous areas which represent local centres of biodiversity. The southern limit (at 20°35'S) is in the central State of Minas Gerais south-east of Belo Horizonte in the Serra do Ouro Branco; six other important centres in this state are Serra da Piedade, Serra do Caraça, Serra do Cipó, Diamantina Plateau, Serra do Cabral and Serra do Grão-Mogol. In the State of Bahia 1100 km to the north (inland from Salvador), the Chapada Diamantina is an extensive elevated area which includes a number of important centres, such as in the west the Serra do Rio de Contas, Pico das Almas and Pico de Itabira and in the east the Serra do Sincorá. Farther north the Chapada dies away (at c. 10°S) in a number of isolated and scattered massifs, such as Morro do Chapéu (Serra do Tombador) and Serra da Jacobina.

The Espinhaço Range was formed intermittently in segments from the Palaeozoic onwards (King 1956). The highest peaks (1800-2107 m) are remnants of older surfaces that have been worn down. The oldest known denudation-surface dates from the Early Cretaceous; at its base are large areas of much older rocks. The structural blocks of the Espinhaço Range date to the Precambrian (Abreu 1984). The large-scale folding along a north-west/south-east axis underwent a long process of erosion and was remodelled by tectonic movement at the beginning of the Tertiary. Relief is highly accentuated, with broad deep valleys, especially in schistose and filite formations, whereas the highest ridges and crests occur in quartzite and sandstone formations (Moreira 1965). Soils are in general shallow and sandy, highly acidic and extremely nutrient poor.

Between the two main areas of the Espinhaço Range (one in Bahia, the other in Minas Gerais) lies lowland with deeper soils, c. 300 km broad from north to south and mostly over 500 m above sea-level, although dissected by large river systems such as the Contas River, with the Pardo and Jequitinhonha rivers farther south. This lowland is a barrier to migration of the northern (Bahian) Espinhaço flora to the south and vice versa, and probably acted as such in the past when it could have been drier or covered by forest (Harley 1988).

The climate is mesothermic (Cwb of Köppen 1931), with mild summers, a rainy season in the summer and average annual temperatures between 17.4°-19.8°C; the average in the hottest month is below 22°C. The rainy season lasts 78 months, while the dry period may last 3-4 months and coincides with winter. Annual rainfall in the southern part of the region is c. 1500 mm. Northwards the dry season increases in duration, until at least in the lowlands it may occupy the major part of the year. Although a strongly seasonal climate prevails, moist clouds provide dew and some rain during the dry season, allowing some soils to remain wet throughout the year.

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Vegetation

Although the Espinhaço Range is entirely within the tropics, its landscape is far removed from most of tropical Brazil. The physiognomy varies greatly according to the locally prevailing physiography, with the largest vegetation formation being "campos rupestres" (rocky fields) (in the terminology of Magalhes 1966).

Various types of forest occur in the Cadeia do Espinhaço. Gallery forests are alongside streams, with a tree stratum 7-15 m high. The widespread dominant trees are Tapirira guianensis, Xylopia emarginata, Richeria grandis, Copaifera langsdorffii, Protium almecega, P. brasiliense, Vismia guianensis, Tibouchina candolleana, Hyeronima alchorneoides, Croton urucurana, Guapira opposita, Humiria balsamifera, Guatteria sellowiana, Cabralea canjerana and Vochysia acuminata. At higher altitudes with increased humidity often occur such species as Drimys brasiliensis, Podocarpus spp., Weinmannia spp. and Lamanonia spp. At the margins and in the lower strata appear small trees (Trembleya parviflora, many Myrtaceae and Rubiaceae) and palms (Geonoma spp.), together with tree ferns (Cyathea, Trichipteris). The gallery forests may extend upwards and link with usually isolated forest clumps ("capões"), which occur in open grasslands at higher altitudes. The passage from gallery forests to capões presents a gradual modification in the floristic composition and the amount of vegetation that is deciduous. At higher altitudes, forests are strongly influenced by atmospheric humidity, clouds, etc. and may be rich in epiphytes, including orchids, bromeliads, Anthurium, Peperomia, ferns and bryophytes.

Where local geological conditions permit, for example on calcareous soils, deciduous or semi-deciduous forests with a very different species composition may occur. On the Bambuí limestone formation scattered through the region dry deciduous forest appears, including Cedrela fissilis, Apeiba tibourbou, Luehea divaricata, Guazuma ulmifolia, Molopanthera paniculata, Phyllanthus acuminatus, Cnidoscolus urens, Hymenaea courbaril, Chorisia sp., Allophylus sericeus, Aloysia virgata, Trema micrantha, Trichilia spp. and the tall columnar cactus Cereus calcirupicola.

