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
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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
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29
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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
. .
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2 . 1 + . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . 11
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44
36
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31
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25
25
25
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17
17
17
17
17
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14
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8
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2
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3
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. . + + . . . . 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
%
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43
29
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100
64
64
50
36
36
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14
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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. The participation of the second author was
possible due to research project number AVOZ 60050516.
References
Alves R.J.V. 1994. Morphological age determination and longevity in some Vellozia populations in Brazil. Folia Geobot.
Phytotax. 29: 55–59.
Alves R.J.V. & Kolbek J. 1994. Plant species endemism in savana
vegetation on table mountains (Campo Rupestre) in Brazil.
Vegetatio 113: 125–139.
Benites V. de M., Caiafa A.N., Mendonça E. de S., Schaefer C.E.
& Ker J.C. 2003. Solos e vegetação nos complexos rupestres
de altitude da Mantiqueira e do Espinhaço. Floresta e Ambiente 10(1): 76–85.
BIODIVERSITAS 2007. Revisão das listas das espécies da
flora e da fauna ameaçadas de extinção do Estado de
Minas Gerais – Relatório final, vol. 2 (RESULTADOS:
Lista Vermelha da Flora de Minas Gerais), Belo Horizonte; http://www.biodiversitas.org.br/listas-mg/Relatorio
Listasmg Vol2.pdf (PDF accessed on April 14, 2008).
Braun-Blanquet J. 1932. Plant Sociology. Koeltz Scientific Books,
Koenigstein, 439 pp.
Brummitt R.K. & Powell C.E. 1992. Authors of plant names.
Royal Bot. Gardens, Kew, 732 pp.
Clayton W.D., Harman K.T. & Williamson H. 2007. GrassBase –
The Online World Grass Flora. www.kew.org/data/grassesdb.html (Accessed 19 January 2008).
Conceição A.A. & Pirani J.R. 2005. Delimitação de habitats em
campos rupestres na Chapada Diamantina: substratos, composição florística e aspectos estruturais. Bol. Bot. Univ. São
Paulo 23: 85–111.
Gaff D.F. 1987. Desiccation tolerant plants in South America.
Oecologia 74: 133–136.
Garcia Q. de S. & Diniz I.S.S. 2003. Comportamento germinativo
de tr˛es espécies de Vellozia da Serra do Cipó, MG. Acta Bot.
Brasil. 17(4): 487–494.
10
R.J.V. Alves & J. Kolbek
Garcia Q. de S., Jacobi C.M. & Ribeiro B. de A. 2007. Resposta
germinativa de duas espécies de Vellozia (Velloziaceae) dos
campos rupestres de Minas Gerais, Brasil. Acta Bot. Brasil.
21(2): 451–456.
GBIF DATA PORTAL. www.gbif.net (Accessed from 4 Oct. 2007
to 20 Jan. 2008).
Giulietti A.M. & Pirani J.R. 1988. Patterns of geographic distribution of some plant species from the Espinhaço Range, Minas Gerais and Bahia, Brazil, pp. 39–69. In: Vanzolini P.E. &
Heyer W. R. (eds), Proceedings of a workshop on neotropical
distribution patterns, Academia Brasileira de Ci˛encias, Rio
de Janeiro.
Giulietti A.M., Menezes N.L., Pirani J.R., Meguro M. & Wanderley M.G. L. 1987. Flora da Serra do Cipó, Minas Gerais:
caracterização e lista das espécies. Bol. Bot. Univ. São Paulo
9: 1–151.
Giulietti A.M., Pirani J.R. & Harley R.M. 1997. Espinhaço Range
Region, Eastern Brazil, pp. 397–404. In: Davis S.D., Heywood
V.H., Herrera-MacBryde O., Villa-Lobos J. & Hamilton A.C.
(eds), Centres of plant diversity – A Guide and Strategy for
their Conservation. World Wildlife Fund & World Conservation Union.
JBRJ. Instituto de Pesquisas Jardim Botânico do Rio de
Janeiro. Jabot – Banco de Dados da Flora Brasileira.
http://www.jbrj.gov.br/jabot (Accessed throughout 2007).
Kolbek J. & Alves R. J. V. 2008. Impacts of cattle, fire and wind
in Rocky Savannas, Southeastern Brazil. Acta Univ. Carol.
Environ. 22:111–130.
Kubitzki K. 1998. Velloziaceae, pp. 459–467. In: Kubitzki K.
(ed.), The families and genera of flowering plants. Vol.
III: Flowering Plants, Lilianae (except Orchidaceae), Berlin,
Springer.
Mello-Silva R. de 1995. Velloziaceae, pp. 772–781. In: Stannard
B.L., Harvey Y.B. & Harley R.M. (eds), Flora of the Pico
das Almas, Chapada Diamantina – Bahia, Brazil. Royal Bot.
Gardens, Kew, London.
Owoseye J.A. & Sanford W.W. 1972. Ecological study of Vellozia
schnitzleinia, a drought-enduring plant of Northern Nigeria.
J. Ecol. 60: 807–817.
Pedralli G., Teixeira M.C.B. & Nunes Y.R. 1997. Estudos
sinecológicos sobre a candeia (Vanillosmopsis erythropappa
Schult. Bip.) na estação ecológica do Tripui, Ouro Preto, MG.
Revista Árvore 21(2): 301–306.
Pedralli G., Teixeira M. do C.B., Freitas V.L. de O., Meyer S.T.
& Nunes Y.R.F. 2000. Florística e fitossociologia da Estação
Ecológica Do Tripuí, Ouro Preto, MG. Ci˛enc. Agrotec. 24:
103–136. (Special Ed.).
Podani J. 2001. SYN-TAX 2000. Computer program for data
analysis in ecology and systematics for Windows 95, 98 &
NT. User’s manual. Scientia Publishing, Budapest, pp. 1–53.
Porembski S. & Barthlott W. 1995. On the occurrence of a velamen radicum in Cyperaceae and Velloziaceae. Nordic J. Bot.
15: 625–629.
Souza A.L. de, Oliveira M.L.R. de, Silva E.F. da & Silva Coelho,
D.J. da 2007. Caracterização fitossociológica em áreas de
ocorr˛encia natural de candeia (Eremanthus erythropappus
(D.C.) MacLeish). Rev. Árvore31(4): 667–677.
Tichý L. 2002. JUICE, software for vegetation classification. J.
Veg. Sci. 13: 451–453.
USDA 2007. USDA ARS, National Genetic Resources Program.
Germplasm Resources Information Network – (GRIN). National Germplasm Resources Laboratory, Beltsville, Maryland. www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?443910
(Accessed from 4 Oct. 2007 to 20 Jan. 2008).
W3 TROPICOS 2007. Missouri Botanical Garden’s VAST (VAScular Tropicos) nomenclatural database. mobot.mobot.org/
W3T/Search/vast.html (Accessed throughout 2007).
Received May 10, 2008
Accepted June 8, 2009
Appendix. Relevé data follow this order: relevé number in bold, date (day, month, year), locality (CCS = Serra de Carrancas, SBP
= Serra Bico de Pedra, SCH = Serra da Chapada, SLN = Serra do Lenheiro, SOB = Serra do Ouro Branco, SOG = Serra do Ouro
Grosso, SSJ = Serra de São José), altitude (m a.s.l.), exposition, inclination (in ◦ ), area (m2 ), cover of woody layers (%), cover of herb
layer (%), cover of moss layer (%), cover of stones (%).
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