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Brazilian Journal of Biology

Print version ISSN 1519-6984On-line version ISSN 1678-4375

Braz. J. Biol. vol.76 no.2 São Carlos Apr./June 2016  Epub Mar 01, 2016

https://doi.org/10.1590/1519-6984.13814 

Articles

What are the most important factors determining different vegetation types in the Chapada Diamantina, Brazil?

Quais são os fatores mais importantes na determinação de diferentes tipos de vegetação na Chapada Diamantina, Brasil?

S. P. S. Nevesa  * 

R. Funchb 

A. A. Conceiçãoa 

L. A. P. Mirandac 

L. S. Funcha 

aPrograma de Pós-graduação em Botânica, Departamento de Ciências Biológicas, Universidade Estadual de Feira de Santana – UEFS, Av. Transnordestina, s/n, Novo Horizonte, CEP 44036-900, Feira de Santana, BA, Brazil

bFundação Chapada Diamantina, Rua Pé da Ladeira, 212, CEP 46960-000, Lençóis, BA, Brazil

cDepartamento de Ciências Biológicas, Universidade Estadual de Feira de Santana – UEFS, Av. Transnordestina, s/n, Novo Horizonte, CEP 44036-900, Feira de Santana, BA, Brazil


Abstract

A transect was used to examine the environmental and biological descriptors of a compact vegetation mosaic in the Chapada Diamantina in northeastern Brazil, including the floristic composition, spectrum of plant life forms, rainfall, and soil properties that defined areas of cerrado (Brazilian savanna), caatinga (seasonally dry tropical forest thorny, deciduous shrub/arboreal vegetation) and cerrado-caatinga transition vegetation. The floristic survey was made monthly from April/2009 to March/2012. A dendrogram of similarity was generated using the Jaccard Index based on a matrix of the species that occurred in at least two of the vegetation types examined. The proportions of life forms in each vegetation type were compared using the chi-square test. Composite soil samples were analyzed by simple variance (ANOVA) to examine relationships between soil parameters of each vegetation type and the transition area. The monthly precipitation levels in each vegetation type were measured and compared using the chi-square test. A total of 323 species of angiosperms were collected distributed in 193 genera and 54 families. The dendrogram demonstrated strong difference between the floristic compositions of the cerrado and caatinga, sharing 2% similarity. The chi-square test did not demonstrate any significant statistical differences between the monthly values of recorded rainfall. The organic matter and clay contents of the soilsin the caatinga increased while sand decreased, and the proportions of therophyte, hemicryptophyte, and chamaephyte life forms decreased and phanerophytes increased. We can therefore conclude that the floristic composition and the spectrum of life forms combined to define the cerrado and caatinga vegetation along the transect examined, with soil being the principal conditioning factor determining the different vegetation types, independent of precipitation levels.

Keywords:  Savanna; seasonally dry tropical forest; biological spectrum; soil; rainfall

Resumo

Foi estabelecida uma transecção para examinar descritores ambientais e biológicos em uma área compacta de vegetação em mosaico na Chapada Diamantina, Nordeste do Brasil. A composição florística, espectro de formas de vida, precipitação e propriedades do solo foram avaliadas na transecção entre cerrado (savana brasileira) e caatinga (floresta tropical sazonalmente seca espinhosa, vegetação arbustivo-arbórea decídua), separados por vegetação de transição cerrado-caatinga. O levantamento florístico foi realizado mensalmente de abril de 2009 a março de 2012. Foi feita análise de agrupamento a fim de determinar a similaridade entre as fisionomias de cerrado, a caatinga e a transição cerrado-caatinga. As proporções de formas de vida foram comparadas utilizando o teste qui-quadrado. Amostras compostas de solo foram analisadas por variância simples (ANOVA) testando a existência de diferenças entre os solos de cada tipo de vegetação. A precipitação mensal em cada tipo de vegetação foi mensurada e os resultados comparados com o teste qui-quadrado. Coletamos 323 espécies de angiospermas pertencentes a 193 gêneros e 54 famílias. A análise de agrupamento demostrou diferença entre a composição florística do cerrado e da caatinga, com apenas 2% de similaridade. O teste qui-quadrado não demonstrou diferença estatística entre os valores registrados para cada mês. À medida que os conteúdos de matéria orgânica e argila aumentaram e o de areia diminuiu na caatinga, a proporção das formas de vida terófito, hemicriptófito e caméfito diminuiu e a de fanerófitos aumentou. Podemos considerar que a composição florística e o espectro de formas de vida delimitaram o cerrado e a caatinga na transecção estudada e que o solo foi o principal fator condicionante para determinação dos diferentes tipos de vegetação, independentemente da precipitação.

Palavras-chave:  Savana; floresta tropical sazonalmente seca; espectro biológico; solo; precipitação

1 Introduction

The Chapada Diamantina, the northern extension of the Serra of the Espinhaço Range in Brazil, is included within the Caatinga biome of northeastern Brazil and hosts a large diversity of vegetation types associated with its physiographic spectrum, including cerrado (neotropical savanna), campo rupestre (open, rocky field grasslands), forest, and caatinga (seasonally dry tropical forest thorny, deciduous shrub/arboreal vegetation) (Harley, 1995; Juncá et al., 2005).

A number of floristic studies have already been undertaken in the Chapada Diamantina, including those considering: campo rupestre (Conceição et al., 2007; Neves and Conceição, 2010), cerrado (Costa et al., 2009), caatinga (Lima and Lima, 1998), submontane seasonal semi-deciduous forest, upper montane forest, and gallery forest (Funch et al., 2008; Nascimento et al., 2010; Couto et al., 2011). Rapid ecological surveys of diverse vegetation types have also been undertaken (Harley et al., 2005; Juncá et al., 2005; Funch et al., 2009), although most of the studies considered only be shrub-arboreal layer and/or made collections on irregular schedules.

Funch et al. (2009) published a vegetation map of the Chapada Diamantina National Park and the surrounding region describing vegetation types, based on Harley (1995), and associated with different soil types distributed contiguously within a relatively limited area. In light of the intriguingly close dispositions of different vegetation types under apparently identical macro-climatic conditions, we investigated cerrado and caatinga vegetation sites and their ecotone along an E-W transect just west of the National Park to contribute to our understanding of the phytogeography of Brazil.

