Diversity, floristic and structural patterns of cerrado vegetation in CentralBrazil

Jeanine Maria Felfili1,*, Manoel Cláudio da Silva Júnior1, Anderson Cássio Sevilha2,Christopher William Fagg1, Bruno Machado Teles Walter2, Paulo Ernane Nogueira1 andAlba Valéria Rezende1

1Departamento de Engenharia Florestal CP 04357 Universidade de Brasília 70 919 970 Brasília-DF;2EMBRAPA-Recursos Genéticos e Biotecnologia, Brasília, DF; *Author for correspondence (e-mail:felfili@unb.br)

Received 28 February 2003; accepted in revised form 31 October 2003

Key words: Alpha and beta diversity, Floristic similarity, Savanna, Brazil

Abstract

The cerrado has been identified as one of the richest and most threatened biomes of the world, but few phyto-geographical studies have been undertaken in the region. A total of 70 land systems based on climate, landscapeand soils have been identified in the region, but it remains to be seen if the distribution and structure of the plantcommunities support these divisions. The aim of this work was to compare the floristic and structural similarityof cerrado sensu stricto within and between three physiographic units, named Pratinha, Veadeiros and São Fran-cisco, which contain six land systems in central Brazil and cover 10 degrees of latitude and five degrees of lon-gitude. The woody vegetation of 15 selected sites of the cerrado sensu stricto physiognomy was surveyed undera standardized methodology. The number of species per site varied from 55 to 97, with most sites having around60 to 70 species, and Shannon´s diversity indices ranged from 3.44 to 3.73, with most sites around 3.5 suggest-ing high alpha diversity. Sørensen´s floristic similarity index was high, with all Figures above 0.5 between thesites in the same land system in each physiographic unit but low between sites in different land systems in theVeadeiros. Czekanowski similarity indices were lower than Sørensen’s in the comparisons due to a high struc-tural differentiation between the sites. There is a large overlap in species occurrence in the sites but the size oftheir populations is very different at each site. Therefore, the high beta diversity is mostly due to differences inabundance of species between sites. The sites were separated by physiographic units, considering the first threedivisions of TWINSPAN classification. The first axis of DCA ordination showed a gradient going from the cer-rado on deep soils in Pratinha, through to those on sandy soils in São Francisco and ending on the shallowersoils of the Veadeiros. Land systems conformed well with the floristic and structural variations of the vegetation,indicating their potential use in designing a network of conservation areas in the cerrado region and as a basis fordecision-making on management.

Introduction

The cerrado vegetation has been identified as one ofthe richest and most threatened in the world �Mitter-meier et al. 2000�, but few phytogeographical studieshave been undertaken in the region. The biome occu-

pies a large latitudinal gradient of over 20 degrees incentral Brazil, and has been extensively replaced byagriculture and cultivated pasture. Estimates point toa 50-60% loss of the natural cover in the cerradobiome �Klink et al. 1996; Mittermeier et al. 2000�,which is probably leading to a high rate of extinction

Plant Ecology 175: 37–46, 2004.© 2004 Kluwer Academic Publishers. Printed in the Netherlands.

37

as the distribution of many plant species is frag-mented �Ratter and Dargie 1992; Ratter et al. 1996;Felfili et al. 1994,1997, Felfili and Silva Júnior 2001�.Even widespread species are endangered on a long-term basis as gene flow occurs mostly at short dis-tances �Collevatti et al. 2001� and the ConservationAreas legally established by the government are frag-mented and cover less than 1% of the territory.

The most recent published account of the flora ofthe cerrado biome, which includes several physiogno-mic units of cerrado sensu lato �savanna� from grass-lands to woodlands, gallery forests, swampy palmformations and dry seasonal forests, describes 6671species, of which c. 2500 occur in the savanna wood-land, called cerrado sensu stricto �Mendonça et al.1998�.

