oliveira filho 1991

26
EDINB. J. BOT. 48(3): 307-332 (1991) A COMPARATIVE STUDY OF FIVE CERRADO AREAS IN SOUTHERN MATO GROSSO, BRAZIL A. T. OLIVEIRA-FILHO* & F. R. MARTINS** Quantitative descriptions of the woody flora and soil analyses are given for five areas of cerrado (savanna woodland) near Cuiaba, Mato Grosso state, Brazil. Comparisons are made of the floristic composition and community structure of these areas using polar ordination and hierarchical classification techniques. Descricoes quantitativas da flora lenhosa e analises de solos sao apresentadas para cinco areas de cerrado nas proximidades de Cuiaba, Mato Grosso. Sao realizadas comparacoes entre estas areas em composicao floristica e estrutura comunitaria utilizando tecnicas de ordenacao polar e classificacao hierarquica. Cerrado is a savanna-like vegetation which covers about 2 million km , representing about one-fifth of Brazil's total area. In the state of Mato Grosso, it occupies about 28.5% (251085km 2 ) of the total area and abuts on the Amazonian forest (Hylaea) to the north and the Pantanal Matogrossense (the low-lying floodplain of the Rio Paraguai) to the south (Fig. 1). It is currently being destroyed very rapidly: in 1980, 12% or 30069km of the Mato Grosso cerrado was already completely cleared for cattle-pasture and crops, particularly soya, and since then this area has probably more than doubled (Cameiro, 1982). The floristic knowledge of this vanishing vegetation is very incomplete. Few surveys of Mato Grosso cerrados are available in the literature. A number of taxonomists who participated in the first scientific expeditions to Mato Grosso (from 1790 to 1915) left some general descriptions and checklists for the regional flora (see Hoehne, 1923 for a review). Pioneer detailed studies of particular areas of cerrado were carried out by Ratter (1971), Ratter et al. (1973, 1977) and Furley et al. (1988) in northeastern Mato Grosso. Oliveira-Filho & Martins (1986) and Oliveira-Filho et al. (1989) provided floristic and phytosociological surveys of cerrados of the region of Cuiaba, the capital of the state. For the seasonally flooded cerrados of the Pantanal Matogrossense, Prance & Schaller (1982) gave a detailed floristic survey while Pott et al. (1986) contributed a detailed checklist and Ratter et al. (1988) a quantitative-floristic study of an important area. The present communication is devoted to a quantitative study of the woody flora of five cerrado areas, on different soils, in the region of Cuiaba and the Chapada dos Guimaraes, in central-southern Mato Grosso. Comparisons are made to evaluate dif- * Departamento de Ciecias Florestais, Escola Superior de Agricultura de Lavras, 37200 Lavras, MG, Brazil ** Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas, 13081 Campinas, SP, Brazil. 307

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A COMPARATIVE STUDY OF FIVE CERRADO AREAS IN SOUTHERN MATO GROSSO, BRAZIL A. T. OLIVEIRA-FILHO* & F. R. MARTINS**

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  • EDINB. J. BOT. 48(3): 307-332 (1991)

    A COMPARATIVE STUDY OF FIVE CERRADO AREAS INSOUTHERN MATO GROSSO, BRAZIL

    A. T. OLIVEIRA-FILHO* & F. R. MARTINS**

    Quantitative descriptions of the woody flora and soil analyses are given for five areas ofcerrado (savanna woodland) near Cuiaba, Mato Grosso state, Brazil. Comparisons are madeof the floristic composition and community structure of these areas using polar ordination andhierarchical classification techniques.

    Descricoes quantitativas da flora lenhosa e analises de solos sao apresentadas para cinco areasde cerrado nas proximidades de Cuiaba, Mato Grosso. Sao realizadas comparacoes entre estasareas em composicao floristica e estrutura comunitaria utilizando tecnicas de ordenacao polare classificacao hierarquica.

    Cerrado is a savanna-like vegetation which covers about 2 million km , representingabout one-fifth of Brazil's total area. In the state of Mato Grosso, it occupies about 28.5%(251085km2) of the total area and abuts on the Amazonian forest (Hylaea) to the northand the Pantanal Matogrossense (the low-lying floodplain of the Rio Paraguai) to thesouth (Fig. 1). It is currently being destroyed very rapidly: in 1980, 12% or 30069kmof the Mato Grosso cerrado was already completely cleared for cattle-pasture and crops,particularly soya, and since then this area has probably more than doubled (Cameiro,1982). The floristic knowledge of this vanishing vegetation is very incomplete. Fewsurveys of Mato Grosso cerrados are available in the literature. A number oftaxonomists who participated in the first scientific expeditions to Mato Grosso (from1790 to 1915) left some general descriptions and checklists for the regional flora (seeHoehne, 1923 for a review). Pioneer detailed studies of particular areas of cerrado werecarried out by Ratter (1971), Ratter et al. (1973, 1977) and Furley et al. (1988) innortheastern Mato Grosso. Oliveira-Filho & Martins (1986) and Oliveira-Filho et al.(1989) provided floristic and phytosociological surveys of cerrados of the region ofCuiaba, the capital of the state. For the seasonally flooded cerrados of the PantanalMatogrossense, Prance & Schaller (1982) gave a detailed floristic survey while Pott etal. (1986) contributed a detailed checklist and Ratter et al. (1988) a quantitative-floristicstudy of an important area.

    The present communication is devoted to a quantitative study of the woody flora offive cerrado areas, on different soils, in the region of Cuiaba and the Chapada dosGuimaraes, in central-southern Mato Grosso. Comparisons are made to evaluate dif-

    * Departamento de Ciecias Florestais, Escola Superior de Agricultura de Lavras, 37200Lavras, MG, Brazil

    ** Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas,13081 Campinas, SP, Brazil.

    307

  • 308 EDINB. J. BOT. 48(3)

    ".^;i;.

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 309

    ferences in floristic composition and community structure of these areas related tovariations in soil properties.

    DESCRIPTION OF THE STUDY AREAS

    The region of Cuiaba and the Chapada dos Guimaraes comprises a wide low plain, theBaixada Cuiabana, and a higher plateau, the Chapada dos Guimaraes (Fig. 1). TheBaixada Cuiabana is a flattish surface with altitudes ranging from c.lOOm, in thesouthern border with the Pantanal Matogrossense, to c.350m, at the base of the Chapadados Guimaraes. This low plateau is the western tip of a large area of tablelands: thePlanalto Central. The Chapada dos Guimaraes is bounded by vertical escarpments of5O-3OOm and the relief at the top is relatively flat with altitudes ranging from 500 to800m. This high plateau is composed of two layers of Devonian sandstone which overliethe Pre-Cambrian phyllite of the Baixada Cuiabana.

    The climate is tropical and continental with high temperatures all year round and arainy summer and dry winter, being classified as Koppen's 'Savanna Subtype' (Aw) ofthe tropical rain climate (A) (Eidt, 1968). Detailed climatic data are available only forCuiaba where the mean annual air temperature is 25.6C, with monthly means rangingfrom 22.8C (July) to 27.2C (October), and the mean annual rainfall of about 1420mmis concentrated during October-May (95% of the total rainfall) (Bombled, s.d.). Thetemperatures of the Chapada dos Guimaraes, however, are 3^C lower than Cuiaba andthe annual rainfall can reach 2000mm (EDIBAP, 1979).