On slopes between 860-1000 m occur areas of "campo cerrado", in which the predominance of arboreal-shrubby forms declines with increase in altitude and greater development of the soil profile, and gradually gives way to "campo sujo" and campo rupestre. These cerrado patches have a peculiar physiognomy, marked by sparse tortuous treelets and shrubs often with thick bark, mainly Vochysia thyrsoidea, Qualea cordata, Q. parviflora, Byrsonima verbascifolia, Neea theifera, Caryocar brasiliense, Kielmeyera coriacea, Didymopanax vinosus, Hymenaea stigonocarpa, Dalbergia miscolobium, Campomanesia adamantium and C. pubescens. These species are scattered through a continuous ground cover of a variety of herbaceous species, particularly the grasses Aristida, Paspalum stellatum and Axonopus brasiliensis.

In the northern (Bahia) part of the Espinhaço Range, a form of campo cerrado occurs at these altitudes that is locally known as "campos gerais", in which the tree stratum is usually absent and the ground stratum rich in stemless palms and with fewer grasses. The "campo rupestre", which forms the bulk of the vegetation on the higher slopes (1000-2000 m) or even from 700-900 m in some areas, is primarily composed of a more or less continuous herbaceous stratum and sclerophyllous evergreen small shrubs and subshrubs often with imbricate leaves, showing much morphological convergence in various families (Menezes and Giulietti 1986; A. Giulietti et al. 1987). However, rather than being a homogeneous vegetation type, these campos are an assemblage of communities forming a rich mosaic, under control of local topography, the nature of the substrate, and the microclimate.

There are patches of pure sand intermixed with patches of pebbles or gravel. Here and there stand rocky outcrops of different sizes that provide shade or shelter from prevailing winds and hold precious moisture in crevices, although the outcrops may increase heat by reflection. The interaction of all these factors is instrumental in the formation of a huge variety of micro-habitats. The bare rock surfaces and crevices are colonised by many lichens and other epilithic plants adapted to exploit dew and mist.

The adaptations include specialized roots and pseudobulbs in orchids, water tanks and specialized hairs in bromeliads and pseudo-trunks covered with the remains of leaf-sheaths between which are numerous adventitious roots in Velloziaceae. Other adaptive strategies for survival in this harsh landscape can be seen in species of many different families. These include waxy leaf surfaces which reflect heat, thick layers of hairs which protect against radiation and reduce transpiration, tightly furled rosettes of leaves and chemical secretions which protect against fire, and many anatomical and physiological modifications that give greater water-storage capacity.

Certain families, genera and species are very typical of the sandy or rocky substrates that form much of the campos rupestres, often occurring in large and sometimes showy populations and contributing to the general physiognomy of the vegetation. These taxa include Velloziaceae (Barbacenia, Vellozia, Pleurostima); Eriocaulaceae (Leiothrix, Paepalanthus, Syngonanthus); Xyridaceae (Xyris); Compositae (Lychnophora, Proteopsis, Wunderlichia, many other genera); Melastomataceae (e.g. Cambessedesia, Microlicia, Marcetia); Ericaceae (Gaylussacia, Leucothoe); Labiatae (Hyptis, Eriope); Leguminosae (Chamaecrista, Calliandra, Mimosa, Camptosema); Rubiaceae (Declieuxia); Lythraceae (Cuphea, Diplusodon); Malpighiaceae (Camarea, Byrsonima); Verbenaceae (Lippia, Stachytarpheta); Myrtaceae (Campomanesia, Myrcia); Euphorbiaceae (Croton, Phyllanthus); Orchidaceae (Laelia, Cleistes, Oncidium); Bromeliaceae (Dyckia, Vriesea, tank species of Tillandsia); Cyperaceae (Bulbostylis, Lagenocarpus); Gramineae (Aristida, Panicum, Paspalum, Axonopus and many other genera, including the bambusoid Aulonemia effusa and Chusquea spp.); and certain ferns and their allies (Anemia, Doryopteris, Huperzia).

On plateaux with poor drainage acid bogs may form, having Cyperaceae (Rhynchospora); Rapateaceae (Cephalostemon); Eriocaulaceae (Syngonanthus, Leiothrix); Xyridaceae (Xyris, Abolboda); Orchidaceae (Habenaria, Stenorrhynchus); Lentibulariaceae (Utricularia, Genlisea); and Droseraceae (Drosera).

The Espinhaço Range is crossed by a large number of streams supporting a rich and varied aquatic vegetation, with species of Eriocaulon, Utricularia, Ludwigia, Eleocharis, Mayaca, Nymphoides, Lycopodium, Lycopodiella, Apinagia and Laurenbergia tetrandra.

The striking differences between the Espinhaço vegetation and the surrounding lowlands are clearly determined by the differences in geological and topographic conditions, as well as the upland presence of clouds and dew, which are absent in the lowlands during the dry season. In Bahia, on descending from the serras, the luxuriant campos usually give way to a band of cerrado or campos gerais, soon replaced at lower altitudes by xerophilous deciduous forest known as "caatinga" (CPD Site SA19), which is the dominant vegetation formation of the semi-arid regions of north-eastern Brazil. In Minas Gerais, the upland formations are surrounded by lower regions with deeper latosols occupied by cerrado (see CPD Site SA21), the dominant formation on the Brazilian Central Plateau.

