Biologia 65/2: 1—, 2010 Section Botany DOI: 10.2478/s11756-010-0005-y Vegetation strategy of Vellozia crinita (Velloziaceae) Ruy José Válka Alves & Jiří Kolbek Museu Nacional, Quinta da Boa Vista s. no., São Cristóvão, Rio de Janeiro, RJ 20940-040, Brazil; e-mail: ruyvalka@yahoo.com Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice, Czech Republic; e-mail: jiri.kolbek@ibot.cas.cz Abstract: Plant communities in which the herb layer is dominated by Vellozia crinita were surveyed in seven mountain ranges with a complex comprising open herbaceous or savanna-like montane vegetation overlying quartzite and sand, locally known as “campos rupestres” in the State of Minas Gerais, southeastern Brazil, by means of 92 vegetation relevés (totaling approx. 4629 m2 ). Vellozia crinita displayed a growth strategy which is rare among the Velloziaceae south of the Espinhaço mountain chain. While most species of Vellozia are phanerophytes with erect caudices and grow exclusively on or among rock outcrops, V. crinita is a caespitose chamaephyte with a profusely branched, repent caudex which grows under the ground or very slightly exposed. It forms cushions which stabilize convex mounds of colluvial sand, thus occupying a distinct niche in the “campos rupestres”. One association, two subassociations and two variants of the white-sand vegetation component are described as new. Key words: campo rupestre; cerrado; phytocoenology; Velloziaceae; Minas Gerais; Brazil Introduction In the “campos rupestres” vegetation of southeastern and northeastern Brazil, white sand savannas alternate with quartzite outcrops, forming a mosaic in contact with several zonal vegetation types. Deeper soils in campo rupestre are usually colluvial deposits forming layers and pockets topped with white sand derived from erosion of the adjacent outcrops, and often include milky-quartz gravel. Benites et al. (2003) found that in campos rupestres: “Soils are shallow, sandy, oligotrophic and contain high amounts of exchangeable Aluminum. Humic to organic horizons are common in these soils indicating low degree of soil organic matter decomposition”. They conclude that, in comparison to soils on igneous lithologies, those overlying quartzite are “poorer in carbon and nutrients, exhibiting stronger signs of degradation that indicate a larger fragility of this biome.” These poorly developed soils rarely exceed 2m in depth, and their occurrence is dictated by local topography and vegetation. Over half of the Neotropical species of Velloziaceae are endemic to the campos rupestres of the Espinhaço chain (Giulietti & Pirani 1988; Giulietti et al. 1997). Many, if not all species of Vellozia are longevous and have very slow growth (Alves 1994). In deep and mechanically unstable sandy soils, in which the water table is deeper than 40–50cm, it can be unavailable to the herb layer for prolonged periods. In campo rupestre the herb layer of such areas is often dominated by low and caespitoseVellozia species, while the erect and shrubby ones usually grow on and among rock outcrops. Distinct c
2010 Institute of Botany, Slovak Academy of Sciences species of Vellozia have apparently developed specific strategies in these contrasting environments (Table 1). Vellozia crinita is endemic to a few campo rupestre localities in the State of Minas Gerais. In the white sand component of the seven studied campo rupestre localities, Vellozia crinita always displayed a peculiar, mound-forming growth strategy, which locally blocks erosion, and ultimately dictates vegetation physiognomy. The campos rupestres are naturally subjected to periodic savanna fire and occasional strong winds, to both of which they are well adapted (Kolbek & Alves 2008). However, these singular habitats are under increasing threat of disappearing, especially due to degradation by cattle (mainly grazing, trampling and manure deposition). In that respect it is alarming that, under such pressure, V. crinita is only considered vulnerable, by IUCN criteria, according to the most recent list of endangered species of Minas Gerais (BIODIVERSITAS 2007). The purpose of the present survey was to investigate the peculiar vegetation strategy of Vellozia crinita, which sets it apart from the erect, rock-dwelling species, to describe the vegetation units dominated by it in the white sand component of the southeastern campos rupestres, and to verify the implications of this relatively unique growth-form for habitat conservation. Material and methods White-sand savannas with Vellozia crinita were studied in seven small mountain ranges with campo rupestre in the State of Minas Gerais (Fig. 1): Serra Bico de Pedra 2 R.J.V. Alves & J. Kolbek Table 1. Distinct vegetation strategies of caespitose and erect Vellozia species. Strategy Erect, shrubby species Caespitose species Growing among outcrops where the discontinuous herb layer limits spread of fire. Prostrate caudex at least partly buried in sand, thus insulated from fire. Leaf rosettes of adult plants above flames from burning herb layer. Mounds with densely packed rosettes keep out grasses, reducing spread of fire. Velamehn radicum and poikilohydry. Velamen radicum and poikilohydry. Shallow substrate: roots reach water table more easily. Roots reach the deep water table in both arid and wet season (Fig. 2). Wind Rooting in mechanically stable substrate: outcrops and fissures. Low stature: prostrate mounds with caudices at least partly underground. Competition Adaptation to extreme oscillations of temperature and humidity. Densely packed rosettes in mounds hinder establishment of other herbs. Fire Drought Fig. 1. Relative positions of the study sites in this paper (round dots) and the Cipó range (square) studied by Garcia & Diniz (2003) and Garcia et al. (2007). (20.474◦ S, 43.595◦ W, 1071–1236m alt.), Serra da Chapada (20.471◦ S, 43.588◦ W, 1276–1286m alt.), Serra do Ouro Branco (20.490◦ S, 43.695◦ W, 1420–1529m alt.), the latter representing the southern limit of the Espinhaço chain, and including southern outliers: Serra de São José, site a (21.086◦ S, 44.169◦ W, 1175m alt.), Serra de São José, site b (21.114◦ S, 44.168◦ W, 897–899m alt.), Serra do Lenheiro (21.131◦ S, 44.306◦ W, 990–1194m alt.), Serra do Ouro Grosso (21.307◦ S, 44.638◦ W, 950–992m alt.), and Serra de Carrancas (21.440◦ S, 44.707◦ W, 1260–1302m alt.). The analyzed vegetation spans 256km from northeast to southwest, with 23,317km2 mostly within the Cerrado biome. The habitat delimitations used herein are somewhat simpler than those of Conceição & Pirani (2005). The combinations of substrate and local topography permit the rough distinction of three main types of terrestrial habitats in campo rupestre, each of which can harbor several distinct Fig. 2. Schematic bisect of the ass. Vellozio crinitae-Loudetiopsietum chrysothrici showing soil depths, underground caudices and root systems; A – Vellozia crinita; B – the shrub Hyptidendron asperrimum; C – the geophyte Dipladenia polymorpha; D – the grass Loudetiopsis chrysothrix outside the Vellozia mound; E – Xyris rupicola; F – the sedge Rhynchospora globosa; G – the sedge Rhynchospora tenuis; R – adventive roots of Vellozia crinita; S – summer level of water table; W – winter level of water table. Drawn according to photographs from the Serra de São José and soil profiles. types of vegetation: (1) rock outcrops and boulders, (2) rock crevices and fissures, and (3) deposits of sandy soil. All of these tend to form intricate mosaics, but relatively sharp borders usually separate the patches of vegetation. During over thirty field trips implemented from 1989 to 2006, relevés using the scale of Braun-Blanquet (1932) were systematically used to sample vegetation on white sand colluvial deposits in the studied sites. The sample plots were selected subjectively in what seemed to be homogenous vegetation stands, and had no defined geometric shape. Soil samples were excavated for the study of underground plant structures. Vegetation strategy of Vellozia crinita 3 Fig. 3. Hierarchical clustering of all obtained relevés. A and B –Ass. Vellozio crinitae-Loudetiopsietum chrysothrici; A – Subass. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis; B – Subass. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis; C – Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum consanguineae. The high dissimilarity levels among the three mail clusters were expected, and are due mainly to the heterogeneity of sample plots imposed by local topography. Dendrogram generated by SYN-TAX 2000 software. Relevés were pre-ordinated using JUICE 6.5.7 (Tichý 2002) and statistically analyzed in SYN-TAX 2000 software (Podani 2001), using a hierarchical classification algorithm (complete linkage, Euclidian distance) based on quantitative data, by β-flexible clustering method (β = −0.25), Chord’s coefficient was used. Nomenclature of all species found in the field follows determinations by specialists, for vascular plants subsequently cross checked and updated via the Missouri Botanical Garden’s Vast nomenclatural database W3 TROPICOS (2007). Authors of plant names follow Brummitt & Powell (1992). Distribution data was obtained from literature and complemented by access to the JBRJ (2007) database, W3 TROPICOS (2007), GBIF (2007), USDA (2007), and Clayton et al. (2007). In tables and appendix, numbers 100, 101, 102, 103 are replaced as 00, 01, 02, 03. Voucher specimens were deposited in the herbaria of the Rio Botanic Garden (RB) to 1996, and later in the National Museum, Rio de Janeiro (R). Results In stands with Vellozia crinita, the tree layer is usually less than 3.5 m tall, and very open (hence it was consid- 4 R.J.V. Alves & J. Kolbek Table 2. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis var. typicum. Relevé number 1 9 8 10 6 44 2 3 4 5 7 95 69 03 98 16 78 88 81 82 68 70 83 77 91 99 90 00 85 86 87 79 96 80 89 97 71 73 74 75 76 72 High shrub layer (A) Eremanthus . . . . . . . . . . . . erythropappus Myrsine monticola . . . . . . . . . . . . Hyptis fruticosa . . . . . . . . . . . . Eremanthus crotonoides . . . . . . . . . . . . Kielmeyera coriacea . . . . . . . . . . . . Eupatorium barbacense . . . . . . . . . . . . Microlicia cinerea . . . . . . . . . . . . Low shrub layer (B) Cuphea ericoides 1 1 1 1 1 1 . +1 1 1 + Tibouchina r . . . . . . . r . + 1 heteromalla Ageratum fastigiatum 1 r . . . . r . . . . . Pavonia cf. garckeana . . . . . . . . . . . . Marcetia taxifolia . . . . . . . . . . . . Baccharis calvescens . . . . . . . . . . . . Stachytarpheta . . . . . . . . . . . . reticulata Cuphea thymoides . . . . . . . . . . . . Hyptis conferta . . . . . . . . . . . . Baccharis platypoda . . . . . . . . . . . . Ditassa decussata . . . . . . . . . . . . Comolia sertularia . . . . . . . . . . . . Chresta pinnatifida . . . . . . . . . . . . Herb layer (C) Vellozia crinita 2 2 3 2 2 2 2 3 2 3 2 1 Melinis minutiflora 1 1 + 1 + + ++ . + . . Loudetiopsis chrysothrix. . . . . . . . . . . . Rhynchospora tenuis . . . . . . . . . . . . Lagenocarpus *rigidus 1 1 1 + 1 2 . . . . r . Ageratum fastigiatum 1 . 1 1 + . 2 2 1 1 . + Syngonanthus niveus . . r + 2 . . . . + . Spermacoce . . . . . . . . . . . r brachystemonoides Arthrocereus melanurus . . . . r . . . . r . . Rhynchospora recurvata . . . . . . . . . . . 