    The main vegetation type of the region is cerrado with its characteristic wide physiog-nomic variations. Forests are mostly restricted to galleries along the watercourses andto some patches at the top of the high plateau. Grasslands can be found in rocky and/orin poorly-drained sites. The vegetation is relatively well-preserved considering that thecolonization of the region by Europeans goes back to the beginning of the 18th century:the very infertile soils have restricted the spread of cultivated areas throughout the regionand human damage is caused principally by repeated fires and more localized goldmining and urban growth. In 1989 a part of the Chapada dos Guimaraes was created aNational Park.

    The five areas of cerrado studied lie in three sites along the road (the MT 305) whichconnects Cuiaba to the Chapada dos Guimaraes: Fazenda Sao Pedro-Bandeira (km 14)*,the Rio Claro region (km 42) and the Salgadeira region (km 45) (see Fig. 1).

    Site A: Fazenda Sao Pedro-Bandeira (1530'S, 5602'W, c.l90m alt.) This is a ranchwhere the cerrado is used as natural pasture for cattle at low density. Two areas werestudied here:

    Area 1 - Murundu cerrado: This type of cerrado occurs on round earthmounds (0.2-20.0m diameter, 0.1-2.0m high and usually bearing termitaria) scattered in a regular

    * The kilometre numbers refer to the distance from Cuiaba. They are marked by roadsigns.

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  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 311

    Universidade Estadual de Campinas (UEC) with duplicates at the Escola Superior deAgricultura de Lavras (ESAL) and the Royal Botanic Garden, Edinburgh (E). A list ofall species recorded is given in the Appendix.

    In each of the five study areas, a number of 0.5 litre samples of soil were collectedfrom the depth of 0-20cm and mixed up to form a compound sample. Chemical andgranulometric analyses were made at the Soil Laboratory of the Universidade Federalde Mato Grosso.

    Woody plants were sampled using the point-centred quarter method (PCQ), asdescribed by Mueller-Dombois & Ellenberg (1974), in three areas of cerrado: the pebblecerrado (Area 2) with 140 points distributed in a 4 x 35 systematic grid, the phyllitecerrado (Area 3) with 60 points in a 5 x 12 grid, and the sandstone cerrado (Area 5) with50 points in two 5 x 5 grids. Inter-point distances were of 10m. The interfluvial cerrado(Area 4) was sampled by 100 contiguous plots of 10 x 10m, arranged in a transect of 50paired plots. The murundu cerrado (Area 1) was sampled on 80 earthmounds (0.5-22mdiameter), comprising in total 0.3ha of murundus. In all surveys, species, height, andcircumference at the stem base were recorded for all shrubs, trees and palms with a stemcircumference of at least 9cm. The number of individuals, relative density, total basalarea, relative dominance (based on basal area), relative frequency, importance valueindex (IVI = rel.dens. + rel.dom. + rel.freq.), and cover value index (CVI = rel.dens. +rel.dom.) were calculated for each species. The CVI was also calculated for each plantfamily. The Shannon-Wiener diversity index and the Pielou evenness index (Brower &Zar, 1984) were obtained for each sampled community.

    The five cerrado communities were compared using three dissimilarity measures: thecomplemented coefficient of community (CC), the percentage difference (PD) and theEuclidian distance (ED), following Gauch (1982):

    cc,, = ioo- 2 0 0 s c

    PD p =100-200.

    where si and S2 = number of species in communities 1 and 2; sc = numberof species in common in the two communities; An, Ai2 = abundance ofspecies i in communities 1 and 2; min [An, Ai2] = lowest value ofabundance of species i in communities 1 and 2.

    The abundance values used for the species in the calculations of PDs and EDs were theirCVIs. ED is a measure determined mostly by larger abundance values while CC isdetermined only by the floristic composition. PD is intermediate, with linear weightingof species (Gauch, 1982). The dissimilarity matrices of CCs, PDs and EDs were used to

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  • A COMPARATIVE STUDY OF CERR ADO AREAS IN BRAZIL 313

    TABLE 1. Murundu cerrado: species sampled on 80 earthmounds (0.5-20.0m indiameter; 0.321 ha in total area). n = number of individuals; ba = basal area (m2);Rel.dens. = relative density; Rel.dom. = relative dominance; Rel.freq. = relativefrequency; IVI = importance value index. Species in descending order of IVI.

    1.2.3.4.5.6.7.8.9.

    10.11.12.13.14.15.16.17.18.19.20.21.22.23.24.25.26.27.28.29.30.31.32.33.34.35.36.37.38.39.40.41.42.43.44.45.

    Curatella americanaDipteryx alataAlibertia edulisAndira cuyabensisTabebuia aurcaHymenaea stigonocarpaEugenia aurataErythroxylum suberosumPseudobombax longiflorumSimarouba vcrsicolorVatairea macrocarpaMatayba guianensisTabebuia caraibaSclerolobium aureumByrsonima pachyphyllaJacaranda cuspidifoliaAstronium fraxinifoliumAbuta selloanaCecropia pachystachyaBrosimum gaudichaudiiCopaifera martiiPouteria ramifloraDiospyros coccolobifoliaCybistax antisyphiliticaBowdichia majorCasearia sylvestrisHeteropterys byrsonimifoliaQualea parvifloraCampomanesia eugenioidesErythroxylum citrifoliumSimaba trichilioidesZanthoxylum cinereumBauhinia bongardiMyrcia formosianaHimatanthus obovatusMagonia pubescensRudgea viburnioidesKielmeyera coriaceaVernonia brasilianaOuratea hexaspcrmaMyrcia uberavensisCardiopetalum calophyllumConnarus suberosusAnnona pygmaeaTocoyena formosa

    n

    39CAt8:4

    rl.ii

    1(i

    i

    Totalba

    4.1942.0530.4710.5030.4790.7770.1350.1120.078

    5 0.2250.273

    l 0.022! 0.093

    0.047> 0.054> 0.145) 0.053i 0.0667 0.142i 0.0255 0.0474 0.0404 0.036

    (5 0.0215 0.0284 0.022;

    ;

    5 0.0235 0.0141 0.013I 0.002> 0.002

    0.031I 0.003

    0.0080.0040.0040.0030.0020.0020.0020.0010.0010.0010.0010.001

    Re Rel.dens. dom.

    45.46 40.895.28 20.029.41 4.595.05 4.914.02 4.671.72 7.573.79 1.323.33 1.102.64 0.761.72 2.200.92 2.661.95 0.211.49 0.910.92 0.451.03 0.530.57 1.411.15 0.520.92 0.640.80 1.390.92 0.240.57 0.460.46 0.390.46 0.350.57 0.210.69 0.270.46 0.220.34 0.230.34 0.130.80 0.130.23 0.020.23 0.020.11 0.300.23 0.030.1 1 0.080.11 0.040.1 1 0.030.11 0.030.10.10.10.10.10.10.10.1

    1 0.021 0.021 0.021 0.011 0.011 0.011 0.011 0.01

    Rel.freq.