These factors are important in explaining the composition, affinities and divergences of these adjacent floras, but the patterns also reflect past history, such as former migration routes and the Pleistocene climatic changes in eastern Brazil.

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Flora

The first naturalists who visited eastern and central Brazil in the 19th century were greatly impressed by the wealth of the flora of the Espinhaço Range. Taxonomic and floristic researchers are now providing detailed evidence of the great floristic richness. Teams of the Universidade de São Paulo and the Royal Botanic Gardens, Kew, U.K. are elaborating inventories of the vascular flora of selected sites within the Espinhaço, to produce local Florulas that will provide better understanding of the distribution and dynamics of the species. Inventories have been published for Mucugê, Bahia (Harley and Simmons 1986) and the Serra do Cipó, Minas Gerais (A. Giulietti et al. 1987). Other areas being studied are the Catolés and Pico das Almas massifs in Bahia, and the Serra do Grão-Mogol, Diamantina Plateau and Serra do Ambrósio in Minas Gerais.

The flora of the Espinhaço Range probably has more than 4000 species. About 200 km² of the Serra do Cipó show the extraordinary number of 1590 species (A. Giulietti et al. 1987), in 103 families of dicotyledons (402 genera), 24 families of monocotyledons (118 genera), 1 family of gymnosperms (Podocarpaceae), 10 of ferns and 11 of bryophytes. The best represented families among dicotyledons are Compositae (169 spp.), Melastomataceae (90), Rubiaceae (47), Fabaceae s.s. (47), Myrtaceae (45), Malpighiaceae (42), Caesalpiniaceae (34), Euphorbiaceae (31), Verbenaceae (26) and Mimosaceae (26), whereas in monocotyledons Gramineae are the most speciose family (130 spp.), followed by Eriocaulaceae (84), Orchidaceae (80), Velloziaceae (58), Xyridaceae (47), Bromeliaceae (36) and Cyperaceae (32).

Elsewhere in the Espinhaço zone the same families are dominant, with variations in relative species richness, as observed for Mucugê (Harley and Simmons 1986). Provisional figures for the Pico das Almas are also broadly the same, though the total of species for this smaller area is c. 1200. Compositae is again represented by the most species, with Melastomataceae, Rubiaceae, Leguminosae and Myrtaceae also well represented among dicotyledons; a greater number of Lentibulariaceae is notable. Among monocotyledons Cyperaceae predominate, with relatively fewer Orchidaceae, Gramineae, Eriocaulaceae, Xyridaceae and Velloziaceae.

A number of phytogeographical elements can be found within the flora of the Espinhaço Range. The majority of forest species show a wide distribution through South America, e.g. Tapirira guianensis, Protium heptaphyllum, Cabralea canjerana, Richeria grandis, Hyeronima alchorneoides and Simarouba amara (Giulietti and Pirani 1988). Some genera have a greater number of species in the Andes and reach south-eastern Brazil (e.g. Hedyosmum, Clethra, Weinmannia, Podocarpus), whereas others show a greater concentration in eastern Brazil although occurring also in northern South America and Central America (e.g. Eugenia, Euplassa). The gallery forests, capões and cloud forests may represent relict communities from a period when forest covered much more of eastern Brazil and probably they acted - and still act - as migration routes for tree species. The epiphytic flora of the more humid altitudinal forest shows a clear link with the Atlantic Coast forest (Mata Atlântica) (e.g. with Oncidium crispum, Peperomia spp., Billbergia amoena, B. vittata).

The flora of the campos rupestres is largely composed of an endemic element, both in genera and species. They may show floristic links to other parts of South America, where vicariant taxa occur; in some cases such patterns can be found in individual species, which have a disjunct distribution between the campo rupestre and other regions. Probably the greater portion of the campo-rupestre flora has common elements with the cerrado flora, which in many areas surrounds it. This cannot be said of the caatinga flora, which is often adjacent to campo rupestre in Bahia but has a very different floristic composition, except for widespread genera that occur in a wide range of ecosystems.

Among genera in which a majority of the species are characteristic of either or both cerrado and campo rupestre are Eremanthus, Vanillosmopsis, Qualea, Campomanesia, Camarea, Peixotoa, Hyptis, Ctenium, Aristida, Mimosa, Mandevilla, Macrosiphonia, Diplusodon, Kielmeyera, Trimezia, Jacaranda, Gomphrena, Declieuxia and the nearly stemless Syagrus spp.