2 Rhynchospora globosa + 1 + 1 1 1 . + + 1 . . Doryopteris ornithopus . . . . . . . . . . . . Chamaesyce caecorum . . . . . . . . . . . . Portulaca mucronata . . . . . . . . . . . . Xyris hymenachne . . . . . . . . . . . . Dipladenia polymorpha . . . . . . . . . . . . Xyris asperula . . . . . . . . . . . . Polygala paniculata . . . . . . . . . . . . Fimbristylis dichotoma . . + + . . . . . . . . Bulbostylis paradoxa . . . . . . . +1 . . . Symphyopappus . . . . . . . . . . . . reticulatus Gomphrena agrestis . . . . . . . . . . . . Myrsine monticola . . . r . . . . . . . . Xyris rupicola . . . . . . . . . . r + Schizachyrium tenerum . . . . . . . . . . . . Klotzschia brasiliensis . . . . . . . . . . . . . . . . . . . . . . . . Dyckia tuberosa Spermacoce verticillata . . . . . . . . . . . . Woody seedlings in herb layer (D) Eremanthus + . . . . . . . . . . . erythropappus Eupatorium barbacense . . . . . + . . . . . . Eremanthus speciosus . . . . . . . . . . . . Moss layer (E) Campylopus + . + + . . ++++ . . savannarum Cladonia bahiana . ++ . . . . + . . . . Campylopus pilifer . . . . . . . . . . . . Cladia aggregata . . . . . . . . . . . . Lichenes indet. . . . . . . . . . . . . % 1 r 2 + . . . . 2 . 1 . . 2 . r + . . . . . . . 3 2 2 3 2 2 36 . 1 . . . . + 1 . . . . 17 14 10 7 10 7 1 . r ++ 1 . ++ 2 1 1 1 1 1 1 1 + 2 + . r 1 1 2 1 1 1 1 1 1 1 . ++ 2 2 . . 1 . . 1 + 2 + . . ++ . . ++ 1 1 +++ . 90 57 . . . . . r . . . . . . + . . + . . . + + . + . + . . . . . + . + r 2 1 + r + 1 . . . . . . . . . . . + . . . . 1 . . . + r . . . . . . . . r . + 1 . . . . + . . . . + . . . . . . . r . + . . . . . . + . . . . . . . . . . . . . . . . . . . + . . . . + . . . . + + . . . + + . . . . . . . . . + . . 24 21 19 17 10 . . . . . . . . . . . . . . . . . . . . + . . . . . . + . . . . . . . . . . + + + . + . . + . + + 1 . . . . . r . . . . + 1 . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + . + . . . . . . . . 1 + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . 10 10 7 7 7 7 1 + + . 1 . 2 . 2 . 3 . + . + + 5 . . . . . . . 3 + 1 . . . . + 2 + 2 3 + + . 1 1 + . 2 + + . . 3 + 1 2 r . 1 2 2 1 2 2 + . . 2 2 2 3 1 . . . + 3 + 2 3 . . + + 3 2 . 2 . 2 + . 3 3 2 2 . . . . 3 + 3 2 . 1 . + 3 + 3 1 . . . . 3 2 3 1 + . . . 4 + r r + . r . + 5 2 + . + . 1 + 2 4 + . + . + 2 3 2 1 2 + . r 1 1 3 1 . . . . + + 4 + . . . . + . 2 . + + . + 3 . 3 1 2 . . . + + 2 1 + r . 1 3 1 + 2 1 + 1 . 3 1 + 3 . . 1 . 4 1 + 2 1 . + . 2 2 + 2 2 . 1 + 3 2 . 3 1 . 1 . 3 2 . 2 + . 1 . 100 83 60 60 57 45 38 36 . 1 . r . . . . . . . . . . . . . r . . . . . + . . . 2 . . . . + + . . . 1 . r . + . . + . . . . + . . . 1 . . 1 . . . 1 . . . r . 3 . + . 1 . . + . . . . . 2 . + + + . . . + . . + . + . . + + . . . r . . + 1 . . . + . . . . . . . . 1 . . . . . + . . r . . . r 1 . . . . . + . + . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . r . . . . . r . . . . . . . . . . . . . . . . . . . . . + . . . + . . . . . . . . + . r . 1 . . . . . . + . . . . . . . . + . . . . . . . . . . . . . . . . + . . . + . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . r + . + . + . r . . . . . . . . + . 1 . r + . . . . + . . . . + . . . . . . . + 1 . + . . . . + . . . . + + . + . . . . r . . . . + r . 1 . + . r . . . . . 33 31 29 19 19 19 12 12 12 12 10 10 10 . . . r . . . . . + . . . . . . . . + 1 . . . . . . . . . . . . 1 . . . . . . + . . . . . + . . + + . . . . . . . . . + . . 1 . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . + . . . . . + . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . r . . 10 . 7 . 7 . 7 . 7 . 7 + 7 . . . . . . . . . . . . r . . . + . . . . . + . . . . . . . . . + . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + . + . . . . . 1 . . . . . + 2 . . . + . + . . . + . r . 1 r . . . . . . . . . . . . . . . 1 . + . . . 1 . . . . . 1 . . . . . . + . . . . + + . . . + . . ++ r . . + r . . . . . . . . . . . . . . . 1 + 1 r 1 29 . . . . . . r . . . . 1 . . . . . . . . . . . . 2 2 2 2 2 2 . . . . . . . . . . . . . . . . . . . ++ . . . . . . 1 . . 21 7 . . . 1 1 3 3 2 . . . . . . . . . . . . . . . . . . . 2 . . 31 . 3 . + 19 14 14 10 . 1 . . + . . . . . . . . . . . . . . . . 2 . . . 3 . . . 3 . . . 4 . . . . . + . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . 2 . + . . . . . 2 . + . 2 . . . 3 . . + . . . + . . . . . . 1 . . . 1 . . . 1 . . . . . Vegetation strategy of Vellozia crinita 5 Table 2. (continued) In two relevés only: A: Byrsonima basiloba +:79, 80, Hyptis reticulata +:79, 2:81, Marcetia taxifolia 1:69, +:71. B: Campomanesia pubescens +:68, r:79, Chamaecrista rotundata +:16, r:81, Chamaecrista rotundifolia +:85, 86, Diplusodon buxifolius +:16, 1:03, Gomphrena virgata 1:68, +:72, Hyptis complicata 1:68, +:96, Hyptis fruticosa r:73, +:74, Peixotoa tomentosa +:16, 85. C: Baccharis calvescens +:79, 90, Calopogonium velutinum 1:80, +:81, Dyckia argentea +:79, 80, Erythroxylum tortuosum 1:68, r:96, Hyptis althaefolia +:4, 1:5, Lindsaea stricta 1:81, r:82, Peixotoa catarinensis +:79, 96, Rhynchospora consanguinea +:79, r:96, Stachytarpheta sellowiana r:80, +:83, Syngonanthus gracilis 1:87, +:98, Vellozia crassicaulis 2:96, +:97, Vernonia linearifolia +:78, 81, Xyris bahiana 1:78, 88, X. sp. r:7, +:03, X. tenella 1:88, +:97. D: Clusia arrudea r:75, 76. E: Cladonia miniata +:79, 1:80. In one relevé only: A: Byrsonima bumeliaefolia +:68, Chamaecrista cathartica +:76, Erythroxylum betulaceum 1:98, Hyptidendron asperrimum 1:16, Ilex amara r:80, Lippia lupulina +:79, Microlicia isophylla +:03, Stryphnodendron adstringens +:79, Tillandsia streptocarpa +:72, T. tenuifolia +:72, Mimosa polycephala var. taxifolia +:99, Trembleya laniflora +:98, Vitex polygama +:96. B: Chamaecrista hedysaroides +:75, Chionolaena arbuscula r:98, Chresta scapigera +:03, Declieuxia aspalathoides r:81, Gaylussacia montana +:16, Gomphrena scapigera r:98, Jacaranda paucifoliolata r:80, Lippia florida +:98, Macrosiphonia