Cerrado is a savanna woodland physiognomy, withthe degree of canopy cover varying from 10 to 60%per site �Eiten 1972�, which occupies most of centralBrazil �Ribeiro and Walter 1998�. From here on, theterm ´cerrado´ is used in the strict sense unless other-wise stated. Most studies on cerrado consist of floris-tic and phytosociological surveys of a site. Someauthors have compared floristic surveys using differ-ent methodologies �Castro 1994; Durigan et al. 2003;Ratter and Dargie 1992; Ratter et al. 1996, 2000,2003�. Felfili and Silva Júnior 1993 and Felfili et al.�1994, 1997� compared sites using a standardizedmethodology within the framework of the project“Biogeography of the Cerrado Biome” and “Conser-vation and Management of the Biodiversity of theCerrado Biome”. These studies have shown that thevegetation is rich and diverse and that the spatial dis-tribution is of mosaics. Felfili et al. �1994, 1997� sug-gested that altitude and soil types are the maindeterminants of the floristic composition and structureof the cerrado of the Pratinha and Veadeiros Plateaux.According to Felfili �1998� the connection with thelarge hydrographical basins of Araguaia-Tocantins�Amazonia�, São Francisco and Paraná rivers alsoseems to influence the phytogeographical patterns ofthe cerrado, as has been suggested for the gallery for-ests of the region �Oliveira-Filho and Ratter 1995�.

The phytogeographic patterns under the same cli-matic domain �Allaby 1992� are determined mostlyby physical factors such as soils, relief and topogra-phy. Those factors were overlayed for a land systemzoning of Central Brazil by Cochrane et al. �1985�.They identified a total of 70 land systems within 25physiographic units in the region of cerrado in Cen-tral Brazil. The hypothesis is that the floristics and

structure of the vegetation vary accordingly to theland system classification. If so, a comparison of alarge area containing several land systems will eluci-date patterns of alpha and beta diversity being, there-fore, useful for decision-making on conservation andmanagement of cerrado vegetation.

The aim of this paper is to compare the floristicsand structure of cerrado within and between threephysiographic unit containing six land systems incentral Brazil and covering ten degrees of longitudeand five degrees of latitude and distances of over1000 km, in order to detect patterns of alpha and betadiversity at a regional level.

Methods

Study sites

The woody vegetation of the cerrado physiognomy of15 selected sites in three physiographic units �Co-chrane et al. 1985� was chosen for the comparison.These three physiographic units include six land sys-tems, see Table 1. A land system is an area or groupof areas throughout which there is a recurring patternof climate, landscape, soils and vegetation physiog-nomies.

The study-sites were selected along a gradient of10 degrees of latitude and five degrees of longitude,see Figure 1. Six study-sites were in the Pratinha landunit �Felfili et al. 1994�, five in the Veadeiros �Felfiliet al. 1997� and four in São Francisco �Table 1�. Fivestudy-sites were Conservation Areas and ten were onprivate land. There were three conservation areas inChapada Pratinha, all in the Federal District c. 50 kmdistant from each other, and all located under thesame land system called Pratinha highlands accord-ing to Cochrane et al. �1985�. They were: the BrasíliaNational Park with 30,000 ha; The environmentallyprotected area �APA� Gama e Cabeça de Veado,which includes the Ecological Reserves of the IBGE,the Ecological Reserve of the University of Brasíliaand the Brasília Botanic Gardens, which are contigu-ous and protect a total of c. 10,000 ha; The Ecologi-cal Station of Água Emendadas �ESEC-AE� with10,000 ha. The latter station consists of a watershedfor the three main hydrographical basins in Brazil:Amazon, São Francisco and Paraná river basins.

The Veadeiros National Park in Goiás State was theonly conservation unit on the Veadeiros plateaux, and

38

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39

its 65,000 ha completely occupies one of the threeland systems of the land unit.

Grande Sertão Veredas National Park with 80,000ha is the only conservation unit in São Francisco, inthe Bahia and Minas Gerais States.

All sites have a Aw climate by Köppen´s classifi-cation; location and other physical characteristics�Cochrane et al. 1985� are given in Table 1.

Data collection and analyses

The study-sites were chosen at the extreme and inter-mediate portions of each physiographic unit and alsocovered the land systems within each unit. Maps, sat-ellite images, aerial photos and preliminary field sur-veys were used to selected places still covered bynative vegetation without evidence of disturbance inthe study-sites outside the conservation areas.

In each physiographic unit, most study sites were100 km or more distant from each other but in Prat-inha, the first physiographic unit to be assessed, threesites were about 50 km from each other, and thesewere the conservation areas of that region, all locatedin the Federal District. The distances between theplots in Patrocínio-MG in Pratinha and those in For-mosa do Rio Preto in São Francisco were over 1,000km, showing the large gradient studied.