    16.928.338.848.086.062.275.305.814.293.542.023.282.782.02

    .77

    .26

    .52

    .52).76.77.26.01.01.01

    0.510.760.760.760.250.510.510.250.250.250.250.250.250.250.250.250.250.250.250.253.25

    IVI

    103.2733.6322.8418.0414.7511.5710.4110.237.697.455.605.445.183.393.333.253.183.082.952.932.29

    .86

    .82

    .79

    .46

    .44

    .33

    .23

    .190.750.750.670.510.450.400.400.400.390.390.380.380.380.380.383.38

    Totals 871 10.258 100 100 100 300

  • 314 EDINB. J. BOT. 48(3)

    TABLE 2. Pebble cerrado: species sampled by point-centred quarter method in asystematic grid of 140 points. See Table 1 for abbreviations.

    1.2.3.4.5.6.7.8.9.

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    Qualea parvifloraCuratella americanaSalvertia convallariodoraMyrcia uberavensisVochysia rufaQualea grandifloraDipteryx alataPouteria ramifloraSalacia crassifoliaDa villa ellipticaBowdichia majorPseudobombax longiflorumConnarus suberosusAndira cuyabensisTabebuia caraibaSclerolobium aureumByrsonima coccolobifoliaByrsonima pachyphyllaSimarouba versicolorVatairea macrocarpaErythroxylum suberosumBrosimum gaudichaudiiDiospyros coccolobifoliaAstronium fraxinifoliumHymenaea stigonocarpaDimorphandra mollisTabebuia ochraceaTabebuia aureaOuratea spectabilisAnnona coriaceaHimatanthus obovataLafoensia pacariAgonandra brasiliensisMagonia pubescensOuratea hexaspermaFerdinandus ellipticaStrychnos pseudoquinaMataybaguianensisDilodendron bipinnatumEriotheca gracilipesPseudobombax tomentosumMyrcia pallensKielmeyera coriaceaDiptychandra aurantiacaKielmeyera rubrifloraAcrocomia aculeataDalbergia miscolobiumJacaranda cuspidifoliaPlatypodium elegansAspidosperma macrocarponAlibertia edulisHancornia speciosaEugenia aurataStryphnodendroii obovatumHirtella gracilipesRudgea viburnioides

    n

    81534842272613191717119

    159

    101288748996466655444333232222222

    Totalba

    1.2121.0860.6630.3030.6450.3840.5993.3250.2090.1160.1940.2190.0540.1710.1340.0270.1240.0980.1140.1970.0630.0330.0240.0870.1450.0730.0580.0510.0420.0130.0390.0260.0230.0530.0340.0290.0530.0150.0260.0200.0130.0090.0080.0050.0050.0340.0300.0220.0220.0180.0110.0100.0080.0060.0060.005

    Rel.dens.

    14.469.468.577.504.824.642.323.393.043.04

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    Rel.dom.

    15.1113.558.273.788.044.797.474.062.611.452.422.730.682.131.700.341.541.221.422.460.790.410.301.091.800.910.730.630.530.170.490.330.290.660.420.360.660.190.320.250.160.110.090.060.060.420.370.280.270.220.140.120.100.080.070.06

    Rel.freq.

    13.159.367.576.974.184.781.993.593.192.992.191.792.39

    ;

    (

    (

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    .991.39.59.59.39

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    0.800.800.800.600.600.600.400.600.400.400.400.400.400.400.400.200.200.200.200.200.200.200.200.200.200.20

    IVI

    42.7332.3724.4218.2517.0414.2211.7911.048.837.476.576.135.755.545.484.874.564.244.073.973.813.613.503.363.313.182.992.902.422.052.00

    .84

    .80

    .80

    .56

    .50

    .42

    .32

    .07

    .010.910.870.850.850.820.800.750.660.650.600.510.500.480.460.450.45

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 315

    TABLE2(Cont.)

    57. Coccoloba mollis58. Eugenia uniflora59. Neca theif'era60. Coussarca hydrangcaefolia61. Cardiopctalum calophyllum62. Cybistax antisyphililica63. Acosmium dasycarpum64. Casearia sylvestris65. Tocoycna formosa

    Totals

    Totalba

    0.0050.0040.0030.0030.0030.0020.0020.0020.001

    Rel.dens.

    0.180.180.180.180.180.180.180.180.18

    Rel.dom.

    0.060.050.040.040.030.030.020.020.01

    Rel.freq.

    0.200.200.200.200.200.200.200.200.20

    IVI

    0.430.430.420.410.410.400.400.400.39

    560 8.018 100 100 100 300

    TABLE 3. Phyllite cerrado: species sampled by point-centred quarter method in asystematic grid of 60 points. See Table 1 for abbreviations.

    1.23.4.5.6.7.8.

    9.10.11.12.13.14.15.16.17.18.19.20.21.22.23.24.25.26.27.28.29.30.31.32.33.34.35.36.

    Magonia pubesccnsCallisthenc fasciculataPscudobombax longiflorumDipteryx alalaAstronium fraxinifoliumVatairea macrocarpaJacaranda cuspidifoliaAnadenanthera peregrina

    var. falcataCuratella americanaTerminalia subsericeaLafocnsia pacariAustroplenckia populncaByrsonima pachyphyllaQualea parvifloraBowdichia majorByrsonima coccolobifoliaKielmcyera rubrifloraSclerolobium aurcumDimorphandra mollisAcrocomia aculeataCampomanesia eugenioidesEriotheca gracilipesAspidosperma nobileHymenaea sligonocarpaQualea multifloraMachaerium acutifoliumKielmeyera coriaceaTabebuiaochraceaGuettarda viburnioidesMyrcia uberavensisHimatanthus obovatusConnarus suberosusLuchea speciosaAcosmium dasycarpumHancornia speciosaPlathvmenia rcticulata

    n

    3222201110117

    89

    118576555443333332332z

    221111

    Totalba

    0.3550.3300.1240.2540.2150.1110.185

    0.1350.0900.0310.0330.0780.0260.0290.0360.0240.0200.0380.0220.0320.0310.0210.0210.0200.0200.0410.0080.0100.0180.0100.0090.0060.0230.0150.0150.010

    Rel.dens.

    13.339.178.334.584.174.582.92

    3.333.754.583.332.082.922.502.082.082.081.671.671.251.251.251.251.251.250.831.251.250.830.830.830.830.420.420.420.42

    Rel.dom.

    14.1813.204.94

    10.168.604.427.38

    5.403.621.251.333.111.031.161.420.980.811.540.891.261.250.840.820.810.811.640.300.410.720.400.360.220.930.590.590.39

    Rel.freq.

    11.987.838.764.154.614.613.23

    3.233.693.693.232.303.232.302.302.302.30

    .84

    .84

    .38

    .38

    .38

    .38

    .381.380.921.380.920.920.920.920.920.460.460.46).46

    IVI

    39.5030.2022.0318.8917.3713.6213.53

    11.9511.059.527.897.497.175.965.815.375.205.054.403.903.883.483.463.443.443.402.942.592.482.162.111.971.811.471.471.27

  • 316 EDINB. J. BOT. 48(3)

    TABLE3(Cont.)