Our study focused on the floristic compositions of the herbaceous-subshrub and shrub-arboreal layers, life forms, rainfall, and soil properties in addressing the question of which factors are most important in defining the cerrado and caatinga vegetation along this transect.

We hypothesized that variations in the physiognomy, floristic compositions, and structures of natural ecosystems could occur due to variations in the chemical and physical properties of their soils (Haridasan, 2000; Amorim and Batalha, 2006; Juhász et al., 2007; Silva et al., 2013), and that permeability and the proportions of fine particles would be the parameters most closely associated with species richness (Medinski et al., 2010). We also considered that: 1) climatic factors have a decisive role in determining species and plant community distributions (Terradas, 2001; Lavers and Field, 2006); 2) water availability is one of the principal determinants of vegetation cover (Silva and Batalha, 2008; Cianciaruso and Batalha, 2009), and; 3) life forms are important elements in vegetation descriptions (Raunkiaer, 1934).

2 Material and Methods

2.1 Study area

The Chapada Diamantina National Park has a much accentuated topography of low mountains with rocky peaks and steep slopes, narrow valleys with extensive plains to the west that are separated by long chains of hog-back mountains. Altitudes in the region vary from 400 to more than 2,000 m a.s.l. The isolated peaks and extended mountain chains have extensive rock outcrops and litholic neosols (thin, rocky soils of low fertility), while soils on the intermediary plains are predominantly latosols (deep, well-drained, acidic, with low fertility) (Juncá et al., 2005).

The regional climate is mesothermic (Cwb) (Köppen, 1948), with average monthly temperatures varying between 18 and 25 °C, with the lowest temperatures occurring during driest months. The average annual precipitation is 1,218 mm (for the period between 1951-2011). The rainy season usually begins in November and extends until March, while the dry season initiates in June and continues until October (Juncá et al., 2005).

The present study was conducted in two vegetation types: cerrado, represented by three local physiognomies (two sites of open cerrado – OP, rocky outcrop cerrado – ROC, and typical cerrado – TC) and caatinga (CAA). The cerrado-caatinga transition (CCT) between of these both vegetation types also was sampled (Table 1). These vegetation are contiguously distributed along a 10 Km E-W transect sampled (12º 26’ 05” S to 12º 27’ 7” S to 41º 31’ 05” W to 41º 35’ 52” W) just west of the Chapada Diamantina National Park in the Chapada Diamantina, Bahia State, Brazil. The phytophysiognomies of cerrado are included in cerrado stricto sensu (Ribeiro and Walter, 2008). Caatinga is characterized by a predominance of low shrubs and trees that are highly branched, often spiny, and deciduous in the dry season (Pennington et al., 2009).

Table 1 Descriptions of vegetation types physiognomies and vegetation types in the mosaic parameters cerrado-caatinga, Chapada Diamantina, NE Brazil. 

Species more important
Families more rich in species Density
(individuals tree-shrub/ha)
Herbaceous-subshrub IV(%) Shrub-arboreal IVI(%)
Total numbers of species Numbers of exclusive species Numbers of families (Number of genera) Herbaceous-subshrub /shrub-Arboreal Monocotyledons
Dicotyledons
Species
Nº %
cerrado ralo (site I)
12º 26’ 18” S
41º 30’ 53” W
Alt.: 857 m
101 13 (12.87%) 25 (75) 9.10 19 (19%)
82 (81%)
Poaceae
Asteraceae
Fabaceae
Rubiaceae
Euphorbicaeae
14
12
12
8
6
14
12
12
8
6
2.380 Trachypogon spicatus
Bulbostylis fasciculata
Pavonia cancelata
Calea harleyi
Jacquemontia evolvuloides
54 Annona coriacea
Eugenia punicifolia
Duguetia furfuracea
Lippia microphylla Stryphnodendron sp.
98
open cerrado ralo (siteII)
12º 26’ 05” S
41º 31’ 05” W
Alt.: 889 m
93 5 (5.37%) 31 (72) 3.20 22 (24%)
71 (76%)
Poaceae Fabaceae
Rubiaceae Cyperaceae Lamiaceae
15
12
6
5
5
16 13
6,5
5,5 5,5
2.160 Trachypogon spicatusStigmaphyllom paraliasAxonopus polydactylus
Ruellia incompta
Bulbostylis fasciculata
43 Croton velutinus
Annona coriacea
Agarista cf. chapadensis
Hyptis macranta
Eugenia punicifolia
64
rocky outcrop cerrado
12º 26’ 08” S
41º 31’ 04” W
Alt.: 884 m
105 17 (16.19%) 33 (84) 2.82 15 (14%)
90 (86%)
Fabaceae Asteraceae Poaceae Rubiaceae Cyperaceae 14
9
9
7
5
13,5
9
9
7
5
4.960 Trachypogon spicatus
Echinolaena inflexa
Axonopus polydactilus
Stigmaphyllom paraliasSebastiania corniculata
31 Croton velutinus
Annona coriacea
Lippia microphylla
Vochysia thyrsoidea
Marcetia taxifolia
53
typicalcerrado
12º 26’ 16” S e 41º 31’ 11” W
Alt.: 843 m
119 25 (20.66%) 32 (85) 3.95 25 (21%)
94 (79%)
Poaceae Fabaceae Asteraceae Malvaceae Euphorbiaceae 20
19
8
7
6
17
19
7
6
5
5.250 Trachypogon spicatus
Stylosanthes scabra
Pavonia cancellataStigmaphyllom paralias
Mimosa somnians
46 Annona coriacea
Dalbergia miscolobium
Cordia rufescens
Solanum stipulaceum
Croton heliotropiifolius
84
cerrado-caatinga transition
12º 26’ 34” S e 41º 32’ 01” W
Alt.: 736 m
94 23 (24.46) 30 (69) 2.44 25 (26%)
69 (74%)
Poaceae Fabaceae Malvaceae Myrtaceae ConvolvulaceaeEuphorbiaceae Rubiaceae 16
10
7
6
5
5
5
17
11
7,5
6,5
5,5
5,5
5,5
3.680 Hohenbergia cf. caatingae
Panicum sp
Streptostachys robusta
Aristida sp.
Billbergia porteana
29 Senegalia riparia
Croton adamantinus
Bignoniaceae
Croton heliotropiifoliusActinostemon concolor
65
caatinga
12º 27’ 07” S e 41º 35’ 58” W
Alt.: 697 m
117 102 (87.17%) (31) 77 3.03 21 (18%)
96 (82%)
Fabaceae Bignoniaceae Euphorbiaceae Malpighiaceae Cactaceae 16
15
13
8
7
15
13
11
7
6
6.520 Neoglaziovia variegata
Poaceae
Tacinga werneri
Cryptanthussp.
Tacinga funalis
52 Senegalia tenuifolia
Maprounea guianensis
Actinostemon sp.
Erythroxylum oxypetalum
Dalbergia cearensis
38