Another element in the campo-rupestre flora contains those species or groups of species that show a disjunct distribution between the montane campos rupestres and the coastal sands ("restingas"). Both habitats share certain edaphic and climatic factors - a quartzitic substrate in a very open habitat with high insolation and frequent periods of high atmospheric humidity. Species in common include the leafless Phyllanthus angustissimus and P. klotzschianus, Mandevilla moricandiana, Marcetia taxifolia, Mimosa lewisii, Lagenocarpus rigidus, Vellozia dasypus, Leiothrix rufula, Paepalanthus ramosus and most species of Syngonanthus section Thysanocephalus. Certain genera show a marked preference for these two ecosystems, including Gaylussacia, Leucothoe, Allagoptera, Bonnetia and Moldenhawera. A genus very typical of the campo rupestre is Eriope, although E. blanchetii is endemic to the restingas of Bahia and Sergipe.

The "tepui" region of the Guayana Highlands (CPD Site SA2) shares many similarities with the campo rupestre; both are formed in part from Precambrian shields that occur north and south of the Amazon. Both regions contain open areas of rock outcrop surrounded by low-lying vegetation that forms a barrier to migration and both experience high atmospheric humidity. Owing to the great distance that separates these regions, few species are restricted to both. Nonetheless, certain families that have their main centre of diversity in the tepuis have outliers or secondary centres in the campo rupestre of the Espinhaço Range. Examples include Cottendorfia, with C. florida in the campos rupestres of Bahia; Bonnetia, with B. stricta in the campo rupestre of Bahia and also occurring in the restinga. In the Rapateaceae, which is centred in the Guayana Highlands (16 genera, c. 100 spp.), Cephalostemon riedelianus only occurs in the campo rupestre of Minas Gerais. Families such as Xyridaceae and Eriocaulaceae have two main centres of diversity - in the campos rupestres of Brazil and the Guayana Highlands. In Xyridaceae, the major generic diversity occurs in the latter region with three small endemic genera, although the number of species in the largest and most widespread genus Xyris is fewer there than in the campos rupestres of Brazil. In Eriocaulaceae, the greatest morphological diversity also can be found in the Guayana Highlands, with up to two or perhaps three endemic genera, although the number of species, notably in Leiothrix but also Paepalanthus and Syngonanthus, is much higher in the campos rupestres. Marcetia taxifolia shows disjunction (reflecting that found in Bonnetia), with a distribution not only in the campos rupestres but also in the restingas of Brazil and the tepuis. For Marcetia taxifolia, however, the isolated occurrence is in the Guayanas, with the main centre of diversity of the genus in the Espinhaço Range.

Elements with an Andean or a southern Brazilian connection occur sporadically within the campo-rupestre flora. In Paepalanthus subgenus Platycaulon, with 46 species, 28 are restricted to the campos rupestres of Minas Gerais, whereas 12 are endemic to the mountains of Colombia. Vellozia section Radia has centres of diversity in the Brazilian campos rupestres and in some mountains of Venezuela and Colombia. Similar links can be found in Labiatae: Hyptis irwinii and H. stachydifolia have their closest relations in the Andes, and Eriope macrostachya, which occurs in campo rupestre along forest margins, also occurs (as Eriope macrostachya var. platanthera) in the Venezuelan Andes.

Endemism can be extremely high in the groups with a primary or important centre of diversity in the campos rupestres of the Espinhaço Range. The very characteristic Velloziaceae has c. 250 species in South America and Central America and only 29 in Africa. More than 70% of the species (173 spp.) are concentrated in Minas Gerais, essentially on the Diamantina Plateau and Serra do Cipó (Mello-Silva 1989). Barbacenia has c. 65 species mostly in the southern sector of the Espinhaço (Minas Gerais), with only three in Bahia. Burlemarxia, described by Menezes and Semir (1991) with three (two new) species, is endemic to the Diamantina Plateau. The data presented by A. Giulietti et al. (1987) show that among the 59 species of Velloziaceae found on the Serra do Cipó, 46.5% are restricted to that range, and 91.5% are endemic to the Espinhaço; 27 have been described in the last 23 years. In the Grão-Mogol area, 19 species of Velloziaceae include nine endemics and three new taxa (Mello-Silva 1989).

In Eriocaulaceae, Paepalanthus subgenus Xeractis (27 spp.) and Leiothrix subgenus Leiothrix (13 spp.) are wholly restricted to the Espinhaço Range in Minas Gerais. For the Serra do Cipó, A. Giulietti et al. (1987) refer to 84 species in this family - 68.7% being endemic to the Espinhaço and 32.5% restricted to the Serra do Cipó. In the Serra do Ambrósio (near Rio Vermelho) occur three endemic and related species of Syngonanthus, with high economic value, which are only now being described. In Xyridaceae, Wanderley (1992) pointed out that of the 152 species of Xyris found in Brazil, 46 are endemic to the Espinhaço Range and 14 exclusive to the Serra do Cipó.