The data were obtained by the sampling of ten�20�50m� plots randomly placed at each study-site.A total of 150 sampled �20�50 m�-plots was includedin this study and 15 ha were measured.

All woody stems equal or greater than 5 cm of di-ameter at 0.30 m from ground level were measuredin each plot with a caliper, and if the stem had an ir-regular shape, two measures, at right angles to eachother, were taken and an average used. The plantswere identified at species level and their heights, con-sidered as the vertical projection from the top of thecrown to the soil, were measured. Lianas were notincluded in this study, especially because they are rarein the cerrado and do not reach the minimum diam-eter for sampling. The definition of individual plantsis sometimes difficult in the cerrado since many spe-cies can have vegetative reproduction from the rootsystem. However, most stems stand alone above theground, and the few cases with multiple or bifurcatedstems were registered in the forms and considered asa single individual when counting density but twostems for calculating basal area.

Vouchers were collected during the survey and inother excursions carried out during the dry and thewet seasons, and were deposited in the IBGEherbarium. The species list of those sites is published

Figure 1. Study area within the cerrado region in Central Brazil.

40

in Felfili et al. 1994, Mendonça et al. 1998 and Felfiliand Silva Júnior 2001.

Community composition and organization wereevaluated at each site. Shannon and Wiener diversityindex and Pielou’s evenness index were used toevaluate alpha diversity, which is related to the num-ber of species and the distribution of individuals perspecies in a community. The beta diversity, related tothe differences in species composition and abundancebetween sites �Kent and Coker 1992�, was evaluatedby similarity indices and also by multivariate analy-ses.

Shannon´s diversity index �nats.individual–1� variesfrom 0 to positive values, depending on the chosenlogarithmic scale, but rarely surpasses 5. The lesseven the distribution of the number of individuals perspecies the closer to zero the Pielou’s evenness index�Margurran 1988�.

Sørensen similarity index, based on presence-absence of the species and Czekanowski similarly in-dicates, also based on the number of individuals perspecies �Kent and Coker 1992�, were used for thecomparison between sites.

The complete data set was also classified byTWINSPAN and ordinated by DCA �Kent and Coker1992�. The variable density �number of trees� of spe-cies per hectare for each site was used for the con-struction of the matrix for the multivariate analyses,with cut levels of 0, 2, 5 and 10 for TWINSPANclassification.

Results

Species richness and alpha-diversity

The number of species per site varied from 55 in theBrasília National Park, in the Pratinha physiographicunit, to 97 in Serra Negra, in the Veadeiros. Thenumber of species was very similar at all the sites inSão Francisco, varying from 66 in Correntina-BA to68 in Formosa do Rio Preto-BA. This is probably areflection of the physical similarities between the sitesin the physiographic unit that contains just one landsystem. The number of species varied from 82 to 97in Veadeiros but a smaller range of variation was de-tected between Serra Negra �97� and Serra da Mesa�91�, both in the third land system of this land unit.However, in the first land system Alto Paraíso �92�and Vila Propicio �82� had a larger difference innumber of species. In Pratinha, the number of species

varied from 55 to 73 with most sites having c. 60species regardless of the two land systems, see Table2.

The number of species was generally higher inVeadeiros but diversity did not follow this pattern dueto the distribution of individuals per species, seePielou’s evenness index, Table 2. The distribution ofindividuals per species was sometimes less even insome of the richest sites, such as Serra Negra, SerraMesa and Alto Paraíso.

Community structure

Density per site varied from 628 individuals per ha inFormosa do Rio Preto-BA to 1348 in Silvânia-GO,with 80% of the sites presenting a density above 800.Basal area per site ranged from 5.89 m2ha–1 in Para-catu-MG, to 11.30 in Silvânia-MG. Around 80% ofthe sites presented basal area above seven, see Table2. Dead standing stems ranged from 3 to 10%, mostsites presenting 5%. The phytosociological structureof the vegetation is published �Felfili et al. 1994,1997, 2001�. The main pattern for species abundancein the communities, considering both density andbasal area, is of a small number of species, c. 10,comprising more than 50% of the density and basalarea in each site. The most abundant species in mostsites contained around 10% of the total values. Diam-eter distributions �Felfili and Silva Júnior 1988, Felfiliet al. 2001� showed that more than 80% of the indi-viduals at the communities are under 10 cm, with afew larger individuals reaching up to c. 50 cm.