2.2 Floristic composition and life forms

The vegetation types were selected along a 10 Km east-west transect based on the vegetation map of Funch et al. (2009) and identified in the field as distinct areas of well-conserved cerrado and caatinga. We established ninety 100 m2 plots (15in each of the four vegetation types examined: the two types of cerrado, the cerrado/caatinga transition, and caatinga) for a total of 0.9 ha. Plant collections were performed monthly in all sites from April/2009 to March/2012 of both fertile and sterile plant material of herbaceous-subshrub and shrub-arboreal species (this period included three rainy seasons and three dry seasons).The voucher specimens were deposited in the Herbarium of the State University at Feira of Santana (HUEFS) and identified with the aid of specialists and by comparison with previously identified material. The classifications of the plant families followed the proposal of the APG III (2009).

The life forms of the specimens were classified according to Raunkiaer (1934), excluding epiphytes and considering succulents separately from phanerophytes. The proportions of herbaceous-subshrub/shrub-arboreal (epiphytes, succulents, chamaephytes, hemicryptophytes, and therophytes /phanerophytes) species were calculated for each of the vegetation types, for cerrado-caatinga transition, and for the different cerrado physiognomies. The importance values (IV) of the herbaceous-subshrub species (the sum of coverage and relative frequency) and shrub-arboreal layer (the sum of the density, dominance and relative frequency) were used to identify the five most important species of each layer in each vegetation type (Neves, 2013).

2.3 Rain fall and soils properties

Rainfall gauges were installed in each vegetation type and in the ecotone. Monthly precipitations levels were measured between January/2010 and March/2012 (Figure 1). Three composite soil samples (0-20 cm) were collected in each vegetation type and in the ecotone. Each composite sample was taken by mixing soils from three random points, totaling 18 composite samples (12 in the cerrado, three in the caatinga and three in the cerrado-caatinga transition). The physical and chemical properties of these samples were analyzed in an agricultural laboratory (Embrapa Semi-Árido - PE), according Donagema et al. (2011).

Figure 1 Rainfall (mm) in caatinga, cerrado, and their transition vegetation, Chapada Diamantina, NE Brazil. 

2.4 Statistical analyses

To determine the floristic similarities between the cerrado, caatinga and ecotone sites (considering the different cerrado physiognomies), clustering analysis was performed based on their Jaccard similarity (JS) as derived from a binary matrix, using the Unweighted Pair Group Method with Arithmetic Mean (UPGMA) run on Past software (Hammer et al., 2001). The cut-off used in this study was 25% following Mueller-Dombois and Ellenberg (1974). We compared the proportions of species in each life form class among the different vegetation types and the three cerrado physiognomies using the chi-square test (ᵡ2). The average values and standard errors of each physical and chemical component of the soils in the different vegetation types were calculated and simple analysis of variance (ANOVA) was performed to detect statistical differences between them, utilizing Sisvar software (Ferreira, 2000). The Tukey test (P<0.05) was used to compare the observed averages. The rainfall indices in the vegetation areas were compared using the chi-square test (2).

3 Results

3.1 Floristic composition

The numbers of species, genera and families, as well as the proportions of monocotyledons and eudicotyledon species, and the ratio of herbaceous-subshrub/shrub-arboreal species of cerrado phytophysiognomies, caatinga vegetation and cerrado-caatinga transition are presented in Table 1. A total of 323 species of angiosperms belonging to 193 genera and 54 families were encountered (Appendix), with 253 species (78%) occurring in only one of the vegetation types, 68 species (21%) in two types, and only four species (1%; Casearia sylvestris [Salicaceae], Croton heliotropiifolius [Euphorbiaceae], Panicum sp. [Poaceae] and Syagrus sp. [Arecaceae]) occurring in all sampled vegetation. Of the 325 of species, 260 species (80%) were eudicotyledons and 65 (20%) monocotyledons. The cerrado vegetation type showed the greatest richness (190 spp.; 58%) (Appendix). The cerrado and cerrado-caatinga transition shared 57 (18%) species belonging to 20 families (37%) and 43 genera (22%). The cerrado-caatinga transition and the caatinga shared 15 (5%) species belonging to 10 (19%) families and 15 (8%) genera. The families more rich in species are in the Table 1.

3.2 Life forms

The floristic spectra of life forms in the sampled vegetation are presented in Figure 2. Chi-Square tests indicated significant differences in the percentages of chamaephytes, therophytes and phanerophytes species between the caatinga and the cerrado, and between the caatinga and the cerrado-caatinga transition areas. Significant statistical differences (using the same test) were also noted between the cerrado (except the rocky outcrop cerrado) and caatinga, and between the cerrado and the cerrado-caatinga transition for the hemicryptophyte.

Figure 2 Floristic life form spectra in the different vegetation type, Chapada Diamantina, NE Brazil. Ch: chamaephyte; H: hemicryptophyte; Ph: phanerophyte; Th: therophyte; Cr: cryptophyte; Su: “succulent”; Ep: epiphyte; OPI = open cerrado, area I; OP II = open cerrado, area II; ROC = rocky outcrop cerrado; TC = typical cerrado; CCT = cerrado-caatinga transition; and CAA = caatinga. 

3.3 Similarities

The dendrogram demonstrated strong difference between the floristic compositions of the cerrado and caatinga, sharing 2% of similarity (Figure 3). Caatinga is the most distinct physiognomy and the cerrado-caatinga transition area is more similar with cerrado (24%). The cerrado physiognomies form a coherent group with 39% similarity.

Figure 3 Cluster of floristic similarity (UPGMA) in the different vegetation type, Chapada Diamantina, NE Brazil. OPI = open cerrado, area I; OP II = open cerrado, area II; ROC = rocky outcrop cerrado; TC = typical cerrado; CCT = cerrado-caatinga transition; and CAA = caatinga. 

