Tree Community Structure in a Seasonally Dry Tropical Forest Remnant, Brazil
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DOI: http://dx.doi.org/10.1590/01047760.201420021540 TREE COMMUNITY STRUCTURE IN A SEASONALLY DRY TROPICAL FOREST REMNANT, BRAZIL Deborah Mattos Guimarães Apgaua1*, Polyanne Aparecida Coelho1, Rubens Manoel dos Santos1, Paola Ferreira Santos1, Ary Teixeira de Oliveira-Filho2 *Autor para correspondência: [email protected] ABSTRACT: Most studies on Seasonally Dry Tropical Forests (SDTFs) investigate phytogeographic patterns and floristic connections of this disjunct biome. However, little is known about the structural characteristics of SDTFs. We aimed to describe the structure of a SDTF in an ecotonal area between the Cerrado and Caatinga domains. In total, 79 tree species were recorded, and high values of Shannon diversity index (3.6 nats/individual) and equability (0.83) were observed. The diameter distribution for the species with higher cover values and for the entire community did not exhibit a reverse-J shaped distribution, which indicates the occurrence of different growth strategies and ecological adaptations to water stress. The results did not indicate the formation of floristic groups, as the high soil fertility in the study area results in a homogeneous environment. The structural characteristics of the study area associated with the soil composition highlight its importance for conservation and emphasize the need for community structure studies in SDTFs. Keywords: diameter distribution, Caatinga, phytosociology, conservation. ESTRUTURA DA COMUNIDADE ARBÓREA DE UM FRAGMENTO DE FLORESTA TROPICAL SAZONALMENTE SECA, BRASIL RESUMO: A maior parte dos estudos envolvendo as Florestas Tropicais Sazonalmente Secas aborda questões fitogeográficas e conexões florísticas deste bioma disjunto. Porém, pouco se conhece a respeito das características estruturais da FTSS. Devido a esta lacuna existente na literatura, buscamos descrever a estrutura de um fragmento remanescente de FTSS em uma área ecotonal entre domínio do Cerrado e da Caatinga. Ocorreram no estudo grande número de espécies arbóreas, 79, e altos valores para os índices de diversidade e equabilidade (3,6 nats/indivíduos e 0,83, respectivamente). A distribuição diamétrica para os indivíduos que apresentaram maiores valores de cobertura e para a comunidade não resultou em padrão de J-reverso, o que indica adaptações ecológicas ao stress hídrico adotadas em diferentes estratégias de crescimento. Não houve formação de grupos florísticos, já que a alta fertilidade do solo da área de estudo configura um ambiente homogêneo. A característica da área de estudo detectada através da estrutura, associada a dados edáficos, nos permitiu vislumbrar sua importância em termos de conservação e reforçar a importância de estudos com este enfoque nas FTSS. Palavras-chave: distribuição diamétrica, Caatinga, fitossociologia, conservação. 1 INTRODUCTION of plant communities in dry environments compared to more humid ones in Brazil (LIMA et al., 2007; SILVA; The Seasonally Dry Tropical Forest (SDTF) biome SCARIOT, 2004). Even though several studies have has physiognomic, taxonomic, and structural peculiarities focused on the phytogeography and florisitic connections of due to a phytogeographical report in which provided its SDTFs (OLIVEIRA-FILHO et al., 2006; PENNINGTON disjunction along South America according to the refuge et al., 2000; SANTOS et al., 2012), few studies aimed theory (PENNINGTON et al., 2000, 2006; PRADO; to examine their structural characteristics. Vegetation GIBBS, 1993). structure studies are complementary to phytogeographic The longest continuous extent of SDTFs occurs ones as they help understand the environmental and spatial in the Caatinga domain as a mosaic of “ecoregions” characteristics of SDTFs. and “floristic units” (SANTOS et al., 2012; VELLOSO; Vegetation structure varies greatly among SDTF SAMPAIO, 2002). This variability is caused mostly by patches, mostly due to the environmental heterogeneity variations in climate, topography, edaphic conditions, along their distribution. Most vegetation structure and human intervention among the areas (VELLOSO; studies in the SDTF biome were done in the more xeric SAMPAIO, 2002). Nevertheless, there are few studies part of the Caatinga (PEREIRA et al., 2002; RODAL; 1 Universidade Federal de Lavras - Lavras, Minas Gerais, Brazil 2 Universidade Federal de Minas Gerais - Belo Horizonte, Minas Gerais, Brazil Cerne, Lavras, v. 20, n. 2, p. 173-182, abr./jun. 2014 174 Apgaua, D. M. et al. NASCIMENTO, 2006), whereas few ones investigated the 2.3 Environmental variables ecotonal area between the Caatinga and Cerrado domains (SANTOS et al., 2011, 2012). However, SDTF patches To determine the relationships between edaphic have differences in tree species