Floristic similarity and beta diversity

Sørensen floristic similarity index, was high with allFigures above 0.5 between the sites in São Franciscoand also in Pratinha, Table 3. In the latter, the indiceswere high, even in the comparisons between sites indifferent land systems such as Paracatu�Brasília Na-tional Park. In the Veadeiros similarity was high be-tween sites in the same land system, such as SerraNegra�Serra da Mesa in one land system, and AltoParaíso�Vila Propício in another. However, similar-ity between sites in different land systems in thisphysiographic unit was not always high, especially inthe comparisons with Alto Paraíso.

Beta diversity was low at Pratinha and São Fran-cisco sites when presence and absence of species wascompared by Sørensen´s indices, but much higher foralmost all comparisons when the density of species

41

was introduced in the comparison by Czekanowski´sindex. Beta diversity is the reverse of these indices, ifsimilarity is high, beta diversity is low and vice-versa.

Classification and ordination

In the final TWINSPAN classification, the sites weregrouped by physiographic units, see Figure 2. The di-visions were strong with eigen-values near or above0.30 �Gauch 1982�. In the first division, the four SãoFrancisco sites were pooled together with the follow-ing Veadeiros sites: Alto Paraíso, Vila Propício, Serrada Mesa and Serra Negra. In the second division, VilaPropício was separated from the others while in thethird division Alto Paraíso, Serra da Mesa and SerraNegra formed a distinct group.

At the other side of the dichotomy the six sites ofPratinha and the Veadeiros National Park were placedtogether. In the second division, Patrocínio and Para-catu formed a group, while in the third division theconservation Areas in the Federal District andSilvânia were separated from the Veadeiros NationalPark.

Ordination by DECORANA, Figure 3, corrobo-rated the TWINSPAN classification results, indicatinga floristic and structural gradient following the envi-ronmental variation, especially with regard to thephysiographic units determined by Cochrane et al.1985. The first axis of the ordination shows a gradi-ent going from the cerrados on deep Latosols in Prat-inha through to those on sandy soils �Areia Quart-zoza� in São Francisco and ending at the shallowerCambisols of Veadeiros �see Brasil 1982; Cochrane etal. 1985; Haridasan et al. 1997; Haridasan 2001; fora description of the soil types in those regions�.

Discussion

Alpha diversity was high, and the number of woodyspecies per site �1-ha sampled per site� varied from55 to 97, which places the cerrado as one of the rich-est savanna physiognomies in the world.

Richness was at a similar level at all sites in SãoFrancisco, a large and uniform physiographic unit,where the most distant sites Correntina and Formosaare more than 500 km distant from each other. How-ever, it was most variable in the Veadeiros, a physi-ographic unit containing three land systems, wherethe most distant sites, Serra da Mesa and Vila Propí-cio were c. 200 km apart.Ta

ble

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42

Shannon´s indices indicated a highly diverse veg-etation with Figures around 3.5 nats.ind–1 in mostsites, similar to those found in gallery forests in Cen-tral Brazil and in Amazonian forests �Felfili 1995;Silva Júnior et al. 2001�. Only Silvânia and Paracatuin Pratinha had Shannon´s diversity indices below3.4, while in all the other sites these values variedfrom 3.44 to 3.73.

Community structure

Similarity of presence-absence of species at the sitesby Sørensen index, was high, therefore the beta di-

versity of the land systems was low, and even for landsystems within the same physiographic unit, as in SãoFrancisco and Pratinha. But it was high between sitesin different land systems in Veadeiros, see Table 3.