3.4 Rainfall and soil properties

The chi-square test did not demonstrate any significant statistical differences between the monthly values of recorded rainfall. The average values and standard errors of the physical-chemical parameters of the soils of each vegetation type and the different cerrado phytophysiognomies are presented in Table 2. The cerrado soil was predominately sandy, while cerrado-caatinga transition and caatinga areas were sandy clay loams. The soils in all of the vegetation types were dystrophic (base saturation < 50%), acidic (pH < 5, except in the open cerrado, site I) and had relatively low Al contents (Al < 1.3 cmol/dm3, except in the cerrado-caatinga transition) and with exchangeable retention capacities less than 13, with high H+ e Al+3 loads.

Table 2 Averages and standard deviations of the physical and chemical characteristics of the soils in the different vegetation type areas, Chapada Diamantina, NE Brazil. 

OPI OPII ROC TC CCT CAA F
Physical characteristics
DP (Kg/dm–3) 2.58±0.02a 2.59±0.02a 2.58±0.005a 2.55±0.02a 2.55±0.04a 2.58±0.05a 0.90
DS (Kg/dm–3) 1.34±0.02a 1.45±0.05ab 1.40±0.08ab 1.46±0.02b 1.39±0.02ab 1.45±0.017ab 3.43
Porosity (%) 47.99±0.44a 44.01±2.69 ab 44.48±2.22 ab 42.54±1.35b 45.44±0.44 ab 43.68±1.25 ab 3.97
Total sand (%) 88.66±1.14a 93.93±22.84a 91.73±3.21a 91.28±2.89a 71.87±4.07b 67.07±3.91b 42.81
Silt (%) 7.51±1.06a 3.33±1.61a 6.89±2.10a 4.58±2.99a 7.11±2.17a 12.51±7.69a 2.20
Clay (%) 3.82±9.51a 2.74±1.80a 1.37±1.12a 4.13±4.03a 21.01±2.1b 20.42±3.41b 37.20
Chemical characteristics
MO (g/kg) 4.45±1.64a 3.51±2.31a 4.51±1.90a 7.08±1.36ab 15.59±8.34b 8.65±0.24ab 4.36
pH 5.2±0.10a 4.73±0.05bc 4.83±0.05b 4.36±0.11d 4.00±0.1e 4.56±0.05cd 70.29
P (mg/dm3) 0.62±0.11ab 0.52±0.07a 0.87±0.11b 1.24±0.11c 0.77±0.11ab 0.70±0.19b 12.15
Al (cmol/dm3) 0.43±0.07a 0.48±0.07a 0.4±0.08a 0.77±0.15b 1.32±0.07c 0.50±0.10ab 38.12
K (cmol/dm3) 0.05±0.02a 0.05±0.002a 0.03±0.005a 0.04±0.005a 0.12±0.04b 0.12±0.004b 19.01
Ca (cmol/dm3) 0.35±0.05a 0.51±0.13a 0.37±0.05a 0.47±0.15a 0.64±0.14ab 0.92±0.17b 7.96
Mg (cmol/dm3) 0.78±0.02a 0.36±0.24a 0.80±0.68a 0.56±0.18a 0.80±0.10a 0.82±0.42a 2.18
Na (cmol/dm3) 0.01±0.005ab 0.01±0a 0.01±0a 0.01±0.005ab 0.02±0.005b 0.01±0.00a 4.80
Potential acidity (cmolc/dm3) 11.6±1.19b 4.78±1.86a 9.29±1.99b 4.40±1.65a 5.17±0.34a 4.67±1.27a 12.44
Sum of bases (cmolc/dm3) 1.19±0.04ab 0.90±0.15a 0.87±0.12a 1.08±0.14a 1.58±0.07bc 1.87±0.26c 21.19
CEC (cmolc/dm3) 12.80±1.14c 6.01±1.55ab 10.16±2.03bc 5.48±1.75a 6.74±0.41ab 6.55±1.53ab 10.84
Base saturation (%) 9.66±1.15ab 16.67±5.03bc 9.00±1.73a 19.00±2.64c 23.67±0.57cd 29.00±2.64d 25.05

Different letters in the same lines indicate significant differences between the vegetation types (ANOVA, P<0.05). Densities of particles (DP), soil density (DS), porosity and proportions of total sand, silt, and clay and the chemical characteristics (organic matter (MO), phosphorus (P), aluminum (Al), potassium (K), calcium (Ca), magnesium (Mg), sodium (Na), Potential acidity, sum of bases (SB), cationic exchange capacity (CEC), and base saturation. Chapada Diamantina, NE Brazil. OPI = open cerrado I, OP II = open cerrado II, ROC = rocky outcroup cerrado, TC = typical cerrado, CCT = cerrado-caatinga transition, and CAA = caatinga. F values from ANOVA.

4 Discussion

The cerrado and caatinga vegetation sites as well as their transition, were well delimited in terms of their floristic compositions, life form spectra and soil properties. The cerrado vegetation demonstrated plant communities with ample varieties of three-dimensional structures and spatial heterogeneities, as reflected in the three different phytophysiognomies identified. Pennington et al. (2009) noted that cerrado vegetation physiognomies can be found under the same (or slightly more humid) climatic conditions as seasonal tropical dry forests, and that these vegetation types are often be found relatively close to one another.

Changes in the floristic compositions of the cerrado, cerrado-caatinga transition, and caatinga were accompanied by changes in the proportions of total sand, clay, and organic matter of their soils. Although high species-richness is often found in ecotone areas due to their greater environmental heterogeneity (Rodrigues and Nave, 2000), the transition zone studied here was less richness than the cerrado and caatinga surveyed. It is probable that the environmental conditions in that area are more rigorous and limit the occurrence of larger numbers of species (Mota et al., 2011).

The cerrado-caatinga transition area shared a greater percentage of species with the cerrado (approximately 70%) than with the caatinga (approximately 16%), indicating a high floristic similarity between them. More than 50% of the life forms shared between these two sites were therophytes –thus making essentially insignificant contributions to the overall differences in physiognomies between those vegetation types. Approximately 60% of the species shared by these two vegetation types were phanerophytes, which demonstrated deciduousness as a strategy to reduce water losses during the dry period (Neves, 2013).