Many other groups of flowering plants show numbers of endemic taxa in the Espinhaço Range that are as high. Striking examples are Pseudotrimezia (Iridaceae) and Cipocereus (Cactaceae), genera endemic to the southern (Minas Gerais) sector of the Espinhaço. The monotypic genera Raylea (Sterculiaceae), Morithamnus and Bishopiella (Compositae) are confined to the Bahian sector.

The species composing these and many other genera in the campos rupestres of the Espinhaço are generally found as disjunct populations, often restricted to single small serras isolated geographically and in gene exchange by natural barriers. This situation is the outcome of a long phase of climatic fluctuation during the Quaternary, when populations expanded and contracted as conditions improved or deteriorated, with resultant irregular gene exchange between neighbouring populations and the evolution of new species, often with very limited present distributions.

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Useful plants

The Espinhaço Range has a long history of occupation and exploitation of minerals. The plant resources have barely been utilized for food or medicinals on a large scale. The large number of taxa in the region undoubtedly represents a vast genetic resource. Plants used locally for food mainly include many fruits, e.g. "goiabinha" and "araçá" (Psidium spp.), "guabiroba" (Campomanesia spp.), "puçá" or "mandapuçá" (Mouriri glazioviana), "jatobá" (Hymenaea spp.), "pequi" (Caryocar brasiliense), "mangabá" (Hancornia speciosa), "murici" (Byrsonima spp.), "cajuzinho-do-campo" (Anacardium spp.) and nuts of a number of palm species (e.g. "licuri" - Syagrus coronata, "catolé" - Attalea spp., "macaúba" - Acrocomia aculeata).

Local people prepare traditional remedies from many species, for instance: dried leaves of arnica (Lychnophora pinaster) are used to prepare a tea with hepatic properties; fresh leaves of L. ericoides in alcohol are said to heal wounds; shoots and leaves of "chapé" (Hyptis lutescens) are believed to cure flu; leaves of "paneira" (Norantea adamantium) provide a tea for kidney problems; roots of "quina-de-vaca" (Remijea ferruginea) are used for stomach diseases; "sambaibinha" (Davilla rugosa) is for baths; roots and leaflets of "caroba" and "carobinha" (Jacaranda spp.) are put into "pinga" (an alcoholic liquor) to cleanse the blood; and "catuaba" (Anemopaegma arvense) is said to be an effective tonic and aphrodisiac.

Other products obtained in the region include gums from jatobá (Hymenaea spp.), resins from "pau-d'óleo" or "copaíba" (Copaifera langsdorffii) and tannins from murici (Byrsonima spp.). The resinous stems of the larger species of Vellozia ("canela-de-ema") were used as fuel for locomotives on the Curvelo-Diamantina railway and these days are stacked to sell as an effective fire lighter for wood stoves and to make torches. The densely woolly leaves of some Lychnophora species provide combustibles for home fires and stuffing for pillows and cushions. The powdered roots of Palicourea marcgravii are a very effective rat poison.

A fairly recent development has been expansion of the trade in everlasting plants ("sempre-vivas") - collection and trade in dried inflorescences of some 40 wild species, several of them rare endemics, that are in demand particularly for export to U.S.A., Japan and Europe (N. Giulietti et al. 1987). About 300 metric tonnes dry weight are being obtained yearly. The species most prized for their beauty are the Eriocaulaceae Syngonanthus elegans ("sempre-viva pé-de-ouro"), S. venustus ("brejeira"), S. xeranthemoides ("jazida"), S. magnificus ("sempre-viva gigante"), S. suberosus ("margarida"), S. brasiliana ("brasiliana"), Paepalanthus macrocephalus ("botão-branco"), Leiothrix flavescens ("botão-bolinha"); the grasses Aristida riparia ("rabo-de-raposa"), Aulonemia effusa ("andrequicé"), Axonopus brasiliensis ("pingo-de-neve"), Gynerium sagittatum ("cana-brava"); the Xyridaceae Xyris nigricans ("coroinha"), X. cipoensis ("abacaxi-dourado"), X. coutensis ("cacau"); and the sedges Rhynchospora globosa ("espeta-nariz"), R. speciosa ("capim-estrela").

Many showy species are uprooted by horticulturists and tourists, especially orchids (e.g. Laelia, Cattleya, Oncidium, Encyclia advena - "sumaré"), bromeliads (e.g. Dyckia, Orthophytum, Vriesea), cacti (e.g. Uebelmannia gummifera, Melocactus, Micranthocereus auri-azureus) and "lilies" (mainly Hippeastrum).

Many other species in a wide range of families have potential as ornamentals. Many endemic species have never been investigated for possible uses - this a rich field for future research.