Sites in the same land system, although around 500km distant from each other, were similar, such asCorrentina and Formosa, while sites only 50 km dis-tant in different land systems, such as Alto Paraísoand Veadeiros National Park, had a low similarity bySørensen’s index. This indicates that the floristiccomposition of the sites conforms well with the land-system zoning. There was a large overlap of specieswithin the same physiographic unit especially if in the

Table 3. Similarity of the woody flora, including plants from 5 cm diameter at 0.30 m from the ground level, in 15 sites at three physi-ographic units, Pratinha , Veadeiros and São Francisco in Central Brazil �FOR – Formosa; SDE – São Desidério; COR – Correntina; PGS –Parna Grande Sertão; SNE – Serra Negra; SM – Serra da Messa; PVE – Parna Veadeiros; ALP – Alto Paraíso; VPR – Vila Propício; AGA– APA Gama; PBR – PARNA Brasília; AEM – ESEC Águas Emendadas; SIL – Silvânia; PAR – Paracatu; PAT – Patrocínio.PARNA�National Park; ESEC�Ecological Station; APA�Environmentally protected area�.

FOR SDE COR PGS SNE SM PVE ALP VPR AGA PBR AEM SIL PAR PAT

Formosa – 43.51 44.19 37.36 24.58 30.02 29.46 22.09 34.92 20.32 22.32 20.11 19.29 23.69 20.05São Desidério 0.61 – 47.58 46.75 34.91 40.56 37.14 28.19 42.73 31.53 30.82 33.91 32.93 21.67 16.87Correntina 0.61 0.66 – 33.70 24.12 31.22 29.66 24.72 31.54 26.24 28.65 25.82 24.79 21.44 14.57PARNA G. S. 0.59 0.57 0.65 – 38.26 40.53 37.80 29.36 59.85 31.58 38.48 42.93 43.56 26.19 31.09Serra Negra 0.49 0.52 0.50 0.56 – 56.30 32.37 31.00 52.54 31.42 27.61 29.30 35.97 30.69 23.75Serra Mesa 0.49 0.42 0.47 0.52 0.61 – 48.83 41.15 56.71 32.80 32.08 32.64 40.11 23.84 20.33PARNA Veadeiros 0.45 0.49 0.53 0.53 0.48 0.51 – 47.35 43.20 36.12 29.14 39.11 35.22 19.49 16.03Alto Paraíso 0.41 0.38 0.37 0.43 0.44 0.47 0.47 – 38.16 24.00 23.73 27.89 22.41 24.61 16.11Vila Propício 0.53 0.50 0.55 0.67 0.57 0.60 0.60 0.50 – 38.72 40.06 43.28 47.14 35.16 30.87APA Gama 0.45 0.51 0.50 0.57 0.45 0.52 0.61 0.47 0.59 – 59.94 55.27 43.87 22.05 35.55PARNA Brasília 0.40 0.46 0.46 0.56 0.39 0.46 0.57 0.46 0.58 0.83 – 55.31 45.57 26.10 33.16ESEC Águas Emendadas 0.41 0.50 0.46 0.53 0.50 0.51 0.59 0.53 0.61 0.73 0.73 – 49.90 23.19 29.63Silvânia 0.52 0.56 0.55 0.60 0.50 0.52 0.58 0.47 0.63 0.69 0.70 0.71 – 22.10 31.35Paracatu 0.43 0.45 0.45 0.52 0.49 0.47 0.46 0.47 0.56 0.52 0.53 0.57 0.59 – 25.11Patrocínio 0.42 0.38 0.39 0.46 0.34 0.40 0.45 0.39 0.48 0.59 0.56 0.58 0.52 0.51 –

not italic: Sørensen index �Qualitative, varies from 0 to 1�, bottom-left side; italic: Czekanowski index �Quantitative, varies from 0 to 100�,top-right side.

Figure 2. Classification by TWINSPAN of the cerrado sensu stricto at 15 study-sites in Central Brazil. �PARNA � National Park, APA �Environmentally protected site, ESEC � Ecological Station�.

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same land system, while the floristic distinctivenesswas built by a large number of species preferential toeach land system. Nine species were found to bewidespread throughout the three physiographic units.

The similarity indices between sites in the first landsystem of Pratinha were the highest with Sørensen’sindices above 0.7. The similarity indices between thesites in São Francisco were also high but of interme-diate values if compared to the lower indices foundbetween sites in the different land systems in Vead-eiros.