The best represented life-form in the cerrado vegetation studied here was that of therophytes – a situation different from other cerrado sensu stricto areas that generally show high proportions of phanerophytes (Batalha and Martins, 2004; Costa et al., 2004). Phanerophytes were the most representative life forms in the caatinga vegetation studied – which was also different from the findings of Costa et al. (2007) in shrub caatinga communities growing in “warm and low latitude and altitude steppe climates (Bsh, according to the Köppen (1948) system)” that were well-represented by therophytes. While the principal strategy used in Caatinga phytoclimates to avoid water loss during the dry season is a predominance of therophytes, the caatinga studied here adapted a phanerophyte strategy of leaf-fall (Larcher, 2000).

The systematic collections undertaken in the present study, which covered three rainy seasons, three dry seasons, and three intermediate seasons, may have been responsible for the contrast between our life form data and those of studies by Batalha and Martins (2004) and Costa et al. (2004, 2009) who all made shorter and less frequent collections. Batalha and Martins (2004) noted that species whose buds are not exposed to the open air (i.e. hemicryptophytes, cryptophytes, and therophytes) tend to be under-represented among collections undertaken during just a single period. Consecutive collections during a 36 month period (in the present study) allowed for ample sampling of the floristic composition.

No significant differences were observed between the three different cerrado phytophysiognomies in terms of their soil properties (principally in relation to physical parameters) and their respective floristic spectrums of life forms. Dantas and Batalha (2011) encountered relationships between edaphic conditions and floristic composition and richness in a fine-scale study of a cerrado area. Marimon Junior and Haridasan (2005) noted that soil fertility did not support the hypothesis of the contiguous occurrence of cerradão vegetation and cerrado sensu stricto vegetation. The same authors reported that the greater percentage of clay in the cerradão site allowed the establishment of greater densities of trees due to greater soil-water availability. Ruggiero et al. (2002) reported that campo cerrado, cerrado sensu stricto and cerradão could not be distinguished based on soil chemical parameters. The greater species richness and densities of shrub-arboreal individuals in the typical cerrado site in the present study appeared to be associated with the greater proportions of clay and organic material there that would presumably increase water retention, aid the growth of soil microorganisms, and increase nutrient availability for plant growth (Motta et al., 2002). The presence of blocks of stone in the rocky outcrop cerrado, the occurrence of wild-fires in the typical cerrado, and the densities of arboreal individuals and the interactions between different species in each phytophysiognomy appear to be factors generating the distinct cerrado phytophysiognomies.

While low precipitation levels (less than 700 mm/year) concentrated into as few as three consecutive months during the summer in the southern hemisphere and associated with average temperatures of 26 °C (Nimer, 1989) are considered typical conditions for the establishment of caatinga vegetation (Andrade-Lima, 1981; Sampaio, 1995), rainfall was not a useful factor for delimiting cerrado and caatinga vegetation, such as their transition, in the present study. The soil-water relationships in these vegetation sampled may be influenced by their respective proportions of total sand and clay, because although the cerrado-caatinga transition and caatinga sites had rainfall levels slightly less than that of the cerrado, the water retentions of their soils would presumably be greater due to their larger proportions of clay and organic matter – thus favoring the establishment of greater plant abundances and species richnesses in the shrub-arboreal layer (Motta et al., 2002).

This study demonstrates that the floristic compositions, floristic life form spectra, and the heterogeneity of the soil were elements that delimited cerrado, cerrado-caatinga transition, and caatinga. The soil was the principal conditioning factor determining the different vegetation types, independent of precipitation levels. As the organic matter and clay contents of the soil increased and the percentages of sand decreased the proportions of therophytes, hemicryptophytes and chamaephytes diminished, while phanerophytes increased. The importance of edaphic conditions, and not just the scarcity and irregularity of rainfall, as is usually supposed, must be considered as a factor in establishing caatinga vegetation.

It must also be emphasized that no single variable determined the occurrence of each vegetation type, but rather the interactions between them. Species interactions themselves likewise influence the establishment of vegetation formations, as species richness depends on a mix of factors that can alter the effects of other variables.

Appendix

Species list in the vegetation types and cerrado physiognomies, such as in the cerrado-caatinga transition, Chapada Diamantina, NE Brazil.