    37.38.39.40.41.42.43.44.45.46.47.48.

    Peltogyne confertifloraPouteria ramifloraPseudobombax tomentosumAnnona coriaceaQualea grandi floraDiptychandra aurantiacaKielmeyera grandifloraAspidosperma subincanumRoupala montanaEugenia aurataCasearia sylvestrisTocoyena formosa

    n

    111111111111

    Totalba

    0.0080.0070.0070.0050.0050.0050.0050.0050.0030.0030.0020.001

    Rel.dens.

    0.420.420.420.420.420.420.420.420.420.420.420.42

    Rel.dom.

    0.310.290.270.220.200.200.200.180.110.110.090.05

    Rel.freq.

    0.460.460.460.460.460.460.460.460.460.460.460.46

    IV!

    1.18.16.15.09.08.08.08.06

    0.990.990.97 +0.92

    Totals 240 2.501 100 100 100 300

    TABLE 4. Interfluvial cerrado: species sampled in 100 quadrats (10 x 10m). See Tablefor abbreviations.

    1.23.4.5.6.7.8.9.

    10.11.12.13.14.15.16.17.18.19.20.21.22.23.24.25.26.27.28.29.30.31.32.33.34.35.

    Syagrus comosaQualea parvifloraMyrcia albotomentosaDavilla grandifloraQualea grandifloraByrsonima verbascifoliaHymenaea stigonocarpaKielmeyera rubrifloraPouteria ramifloraLicania humilisRourea indutaVochysia rufaAndira cuyabensisCouepia grandifloraTabebuia caraibaAspidosperma tomentosumTerminalia brasiliensisMyrcia uberavensisDalbergia miscolobiumPsidium widgrenianumMouriri pusaSalacia crassifoliaLafoensia pacariVatairea macrocarpaKielmeyera coriaceaEugenia punicifoliaCaryocar brasilienseMouriri ellipticaPeltogyne confertifloraRoupala montanaByrsonima coccolobifoliaBowdichia virgilioidesGuapira noxiaAnnona crassifloraAndira inermis

    n

    355211153152709550733936514533282532233519241427171918187

    181512148

    101112

    Totalba

    1.9522.6891.2690.7361.5840.5391.2100.7421.0900.8810.1820.3830.5320.5380.5970.2950.5150.2280.4830.3530.5860.2090.2730.2360.1580.0900.3810.0750.1280.1310.0870.2240.1790.1430.091

    Rel.dens.

    18.8011.188.108.053.715.032.653.872.071.912.702.38

    .75

    .48

    .32

    .70

    .22

    .85

    .011.270.741.430.901.010.950.950.370.950.790.640.740.420.530.58().64

    Rel.dom.

    9.2812.786.033.507.532.565.753.535.184.190.861.822.532.562.841.402.451.082.301.682.780.991.301.120.750.431.810.360.610.620.411.060.850.680.43

    Rel.freq.

    6.255.616.566.664.285.044.004.382.952.573.622.472.382.382.002.761.622.091.621.71

    (

    .14

    .90

    .52

    .24

    .43

    .52).67.52.14.05

    1.140.760.860.950.95

    IVI

    35.0229.5720.7018.2115.5212.6412.3911.7710.208.667.186.686.666.426.075.865.285.034.924.664.674.333.723.363.132.902.852.832.542.312.302.252.242.212.02

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 317

    TABLE4(Cont.)

    36.37.38.39.40.41.42.43.44.45.46.47.48.49.50.

    51.52.53.54.55.56.57.58.59.60.61.62.63.64.65.66.67.68.69.70.

    Mczilaurus aff. lindavianaEriothcca gracilipesEugenia aurataDiospyros coccolobifoliaEugenia chrysanthaSyagrus petraeaTabebuia aureaPterodon pubescensCasearia sylvestrisDimorphandra mollisEretnanthus mattogrossensisBocageopsis mattogrossensisHimatanthus phagendaenicusOuratea hexaspermaSclerolobium paniculatum

    var. rubiginosumMiconia albicansAndira vermifugaTabebuia ochraceaDiptychandra glabraCuratclla americanaHancornia speciosaAspidosperma macrocarponConnarus suberosusManihot tripartitaAlibertia edulisStrychnos pscudoquinaPlathymcnia rcticulataEmmotum nitensMiconia rubiginosaAgonandra brasiliensisHimatanthus obovatusXylopia aromaticaErythroxylum tortuosumPalicourea rigidaTocovena formosa

    nTotal

    ba

    -) 0.20310 0.114

    (c

    ) 0.0667 0.076-> 0.089) 0.0245 0.0684 0.098(

    .

    L

    I

    3 0.0275 0.0375 0.021? 0.024i 0.025i 0.012

    ? 0.036i 0.0295 0.015? 0.010? 0.020

    0.057> 0.014

    0.044> 0.011> 0.004

    0.0350.033

    > 0.0190.0120.0050.0070.0070.0060.0060.0010.001

    Rel.dens.

    0.320.530.480.370.320.480.260.210.320.260.260.160.210.21

    0.160.210.160.160.160.050.110.050.110.110.050.050.110.050.050.050.050.050.050.050.05

    Rel.dom.

    0.960.540.310.360.420.110.320.470.130.180.100.110.120.06

    0.170.140.070.050.100.270.070.210.050.020.170.160.090.060.050.030.030.030.030.010.01

    Rel.freq.

    0.570.760.860.670.570.670.480.380.570.480.480.280.380.38

    0.280.190.280.280.190.100.190.100.190.190.100.100.100.100.100.100.100.100.100.100.10

    IVI

    .85

    .83

    .65

    .40

    .31

    .26

    .06

    .06

    .020.920.840.750.710.65

    0.620.540.520.490.440.420.360.360.350.320.310.310.290.200.200.180.180.180.180.15().15

    Totals 1888 21.044 100 100 100 300

    TABLE 5. Sandstone cerrado: species sampled by point-centred quarter method in asystematic grid of 50 points. See Table 1 for abbreviations.

    1.23.

    4.5.6.7.8.9.

    10.

    Teminalia fagifoliaVochysia petraeaSclerolobium paniculatum

    var. subvelutinumKielmeyera rubrifloraVcllozia flavicansPterodon pubescensPouteriarami floraQualca parvifloraEmmotum nitensDavilla grandiflora

    n

    2341

    21181688776

    Totalba

    0.6810.187

    0.3410.0640.0860.1810.1470.1610.1000.035

    Rel.dens.

    11.5020.50

    10.509.008.004.004.003.503.503.00

    Rel.dom.

    23.738.17

    14.882.813.747.916.437.014.381.54

    Rel.freq.

    11.9017.26

    10.718.936.553.574.762.983.573.57

    IVI

    53.1445.93

    36.0920.7418.2815.4815.1913.4911.468.11

  • 318 EDINB. J. BOT. 48(3)

    TABLE5(Cont . )

    11.12.13.14.15.16.17.18.19.20.21.22.23.24.25.26.27.28.29.30.31.32.