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Social and environmental values

Up to 1890 the greatest part of the population of the south-east was concentrated in Minas Gerais. More than a million gold and diamond prospectors lived in the region, principally distributed from Ouro Preto to Diamantina. In Bahia on the Chapada Diamantina, the population was large for the same reasons, with the mines principally from Rio de Contas to Catolés in the west and Mucugê to Lençóis in the east, with the road at that time extending northward to Jacobina. This period of mineral extraction, with indiscriminate exploitation of the natural resources, coincided with expansion of small towns. With the decline of this industry at the end of the last century, the towns fell on hard times. In recent years as means of transportation and communication have improved, many of these towns have become attractive centres of tourism, due to their picturesque surroundings and old colonial-style houses, as well as the rich diversity of the local fauna and flora. Ouro Preto and Diamantina in Minas Gerais and Lençóis in Bahia are examples where the main income is from tourism. In spite of the decline of gold and diamond mining, this activity still occurs on a small scale in the whole length of the Espinhaço Range; for example in Diamantina, diamonds still remain as the principal mining resource of the municipality. In the last few decades the extraction of quartz crystals for the electronics industry has developed in a number of areas, such as Serra do Cabral in Minas Gerais and Barra da Estiva and near Catolés in Bahia.

Various rivers have their source in the mountains of the Espinhaço Range and are of vital importance for the economic development of north-eastern and south-eastern Brazil. Among those flowing to the Atlantic are the Itapicuru, Jacuípe, Paraguaçu and Contas rivers in Bahia and Jequitinhonha, Mucuri and Doce rivers in Minas Gerais, and those draining west that are tributaries on the east bank of the São Francisco River: Salitre, Jacaré, Verde and Paramirim rivers in Bahia and Verde Grande, Pacuí and Velhas in Minas Gerais. Protection of their mountain sources is essential to maintain water quality and avoid excessive erosion and silting.

The Espinhaço Range is not a region of high soil fertility, being mostly suitable for subsistence agriculture with such crops as beans and cassava, and coffee production in some forested areas, principally the Chapada Diamantina. Cotton is cultivated in many areas, particularly the Rio das Velhas Basin to the north of Belo Horizonte. Various areas of campo rupestre have experienced the introduction (in the last two decades) of exotics such as Eucalyptus and Pinus. These species have not proved profitable - which was completely predictable, given the soil characteristics.

Throughout the region there is extensive cattle-ranching, using natural pastures which are annually burnt to stimulate young growth of herbs and grasses. In Minas Gerais a total of c. 80,000 km², principally with cerrados, are now used for this purpose, producing both milk and meat; in Bahia the occurrence is on a smaller scale. Upland pastures that retain forage during the dry season are invaded by grazing cattle from the lowland areas.

Over the last 15 or so years a large number of local communities in the Espinhaço Range have become involved in the collection and commercialization of sempre-vivas, especially in the neighbourhoods of the Serra do Cipó, Diamantina and the Serra do Cabral in Minas Gerais and from Mucugê and Piat in Bahia. This activity is seasonal, almost entirely restricted to January to August when the species of greatest commercial value can be found in flower. The flowering stems are collected, dried, sorted into bundles and sold for decoration in homes and floral displays. In the Diamantina district, this industry has become the second greatest source of income (after diamonds) from natural resources in the municipality. The entire city of Mucugê turns its attention during July to the collection and drying of the sempre-viva de Mucugê, with the drying carried out in the streets. This species became scientifically recognized only recently, as Syngonanthus mucugensis. On upper slopes of the Chapada do Couto (Minas Gerais), temporary settlements sometimes of over 900 inhabitants occur during April-May solely to collect and dry three species: Syngonanthus elegans, S. cf. bisulcatus ("sempre-viva chapadeira") and Xyris coutensis - the latter two are endemic to the Chapada do Couto. In the Serra do Ambrósio there are also many endemic species, four of which are commercially important: Uebelmannia gummifera, a globose cactus that has been much exported to Europe, and Syngonanthus brasiliana, S. magnificus and S. suberosus. The latter three species, which are also new to science, have the greatest wholesale value among all those used and are found only in that limited area. Species of sempre-vivas unknown to science have been discovered in local storage.

This region is the main part of the Bahia-Minas Gerais tablelands Endemic Bird Area (EBA B50), which has five restricted-range bird species confined to it: the hooded visorbearer (Augastes lumachellus), hyacinth visorbearer (A. scutatus), Cipó canastero (Asthenes luizae), grey-backed tachuri (Polystictus superciliaris) and pale-throated pampa-finch (Embernagra longicauda). The Cipó canastero was discovered as recently as 1990 and is endemic to a small area in the Serra do Cipó. It is the only one of these species currently listed as threatened, but the others are listed as near-threatened. Habitats are decreasing throughout this EBA.

Economic assessment

The Cadeia do Espinhaço has contributed its natural resources and services to the national economy in various ways. At present, current values on plant products have either not been calculated or may be difficult to extricate from the current literature. (No attempt is made here to produce figures for mineral extraction and other general forms of activity.)