Species Voucher LF OP I OP II ROC TC CCT CAA
Acanthaceae
Ruellia incomta Lindau 230 Ch x x x x
Thyrsacanthusramosissimus (Moric.) V.M.Baum 229 H x x
Agavaceae
Herreria sp. Th x
Amaranthaceae
Alternanthera cf. brasiliana Kuntze 327 Th x x x x
Gomphrena agrestis Mart. 214 Th x x x x x
Gomphrena mollis Mart. 202 Th x
Pfaffia acutifolia O. Stützer Th x
Anacardiaceae
Astronium sp. Ph x
Spondias sp. 439 Ph x
Annonaceae
Annona coriacea Mart. 199 Ph x x x x
Duguetia furfuracea (A. St.-Hil.) Saff. 210 Ch x x x x
Apocynaceae
Blepharodon nitidum (Vell.) Macbr. 397 Th x
Ditassa retusa Mart. 231 Ph x x
Hancornia speciosa Gomez 265 Ph x x x
Oxypetalum capitatum Mart. 433 Th x
cf. Prestonia coalita (Vell.) Woodson Th x
Secondatia floribunda A.DC. Th x
Araceae
Philodendron lundii Warm. H x
Arecaceae
Syagrus sp. Ph x x x
Asteraceae
Acanthospermum australe Kuntze Ch x x x
Aspilia hispidantha H. Rob. 398 Th x x x
Calea harleyi H.Rob. 377 Ch x x x x
Conocliniopsis prasiifolia (DC.) R.M. King & H. Rob. 381 Ch x x x x x
Chaptaliaintegerrima (Vell.) Burkart 302 Th x
Chaptalia sp. 2 Th x
Chromolaena sp. 301 Th x
Elephantopus hirtiflorus DC. 376 H x x x
Eremanthus capitatus (Spreng.) MacLeish 383 Ph x
Lasiolaena blanchetti (Baker) R.M.King & H.Rob. 379 Th x
Lepidaploa chalybaea (Mart. ex. DC.) H.Rob. 375 Ch x x x x
Lepidaploa cotoneaster Willd ex. Spreng. 374 Ch x x x
Platypodanthera melissifolia (DC.) R.M.King & H.Rob. 473 Th x
Porophyllum ruderale (Jacq.) Cass. 384 Th x
Stilpnopappus tomentosus Gardner 380 Th x x x
Trichogonia salviifolia Gardner 473 Th x
Tridax procumbens L 474 Th x
Bignoniaceae
Arrabidaea selloi (Spreng.) Ph x
Bignonia binata Thunb. 457 Ph x
Bignonia convolvuloides (Bureau & K.Schum.) L.G.Lohmann 466 Ph x
Bignonia erubescens S.Moore Ph x
Fridericia chica (Bonpl.) L.G.Lohmann 465 Ph x
Fridericia cinerea (Bureau ex K.Schum.) L.G.Lohmann 308 Ph x
Fridericia platyphylla (Cham.) L.G.Lohmann 313 Ph x
Mansoa hirsuta DC. Ph x
Neojobertia candolleana Bureau & K.Schum. Ph x
Proterantha glandulosa A.H. Gentry 310 Ph x
Tabebuia spongiosa Rizzini 463 Ph x
Bignoniaceae 1 Ph x
Bignoniaceae 2 Ph x
Bignoniaceae 3 Ph x
Bignoniaceae 4 Ph x
Bignoniaceae 5 Ph x
Bignoniaceae 6 Ph x
Boraginaceae
Cordia cf. rufescens A.DC. 24 Ch x x
Varronia leucomalloides (Taroda) J.S.Mill 410 Th x
HeliotropiumscorpioidesWilld. ex Roem. & Schult. 246 Th x
Bromeliaceae
Billbergia porteana Brongn. ex. Beer 469 H x
Cryptanthus sp. 343 H x
Hohenbergia cf. caatingae var. eximbricata L.B. Smith & Read 470 Cr x
Neoglaziovia variegata Mez 342 H x
Tillandsia loliacea Mart. Ex Schult.f. Ep x
Tillandsia streptocarpa Baker Ep x
Tillandsia cf. tenuifolia L. 309 Ep x
Burseraceae
Commiphora leptophloeos (Mart.) J.B. Gillett 462 Ph x
Cactaceae
Arrojadoa penicillata (Gürke) Britton & Rose 471 Su x x
Cereus albicaulis Luetzelb. Su x
Species Voucher LF OP I OP II ROC TC CCT CAA
Pilosocereus glaucochrous (Werderm.) Blyles & G.D.Rowley Su x x
Stephanocereus leucostele A.Berger Su x
Tacinga funalisBritton & Rose Su x
Tacinga palmadora (Britton & Rose) N.P.Taylor & Stuppy Su x
Tacinga werneri (Eggli) N.P.Taylor & Stuppy Su x x
Capparaceae
Cynophalla flexuosa (L.) J.Pres Ph x
Cynophalla jacobinaeMoric. ex Eichler Ph x
Neocalyptrocalyx longifolium (Mart.) X.Cornejo & H.H.Iltis Ph x
Caricaceae
Jacaratia corumbensis Kuntze Ph x
Caricaceae 1 Ph x
Combretaceae
Terminalia sp. Ph x
Commelinaceae
Commelina erecta L. 200 Th x x x x
Convolvulaceae
Bonamia burchellii Hallier f. 369 Ph x
Evolvulus elegans Moric. 314 Ch x x
Evolvulus elaeagnifolius Dammer 311 Th x
Evolvulus glomeratus Choisy 227 Ch x x x x x
Ipomoea subtomentosa (Chodat & Hassl.) O´Donell 221 Ph x x x x x
Jacquemontiaagrestis Meisn. 209 Ch x x x x
Jacquemontia montana Meisn. 226 Ph x x x x
Jacquemontia nodiflora G.Don 370 Ph x
Cyperaceae
Bulbostylis capillaris Nees 442 H x x x
Bulbostylis fasciculata Cherm. 315 H x x x x
Bulbostylis junciformis C.B.Clarke. 316 H x x x
Cyperus agregatus (Willd.) Endl. 317 Th x x x x x
Rhynchospora sp. 351 H x x x
Cyperaceae 444 Th x
Ericaceae
Agarista cf. chapadensis (Kin.-Gouv.) Judd 332 Ph x
Gaylussacia brasiliensis Meisn. 232 Ch x x
Erythroxylaceae
Erythroxylum betulaceum Mart. 335 Ch x x
Erythroxylum oxypetalum O.E. Schulz 338 Ph x
ErythroxylumsuberosumA.St.-Hill. 339 Ch x x x
Erythroxylum sp. Ch x
Euphorbiaceae
Actinostemonconcolor Mül.Arg. 271 Ph x x
Actinostemonsp. Ph x
Croton adamantinus Mull. Arg. 236 Ch x
Croton argyrophyllus Kunth Ch x
Croton compressus Lam. Ph x
Croton echioideus Baill. 329 Ph x
Croton glandulosus L. Th x
Croton heliotropiifolius Kunth. 234 Ph x x x
Croton limae A.P. Gomes, M.F. Sales & P.E. Berry 331 Ph x
Croton velutinus Baill. 233 Ch x x