Veadeiros is a complex land unit having both flatand steep relief, infertile and shallow soils mingledwith limestone and quartz outcrops and some patchesof deep Latosols, while Pratinha mostly has on deepdystrophic Latosols and São Francisco has a predom-inantly plain relief with sandy soils �Areia Quartzoza�and Latosols to a much lesser extent. Cerrado is adominant vegetation type at Pratinha and São Fran-cisco while grassland dominates Veadeiros, with cer-rado occurring mostly on patches of Latosols.

Sørensen’s similarity indices were high when com-paring Grande Sertão Veredas National Park to 12 ofthe 15 sites. The other two National Parks, Brasíliaand Veadeiros had high similarity, respectively, withseven and eight of the 15 areas compared. The Eco-logical Stations of Águas Emendadas and APA Gamae Cabeça de Veado in Brasília also had a high simi-larity, with 12 of the 15 sites compared, in spite oftheir small size in relation to the National Parks.

The five conservation areas had a high similaritybetween themselves, which is positive for the conser-

vation of the species populations. A network of con-servation areas containing overlapping speciesthroughout the cerrado is especially important sinceDNA studies have shown that breeding happens atshort distances, even for widespread species in thecerrado biome �Collevatti et al. 2001�.

Felfili and Silva Júnior �1993�, comparing the sixsites of Chapada Pratinha, suggested that the densityof species was one of the most important factors dif-ferentiating the cerrado sites. This study of 15 sitesconfirms their finding, suggesting that density of spe-cies should be an important parameter for decisionmaking on selecting sites to protect cerrado commu-nities. A species may occur in a conservation unit butmay not be protected on a long term basis if it occursat a low density and is isolated from other popula-tions.

Czekanowski similarity indices were lower thanSørensen´s in the comparisons, suggesting a highstructural differentiation between the sites and there-fore a high beta-diversity. There is a large overlap inspecies occurrence in the sites but the size of theirpopulations are very different at each site, reinforcingthe urgent need for new conservation areas in theCerrado biome.

Only nine species were common to all sites, Acos-mium dasycarpum, Aspidosperma tomentosum,Bowdichia virgilioides, Byrsonima coccolobaefolia,Erythroxylum suberosum, Kielmeyera coriacea, Ou-ratea hexasperma, Qualea grandiflora and Tabebuiaochracea. These species are widespread in the Cer-rado biome, being listed amongst the 38 most widely

Figure 3. Positioning of the 15 cerrado sensu stricto study-sites in the DECORANA ordination axes �PARNA � National Park, ESEC �Ecological Station and APA � Environmentally Protected Area�.

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distributed species by Ratter et al. 2003 who havecompared rapid surveys and published lists for 376areas of cerrado sensu lato within the region. The lownumber of species common to all sites, in spite of thehigh floristic similarity for c. 60% of the comparisonsbetween the sites, confirms previous findings �Ratterand Dargie 1992; Felfili and Silva Júnior 1993� thatspecies distribution is mosaic.

TWINSPAN classification of the cerrado at thethird level of the divisions agreed completely withland system zoning Cochrane et al. �1985� for thecerrado in physiographic units. The subdivision ofthose units in land system perfectly matched the flo-ristic and structural classification produced by TWIN-SPAN for Pratinha and São Francisco. Only one landsystem �Alto Paraíso-Vila Propício� of the complexVeadeiros physiographic unit did not fit well into theclassification of Cochrane et al. �1985�.

DECORANA ordination suggested a higher betadiversity between the sites of Veadeiros, especiallycomparing Alto Paraíso, Serra Mesa and Serra Negrawith the others. This is due to the complex landscapeof that area, with a wider range of soil types �Hari-dasan et al. 1997� and relief variation �BRASIL 1982,Table 1�.

Grande Sertão Veredas National Park was at an in-termediate position along the first axis of DECO-RANA ordination with Brasília National Park, and theother conservation areas in Pratinha at one extremeand Veadeiros at the opposite side. This agrees withthe similarity indices that showed Grande Sertão asthe site with a higher similarity with other sites.

Brasilia National Park and the other conservationareas of Pratinha represented well the land system inwhich they were placed, being characterized by high-lands with an altitude of around 1000 m �Felfili andSilva Júnior 1993�.

Veadeiros National Park occupied a land systemdistinct from the other sites in Veadeiros physio-graphic unit, suggesting the need to create a networkof conservation areas and corridors to ensure the con-servation of that variable landscape.

