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Evaluation of Vegetation after Four Years in a Caatinga Fragment in the State of Sergipe, Brazil

Abstract

The comparison of vegetation at two different moments allows for recognizing the stability of plant communities. The structure and floristic composition of a Caatinga fragment in the municipality of Poço Verde, state of Sergipe, were evaluated after four years (2011-2015). Sampling was performed through 30 plots of 20 m × 20 m, considering individuals with circumference at breast height ≥ 6 cm. Despite the increase in density (0.79%) and basal area (4.82%), changes in floristic composition, in Shannon-Wiener diversity (from 3.33 nats.ind-1 to 3.30 nats.ind-1) and in Pielou equability (from 0.78 to 0.80) were small. None of the structural parameters evaluated (richness, density, basal area, importance value and Shannon-Wiener diversity) significantly differed between evaluation periods. The vegetation remained stable over time, tolerating current anthropization levels and enabling the use of its natural resources through planned management.

Keywords:
floristic survey; phytosociology; stability

1. INTRODUCTION AND OBJECTIVES

Studies comparing vegetation at different times can detect floristic and structural variations in populations and plant communities, supporting the understanding of factors that affect changes in these communities (e.g., Carvalho et al., 2010Carvalho FA, Fagg CW, Felfili JM. Population dynamics of Acacia tenuifolia (L.) Willd. in a dry forest on limestone outcrops in Central Brazil. Scientia Forestalis 2010; 38(86): 297-306.; Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
; Mews, Marimon, Pinto et al., 2011Mews HA, Marimon BS, Pinto JRR, Silverio DV. Dinâmica estrutural da comunidade lenhosa em Floresta Estacional Semidecidual na transição Cerrado-Floresta Amazônica, Mato Grosso, Brasil. Acta Botanica Brasilica 2011; 25(4): 845-857. 10.1590/S0102-33062011000400011
https://doi.org/10.1590/S0102-3306201100...
). In addition, these studies allow for inferences about vegetation dynamics; short-term assessments are more sensitive in detecting changes in a community (Braga & Rezende, 2007Braga FMS, Rezende AV. Dinâmica da vegetação arbórea da Mata de Galeria do Catetinho, Brasília-DF. Cerne 2007; 13(2): 138-148.).

Adopting a temporal scale in structural and floristic evaluations enables recognizing the stability of plant communities; changes in the community are inevitably continuous and time-dependent (Condit et al., 1992Condit R, Hubbell SP, Foster RB. Short-term dynamics of a neotropical forest. BioScience 1992; 42(11): 822-828. 10.2307/1312081
https://doi.org/10.2307/1312081...
). Data obtained from these evaluations are fundamental to subsidize conservation actions and programs for the recovery of degraded areas, providing subsidies for the sustainable vegetation management (Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
; Mews, Marimon, Maracahipes et al. 2011Mews HA, Marimon BS, Pinto JRR, Silverio DV. Dinâmica estrutural da comunidade lenhosa em Floresta Estacional Semidecidual na transição Cerrado-Floresta Amazônica, Mato Grosso, Brasil. Acta Botanica Brasilica 2011; 25(4): 845-857. 10.1590/S0102-33062011000400011
https://doi.org/10.1590/S0102-3306201100...
; Rolim et al., 1999Rolim SG, Couto HTZ, Jesus RM. Mortalidade e recrutamento de árvores na Floresta Atlântica em Linhares (ES). Scientia Forestalis 1999; (55): 49-69.).