Euphorbia potentilloides Boiss. 208 Ch x x x x
Jatropha sp. Ph x
Maprounea guianensis Aubl. 235 Ph x x
Microstachys daphinoides Müll.Arg. 333 Ch x x
Microstachys serrulata Müll.Arg. 401 Ph x x
Sapium glandulosum Druce 394 Ph x
Sebastiania corniculata Pax 401 Ch x x x x
Sebastiania salicifolia Pax Ch x x x
Stillingia saxatilis Müll.Arg 307 Ch x
Tragia friesii Pax & K.Hoffm. 303 Th x x x
Icacinaceae
Emmotum nitens (Benth.) Miers 238 Ph x
Iridaceae
Sisyrinchiumluzula Klotzsch ex Klatt 440 Th x
Lamiaceae
Eriope hypenioides Mart. Ex Benth. 321 Ph x x
Raphiodon echinus (Nees & Mart.) Schauer. 363 H x x x
Hyptis macrantha A.St.-Hil. 197 Ch x
Hyptisplatanifolia Mart. 320 Ch x x x x x
Hyptis rugosa Benth. 409 Ch x
Hypenia vitifolia Pohl. 322 Ch x x x x
Fabaceae
Aeschynomene brevipes Benth. 247 Ph x x x x
Aeschynomene histrix Poir. 282 Ch x x x x x
Aeschynomene martii Benth. Ph x
Andira humilis Mart. 276 Th x x x x
Bauhinia catingae Harms Ph x
Centrolobium tomentosum Guill. ex Benth. Ph x
Chamaecrista flexuosa Greene 248 Ch x x x x
Chamaecrista repens var. multijuga (Benth.) H.S.Irwin & Barneby. 249 Ch x x x x x
Chamaecrista rotundifolia Greene 228 Ch x x
Crotalaria holosericea Nees & Mart. 275 Ph x
Dalbergia cearensis Ducke Ph x
Species Voucher LF OP I OP II ROC TC CCT CAA
Dalbergia miscolobium Benth. 203 Ph x x x x
Dimorphandra gardneriana Tul. 253 Ph x
Hymenaea stigonocarpa Mart. Ex Hayne 212 Ph x x x
Lonchocarpus obtusus Benth. Ph x
Machaerium acutifolium Vogel Ph x
Machaerium brasiliense Vogel Ph x
Machaerium nictitans Benth. 461 Ph x
Mimosa hirsutissima Mart. 287 Th x
Mimosa paludosa Benth. (P1) 286 Th x x x
Mimosa quadrivalvis L. 274 Th x
Mimosa somnians Humb. & Bonpl. ex.Willd. 213 Ph x
Periandra mediterranea (Vell.) Taub. 349 Ph x
Phanera microstachya (Raddi) L.P.Queiroz Ph x
Piptadenia irwinii G.P.Lewis Ph x
Poeppigia procera C.Presl 281 Ph x
Pseudopiptadeniabrenanii G.P.Lewis & M.P.Lima Ph x x
Senegalia langsdorffii (Benth.) Bocage & L.P.Queiroz 425 Ph x
Senegalia piauhiensis (Benth.) Bocage & L.P.Queiroz 428 Ph x
Senegalia riparia (Kunth)Britton & Rose in Britton & Killip 254 Ph x x
Senegalia tenuifolia Britton & Rose 426 Ph x
Senna esplendida (Vogel) H.S.Irwin & Barneby Ph x
Senna macranthera (Coll) H.S.Irwin & Barneby 399 Ph x
Senna rugosa (G.Don) H.S.Irwin & Barneby 400 Ph x
Stryphnodendron rotundifolium Mart. Ph x x
Stylosanthes gracilis Kunth 262 Ch x x x
Stylosanthes macrocephala M.B.Ferreira & Sousa Costa 250 Ch x x x x
Stylosanthes scabra Vogel. 251 Ch x x x x x
Stylosanthes seabrana B.L.Maass & ´t Mannetje 371 Ch x
Tachigali sp. Ph x x
Zornia gemella Vogel. 252 Th x x x
Zornia sericea Moric. 284 Th x x x
Lythraceae
Cuphea sessilifolia Mart. 225 Ch x x x x
Loganiaceae
Antonia ovata Pohl 218 Ph x
Spigelia cf. anthelmia L. Th x x
Loranthaceae
Struthathus cf. flexicaulis Mart. 319 Th x x x x
Malpighiaceae
Banisteriopsis stellaris (Griseb.) B.Gates 293 Th x x x
Byrsonima sericea DC. 292 Ph x x x
Byrsonima correifolia A.Juss. 337 Ph x
Byrsonima sp. (caatinga) Ph x
Heteropterys sp. Ph x
Mascagnia sp. Ph x
Stigmaphyllon auriculatum A.Juss. 456 Th x
Stigmaphyllon paralias A.Juss. 211 Th x x x x x
Tetrapterys sp. 352 Ph x
Malpighiaceae 1 458 Ph x
Malpighiaceae 2 Ph x
Malpighiaceae 3 Ph x
Malpighiaceae 4 Ph x
Malvaceae
Ceiba erianthos (Cav.) K.Schum. 344 Ph x
Helicteres velutina K.Schulz. 435 Ph x
Herissantia crispa (L.) Briz. 396 Th x x
Luehea sp. Ph x
Gossypium sp. 475 Ph x
Pavonia cancellata Cav. 360 Th x x x x x
Pavonia glazioviana Gürke 387 Th x
Pavonia martii Mart. ex Colla 391 Th x
Sida angustissima Miq. 390 Th x x x x
Sida cordifolia L. 389 Th x
Sida galheirensis Ulbr. 395 Th x x
Sida linifolia Cav. 392 Th x x x
Sida spinosa L. 387 Th x
Waltheria indica L. 362 Ch x x x
Waltheria macropoda Turcz. 361 Ch x x
Wissadula hirsuta C. Presl 386 Ch x x x
Melastomataceae
Marcetia macrophylla DC. 240 Ph x
Miconia cf. albicans Steud. 273 Ph x
Miconia alborufescens Naudin 241 Ph x
Tibouchina blanchetiana Cogn. 239 Ph x x
Moluginaceae
Mollugo verticillata L. 207 Th x x
Myrtaceae
Campomanesia guaviroba Benth. & Hook.f. Ph x
Campomanesia cf. sessiliflora var. lanuginosa (Chodat. & Hassl.) Landrum 345 Ph x x
Eugenia cf. cerasiflora Miq. 406 Ph x
Eugenia cf. punicifolia DC. 204 Ph x x x x x
Eugenia sonderiana O.Berg 405 Ph x x
Myrciablanchetiana (O.Berg) Mattos 404 Ph x
Myrcia cf. rufipes DC. 346 Ph x x
Species Voucher LF OP I OP II ROC TC CCT CAA
Myrcia splendes O.Berg. 270 Ph x x x
Myrciaria floribunda O.Berg. Ph x x
Psidium cf. brownianun Nied. 216 Ph x
Psidium grandifolium Mart. 359 Ph x
Psidiumschenckianum Kiaersk. 403 Ph x
Ochnaceae
Ouratea cf. floribunda Engl. 245 Ph x x x
Orchidaceae
Campylocentrum micranthum (Lindl.) Rolfe Ch x