For the sempre-vivas industry, which is almost exclusive to the Espinhaço Range, some figures have been supplied (N. Giulietti et al. 1987). Cases showing the situation in 1984 can be given: Syngonanthus elegans - 40,000 kg/year; S. venustus plus S. dealbatus ("brejeira") - 20,000 kg/yr; S. xeranthemoides - 12,000 kg/yr; Paepalanthus macrocephalus - 10,000 kg/yr; S. magnificus and S. suberosus each - 4000 kg/yr; Aristida jubata ("barba-de-bode") - 40,000 kg/yr; A. riparia - 20,000 kg/yr; Aulonemia effusa and Axonopus brasiliensis - each 12,000 kg/yr; Xyris nigricans - 5000 kg/yr; X. cipoensis - 1000 kg/yr; Rhynchospora globosa - 20,000 kg/yr; R. speciosa - 5000 kg/yr. Of 40 species with high economic value, those in Syngonanthus command the highest wholesale prices. Between 1974-1986, an average of 645,000 kg of sempre-vivas were exported annually. However, whereas 1,007,449 kg were exported in 1978, there were only 320,003 kg in 1986. This might be explained by a fall in product value and/or a reduction in the populations. In the same period the price per kg varied from just US$4.54-4.97, showing that an instability of prices cannot be the reason for the dramatic fall in trade.

The sempre-vivas industry may be an important factor in ensuring the survival of many local communities in the Espinhaço zone and reducing emigration to the towns. Usually whole families are involved in harvesting from the wild. To support the populace, the temporary settlements include markets and shops to sell food and other necessities to aid those working in the field. New strategies must be developed to rationalize the conservation of threatened species and the needs of the local communities. One solution may be cultivation of suitable species, such as Syngonanthus elegans in the Diamantina district where it is native. This species has already proved very productive in experimental cultivation, but most species of Eriocaulaceae appear to have very special growing requirements. Cultivation of suitable species, together with a policy for the sustainable collection of the wild plants, could go far in ensuring the survival of their native populations while having a beneficial effect on the local economies.

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Threats

Familiar mining activities, once carried out on a large scale in the Espinhaço Range, still persist localized to some areas, with development of new industries in others (manganese, etc.). The impact of major soil disturbances in the search for gold and diamonds is notorious, as long-abandoned mining areas remain bare and devoid of their former vegetation for decades and in some cases even longer. Soil erosion is greatly accentuated by construction of badly planned highways; frequently soil has to be brought from other areas for their construction, encouraging invasive weeds.

Today the main activity throughout the Espinhaço Range is cattle-raising, which is favoured by the existence of natural grassland. The destruction of these areas is accentuated by the trampling of cattle and annual burning carried out by local farmers to stimulate regrowth of the herbaceous vegetation. This has a selective effect on the species diversity of the grasslands, because although many species have adaptations to resist fire, its frequent and regular application eventually favours few species of animals and plants.

Due to the low fertility and shallowness of the soils, the impact of agricultural activities is greatly restricted in the mountains of the Espinhaço Range, compared to other areas such as the cerrado. Where the soils are deeper and more fertile, however, subsistence agriculture can be practised, mainly with sugarcane, beans, rice and maize. In a few places in the Chapada Diamantina on more extensive areas of level ground where water is available, cash crops also may be raised on a small scale. Particularly where deeper soils support forest, coffee and other crops have been planted after removal of the natural vegetation.

Forest and cerrado formations in the Espinhaço Range have been intensively exploited for fuelwood for domestic use and especially in production of charcoal. This is an important material in the iron and steel industries of Minas Gerais. Even in the campo rupestre, which is much less rich in woody plants, the collection of the resinous branches of various species of Vellozia (e.g. V. sincorana – "candombá", in the Serra do Sincorá) has caused a drastic reduction in the size of natural populations of these species, which probably take decades to reach maturity.

The indiscriminate collection of whole plants for horticultural use, particularly orchids, bromeliads, ferns and cacti, undoubtedly threatens the survival of a number of species. Those species that are the rarest, with small populations endemic to restricted areas, are most sought after for that very reason, as they will fetch the highest prices in the horticultural market. An example is the small orchid Constantia cipoensis, endemic to the Serra do Cipó and occurring as an epiphyte only on an undescribed species of Vellozia. As this Vellozia (also endemic to the mountain) is extremely tall, collectors not only strip the orchids from its branches, but also destroy it in the process.

Finally, the constant collection of sempre-vivas, which completely removes the inflorescences, is reducing the size of natural populations. For example, whereas the volume of Syngonanthus magnificus exported in 1984 was 4000 kg, in 1986 it fell to 1500 kg. This spectacular species, which fetched the highest prices, has a very restricted range, being found only in the Serra do Ambrósio, and is now greatly diminished in the wild.