In Brazil, publications comparing vegetation composition and/or structure between two distinct periods were carried out in Cerrado areas (e.g., Mews, Marimon, Pinto et al., 2011Mews HA, Marimon BS, Pinto JRR, Silverio DV. Dinâmica estrutural da comunidade lenhosa em Floresta Estacional Semidecidual na transição Cerrado-Floresta Amazônica, Mato Grosso, Brasil. Acta Botanica Brasilica 2011; 25(4): 845-857. 10.1590/S0102-33062011000400011
https://doi.org/10.1590/S0102-3306201100...
; Silva Neto et al., 2017Silva Neto AJ, Mello JM, Fontes MAL, Santos RM, Batista APB, Scolforo JRS. Dinâmica da comunidade arbórea em um fragmento de cerrado sensu stricto em Minas Gerais, Brasil. Scientia Forestalis 2017; 45(113): 21-29. 10.18671/scifor.v45n113.02
https://doi.org/10.18671/scifor.v45n113....
), galley forests (e.g., Braga & Rezende, 2007Braga FMS, Rezende AV. Dinâmica da vegetação arbórea da Mata de Galeria do Catetinho, Brasília-DF. Cerne 2007; 13(2): 138-148.; Guimarães et al., 2008Guimarães JCC, Van Den Berg E, Castro JC, Machado ELM, Oliveira-Filho AT. Dinâmica do componente arbustivo-arbóreo de uma floresta de galeria aluvial no planalto de Poços de Caldas, MG, Brasil. Brazilian Journal of Botany 2008; 31(4): 621-632. 10.1590/S0100-84042008000400008
https://doi.org/10.1590/S0100-8404200800...
), ombrophilous forests and/or montanes (e.g., Chagas et al., 2001Chagas RK, Oliveira-Filho AT, Van Den Berg E, Scolforo JRS. Dinâmica de populações arbóreas em um fragmento de floresta estacional semidecidual montana em Lavras, Minas Gerais. Revista Árvore 2001; 25(1): 39-57.; Gomes et al., 2003Gomes EPC, Mantovani W, Kageyama PY. Mortality and recruitment of trees in a secondary montane rain forest in Southeastern Brazil. Brazilian Journal of Biology 2003; 63(1): 47-60. 10.1590/S1519-69842003000100007
https://doi.org/10.1590/S1519-6984200300...
; Oliveira Filho et al., 2007Oliveira Filho AT, Carvalho WAC, Machado ELM, Higuchi P, Appolinário V, Castro GC et al. Dinâmica da comunidade e populações arbóreas da borda e interior de um remanescente florestal na Serra da Mantiqueira, Minas Gerais, em um intervalo de cinco anos (1999-2004). Revista Brasileira de Botânica 2007; 30(1): 149-161. 10.1590/S0100-84042007000100015
https://doi.org/10.1590/S0100-8404200700...
; Schaaf et al., 2005Schaaf LB, Figueiredo-Filho A, Sanquetta CR, Galvão F. Incremento diamétrico e em área basal no período 1979-2000 de espécies arbóreas de uma floresta ombrófila mista localizada no sul do Paraná. Floresta 2005; 35(2): 271-290. 10.5380/rf.v35i2.4615
https://doi.org/10.5380/rf.v35i2.4615...
) and seasonal and semideciduous forests (e.g., Mews, Marimon, Maracahipes et al., 2011Mews HA, Marimon BS, Maracahipes L, Franczak DD, Marimon BH Jr. Dinâmica da comunidade lenhosa de um cerrado típico na região nordeste do estado de Mato Grosso, Brasil. Biota Neotropica 2011; 11(1): 73-82. 10.1590/S1676-06032011000100007
https://doi.org/10.1590/S1676-0603201100...
; Nascimento et al., 1999Nascimento HEM, Dias AS, Tabanez AAJ, Viana VM. Estrutura e dinâmica de populações arbóreas de um fragmento de floresta estacional semidecidual na região de Piracicaba, SP. Revista Brasileira de Biologia 1999; 59(2): 329-342. 10.1590/S0034-71081999000200015
https://doi.org/10.1590/S0034-7108199900...
; Paula et al., 2002Paula A, Silva AF, Souza AL, Santos FAM. Alterações florísticas ocorridas num período de quatorze anos na vegetação arbórea de uma Floresta Estacional Semidecidual em Viçosa-MG. Revista Árvore 2002; 26(6): 743-749. 10.1590/S0100-67622002000600010
https://doi.org/10.1590/S0100-6762200200...
; Silva & Araujo, 2009Silva APN, Moura GBA, Giongo PR, Silva AO. Dinâmica espaço-temporal da vegetação no semi-árido de Pernambuco. Revista Caatinga 2009; 22(4): 195-205.). Despite the existence of publications for Brazilian seasonally dry tropical forests (SDTF) (e.g., Werneck & Franceschinelli, 2004Werneck MS, Franceschinelli EV. Dynamics of a dry forest fragment after the exclusion of human disturbance in southeastern Brazil. Plant Ecology 2004; 174(2): 337-346. 10.1023/B:VEGE.0000049112.06259.4c
https://doi.org/10.1023/B:VEGE.000004911...
), studies of this type are scarce for the Caatinga biome (Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
), which contributes to the little knowledge about its dynamics (Pereira et al., 2001Pereira IM, Andrade LA, Costa JRM, Dias JM. Natural regeneration in a Caatinga fragment under different disturbance levels. Acta Botanica Brasilica 2001; 15(3): 413-426. 10.1590/S0102-33062001000300010
https://doi.org/10.1590/S0102-3306200100...
).