    Micropholis rigidaByrsonima pachyphyllaMyrcia uberavensisConnarus suberosusHeteropterys byrsonimifoliaSyagrus comosaByrsonima coccolobifoliaMimosa xanthocentraIlex asperulaMiconia albicansCaryocar brasilicnseNorantea goyazensisErythroxylum ambiguumFerdinandusa ellipticaOuratca spectabilisLadenbcrgia chapadensisEsenbeckia pumilaAspidosperma macrocarponEschweilera nanaBredemeyera altissimaStyrax ferrugineusMouriri pusa

    Totals

    n

    i

    Totalba

    5 0.0515 0.036\ 0.0104 0.009:\ 0.0205 0.015I 0.028\ 0.003> 0.023> 0.021

    0.0230.0140.0140.0090.0070.0050.0040.0040.0030.0030.0030.003

    200 2.291

    Rel.dens.

    2.502.502.002.00

    .50

    .50

    .00

    .50

    .00

    .000.500.500.500.500.500.500.500.500.500.500.500.50

    100

    Rel.dom.

    2.231.570.450.390.880.641.230.131.040.931.010.610.610.380.290.220.150.150.130.130.110.11

    100

    Rel.freq.

    2.982.982.38

    .79

    .79

    .79

    .19

    .79

    .19

    .190.600.600.600.600.600.600.600.600.600.600.600.60

    100

    IV!

    7.717.054.834.184.173.923.423.413.233.122.111.711.711.471.391.311.251.251.221.221.211.21

    300

    TABLE 6. Synthetic data for the vegetation of the five areas of cerrado studied. n = totalnumber of individuals; S = number of species; TAD = total absolute density (indiv./ha);H = average height (m); D = average diameter (cm); TADo = total absolute dominance(m2/ha); H, = Shannon-Wiener diversity index (with logs, in base 2); J, = Pielouevenness index (logs, in base 2).

    Cerrado AreaMurunduPebblePhylliteInterfluvialSandstone

    nN871560240

    1888200

    S4565487032

    TAD2713.71049.91546.41888.01187.0

    H2.753.594.243.063.38

    D9.87

    11.9110.3911.9010.36

    TADo31.9615.0316.1221.0413.60

    H,3.4004.8994.8504.6644.057

    J,0.6100.8130.8680.7610.811

    TABLE 7. Analytical data for surface horizon (0 20cm) of soils of the five areas of cerradostudied.

    Cerrado Area

    Murundu**Pebble**Phyllite**Interfluvial*Sandstone*

    pH inH,O

    4.75.55.34.95.2

    ChemicalP

    ppm

    5.0 110.0

    1.21.60.6

    PropertiesK

    105964138

    Ca + Mgmeq./

    1.02.21.50.30.4

    AllOOg

    1.90.51.00.80.2

    O.M.0 /

    2.32.02.01.90.6

    Texturesand

    4777628789

    silt0 /

    267

    1422

    da

    271624119

    Parent material: *Sandstone; **Phyllite. O.M.=Organic Material

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 319

    XUJQ

    40

    20

    40

    20

    4 0 -

    20

    UJ

    OC 6 0UJsO 40

    20

    80

    6CH

    20

    O lCD

    ure

    E

    mureorenEo

    CDEoU

    0>re0>orec

    igno

    o>re01ure5

    nacc

    orej zOlare2

    TOre_QO(/i>.

    C

    O

    01re01oreN

    0)>

    UJ

    i PD-> ED diagrams) the more these cerrados are polarized.In both the polar ordination diagram and the dissimilarity dendrogram based on CCmeasures (first pair in Fig. 3; emphasis on floristic composition) the cerrados of theBaixada Cuiabana (murundu, pebble and phyllite cerrados) form a cohesive group,separated from the cerrados of the Chapada dos Guimaraes (interfluvial and sandstonecerrados). In both cases the sandstone cerrado is the most different in terms of floristiccomposition. On the other hand, in terms of community structure, which is emphasizedby EDs (lowest pair in Fig. 3), the murundu cerrado is shown quite isolated from theother cerrados in both the diagram and the dendrogram.

    The main floristic dissimilarity between the cerrados of the Chapada dos Guimaraes(interfluvial and sandstone cerrados) and those of the Baixada Cuiabana (murundu,pebble and phyllite cerrados) is probably due mostly to differences in soil properties.The sandstone bedrock of the Chapada dos Guimaraes gives rise to coarse and very

  • 322 EDINB. J. BOT. 48(3)

    infertile sediments while the soils of the Baixada Cuiabana are mainly influenced by thephyllite bedrock, which gives rise to finer and more fertile sediments.

    The phyllite cerrado presents many species considered as indicators of the mesotro-phic facies cerradao first described by Ratter (1971) in northeastern Mato Grosso andrecognized later in many parts of Central Brazil (Ratter et al, 1973,1977,1978,1988).This type of cerradao (augmentative of cerrado in Portuguese) occurs mainly on soilswith relatively high levels of Ca and Mg. The authors termed it 'Magonia pube-scenslCallisthene fasciculata cerradao', from the two main indicator species which arealso the species with the highest importance values in the phyllite cerrado. Some otherindicator species that could be found in abundance in this cerrado are Pseudobombaxlongiflorum, Dipteryx alata, Astroniumfraxinifolium, Jacaranda cuspidifolia and An-adenanthera peregrina var.falcata. Some species occurring in small numbers - such asPseudobombax tomentosum, Acrocomia aculeata, Aspidosperma subincanum, Lueheaspeciosa and Guettarda viburnioides - are also absolutely typical of this community.

    The pebble cerrado is a widely dispersed type of cerrado that can be found throughoutthe Baixada Cuiabana in all sites where the soils are shallow and contain the layer ofpebbles. These aspects favour higher rates of seepage and runoff in the groundwaterregime and also reduce strongly the available volume for root growth and feeding.Despite having the highest levels of Ca+Mg and P, the apparent richness of these soilsis due mainly to the high proportion of pebbles (c.75% of volume) which are removedbefore the analyses. Therefore, the figures in Table 7 really refer to a soil volume aboutfour times greater than in the other cerrados. The plants therefore have available onlyabout one-quarter of the apparent Ca+Mg content, which explains the presence of fewindicator species of mesotrophic facies cerradao. Dipteryx alata, Pseudobombax longi-florum and Astroniumfraxinifolium, however, do occur but only the first is among the10 species with highest importance values. Four of the most important species are inthe Vochysiaceae: Qualea parviflora, Qualea grandiflora, Salvertia convallariodora,and Vochysia rufa. These species, also common in wide areas of cerrado in all CentralBrazil can be considered as amongst the most characteristic species of cerrado in a broadsense. Curatella americana (Dilleniaceae) which is very common in most cerrados ofthe Baixada Cuiabana is usually an indicator of soils with seasonally high water-table(Oliveira-Filho & Martins, 1986; Oliveira-Filho et al., 1989: Ratter et al., 1988).