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Conservation

The Espinhaço Range has two national parks. Chapada Diamantina National Park (Bahia), created in 1985, comprises 1520 km² already delimited; it is still in the initial phase of organization. Serra do Cipó NP (338 km²) (Santana do Riacho and Jaboticatubas, Minas Gerais) was created in 1984, and is delimited and in operation; it is surrounded by a buffer zone c. 30 km wide. However, the park's infrastructure is not yet sufficient for effective protection.

Other variously conserved areas include Gruta dos Brehões APA (Environmental Protection Zone), 119 km² in Bahia; and in Minas Gerais, Cavernas do Peruaçu APA, 1500 km² (Januária and Itacarambi); Cachoeira Andorinhas APA, 187 km²; Rio Doce State Park, 360 km²; Itacolomi State Park, 70 km² (Mariana and Ouro Preto); and Tripuí Ecological Station, 7 km² (Ouro Preto).

Considering the high endemism of the flora and fauna, these variously designated areas form only part of what needs to be conserved in the region. Areas in Bahia, such as Pico das Almas (1850 m) and Serra de Barbado (2107 m) – the highest point in the Espinhaço Range, and areas in Minas Gerais, such as in the vicinity of Diamantina in the central part of the range and Grão-Mogol in the north, are a few of the centres of extremely high diversity that are unprotected. Such natural areas must be included in a more coordinated plan for the conservation of the biodiversity in the region. Such a strategy needs a much greater knowledge of the diversity and distribution of the fauna and flora and a better understanding of the dynamics of the campo-rupestre ecosystem.

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Map 56. Espinhaço Range Region, Brazil (CPD Site SA20), showing land above 1000 m in the Espinhaço range, in the states of Minas Gerais and Bahia

References

Abreu, A.A. (1984). O Planalto de Diamantina: um setor da Serra do Espinhaço em Minas Gerais. Orientação - Instituto de Geografia Univ. São Paulo 5: 75-79.

Giulietti, A.M. and Pirani, J.R. (1988). Patterns of geographic distribution of some plant species from the Espinhaço Range, Minas Gerais and Bahia, Brazil. In Vanzolini, P.E. and Heyer, W.R. (eds), Proceedings of a workshop on neotropical distribution patterns. Academia Brasileira de Ciências, Rio de Janeiro. Pp. 39-69.

Giulietti, A.M., Menezes, N.L., Pirani, J.R., Meguro, M. and Wanderley, M.G.L. (1987). Flora da Serra do Cipó, Minas Gerais: caracterização e lista das espécies. Bol. Botânica, Univ. São Paulo 9: 1-151.

Giulietti, N., Giulietti, A.M., Pirani, J.R. and Menezes, N.L. (1987). Estudos em sempre-vivas: importância econômica do extrativismo em Minas Gerais, Brasil. Acta Bot. Bras. 1(2) (supl.): 179-193.

Harley, R.M. (1988). Evolution and distribution of Eriope (Labiatae), and its relatives, in Brazil. In Vanzolini, P.E. and Heyer, W.R. (eds), Proceedings of a workshop on neotropical distribution patterns. Academia Brasileira de Ciências, Rio de Janeiro. Pp. 71-120.

Harley, R.M. and Simmons, N.A. (1986). Florula of Mucugê. Chapada Diamantina – Bahia, Brazil. Royal Botanic Gardens, Kew. 227 pp.

King, L.C. (1956). A geomorfologia do Brasil Oriental. Rev. Brasil. Geogr. 18: 147-265.

Köppen, W. (1931). Climatología. Fondo de Cultura Económica, Buenos Aires.

Magalhães, G.M. (1966). Sobre os cerrados de Minas Gerais. An. Acad. Brasil. Ciênc. 38 (supl.): 59-70.

Mello-Silva, R. (1989). Velloziaceae de Grão-Mogol, Minas Gerais, Brasil. M.S. thesis. Universidade de São Paulo, São Paulo.

Menezes, N.L. and Giulietti, A.M. (1986). Campos rupestres. Paraíso botânico na Serra do Cipó. Ciência Hoje 4(26): 38-44.

Menezes, N.L. and Semir, J. (1991). Burlemarxia, a new genus of Velloziaceae. Taxon 40: 413-426.

Moreira, A.N. (1965). Relevo. In Instituto Brasileiro de Geografia e Estatística (IBGE), Geografia do Brasil, vol. 5. Grande Região Leste. IBGE, Rio de Janeiro. Pp. 5-54.

Wanderley, M.G.L. (1992). Estudos taxonômicos no gênero Xyris L. (Xyridaceae) da Serra do Cipó, Minas Gerais, Brasil. Ph.D. thesis. Universidade de São Paulo, São Paulo.

Authors

This Data Sheet was written by Dra. Ana Maria Giulietti and Dr José R. Pirani (Universidade de São Paulo, Departamento de Botânica, Caixa Postal 11461, 05499 São Paulo, SP, Brazil) and Dr Raymond M. Harley (Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England, U.K.).

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