In Caatinga, anthropization and climatic variations are the main responsible for vegetation changes (Alves, 2009Alves JAA. Caatinga do cariri paraibano. Geonomos 2009; 17(1): 19-25. 10.18285/geonomos.v17i1.80
https://doi.org/10.18285/geonomos.v17i1....
; Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
). In this biome, disturbed environments have shown declines in diversity (Kauffman et al., 1993Kauffman JB, Sanford RL Jr, Cummings DL, Salcedo IH, Sampaio EVSB. Biomass and nutrient dynamics associated with slash fires in neotropical dry forests. Ecology 1993; 74(1): 140-151. 10.2307/1939509
https://doi.org/10.2307/1939509...
), richness, basal area and plant distribution in diameter classes and an increase in mortality rates of individuals (Cavalcanti & Rodal, 2010Cavalcanti ADC, Rodal MJN. Efeito de borda e dinâmica de plantas lenhosas em áreas de Caatinga em Carnaubais, RN. Revista Caatinga 2010; 23(2): 41-50.; Pereira et al., 2001Pereira IM, Andrade LA, Costa JRM, Dias JM. Natural regeneration in a Caatinga fragment under different disturbance levels. Acta Botanica Brasilica 2001; 15(3): 413-426. 10.1590/S0102-33062001000300010
https://doi.org/10.1590/S0102-3306200100...
; Santos et al., 2009Santos MFAV, Guerra TNF, Sotero MC, Santos JIN. Diversidade e densidade de espécies vegetais da caatinga com diferentes graus de degradação no município de Floresta, Pernambuco, Brasil. Rodriguésia 2009; 60(2): 389-402. 10.1590/2175-7860200960211
https://doi.org/10.1590/2175-78602009602...
). In addition, longer drought periods have caused higher mortality rates on individuals’ ingress, as well as decreases in density, diameter and height of these individuals (Albuquerque, 1999Albuquerque SG. Caatinga vegetation dynamics under various grazing intersites by steers in the semi-arid Northeast, Brazil. Journal Range Management 1999; 52(3): 241-248.; Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
).

Given the importance of temporal evaluations to understand the functioning of plant communities (Condit et al., 1992Condit R, Hubbell SP, Foster RB. Short-term dynamics of a neotropical forest. BioScience 1992; 42(11): 822-828. 10.2307/1312081
https://doi.org/10.2307/1312081...
; Libano & Felfili, 2006Libano AM, Felfili JM. Mudanças temporais na composição florística e na diversidade de um cerrado sensu stricto do Brasil Central em um período de 18 anos (1985-2003). Acta Botanica Brasilica 2006; 20(4): 927-936. 10.1590/S0102-33062006000400016
https://doi.org/10.1590/S0102-3306200600...
) and the absence of data for Caatinga communities (Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
), this study was carried out with the objective of assessing the structure and the floristic composition after a four-years period (2011-2015) in a Caatinga fragment in the municipality of Poço Verde, state of Sergipe. It is expected that factors observed in the study area, such as anthropization and interannual climatic variability, cause structural and floristic changes in the vegetation between evaluation moments.

2. MATERIALS AND METHODS

2.1. Study area

The study was conducted on a 71.42-hectare Hypoxero-philic Caatinga fragment, located in the settlement of Santa Maria da Lage (10°44’31’’S and 38°05’53’’W), in the municipality of Poço Verde, mid-southern region of Sergipe, Brazil (Figure 1). This fragment is characterized by dense and dry forest vegetation, with closed sub-forest, presenting the record of 170 species, 129 genera and 46 families (Ferreira et al., 2013Ferreira EVR, Prata APN, Mello AA. Floristic list from a Caatinga remnant in Poço Verde, Sergipe, Brazil. Check List 2013; 9(6): 1354-1360. 10.15560/9.6.1354
https://doi.org/10.15560/9.6.1354...
).

Figure 1
Location of the study area and distribution of georeferenced plots for the Caatinga fragment in the municipality of Poço Verde, Sergipe, Brazil.