Grande Sertão Veredas showed a high similarity toall sites in São Francisco, being very representativeof that region. The creation of other smaller conser-vation areas and good management in the regioncould ensure the protection of these north-easterncerrados.

Conclusions

Cerrado is a rich and diverse physiognomy with ahigh beta-diversity, especially due to the variation indensity of species between sites. Even when there isa large number of common species between sites, thesize of the species populations is very variable fromplace to place.

The floristic and structural variations conformedwell with the variation in climate, landscape and soildevised in the land-system zoning proposed by Co-chrane et al. �1985� for the Cerrado biome. Land-sys-tem zoning should be used as a tool to set upconservation strategies at a regional level to maxi-mize the protection of the biodiversity.

The most critical sites for preservation due to theirlow similarity to other sites were Alto Paraíso deGoiás, Serra da Mesa e Serra Negra in Veadeiros andParacatu and Patrocínio in the Pratinha physiographicunit.

Acknowledgements

We are grateful to the Ministry of Environment-FNMA �Projeto Biogeografia do bioma Cerrado�, toDFID �Conservation and Management of the Biodi-versity of the Brazilian Cerrado Biome Project� andto the Brazilian Research Council �CNPq� for supportto this project, and also to the field and herbariumteams �especially Roberta Mendonça, Newton Rod-rigues and Edson Cardoso�.

References

Allaby M. 0000. The concise oxford dictionary of botany. OxfordUniversity Press, Oxford, UK. 442 p.

Brasil, Ministério de Minas e Energia, Secretaria Geral. �ed.� 1982.Projeto RADAMBRASIL. Folha SD. 23. Brasília. Geologia,geomorfologia, pedologia, vegetação e uso potencial da terra.IBGE, Rio de Janeiro, Brazil. 660 p. �Levantamento de Recur-sos Naturais, 29�.

Castro A.A.J.F. 1994. Comparação florístico-geográfica (Brasil) efitossociológica (Piauí-São Paulo) de amostras de cerrado.UNICAMP, Campinas, Brazil. 520 p. �Doctorate Thesis�.

Cochrane T.T., Sanchez L.G., Azevedo L.G., Porras J.A. andGarver C.L. 1985. Land in tropical america. CIAT/EMBRAPA-CPAC, Cali. 3 vols.

Collevatti R.G., Grattapaglia D. and Hay J.D. 2001. High resolu-tion microsatellite based analysis of the mating system allowsthe detection of significant biparental inbreeding in Caryocarbrasiliense, an endangered tropical tree species. Heredity 86�1�:60–67.

45

Durigan G., Ratter J.A., Bridgwater S., Siqueira M.F. and FrancoA.D.C. 2003. Padrões fitogeográficos do cerrado paulista sobuma perspectiva regional. Hoehnea 30�1�: 39–51.

Eiten G. 1972. The cerrado vegetation of Brazil. Botanical Review38: 201–341.

Felfili J.M. and Silva Júnior M.C. 1988. Distribuição dos diãmetrosnuma faixa de cerrado na Fazenda Água Limpa �FAL� EMBrasília-DF. Acta Botânica Brasílica 2: 85–104.

Felfili J. M. and Silva Júnior. M.C. 1993. A comparative study ofcerrado �sensu stricto� vegetation in Central Brazil. Journal ofTropical Ecology 9: 277–289.

Felfili J.M., Filgueiras T.S., Haridasan M., Silva Júnior M.C.,Mendonça R. and Rezende A.V. 1994. Projeto biogeografia dobioma cerrado: Vegetação e solos. Cadernos de geociências doIBGE 12: 75–166.

Felfili J.M. 1995. Diversity, structure and dynamics of a galleryforest in central Brazil. Vegetatio 117: 1–15.

Felfili J.M., Silva Júnior M.C., Rezende A.V., Nogueira P.E., WalterB.W.T., Silva M.A. and Encinas J.I. 1997. Comparação florísticae fitossociológica do cerrado nas Chapadas Pratinha e dos Vead-eiros.. In: Leite L. and Saito C. �eds�, Contribuição ao conheci-mento ecológico do cerrado. UnB, Brasília-DF. pp. 6-11.