Epidendrum sp. Ch x
Oncidium sp. Ch x
Trichocentrum cebolleta (Jacq.) M.W.Chase & N.H.Williams 328 Th x
Oxalidaceae
Oxalis frutescens sub. frutescens Linnaeus 242 Th x x x x x
Arecaceae
Syagrus sp. Ph x x x
Passifloraceae
Passiflora cincinnata Mast. 295 Th x x
Passiflora edmundoi Sacco 325 Th x x x
Piriqueta duarteana (A. St.-Hil., A. Juss & Cambess) Urb. 452 Th x x
Piriqueta sidifoliavar.multiflora Urb. 289 Th x x
Turnera blanchetiana Urb. 434 Ph x
Turnera melochioides var.latifolia Urb. 288 Th x x x x
Phyllanthaceae
Astrocasia sp. Ph x
Phyllanthus orbiculatus Rich. 353 Ch x
Phytolacaceae
Microtea paniculata Moq. 201 Th x x x x
Picramniaceae
Picramnia sp. Th x
Poaceae
Aristida sp. 1 421 H x x x x
Axonopus barbigerus Hitchc. 260 H x x
Axonopus polydactylus (Steud.) Dedecca 259 H x x x x
Dichanthelium sciurotis Trin. H x x x
Digitaria insularis (L.) Fedde 258 H x
Echinolaena inflexa Chase 415 H x x x
Eragrostis polytricha Nees 257 H x x x
Eragrostis solida Nees H x x
Eragrostis sp. 300 H x x
Eragrostis sp. 418 H x
Gymnopon sp. 1 H x
Ichnanthus zehntneri Mez 298 H x
Melinis minutiflora P.Beauv. 414 H x
Melinis repens (Willd.) Zizka. 261 H x x x
Mesosetum loliiforme Chase 255 H x x x x
Mesosetum sp. 1 297 H x
Mesosetum sp. 2 417 H x
Panicum sellowii Nees 256 H x x
Panicum sp. 1 416 H x
Panicum sp. 2 419 H x x x x
Paspalum arenarium Schrad. 423 H x x
Raddia portoi Kuhlm. 413 H x
Schysachirium sp. 1 422 H x x
Setaria gracilis Kunth 412 H x
Setaria parviflora Poir. Kerguélen 424 H x
Setaria setosa P.Beauv. 411 H x x
Setaria tenax Desv. H x x
Setaria sp. 1 296 H x x
Setaria sp. 2 299 H x
Setaria sp. 3 H x
Streptostachys ramosa Zuloaga & Soderstr. 420 H x x
Streptostachys robusta Renvoize 347 H x x x x
Trachypogon spicatus Kuntze 156 H x x x x x
Poaceae sp. 1 H x x
Poaceae sp. 2 H x
Poaceae sp. 3 H x x x x
Polygalaceae
Polygala decumbens A.W.Benn. 205 Th x x x x x
Polygala obovata A.St.-Hil. 206 Th x x x x
Polygala trichosperma Jacq. 326 Th x x x x
Polygala sp. nova 220 Th x
Portulacaceae
Portulaca hirsutissima Cambess. 215 Th x x x x x
Portulaca mucronata Link 318 Th x x x
Primulaceae
Myrsine guianensis (Aubl.) Kuntze. 336 Ph x x
Rubiaceae
Alseis floribunda Schott 323 Ph x
Borreria sp. 453 Th x
Chomelia sp. 1 455 Ph x
Chomelia sp. 2 459 Ph x
Diodella apiculta (Willd. ex Roem. & Schult.) Delprete 454 Th x
Declieuxia fruticosa Kuntze 224 Th x x
Mytracarpus salzmannianus DC. 223 Th x x x x x
Species Voucher LF OP I OP II ROC TC CCT CAA
Mytracarpus sp. 431 Th x x x x
Palicourea marcgravii A.St.-Hil. 460 Ph x
Psyllocarpus asparagoides Mart. 222 Th x x x x x
Psyllocarpus sp. 334 Th x x x
Randia armata DC. Ph x x
Richardia grandiflora Steud. 432 Ch x x x x x
Simira sp. Ph x
Rubiaceae 1 430 Th x x
Rubiaceae 2 Ph x
Rutaceae
Zanthoxylum nannophyllum (Urb.) Alain Ph x
Zanthoxylum sp.2 Ph x
Salicaceae
Casearia cf. sylvestris Sw. 237 Ph x x x
Sapindaceae
Cupania oblongifoliaMart. Ph x
Diatenopteryx grazielae Vaz & Andreata 368 Ph x
Serjania lethalis A.St.-Hil. 268 Ph x x
Serjania pinnatifolia Radlk. 269 Ph x
Sapindaceae Ph x
Smilacaceae
Smilax elastica Griseb. 324 Ph x
Solanaceae
Schwenkia sp. 355 Th x x x x x
Solanum buddlejifolium Sendtn. 266 Ph x x x x
Solanum stenandrum Sendtn 358 Th x x x x x
Solanum cf. stipulaceum Brouss. Roem. & Schult. 357 Ph x
Verbenaceae
Aegiphila verticillata Vell. 354 Ph x x
Lippia microphylla Cham. 198 Ph x x x
Lippia sp. 467 Ph x
Stachytarpheta crassifolia Schrad 365 Ch x x x
Vitex sp. Ph x
Vitaceae
Cissus bahiensis Lombardi 450 Th x
Vochysiaceae
Vochysia thyrsoidea Phol 290 Ph x
Undetermined
Undetermined 1 437 Th x
Undetermined 2 438 Th x
Undetermined 3 Ph x
Undetermined 4 Ph x
Undetermined 5 Ph x
Undetermined 6 Ph x
Undetermined 7 Ph x
Undetermined 8 Ph x
Undetermined 9 Ph x
Undetermined 10 Ph x
100 93 105 121 94 117

LF = Life form; Ch: chamaephyte; H: hemicryptophyte; Ph: phanerophyte; Th: therophyte; Cr: cryptophyte; Su: “suculenta”; Ep: epiphyte; OPI: open cerrado (site I); OPII (site II): open cerrado II; ROC: rocky outcrop cerrado; TC: typical cerrado; CCT: cerrado-caatinga transition; CAA: caatinga.

Acknowledgements

The authors would like to thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico for financing this research project (CNPq 480508/2008-09) and for the productivity grants to the third and final authors, and the Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB 5303/2009) and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior for the doctoral study grant awarded to the first author.

(With 3 figures)

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Received: July 28, 2014; Accepted: March 01, 2015

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