velame r:98, Microlicia euphorbioides +:71, Peixotoa catarinensis r:80, Polygala pseudoerica +:82, Spigelia heliotropoides +:80, Tocoyena formosa r:95. C: Anemia villosa r:16, Buchnera lavandulacea r:79, Chaptalia runcinata r:71, Commelina erecta +:88, Danthonia montana +:72, Diplusodon buxifolius +:16, Dyckia minarum r:71, Hippeastrum glaucescens +:98, H. morelianum r:83, Hypogynium virgatum +:1, Leiothrix prolifera 2:69, Lessingianthus simplex +:8, Paepalanthus elongatus 2:97, Pfaffia jubata +:96, Phyllanthus niruri r:82, Polypodium hirsutissimum +:72, Psidium incanescens +:80, Rhynchospora albiceps r:6, Skeptrostachys congestiflora r:82, Spermacoce capitata +:85, Trachypogon vestitus 1:72, Trimezia juncifolia r:80, Vitex polygama 2:97. D: Dasyphyllum brasiliense +:69, Gaylussacia riedelii +:79. ered part of the tall shrub layer in the tables). Eremanthus erythropappus is the most constant tree species, in this study with a mean height of 126 cm (8–307 cm), while in forests and among outcrops this species can reach 12 m (Souza et al. 2007). Eremanthus erythropappus has been considered a “pioneer” species, due to its capacity of colonizing forest clearings (Pedralli et al. 1997, 2000). In the studied localities, seedlings of E. erythropappus have been found almost in every investigated campo rupestre vegetation subtype. Other tree species with high constancies are Tibouchina heteromalla (9–180 cm, mean = 48 cm) and Hyptidendron asperrimum (18–296 cm). In the studied communities, Vellozia crinita dominates the herb layer, in which it has no co-dominant species. The herb layer with V. crinita consists of a series of regularly spaced mounds separated by sparse herbs and small subshrubs (Fig. 2). Unlike most erect species in its genus, the caespitose V. crinita does not depend directly on the presence of a rock outcrop. Individual plants have repent, horizontally growing, halfburied, profusely branched caudices and adventitious roots projected downwards at relatively regular intervals. They live very long and form rounded mounds in mildly sloping white sand deposits. Each mound may bear over 100 leaf rosettes/m2 . Given time the cushions accumulate sand more than the surrounding vegetation, and thus become slightly convex, but usually not taller than 10 cm. Small trees and shrubs grow sparsely among the mounds, as do tussock grasses or sedges and geophytes, with a few therophytes. Populations observed in the all ranges are identical in physiognomy. The moss layer is sparse and usually poorly developed. The influence of overgrazing by cattle is indicated by frequent occurrence of the invasive Melinis minutiflora, Spermacoce brachystemonoides and Ageratum fastigiatum (Kolbek & Alves 2008). The synthesis of vegetation with Vellozia crinita revealed three clusters (Fig. 3). Despite the considerable species richness, not many species are common to all three clusters and, if they are, their constancies tend to differ. Species in the shrub layer common to all of these are only Eremathus erythropappus, E. crotonoides, and Myrsine monticola (all very frequent throughout the southern campos rupestres). In the subshrub layer, this is also true for Cuphea ericoides and Tibouchina heteromalla and in the herb layer for Lagenocarpus rigidus subsp. rigidus, Loudetiopsis chrysothrix, and the invasive ruderalAgeratum fastigiatum, Melinis minutiflora and Spermacoce brachystemonoides. The three clusters (Table 2–4) include the characteristic Vellozia crinita, along with V. crassicaulis, Cuphea thymoides, Peixotoa tomentosa, Arthocereus melanurus, Bulbostylis paradoxa, Dipladenia polymorpha, Schizachyrium tenerum, and Xyris rupicola. Vellozio crinitae-Loudetiopsietum chrysothrici ass. nova hoc loco Nomenclatural type: Table 2, rel. 81 Characteristic species: Vellozia crinita, Loudetiopsis chrysotrix, Peixotoa tomentosa, Arthrocereus melanurus subsp. melanurus, Bulbostylis paradoxa, Dipladenia polymorpha, Schizachyrium tenerum, Xyris rupicola. A predominantly herbaceous terrestrial community on white quartzitic sand in which herbs cover a mean of 65% while the shrub layer is kept open by fires which occur once every 2–3 years, reaching a mean cover of 8%. The moss layer has a mean cover of 11%. The longevous, cushion-forming V. crinita dominates the herb layer in most of the sampled stands. Bulbostylis paradoxa is a pyrophyte widespread throughout Neotropical savannas on latossol, it is only occasionally found in campo rupestre on quartzitic sand, but in this community it is considered a diagnostic species. Loudetiopsis chrysothrix and Eremanthus erythropappus also pene- 6 R.J.V. Alves & J. Kolbek Table 3. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis var. Xyris bahiana and Syngonanthus gracilis. Relevé number 17 18 31 41 40 43 19 42 20 33 37 34 38 35 36 39 45 46 47 49 50 61 62 48 84 60 57 58 59 51 52 63 64 65 66 67 % High shrub layer (A) Eremanthus erythropappus . . Vernonia crotonoides . 2 Myrsine monticola . . Low shrub layer (B) Cuphea ericoides 2 . Tibouchina heteromalla + 2 Comolia sertularia + . Croton gnidiaceus . . Marcetia taxifolia + . Pavonia cf. garckeana . . Microlicia euphorbioides . . Lychnophora trichocarpha . . Herb layer (C) Rhynchospora tenuis 1 . Melinis minutiflora 1 1 Ageratum fastigiatum . . Xyris bahiana 1 + Loudetiopsis chrysothrix 2 1 Spermacoce + + brachystemonoides Syngonanthus gracilis . . Lagenocarpus *rigidus . . Leiothrix prolifera 2 + Senecio pohlii 1 . Rhynchospora recurvata . + Paepalanthus planifolius r 3 Xyris asperula + . Xyris rupicola . . Andropogon leucostachyus . . Syngonanthus niveus . . Rhynchospora globosa 1 . Xyris fallax . 