composition, abundance, variables and vegetation, three 0.5 l surface samples (0–20 and individual size (SANTOS et al., 2007). cm depth) were taken per plot for analysis of particle size Because SDTFs occur in areas of high soil distribution and chemical composition of soils. The soil fertility, they are subjected to disturbance by farming samples were stored in plastic bags and taken to the Federal (PENNINGTON et al., 2000, 2006), and only few remnants University of Lavras Soil Analysis Laboratory (Laboratório with primary vegetation can be found in the area. The de Análise de Solos da Universidade Federal de Lavras), settlement and development of the region caused the where the following variables were measured: pH in water; reduction and alteration of natural vegetation, and diversity levels of potassium (K), phosphorus (P), calcium (Ca), studies of these communities are fundamental to design magnesium (Mg), and aluminum (Al); potential acidity conservation strategies for the remaining forests. (H+Al), sum of bases (Sb), effective CEC (t), CEC at pH This study aimed to investigate the structure of 7.0 (T), aluminum saturation (m), base saturation (V), the tree community of a Seasonally Dry Tropical Forest carbon (C), organic matter (OM), and sand, silt, and clay remnant in an ecotonal area between the Cerrado and fractions. Laboratory procedures followed the Brazilian Caatinga domains in northern Minas Gerais, Brazil. Agricultural Research Corporation protocols (EMPRESA BRASILEIRA DE PESQUISA AGROPECUÁRIA - 2 MATERIAL AND METHODS EMBRAPA, 2006). 2.1 Study area 2.4 Tree community descriptive parameters The study was done in a SDTF remnant of To describe tree community structure, the traditional approximately 40 ha located in the municipality of quantitative parameters proposed by Mueller-Dombois and Juvenília (14º26’04’’ S and 44º10’67’’ W). The area has Ellenberg (1974) were calculated per species: absolute a gentle slope with the soil depth gradient characterized density (AD, ind/ha); absolute frequency (AF, %); absolute by shallower soils in its lowest part, rocky outcrops in dominance (ADo) expressed by basal area (m²/ha); and its intermediate section, and deeper soils in its highest cover value (CV). The Shannon diversity index (H’) and part. The average annual rainfall is 1000 mm and the Pielou equability (J’) measures of diversity (BROWER; area experiences more rain from November to January ZAR, 1984) were also calculated. Tree density distributions (SANTOS et al., 2011). per diameter class were analyzed for the community and for the 11 species of highest cover value. Class intervals 2.2 Tree community survey with growing ranges were used to compensate for the significant decrease in the number of individuals in Sixty 20 x 20 m (400 m2) plots, totaling 2.4 ha, larger diameter size classes, typical of a reverse-J shaped were distributed along three transects 50 m apart from negative exponential distribution (BOTREL et al., 2002). each other, parallel to the largest dimension of the remnant The Kruskal-Wallis test was used to assess if the number and following the soil depth gradient. At each point of the of individuals in each diameter class was similar among transect, two plots were established 10 m apart from each the three transects (ZAR, 2010). other, and the same procedure was repeated every 20 m. All individuals with DBH (diameter at breast height) ≥ 10 2.5 Analysis of environmental variables cm at 1.30 m from soil were recorded. The botanical material was herborized and To correlate the environmental variables with the control collection deposited in the Montes Claros vegetation variables, a Canonical correspondence herbarium of Universidade Estadual de Montes Claros Analysis (CCA) was performed using Pc-ord-5 software (UNIMONTES). The Angiosperm Philogeny Group II (MCCUNE; MEFFORD, 2006). The abundance values (ANGIOSPERM PHYLOGENY GROUP - APG, 2003) were log10 (a+1) transformed to compensate for deviations classification system was adopted. caused by outliers (TER BRAAK, 1995). Cerne, Lavras, v. 20, n. 2, p. 173-182, abr./jun. 2014 Tree community structure ... 175 Fifteen soil parameters [P, pH, K, Ca, Mg, H+Al, (five species each); Anacardiaceae and Rutaceae (four SB, (t), (T), V, OM, P-rem, sand, silt, and clay] were species each); Nyctaginaceae (three); followed by investigated, but after a preliminary analysis, 10 parameters Cordiaceae, Cactaceae, Lamiaceae, Malpighiaceae, were removed as they were weakly correlated with the Meliaceae, Moraceae, Myrtaceae, Polygonaceae, and environmental data or redundant, which could result in high Rubiaceae (two species each), representing 55.2% of collinearity. Thus, only the five variables [K, Ca, (T), OM, the sampled species. The other families (44.8%) were and P-rem] most strongly correlated with the ordination