    The murundu cerrado is a very different community because of the special environ-mental conditions: the restricted growth-space permanently free of flooding on theearthmounds, which constitute true ecological islands (Diniz de Araujo Neto et al., 1986;Oliveira-Filho, 1988). Although the flora of the murundus shows most species incommon with the neighbouring cerrado (the pebble cerrado), its community structure isvery different, with a strong concentration of the importance values in a few species,particularly Curatella americana. This species, as already mentioned, is an indicator ofseasonal high water levels (floods or seepage) and is also typical of murundus of theSerra do Roncador (Ratter et al., 1973), of the Ilha do Bananal (Ratter, 1987) and of thePantanal Matogrossense (Prance & Schaller, 1982; Ratter et al., 1988). Other cerradospecies which commonly occur in many areas of murundus are Dipteryx alata, Andira

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 323

    cuyabensis, Simarouba versicolor and Matayba guianensis. For typical cerrado to occur,the soils must be freely drained since the majority of species of the cerrado flora do nottolerate soil waterlogging, even during relatively short periods (Eiten, 1972: Oliveira-Filho et al., 1989). So, the cerrado species successfully colonizing the murundus arethose best able to tolerate high soil moisture. The less tolerant species do not succeedin colonizing the murundus and this seems to happen to the Vochysiaceae whose speciesare almost entirely absent in the murundu cerrado despite their abundance in theneighbouring pebble cerrado. The absence or rarity of the Vochysiaceae were alsonoticed by Ratter et al. (1988) in some areas of inundable cerrado of the PantanalMatogrossense, although the same author (Ratter et al., 1973) and Oliveira-Filho et al.(1989) noted Qualea parviflora as characteristic of the cerrado/wet campo transition inMato Grosso.

    The environmental limiting factors of the sandstone cerrado, at the top of the Chapadados Guimaraes represent the other extreme from those of the murundu cerrado. Thevegetation can grow only where some soil accumulates within cracks or in the spacesamong the rocks. The paucity of real soil is aggravated by its coarse texture, by the verylow content of mineral nutrients and organic matter, and by the low water storagecapacity. The species living in such a rigorous environment must have specializedadaptations. Thus, the species with the highest importance values - Vochysia petraea,Terminalia fagifolia Sclerolobium paniculatum var. subvelutinum and Vellozia flavi-cans - are characteristic of drier cerrado locations and are absent or very rare in all theneighbouring cerrados. However, species such as Kielmeyera rubriflora, Pterodonpubescens and many others with lower importance values also occur in the interfluvialcerrado 300m below, at the base of the escarpments.

    The interfluvial cerrado also occurs on coarse-textured infertile soils which arose fromthe sandstone. Nevertheless, the soils are deeper and with higher water storage capacitythan those of the sandstone cerrado. The environmental limitation of this cerrado isrelated principally to water table levels which rise in the rainy season so that waterflushes on the valley sides. The highest level reached by seepage and runoff on the valleyside corresponds to the border line between the cerrado and the marshy campo. Theinfluence of water-table level within this cerrado is described in details by Oliveira-Filhoetal. (1989).

    ACKNOWLEDGEMENTS

    We are indebted to the Universidade Federal de Mato Grosso for support received frommany undergraduate students who participated in field activities: in special Silvio A.Rodrigues, Julio C. Dalponte, Beatriz Schwantes, Cecilia B. Rocha, Susan H. Damas-ceno, Nelson Laturner, Paulo B. Andrade, and Lyria C. A. Oliveira. We also acknow-ledge with gratitude the identification of botanical specimens by the taxonomists of theUniversidade Estadual de Campinas and the critical reading of this text by Dr James A.Ratter, Royal Botanic Garden, Edinburgh, and Dr Paulo S. Oliveira, the UniversidadeEstadual de Campinas.

  • 324 EDINB. J. BOT. 48(3)

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    POTT, V.J., POTT, A., RATTER, J.A. & VALLS, J.F.M. (1986). Flora da FazendaNhumirim, Nhecolandia, Pantanal. Relacao preliminar. Pesquisa em Andamento(EMBRAPA CPAP) No. 5, 26 pp.

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 325

    PRANCE, G.T. & SCHALLER, G.B.(1982). Preliminary study of some vegetation types ofthe Pantanal, Mato Grosso, Brazil. Brittonia 34: 228-251.

    RATTER, J.A. (1971). Some notes on two types of cerradao occurring in northeasternMato Grosso. /// Simposio sobre o cerrado: 100-102. Universidade de SSo Paulo.

    , (1987). Notes on the vegetation of the Parque Nacional do Araguaia (Brazil). Notesfrom the Royal Botanic Garden Edinburgh 44: 311-342.

    , RICHARDS, P.W., ARGENT, G. & GIFFORD, D.R. (1973). Observations on thevegetation of northeastern Mato Grosso. I. The woody vegetation types of theXavantina-Cachimbo Expedition area. Philosophical Transactions of the RoyalSociety of London (B) 226: 449-492.

    , ASKEW, G.P., MONTGOMERY, R.F. & GIFFORD, D.R. (1977). Observacoes adicionaissobre o cerradao de solos mesotroficos no Brasil Central. TV Simposio sobre oCerrado: 306-316. Universidade de Sao Paulo.

    , , & (1978). Observations on forests of some mesotrophic soils in CentralBrazil. Revista Brasileira Botdnica 1: 47-58.

    , POTT, A., VALI, J.P., CUNHA, C.N. & HARIDASAN, M. (1988). Observations onwoody vegetation types in the Pantanal and at Corumba, Brazil. Notes from the RoyalBotanic Garden Edinburgh 45: 503-525.

    APPENDIX

    Species recorded in the study areas of the Baixada Cuiabana (BC) and the Chapada dosGuimaraes (CG)

    +, present; -, not observed; br, bromeliad; li, liana; pi, palm; sc, succulent;sh, shrub; ss, sub shrub; tr, tree

    The list contains all species collected and/or observed and not only those recorded inthe sample areas. It is confined to cerrado vegetation (gallery forests and grasslands areexcluded).

    growth regionsform

    BC CGANACARDIACEAE

    Anacardium humile St. Hil.Astronium fraxinifolium SchottA. urundeuva (Fr. All.) EnglerTapirira guianensis AubletT. marchandii Engler

    ANNONACEAEAnnona cohacea Mart.A. crassiflora Mart.A. dioica St. Hil.A. pygmaea Warm.Bocageopsis mattogrossensis R. E. Fr.Cardiopetalum calophyllum SchldlDuguetia furfuracea (St. Hil.) Benth. & Hook.D. lanceolata St. HU.Xylopia aromatica (Lam.) Mart.X. sericea St. Hil.

    ss

    trtrtrtr

    shtrss

    ss

    trtr

    sh-sssh-ss

    trtr

  • 326 EDINB. J. BOT. 48(3)

    APOCYNACEAEAspidosperma cylindrocarpon MiilL Arg.A. macrocarpon Mart.A. nobile MiilL Arg.A. subincanum Mull. Arg.A. tomentosum Mart.Hancornia speciosa GomezHimatanthus obovatus (M. Arg.) WoodsonH. phagendaenicus (Mart.) WoodsonOdontadenia sp.Rauwolfia sp.