Felfili J.M. 1998. Águas Emendadas no contexto fitogeográfico docerrado.. In: Seminário sobre unidades de conservação. ,SEMATEC, Brasília-DF. pp. 71-87.

Felfili J.M. and Silva Júnior M.C. , �ed.� 2001. Biogeografia doBioma Cerrado. Estudo fitofisionômico na Chapada do espigãoMestre do São Francisco. UnB, Brasília-DF. 152 p.

Gauch H.G. 1982. Multivariate analysis in community ecology.Cambridge University Press, Cambridge, UK. 298 p.

Haridasan M., Silva Júnior M.C., Felfili J.M., Rezende A.V., SilvaP.E.N. 1997. Gradient analysis of soil properties and phytoso-ciological parameters of some gallery forest the Chapada dosVeadeiros.. In: Preceedings of the international symposium onassessment and monitoring of forests in tropical dry regions.UnB, Brasília- DF. pp. 259-275.

Haridasan M. 2001. Solos.. In: Felfili J.M. and Silva Júnior M.C.�ed.�, Biogeografia do Bioma Cerrado. Estudo fitofisionômico naChapada do espigão Mestre do São Francisco. UnB, Brasília,DF. 152p. pp. 12-17.

Kent M. and Coker P. 1992. Vegetation description and analysis.Belhaven Press, London, UK.

Klink C.A, Macedo R.H. and Mueller C.C. 1995. De grão em grãoo cerrado perde espaço. Cerrado – Impactos do processo de ocu-pação. WWF/PRÓ-CER, Brasília. 66 p.

Margurran A.E. 1988. Ecological diversity and its measurement.Chapman and Hall, London, UK. 179 p.

Mendonça R., Felfili J.M., Walter B.M.T., Silva Júnior M.C.,Rezende A.V., Filgueiras T.S. and Nogueira P.E. 1998. Floravascular do Cerrado.. In: Sano S. and Almeida S. �eds�, Cerrado:ambiente e flora. EMBRAPA-CPAC, Planaltina-DF. pp. 287-556.

Mittermeier R.A., Myers N. and Mittermeier C.G. 1999. Hotspots:Earth´s biologically richest and most endangered terrestrialecoregions. Conservation International-CEMEX, New York.,USA, 430 p.

Oliveira-Filho A.T. and Ratter J.A. 1995. A study of the origin ofcentral brazilian forests by the analysis of plant species distribu-tion patterns. Edinburgh Journal of Botany 52 �2�: 141–194.

Ratter J.A. and Dargie T.C.D. 1992. An analysis of the floristiccomposition of 26 cerrado areas in Brazil. Edinburgh Journal ofBotany 49: 235–250.

Ratter J.A., Bridgewater S., Atinkinson R. and Ribeiro J.F. 1996.Analysis of the floristic composition of the Brazilian cerradovegetation II: Comparison of the wood vegetation of 98 areas.Edinburgh Journal of Botany 53: 153–180.

Ratter J.A., Bridgewater S., Ribeiro J.F., Dias T.A. B. and SilvaM.A. 2000. Estudo preliminar da distribuição das espécies len-hosas da fitofisionomia cerrado sentido restrito nos estados com-preendidos pelo bioma cerrado. Boletim do Herbário EzechiasPaulo Heringer 5: 5–43.

Ratter J.A., Bridgewater S. and Ribeiro J.F. 2003. Analysis of thefloristic composition of the Brazilian cerrado vegetation III:Comparison of the wood vegetation of 376 areas. EdinburghJournal of Botany 60: 57–109.

Ribeiro J.F. and Walter B.M.T. 1998. Fitofisionomias do BiomaCerrado.. In: Sano S. and Almeida S. �eds�, Cerrado: ambientee flora. EMBRAPA-CPAC, Planaltina. pp. 89-166.

Silva Júnior M.C., Felfili J.M., Walter B.M.T., Nogueira P.E.,Rezende A.V., Morais R.O. and Nóbrega M.G.G. 2001. Análiseda flora arbórea de Matas de Galeria no Distrito Federal: 21 le-vantamentos.. In: Ribeiro J.F. �Eds.�. Cerrado Caracterização eRecuperação de Matas de Galeria. EMBRAPA-CPAC, Planal-tina-DF. pp. 143-194.

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