1 Paepalanthus elongatus . . Xyris hymenachne . . Dicranopteris flexuosa . . Vellozia kolbekii . . Vernonia crotonoides r . Xyris tenella . + Sauvagesia rubiginosa . . Cuphea balsamona . . Schizachyrium tenerum 2 . Doryopteris ornithopus . . Polygala paniculata . . Vellozia crinita . . Actinocephalus bongardii + . Chamaesyce caecorum + . . . Commelina erecta Sisyrinchium vaginatum . . Chaetostoma cupressinum . . Utricularia simulans . . Aristida recurvata . . Schultesia gracilis . . Trimezia juncifolia . . Skeptrostachys congestiflora . . Woody seedlings in herb layer (D) Eremanthus erythropappus . . Stryphnodendron adstringens . . Moss layer (E) Campylopus savannarum 1 2 Sphagnum magellanicum . . . . + . . + . . . . 2 . . . . . . . . . . . . . . . . . . r 1 . . . 14 . . + . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 2 . . 11 2 . 1 + . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . 11 + 2 1 . . . . . + 2 . . . . . . + 1 1 . . + . . . 1 . . . . . . . . . . . . . . . . + . . . . . + r . . . . 2 . 2 1 2 1 . . . . 1 1 1 . . . . . + 2 1 + . . . 1 + + 1 1 + . . 2 + 2 1 2 . . . . . . r . . . . . 2 + . . . . + . 2 . . . . . . . 3 + . . . . + . 2 1 . . . . . . 2 + . . . . . . 2 + . . . 1 . . 1 . . . . 1 . . + . . . . . . . 2 + . . . . . . + + . . . . . . + . . . . . . . 1 + . . . r . . 2 + . . . + . . 1 . . . . . . . 2 . . . . . . . 1 + . 3 2 . . . 1 2 . 1 r . . . 1 . . 1 2 . . . 2 . . . . . . . 1 . . . . . . . 86 67 31 22 14 14 8 8 1 1 + . 2 . + 3 + . 1 . 1 3 1 . 4 1 + . . . 3 1 . . + . 2 . . 2 . 1 1 + . 2 1 + . 1 1 + . . 1 + + + 1 2 . . . . 1 1 . . . 1 1 . 1 + + + . + . . 1 . + . . + 3 . . . r 3 . + + . + 3 . 2 + . + + . 2 . 1 . 2 2 3 1 . + 4 1 2 + . r 3 1 2 + + . 3 + 2 . . . 2 . + 1 2 . . 1 2 2 1 + 2 + . 2 2 . 2 3 . + 2 . 2 2 . + 3 r 2 . . . 2 . . 1 1 . . 1 . 1 . 1 . 1 . . . . . . + . 2 . . . . . . . . . . . + . . . . . 1 . . . 64 61 58 53 50 47 . . . . 2 3 . . . . . . 2 2 . . . . . . . . . . . . . . . . . . . . . . . + . 3 . . . . . . . . 1 . . . . . 1 r . . . . . . . . . . . . . + . 1 . 1 . . . . . . + . . . r . + . . + . . 1 + r + . . . . . . . + . + + + . . . . . . + . . . + . . . + + . . 1 . + . . . . . . . . . . . . 2 . . . . . . . + 1 . . . + . . . . . . . . . . . . . . . . + . . . 2 . 2 . . . + 3 . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . + 3 2 + 2 . 1 . . . . . . . r . r . + + . . . . . + . . 1 . . . . . + 2 1 . 2 . + . 1 . . . . . . . . . . . 1 . . . . . . . . . . . . . . 1 3 r 2 . . . . 1 . . . . . . . . . 1 . + . . . . . . 1 . . . . . . . 3 . 1 . . . . . . . . . + + . . . . . . . . . . . . . . . . . . 2 3 3 + . . . . . . . . . . . . . 1 + . + 1 . 2 1 + . . . + . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + + . 1 + . . + . . . . . + . . + . . . 1 1 . + . . . . 2 . . . . . . . . . . . . . . . + + . + . . r . . + 2 + . . . . . . 1 . . . . . . . . . . . . . . . + . . . . . . . . . . 2 . r . . . . . . + + . . . . . . . . . . . . . . + + . . . . . . . + . . . . . . . . . 1 . . . + . . . . . . . 3 . . + 1 . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . r . 2 . . . . . . . . . . . . 1 . . . . . . . . . 3 . . . . . . . . 1 . . . . . . . . . 1 . . . . . . r . 1 . . . . + . . . . . . . . . 1 2 . . . . . 1 . . + . . . . . . . . . . . . + . . . + . . . . . . . 1 1 + r . . . 1 . + 1 . + + . . . . . . . . 1 + . . . + . . . . . . 2 + + . . . 1 + . . + . + + . . . 1 . . . . + 2 . . . . . . . . . r 1 . . . . . . . . 3 . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . 3 . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . 3 . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 + 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . 1 . . . . . . . . . . . . . . . . . . . . . . 44 36 31 31 28 25 25 25 25 25 22 17 17 17 17 17 14 14 14 14 11 11 11 11 8 8 8 8 8 8 8 8 8 8 1 2 2 . 2 . 1 . . . . . . . . + . 1 . 1 . . . . . . . . . . . . . . 2 1 + 1 . . . + . 2 2 1 2 . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 2 r 3 . . . . . . . . . + + r . . 1 . . . . . . . . r . . . . . . . + + . . . . 1 . 1 . . . . . . . . . . . . . . . . . . . . 11 . r + . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . 8 . . + . . . 1 1 1 2 2 1 . . 2 3 . . . . . . . 1 . . . . 3 3 3 3 3 47 . . . . 2 3 . . . . 2 2 3 . . . . . . . . . . . . . . . . . . . . . 14 In two relevés only: A: Myrcia mutabilis r:40, +:43, Eupatorium barbacense +:18, 43, Stryphnodendron adstringens +:18, 40, Tibouchina frigidula 2:20, +:43. B: Aristolochia gracilis +:40, 43, Baccharis platypoda +:40, 43, Campomanesia pubescens +:40, 43, Chamaecrista rotundifolia r:50, 61, Cuphea thymoides +:59, 61, Hyptis fruticosa +:38, 1:40, Palicourea rigida r:40, +:43, Peixotoa tomentosa +:43, 61, Smilax seringoides +:42, 43. Vegetation strategy of Vellozia crinita 7 Table 3. (continued) C: Ageratum conyzoides r:17, +:62, Alstroemeria cf. cunha 2:51, r:52, Aspilia foliacea +:17, 1:34, Bulbostylis paradoxa +:45, 60, Clusia arrudea r:36, +:43, Coccocypselum lanceolatum +:39, 43, Cyperus lanceolatus r:39, +:47, Emilia sonchifolia +:61, 62, Gomphrena agrestis r:17, +:43, Hypoxis decumbens +:61, 62, Inulopsis scaposa +:42, 62, Klotzschia brasiliensis r:38, 84, Lycopodiella camporum 2:19, +:42, Microlicia isophylla 2:34, +:60, Smilax seringoides +:40, r:41, Xyris sp. 2:57, 58. D: Eupatorium barbacense 2:36, 1:59; Miconia ligustroides r:36, +:39; Myrsine monticola +:40, 43. E: Lichenes indet 3:49, 50. In one relevé only: A: Byrsonima basiloba 1:20, Byrsonima variabilis 1:36, Cecropia lyratiloba +:40, Kielmeyera coriacea +:40, Pseudobombax longiflorum 1:40, Gomidesia eryocalyx +:43, Jacaranda paucifoliolata 1:40, Miconia ligustroides +:19, M. peppericarpa 1:39, M. theaezans +:19, Microlicia fasciculata +:18, Plenckia populnea +:40, Vellozia kolbekii 2:17. B: Lychnophora passerina +:60, Microlicia fulva r:20, Sauvagesia glandulosa +:17. C: Aechmea nudicaulis +:43, Alstroemeria sp. +:43, Andropogon selloanus +:59, Anemia villosa r:42, Arthrocereus melanurus +:43, Aylthonia tomentosa +:43, Ayapana amygdalina +:17, Baccharis trimera 1:20, Bulbophyllum ipanenensis +:43, Cuphea thymoides 1:31, Declieuxia cordigera r:40, Dipladenia polymorpha +:33, Erythroxylum tortuosum +:40, Guapira opposita r:40, Hippeastrum morelianum +:43, Hydrocotyle quinqueloba 1:42, Koellensteinia tricolor +:42, Lagenocarpus polyphyllus +:37, Lavoisiera bergii +:42, Lessingianthus simplex +:35, Lindsaea stricta 1:17, Lycopodiella alopecuroides +:42, Microlicia fasciculata 1:19, M. fulva +:42, Phlebodium pseudoaureum r:43, Phyllanthus niruri +:43, Plantago australis ssp. hirtella 2:20, Poaceae indet. 