In the fragment under study, past records point to the extraction of medium-impact vegetation inside and around this fragment (Fernandes et al., 2015Fernandes MRM, Matricardi EAT, Almeida AQ, Fernandes MM. Mudanças do uso e de cobertura da terra na região semiárida de Sergipe. Floresta e Ambiente 2015; 22(4): 472-482. 10.1590/2179-8087.121514
https://doi.org/10.1590/2179-8087.121514...
; Ferreira et al., 2013Ferreira EVR, Prata APN, Mello AA. Floristic list from a Caatinga remnant in Poço Verde, Sergipe, Brazil. Check List 2013; 9(6): 1354-1360. 10.15560/9.6.1354
https://doi.org/10.15560/9.6.1354...
). Currently, anthropic impacts are restricted to the selective extraction of wood, animal husbandry (causing herbivory and overgrazing) and to fires (personal observation).

The study area, located in the backcountry pediplane, presents dissected surfaces, residual mountains at an altitude of 273 meters (Sergipe, 2014Sergipe, Secretaria de Estado do Planejamento Orçamento e Gestão. Enciclopédia dos municípios sergipanos. Aracaju: Seplag; 2014.) and Planasol, Halomorphic and Regosol soils (Sergipe, 1997Sergipe, Secretaria de Estado do Planejamento e da Ciência e Tecnologia. Perfis municipais: Poço Verde. Aracaju: Supes; 1997.).

The climate of the region is characterized as As type, according to the Köppen climatic classification (Alvares et al., 2013Alvares CA, Stape JL, Sentelhas PC, Moraes Gonçalves JL, Sparovek G. Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift 2013; 22(6): 711-728. 10.1127/0941-2948/2013/0507
https://doi.org/10.1127/0941-2948/2013/0...
; Sergipe, 2011Sergipe, Secretaria de Estado do Planejamento Orçamento e Gestão. Sergipe em dados. Aracaju: Seplag; 2011.). The average annual temperature is 23.7°C, with average annual precipitation of 780 mm (Sergipe, 1997Sergipe, Secretaria de Estado do Planejamento e da Ciência e Tecnologia. Perfis municipais: Poço Verde. Aracaju: Supes; 1997., 2011Sergipe, Secretaria de Estado do Planejamento Orçamento e Gestão. Sergipe em dados. Aracaju: Seplag; 2011.).

2.2 Data collection

Initially, annual precipitation, temperature and mean maximum temperature data for the study region were obtained for the period from 1960 to 2007 (historical average) and from 2008 to 2015, using online databases (INMET, c2016Instituto Nacional de Meteorologia - INMET (BR). Homepage [Internet]. c2016 [cited 2016 Aug. 7]. Available from: Available from: http://www.inmet.gov.br/
http://www.inmet.gov.br/...
; Sergipe, c2016Sergipe, Secretaria de Meio Ambiente e de Recursos Hídricos. Centro de Meteorologia de Sergipe [Internet]. c2016 [cited 2016 Aug. 7]. Available from: www.semarh.se.gov.br/meteorologia).

The evaluation of vegetation structure and composition was performed by comparing two surveys performed in 2011 (t0) and 2015 (t1), and the same plots and methodology were used in both surveys. In t0, plots were installed according to the fixed area method, and 30 square plots of 20 m × 20 m (400 m²) were randomly distributed (Figure 1), which is the recommended size for inventory standardization in Caatinga (RMFC, 2005Rede de Manejo Florestal da Caatinga - RMFC. Protocolo de medições de parcelas permanentes. Recife: Associação de Plantas do Nordeste; 2005.). In t1, after relocating the demarcated plots in t0, each plot was re-sampled (further details on the methodology in Oliveira et al., 2018Oliveira EVS, Prata APN, Pinto AS. Caracterização e atributos da vegetação herbácea em um fragmento de Caatinga no estado de Sergipe, Brasil. Hoehnea 2018; 45(2): 159-172. 10.1590/2236-8906-70/2017
https://doi.org/10.1590/2236-8906-70/201...
).