    AQUIFOLIACEAEIlex asperula Mart.I. cerasifolia Reiss.

    ARACEAETaccarum weddellianum [Brong. ex] Schott

    ARALIACEAEDidymopanax macrocarpum (Cham. & Schldl.) Seem.D. vinosum (Cham. & Schldl.) Marchand

    BIGNONIACEAEAnemopaegma glaucum Mart.Arrabidaea brachypoda (DC) Bureau. & K. Schum.A. corrallina (Jacq.) Sandw.Cybistax antisyphilitica Mart.Jacaranda cuspidifolia Mart.J. decurrens Cham.Memora pubescens (Sprengel) K. Schum.Tabebuia aurea Benth. & Hook.T. caraiba (Mart.) BureauT. ochracea (Cham.) Standl.T. roseoalba (Ridley) Sandw.Zeyheria digitalis (Veil.) Hoehne

    BOMBACACEAEEriotheca gracilipes (K. Schum.) A. RobynsPseudobombax longiflorum (Mart. & Zucc.) A. RobynsP. tomentosum (St Hil.) A. Robyns

    BORAGINACEAECordia alliodora (Ruiz & Pav.) OkenC. insignis Cham.C. glabrata (Mart.) A. DC.

    BROMELIACEAEAnanas sp.Bromelia balansae Mez

    BURSERACEAEProtium almecega MarchandP. heptaphyllum (Aublet) MarchandP. ovatum Engler.

    CACTACEAECereus peruvianus (L.) MillerMelocactus sp.

    growthform

    trtrtrtrtr

    tr-shtr-sh

    trliss

    trtr

    sc

    tr-shsh

    ss

    shli

    tr-shtrss

    ss

    trtrtrtr

    sh-ss

    trtr-shtr-sh

    trss

    tr

    brbr

    trtrss

    sc

    sc

    regions

    BC CG

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 327

    growth regionsform

    BC CGCARDYOCARACEAE

    Caryocarbrasiliense Cambess. tr - +CELASTRACEAE

    Austroplenckia populnea (Reisseck) Lundell tr - +CHRYSOBALANACEAE

    Couepia grandiflora (Mart. & Zucc.) Benth. tr - +Hirtella glandulosa Mart. & Zucc. tr + +H. gracilipes (Hook.) Prance tr + +Iicania hoehnei Pilger tr - +L. kunthiana Hook. tr - +L. humUis Cham. & Schldl. tr - +

    COCHLOSPERMACEAECochlospennum regium (Mart. & Zucc.) Pilger sh-ss + +

    COMBRETACEAEBuchenavia tomentosa Eichler tr - +Combretum discolor Taub. ss + +T. brasiliensis (Cambess.) Eichler tr + +T. fagifolia Mart. & Zucc. tr - +T. subsericeaa Eichler tr + -

    COMPOSITAEEremanthus mattogrossensis Kuntze tr-sh - +Ichthyothere cunabi Mart. ss - +Piptocarpha rotundifolia (Less.) Baker tr - +Vernonia brasiliana (L.) Drude sh + -V. ruficoma Schldl. sh +

    CONNARACEAEConnarus suberosus Planch, var. fulvus (Planchon) Forero tr-sh + +Rourea induta Planch. sh + +

    DILLENIACEAECuratella amencana L. tr + +Davilla elliptica St. Hil. sh + +D. grandiflora St. Hil. & Tul. sh + +

    EBENACEAEDiospyros coccolobifolia Mart tr + +

    ERYTHROXYLACEAEErythroxylum ambiguum St. Hil. tr-sh + +E. campestre SL Hil. ss + +E. citrifolium St Hil. tr-sh + +E. cuneifolium (Mart.) E. Schulz ss - +E. deciduum St. Hil. tr-sh +E. suberosum St. Hil. sh + +E. tortuosum Mart. sh + +

    EUPHORBIACEAECnidoscolus urens (L.) Arthur li +Dalechampia sp. li + -Manihot tripartita Mull. Arg. sh-ss - +

    FLACOURTIACEAECasearia sylvestris Sw. tr-sh + +

    GUTTIFERAEKielmeyera coriacea (Spr.) Mart tr-sh + +K. grandiflora (Wawra) Saddi tr-sh + +

  • 328 EDINB. J. BOT. 48(3)

    GUTTIFERAE (contd.)K. rubriflora Cambess.

    HIPPOCRATEACEAEPeritassa campestris (Cambess.) A.C. SmithSalacia crassifolia (Mart.) Peyr.

    ICACINACEAEEmmotum nitens (Benth.) Miers

    LAURACEAEMezilauras crassiramea (Meissn.) Taub.M. aff. lindaviana Schw. & MezOcotea sp.

    LECYTHIDACEAEEschweilera nana (Berg) Miers

    LEGUMINOSAE CAESALPINIOIDEAEBauhinia bongardi Steudel.B. cuyabensis Steudel.B. pulchella Mart.B. rufa Steudel.Cassia alata L.C. claussenii Benth.C. desvauxii (Colladon.) KillipC. ragosa G. Don.C. sylvestris Veil.C. velutina VogelCenostigma gardnerianum Tul.Copaifera elliptica Mart.C. langsdorffii Desf.C. martii HayneDimorphandra mollis Benth.Diptychandra aurantaca Tul.D. glabra Benth.Hymenaea stigonocarpa Mart.Peltogyne confertiflora (Hayne) Benth.Sclerolobium aureum (Tul.) Benlh. var. aureum (Tul.) Benth.S. paniculatum Vogel var. rabiginosum (Tul.) Benth.S. paniculatum Vogel var. subvelutinum Dwyer

    LEGUMINOSAE FABOIDEAEAcosmium dasycarpum (Vogel) YakovlevAndira cuyabensis Benth.A. inermis KunthA. vermifuga Mart.Bowdichia major (Mart.) Benth.B. virgilioides KunthCentrolobium tomentosum Benth.Dalbergia miscolobium Benth.Dioclea tricolor Benth.Dipteryx alata VogelEriosema rufum E. MayerErythrina velutina Willd.Galactia glaucescens KunthHarpalyce brasiliana Benth.

    growthform

    tr-sh

    ss

    tr-sh

    tr

    trtr

    sh-ss

    tr-sh

    sh-sssh-sssh-sssh-ss

    shsh-ss

    shsh-sssh-sssh-ss

    shsh-ss

    trtr-shtr-sh

    trtrtrtrtrtrtr

    tr-shtrtrtrtrtrtrtr

    sh-sstrss

    trsh-sssh-ss

    regions

    BC CG

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 329

    LEGUMINOSAE FABOIDEAE (contd.)Machaerium acutifolium VogelPlatypodium elegans VogelPterodon pubescens Benth.Vatairea macrocarpa Benth.

    LEGUMINOSAE MIMOSOIDEAEAcacia plumosa LoweAnadenanthera peregrina (L.) Speg. var. falcata (Benth.)