1:35, Psychotria sp. 1:59, Pteridium aquilinum r:41, Rhynchospora riedeliana 2:19, Sarcoglottis cogniauxiana +:58, Scleria bracteata +:61, Spermacoce verticillata r:41, Stachytarpheta sellowiana +:65, Trachypogon vestitus +:37, Vellozia crassicaulis +:57. D: Baccharis platypoda +:42, Cassia splendida 1:41, Macairea radula 1:42, Miconia theaezans r:43, Tibouchina frigidula +:20. E: Campylopus pilifer 1:84. trate into another association of this alliance (in prep.). Arthrocereus melanurus subsp. melanurus occurs as a characteristic species, but is usually restricted to spots with an outcrop or shallower soil. This community was detected in the Serra São José, Lenheiro, Ouro Grosso (Itutinga), Ouro Branco and Carrancas Ranges in altitudes 950–1536 m alt. on slopes with inclinations of 0–60◦ and various expositions. Within the association, two subassociations were distinguished (see Fig. 3 A–B, Tabs 2, 3): Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis subass. nova hoc loco Nomenclatural type: identical with the type of the association name. Differential species: Comolia sertularia, Rhynchospora globosa, R. recurvata, R. tenuis, Syngonanthus niveus. The subassociation represents the terrestrial, seasonally wet stands with higher dominance of three species of the genus Rhynchospora and of Syngonanthus niveus. Average cover of shrub layer per relevé is 9%, herb layer 65% and moss layer 13%. The number of species varies from 4 to 43, averagely 16, mainly depending on the sample plot size. Stands of this terrestrial community are situated on white quartzitic sand with frequent milky quartz gravel in Serra do Ouro Grosso (Itutinga), Serra do Ouro Branco, Serra de São José, and Serra do Lenheiro. The environmental characteristics are comparable to those of the described association but with seasonally humid, shallower (ca. 30 cm deep) soils. Within the subassociation two variants were distinguished: a) var. typicum (Tab. 2, Fig. 3A), with high dominace and constancy of Vellozia crinita and the highest encountered constancy of Arthrocereus melanurus subsp. melanurus (absent from Serra do Ouro Branco)when compared to other investigated stands of the association. Stands are situated in Serra de São José, Serra do Lenheiro, Serra do Ouro Grosso (Itutinga), Serra do Ouro Branco, in altitudes of 1040–1420m alt. on slopes with inclinations of 0–30◦, with apparent preference for eastward exposition. Average cover of shrub layer per relevé is 8% (0–35%), herb layer 62% (20–95%) and moss layer 11% (0–70%). Number of species varies from 5 to 33, averagely 16. b) var. Xyris bahiana and Syngonanthus gracilis, with Croton gnidiaceus, Leiothrix prolifera, Paepalanthus planifolius, and Senecio pohlii (Tab. 3, Fig. 3B). Stands were investigated in the Serra de São José and Serra do Lenheiro in altitudes of 950–1320m alt. on slopes with inclinations of 0–60◦, with predominantly westward and southward expositions. Average cover of shrub layer per relevé is 11% (0–65%), herb layer 69% (40–95%) and moss layer 16% (0–50%). Number of species varies from 4 to 43, averagely 16. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum consanguineae subass. nova hoc loco Nomenclatural type: Tab. 4, rel. 92 Differential species: Jacaranda paucifoliolata, Clittoria guianensis, Dyckia minarum, Hyptis althaefolia, Rhynchospora consanguinea, Schizachyrium tenerum. The subassociation (Fig. 3C) represents the stands with higher dominance of Rhynchospora consanguinea. This terrestrial community on white quartzitic sand has an average cover of shrub layer per relevé is 5% (0–10%), herb layer 60% (9–95%) and moss layer 2% (0–7%). Number of species varies from 9 to 16, averagely 11. Stands are situated in Serra do Lenheiro, Carrancas, Serra de São José in altitudes of 1175–1394m alt. on slopes with milder inclinations (0–20◦) and deeper soils (>60 cm) than the previous syntaxa. Plots are mainly 8 R.J.V. Alves & J. Kolbek Table 4. Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum consanguineae. Relevé number High shrub layer (A) Eremanthus erythropappus Myrsine monticola Byrsonima basiloba Vernonia crotonoides Jacaranda paucifoliolata Evolvulus helichrysoides Lippia lupulina Vochysia thryrsoidea Low shrub layer (B) Cuphea thymoides Clittoria guianensis Peixotoa tomentosa Tibouchina heteromalla Gomphrena virgata Cuphea ericoides Diplusodon buxifolius Croton antisyphiliticus Anacardium humile Chamaechrista desvauxii Jacaranda decurrens Aristolochia gracilis Herb layer (C) Loudetiopsis chrysothrix Vellozia crinita Melinis minutiflora Rhynchospora consanguinea Hyptis althaefolia Schizachyrium tenerum Dyckia minarum Spermacoce brachystemonoides Vellozia crassicaulis Chaetostoma cupressinum Lagenocarpus *rigidus Senna cf. organensis Dipladenia polymorpha Xyris rupicola Arthrocereus melanurus Myrsine monticola Bulbostylis paradoxa Triumfetta rhomboidea Vernonia linearifolia Ageratum fastigiatum Moss layer (D) Campylopus savannarum 11 12 13 14 15 01 02 92 94 93 53 54 55 56 % . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . 1 . . . . . . . . . . . r + + . . . . . . + + . 1 . . . . . . . + . . . . . . . + . . . . . . . r r r + + . . 2 . r . 1 r . . . . + + . r . . . . r r . r . . + 43 29 21 14 36 14 14 14 . . . . . . . . . . . . . . . . r . . . . . . . . . . . r + . . . . . . . . . . . . 1 . . . . . . . . . . + 1 . . . . . . . . . . . . + + r . . . . . . . . . 1 r + . . . . . 2 . . . . . . + . . . . 1 . . . . . . 1 . . . . 1 . . . . . . . . 1 + r . . . . . . . . + 1 + + . . . . . . . . + 1 r r . . . . . . . . . + + r . . . . . . . . . 29 29 29 21 14 14 14 14 14 14 14 14 4 1 1 . 1 1 r . . . . . . . + 1 . . . . 4 1 1 . 2 1 + . 1 . . . . . . . r . . . 1 1 r . 2 1 1 + 1 . . . . . 1 + 1 . . 2 . . . 1 1 r . r . . . . . . . . . . . 2 1 . . 1 1 . . . . . . . . . . . 1 . . 1 1 . . . . . . . . . . . . . . . . + . + + . . . . . . . . . . . . . . . . + . 2 2 + + . . . + . 3 2 + + + . . . . . . 1 2 1 + . . . + . 2 2 1 + + . . . . . . 2 2 1 + . . . + . 2 2 + + + . . . . . . 4 . + 1 . . . . . . . . . . . + . . . + 4 . + 1 . . . . . . . . . . . . . . . r 4 . + 1 . . . . . . . . . . . . . . . . 5 . . 1 . . . . . . . . . . . . . . . . 100 64 64 50 36 36 29 29 21 21 21 21 21 21 14 14 14 14 14 14 + + + + + . . . . . . . . . 36 In one relevé only: A: Annona crassiflora r:01. B: Campomanesia pubescens r:02, Chresta scapigera +:92, Galactia martii r:01, Jacaranda paucifoliolata 1:53, Stachytarpheta sellowiana +:01. C: Leptocoryphium lanatum r:14, Oxalis hirsutissima +:01, Phyllanthus niruri +:15. oriented toward the East. The species assemblages in table 4 suggest that subdivisions into further vegetation units can be made once sufficient additional relevés are obtained. Discussion Homogenous stands in the studied environment may occupy areas ranging from just a few to several hundred square meters. When the floristic composition and structure of sample plots of similar sizes are compared, the ass. Vellozio-Loudetiopsietum turns out to be relatively homogenous. In the intricate mosaic of campo rupestre plant communities, when smaller stands are registered, the variability described above is the inevitable consequence of species poverty. Despite this fact, the stands are easily distinguishable and separated from surrounding communities (on shallow soil or on outcrops) by sharp borders. The Velloziaceae are among the plants best adapted to campo rupestre through drought-resistance structures, mechanisms which allow for quick uptake of water and mist, and by synchronizing flower production with rainfall (Owoseye & Sanford 1972; Gaff 1987; Alves & Kolbek 1994; Porembski & Barthlott 1995). Several Vellozia species are also notorious for their longevity and resistance to fire (Alves 1994; Kubitzki 1998). Other psammophilous, caespitose and probably mound-forming species of the genus Vellozia are known to occur, especially in localities along the Espinhaço Vegetation strategy of Vellozia crinita chain situated to the north of the study area (MelloSilva 1995). Within sandy grasslands of campo rupestre, these species may well have an analogical function to that of V. crinita, but no studies of their vegetation strategy have been published to date. The laboratory study of germination properties of two erect, outcrop-dwelling and three caespitose, probably psammophilous Vellozia species (Garcia & Diniz 2003; Garcia et al. 2007) revealed that all have very small seeds which germinate best when exposed to light, but require higher temperatures (35–40 ◦C) to germinate in darkness. At 15 ◦C, all species exhibited deficient germination or none at all. Though the influence of fire was not considered by Garcia & Diniz (2003) and Garcia et al. (2007), the described germination properties are consistent with species in which seed-dormancy is broken by sudden increases in temperature by passing fires. A fire would be required to produce the optimum germination temperature for seeds which are insulated by a layer of sandy soil, which is known to be an efficient thermal insulator. Furthermore, germination is only one of the processes necessary for the successful establishment of the plants. Small seeds have very few reserves, which mean that soon after they germinate, the subsequent development of the plants becomes highly dependent on photosynthesis. Effective germination in both species of Vellozia investigated by Garcia et al. (2007) proved to depend on light. It is thus not probable that adult plants of Vellozia spp. could develop from seeds germinated under the soil surface, unless they become exposed by erosion. Two other species of Vellozia (present in relevés), with distinct vegetation strategies, occur sympatrically in the Serra de São José, which is the type locality of V. crinita. Vellozia kolbekii is an erect, to 2 m tall, sparingly branched shrubby species which grows exclusively among exposed outcrops and narrow fissures. An intermediate strategy is exhibited by V. crassicaulis: when young, this species forms prostrate mounds on shallower sandy and stony soils (<10 cm deep) or in outcrop fissures, but older individuals, in localities not affected by fire, can have erect branches up to 50 cm tall. Vellozia crinita and V. kolbekii have distinct habitat preferences: they do not grow together in the same community. Of the 168 sampled vascular species, 109 grow exclusively in stands with V. crinita, 11 grow together with V. kolbekii but not the former, and only 26 grow with both. The species of intermediate strategy, V. crassicaulis, is present in only six relevés, in five of which (Tab. 2: 96, 97; Tab. 3: 57 and Tab. 4: 12, 13) it co-occurs with V. crinita, indicating the presence of