    AltschulCalliandra parviflora Benth.Enterolobium contortisiliquum (Veil.) MorongInga heterophylla Willd.Mimosa hirsuta SprengelMimosa pteridifolia Benth.Mimosa xanthocentra Mart. var. subsericea (Benth.) BarnebyPlathymenia reticulata Benth.Pithecellobium edwallii HoehneStryphnodendron obovatum Benth.

    LOGANIACEAEAntonia ovata PohlStrychnos pseudoquina St. Hil.

    LYTHRACEAELafoensia pacari St. Hil.Physocalymma scaberrimum Pohl

    MALPIGHIACEAEByrsonima coccolobifolia KunthB. coriacea DC.B. crassa Niedenzu.B. dealbata Gris.B. gaultherioides Gris.B. pachyphylla Juss.B. subterranea Brade & MarkB. verbascifolia (L.) Rich, ex Juss.Heteropterys byrsonimifolia A. Juss.Heteropterys sp.

    MALVACEAEPavonia sp.

    MARCGRAVIACEAENorantea goyazensis Camb.

    MELASTOMATACEAEMiconia albicans (Sw.) TrianaM. fallax DC.M. ferruginata (DC.) Cogn.M. guianensis Cogn.M. rubiginosa (Bonpl.) Tr.Mouriri clliptica Mart.M. pusa GardnTibouchina aff. aspera Aublet.T. pogonanthera (Naud) Cogn.

    MELIACEAEGuarea guidonia (L.) Sleumer

    growthform

    trtrtrtr

    trtr

    sh-sstrtrshsh

    sh-sstrtrtr

    trtr

    trtr

    tr-shtr-shtr-sh

    ss

    shtr-sh

    ss

    tr-shtr-shsh-ss

    ss

    sh

    tr-shtr-shtr-sh

    trshshtr

    sh-sssh

    tr

    regions

    BC CG

  • 330 EDINB. J. BOT. 48(3)

    growth regionsform

    BC CGMENISPERMACEAE

    Abuta selloana Eichler sh + +MONJMIACEAE

    Siparuna guianensis Aublet sh - +MORACEAE

    Brosimum gaudichaudii Trecul tr-sh + +Cecropia cinerca Miq. tr - +C. pachystachya Trecul tr + +Ficus guianensis Aublet tr - +F. cnormis (Mart, ex Miq.) Miq. tr +

    MYRISTICACEAEVirola sebifera Aublet ss + +V. sessilis(A. DC.)Warb. tr - +

    MYRSINACEAERapanea guianensis Aublet tr - +

    MYRTACEAECampomanesia eugenioides (Cambess.) Legr. sh + +Eugenia aurata Berg tr-sh + +E. bimarginata DC. sh - +E. chrysantha Berg tr-sh - +E. punicifolia (Kunth) DC. tr-sh - +E. uniflora L. tr-sh +Myrcia albo-tomentosa Cambess. tr-sh + +M. formosiana DC. tr-sh - +M. intermedia (Berg) Kiaersk. sh +M. aff. nivea Berg ss - +M. pallensDC. tr-sh + +M. suffruticosa Berg ss + -M. uberavensis Berg tr-sh + +M. variabilis DC. sh-ss - +Psidium multiflorum Camb. ss +P. widgrenianum Berg tr-sh - +

    NYCTAGINACEAEGuapira noxia (Netto) Lundell tr - +Neea theifera Oersted tr + +

    OCHNACEAEOuratea floribunda (SL Hil.) Engler ss + +O. hexasperma (St. Hil.) Engler tr-sh + +O. spectabilis (Mart.) Engler tr-sh + +

    OPEJACEAEAgonandra brasiliensis Miers tr + +

    PALMAEAllagoptera campestris (Mart.) Kuntze pi - +A. leucocalyx (Drude) Kuntze pi +Acrocomia aculeata (Jacq.) Lodd. ex Mart. pi +Astrocaryum campestre Mart. pi - +Butia leiospatha (Mart.) Becc. pi - +Syagrus comosa (Mart.) Becc. pi - +S. petraea (Mart.) Becc. pi - +

    POLYGALACEAEBredemeyera altissima Benth. sh-ss - +

  • A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL 331

    growth regionsform

    BC CGPOLYGONACEAE

    Coccoloba mollis Casar.C. cf. cordifolia Meissner

    PROTEACEAERoupala montana Aublet

    RHAMNACEAERhamnidium elaeocarpum Reisseck.

    RUBIACEAEAlibertia concolor (Cham.) K. Schum.A. edulis Rich.A. sessilis (Cham.) K. Schum.A. verrucosa MooreChomelia ribesioides Benth.Coussarea hydrangeaefolia Benth. & Hook.Ferdinandusa elliptica PohlGuettarda viburnioides Cham. & Schldl.Ladenbergia chapadensis MoorePalicourea coriacea (Cham.) K. Schum.P. rigida KunthRudgea viburnioides (Cham.) Benth.Sabicea cana Hook.Tocoyena formosa (Cham. & Schl.) K. Schum.

    RUTACEAEEscnbeckia pumila PohlZanthoxylum arenarium Engl.Z. cinereum Engl.

    SAPINDACEAEDilodendron bipinnatum Radlk.Magonia pubescens St. Hil.Matayba guianensis AubletSerjania erecta Radlk.S. sp.Talisia subalbens Radlk.

    SAPOTACEAEChrysophyllum marginatum (Hook. & Am.) Radlk.Micropholis rigida PierrePouteria lateriflora (Benth.) Radlk.P. ramiflora (Mart.) Radlk.

    SIMAROUBACEAESimaba trichilioides St. Hil.Simarouba versicolor St. Hil.

    SMILACACEAESmilax sp.

    SOLANACEAESolanum lycocarpum St. Hil.

    STERCULIACEAEGuazuma tomentosa KunthG. ulmifolia Lam.Helicteres sacarolha St. Hil.Waltheria sp.

    tr-shss

    tr

    tr

    shtr-sh

    shtr-sh

    shtr-shtr-shtr-shtr-sh

    ss

    sh-sstr-sh

    ss

    sh-ss

    tr-shtrtr

    tr-shtr

    tr-shss

    liss

    trtrss

    tr

    sh-sstr

    li

    sh

    trtrss

    ss

  • 332 EDINB.J.BOT.48(3)

    STYRACACEAEStyrax camporum PohlS. fertugineus Nees & Mart.

    TTLIACEAELuehea speciosa Willd.

    ULMACEAETrema micrantha Blume

    VELLO23ACEAEVellozia flavicans Mart.

    VERBENACEAEAegiphila sellowiana Cham.Vitex cymosa Bert.

    VTTACEAECissus sp.

    VOCHYSIACEAECallisthene fasciculata (Spr.) MartQualea grandiflora MartQ. multiflora Mart.Q. parviflora Mart.Salvertia convallariodora St Hil.Vochysia divergens Mart.V. haenkeana Mart.V. petraea Warm.V. pyramidalis (Spr.) Mart.V. rufa (Spr.) Mart.V. tucanorum (Spr.) Mart.

    growthform

    trtr

    tr-sh

    tr

    sh

    trtr

    trtrtrtrtrtrtrshtrtrtr

    regions

    BC CG