spots with shallow soil. In the sixth relevé with V. crassicaulis, the other two species are absent. We can conclude that for the stability of the studied vegetation, Vellozia crinita is the most important species. The caespitose habit and the repent, profusely branched caudices are convenient adaptations to strong wind, while the usually subterranean position of the caudices is an effective defense against fire. Though individuals are longevous and slow-growing, forming 9 mounds with densely packed rosettes allows V. crinita to cover larger areas, thus occupying a distinct niche not disputed by erect shrubby species of Vellozia. However, V. crinita is unable to compete successfully against shading by invasive herbaceous species, which grow profusely in areas affected by cattle. Juvenile individuals have very rarely been observed in all sites surveyed herein (see also Alves 1994). If campo rupestre dominated by Vellozia spp. should be preserved, further investigation of the environmental conditions necessary for the germination and subsequent growth of juvenile Vellozia populations is urgently needed. The classification of V. crinita and other species as vulnerable (viz. BIODIVERSITAS 2007) may be an understatement of the dangers truly faced by Vellozia species due to the use of campos rupestres as grazing areas for livestock Hence not only the species, but also the studied plant communities and associated biota are in peril. Acknowledgements Our paper is dedicated to the late Prof. Johann Becker, an outstanding Brazilian naturalist who took part of the field studies. This survey was only possible thanks to the contributions of more than eighty specialists in many fields of expertise, especially taxonomists, and to the support by people and institutions, the totality of which cannot fully be acknowledged. We thank Ivana Rajznoverová for technical assistance. 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Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis var. typicum 1 9 8 10 6 44 2 3 4 5 7 95 69 03 98 16 78 88 81 82 68 70 83 77 91 99 90 00 85 28 29 29 29 29 12 28 28 28 28 29 1 12 27 31 2 3 22 6 6 12 12 7 20 18 26 18 26 18 10 10 10 10 10 9 10 10 10 10 10 11 9 10 10 10 10 10 10 10 9 9 10 9 10 10 10 10 10 2004 2004 2004 2004 2004 1989 2004 2004 2004 2004 2004 1989 1989 2005 2004 1997 1989 1989 1989 1989 1989 1989 1989 1989 1989 2005 1989 2005 1989 SSJ SLN SLN SLN SSJ SSJ SSJ SSJ SSJ SSJ SLN SSJ SSJ SOG SOB SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ 1178 1178 1178 1178 1178 1150 1178 1178 1178 1178 1178 1230 1150 1175 1536 1400 1300 1270 1410 1410 1140 1150 1230 1040 1300 1175 1300 1175 1200 E E E E E W E E E E E S SW W ENE – SE SSW SSE SSW ESE W E W WSW N WNW E W 5 5 5 5 5 5 5 5 5 5 5 10 5 10 10 0 20 15 20 15 10 5 10 5 10 20 10 5 10 4 4 4 4 4 6 4 4 4 4 4 18 9 32 60 10 50 25 60 50 150 160 25 16 15 10 15 64 50 5 0 3 1 5 0 1 0 0 0 0 <5 5 4 4 4 <5 15 15 <5 10 <5 <5 0 0 10 10 5 <5 25 30 50 30 30 40 20 50 30 40 30 25 60 80 90 60 60 75 60 60 65 70 70 90 90 75 85 80 95 4 0 1 1 0 40 1 2 1 1 0 0 45 4 0 4 <5 30 40 50 30 70 <5 0 0 4 0 1 0 0 0 0 0 0 0 0 0 0 0 0 10 10 0 0 0 5 0 5 10 20 10 10 10 0 4 0 20 0 Vegetation strategy of Vellozia crinita 86 87 79 96 80 89 97 71 73 74 75 76 72 18 18 3 21 5 18 5 15 18 18 18 18 15 10 10 10 9 10 10 10 9 9 9 9 9 9 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ 11 1200 1200 1375 1040 1420 1300 1375 1200 1200 1200 1200 1200 1220 – NW NNW NE N WNW N E E E E E E 0 10 20 15 20 15 30 8 10 10 10 10 10 50 50 80 100 30 15 30 200 100 100 100 100 400 <5 <5 20 <5 <5 0 0 35 20 15 35 20 25 95 80 80 75 30 75 50 70 70 80 75 80 70 0 0 10 0 15 10 40 10 0 <5 10 <5 5 0 0 5 10 5 0 10 10 10 10 10 10 10 Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum tenuis var. Xyris bahiana, Syngonanthus gracilis 17 18 31 41 40 43 19 42 20 33 37 34 38 35 36 39 45 46 47 49 50 61 62 48 84 60 57 58 59 51 52 63 64 65 66 67 2 20 1 13 23 22 23 14 23 1 23 1 23 2 22 19 23 23 23 23 23 23 24 23 3 23 20 20 20 7 7 20 20 20 20 20 10 10 11 9 9 9 10 9 10 10 9 10 9 10 9 10 9 9 9 9 9 9 9 9 7 9 9 9 9 11 11 10 10 10 10 10 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 SSJ SSJ SLN SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ SSJ 1320 1370 1270 1000 1000 1020 1120 1000 1100 1320 1220 1310 1030 1330 1030 1300 1220 1220 1230 1230 1230 1240 1240 1230 1230 1240 1030 1040 1040 950 950 1200 1200 1200 1230 1230 W SE S S WNW WNW N SE – NE WNW NE SSW NW SW NW W W SW W W SW SW SW – S W W SW – – N NW NW – – 15 35 45 35 35 30 10 20 0 15 10 20 10 25 30 60 10 5 25 10 5 15 15 20 0 20 15 15 10 0 0 10 10 10 0 0 40 12 18 25 150 100 20 10 30 36 105 48 120 60 60 1.5 40 24 30 100 48 100 36 35 500 32 32 105 160 12 12 6 12 6 4 4 10 25 20 10 10 5 4 0 50 10 20 5 35 10 30 4 4 0 4 4 0 4 0 0 <5 4 0 4 4 0 5 65 10 30 0 0 50 70 95 85 70 60 95 85 70 55 60 60 75 40 80 50 75 80 70 70 85 90 95 45 90 60 70 55 75 80 75 50 50 60 65 60 4 10 0 10 4 0 10 40 0 4 4 4 40 20 50 0 4 10 50 30 40 20 10 0 5 0 4 10 0 0 0 50 50 30 30 30 80 80 70 0 0 0 90 0 0 0 0 0 0 50 0 50 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 4 4 4 4 32 15 36 36 36 25 25 25 25 0 0 0 10 5 4 4 10 5 5 10 5 4 5 70 60 30 30 40 9 9 95 80 80 80 85 80 90 5 5 7 2 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 10 1 1 1 1 Vellozio crinitae-Loudetiopsietum chrysothrici rhynchosporetosum consanguineae 11 12 13 14 15 01 02 92 94 93 53 54 55 56 29 29 29 29 29 27 27 30 30 30 01 01 01 01 10 10 10 10 10 10 10 10 10 10 11 11 11 11 2004 2004 2004 2004 2004 2005 2005 1989 1989 1989 1989 1989 1989 1989 SLN SLN SLN SLN SLN CCS CCS SSJ SSJ SSJ SLN SLN SLN SLN 1394 1394 1394 1394 1394 1175 1175 1110 1110 1110 1190 1190 1190 1190 E E E E E NE W – – – ESE E E ESE 20 20 20 20 20 10 10 0 0 0 10 10 20 20