In both surveys all individuals present inside the plots (including columnar cacti), with height above 1.30 m and circumference at breast height (CBH) greater than or equal to 6 cm were measured using a tape measure. Fertile and/or vegetative samples were collected from individuals not identified in the field and referred to the Herbarium of the Federal University of Sergipe (ASE). After herborization (according to Mori et al., 1985Mori SA, Silva LAM, Lisboa G, Coradin L. Manual de manejo do herbário fanerogâmico. Ilhéus: Centro de Pesquisas do Cacau; 1985.), the collected material was identified by comparison with exsiccates present in the collection of the ASE herbarium, considering a floristic listing published for the study area (Ferreira et al., 2013Ferreira EVR, Prata APN, Mello AA. Floristic list from a Caatinga remnant in Poço Verde, Sergipe, Brazil. Check List 2013; 9(6): 1354-1360. 10.15560/9.6.1354
https://doi.org/10.15560/9.6.1354...
).

2.3. Data analysis

For the survey of the current horizontal structure (t1), the classical phytosociological parameters (Mueller-Dombois & Ellenberg, 1974Mueller-Dombois D, Ellenberg H. Aims and methods of vegetation ecology. New York: John Wiley & Sons; 1974.) were estimated: density, frequency, dominance and importance value (IV). These calculations were performed with the aid of the Fitopac 2.2 software (Shepherd, 2010Shepherd GJ. Fitopac: versão 2.1.2.85. Campinas: Unicamp; 2010.). Shannon-Wiener diversity index (H’), Pielou-J equability (Magurran, 2004Magurran AE. Measuring biological diversity. Oxford: Blackwell Science; 2004.) and the floristic similarity using the Sørensen index (the latter between the two samplings - t1 and t0) were also calculated by means of the Past 2.17 software (Hammer et al., 2013Hammer Ø, Harper DAT, Ryan PD. Past: palaeontological statistics [Internet]. 2013 [cited 2017 Apr. 2]. Available from: Available from: http://bit.ly/2IkI5RE
http://bit.ly/2IkI5RE...
). The total increment in basal area was calculated by subtracting the values found between t1 and t0; the annual rate of this parameter was obtained by dividing its total increment by four. For the analysis of the diametric structure, individuals were distributed into diameter classes, using the class intervals calculated with formulas present in Spiegel (1976Spiegel MP. Estatística. São Paulo: McGraw-Hill; 1976.).

2.4. Statistical analyses

After verifying data normality using the Shapiro-Wilk test, statistical differences (α = 0.05) between the two periods (t0 and t1) were evaluated for richness, absolute density and basal area; all of these using the paired t test (Vieira, 1980Vieira S. Introdução à bioestatística. Rio de Janeiro: Campus; 1980.; Zar, 2010Zar JH. Biostatistical analysis. New Jersey: Prentice-Hall; 2010.). Differences in the proportion of IV for the three species with higher value of this index were tested using the McNemar’s chi-square test (x²) (Elzinga et al., 1998Elzinga CL, Salzer DW, Willoughby JH. Measuring and monitoring plant populations. Denver: Nature Conservancy; 1998.). The diversity index (H’) was compared between the two periods by the Hutcheson’s t test (Magurran, 2004Magurran AE. Measuring biological diversity. Oxford: Blackwell Science; 2004.). Except for the last test, carried out in the Past 2.17 software (Hammer et al., 2013Hammer Ø, Harper DAT, Ryan PD. Past: palaeontological statistics [Internet]. 2013 [cited 2017 Apr. 2]. Available from: Available from: http://bit.ly/2IkI5RE
http://bit.ly/2IkI5RE...
), all the others were performed with the aid of the R software (R Development Core Team, 2013R Development Core Team. R: a language and environment for statistical computing [Internet]. 2013 [cited 2016 Aug. 7]. Available from: Available from: http://www.R-project.orgS
http://www.R-project.orgS...
).

3. RESULTS

In the analysis period, the study region presented precipitation values below previous values and historical average. An annual increase in precipitation was observed between 2011 and 2014, with a marked reduction in 2015 (Figure 2). The mean temperature and mean maximum temperature values were above the historical average. Except for 2014 and 2015, temperature remained relatively constant in the study period in relation to the previous year (2010), although with higher average maximum temperature variation (Figure 2).

Figure 2
Historical average (1960-2007), total annual rainfall, average temperature and average maximum annual temperature observed for the study region in the municipality of Poço Verde, Sergipe, Brazil.

After four years, there was a 0.74% increase in density, from 2,700.8 to 2,720.8 individuals.ha-1, but it was not significant (df = 29, t = 0.54, p > 0.58). The basal area had an increase of 0.62 m².ha-1 (4.82%) and annual of 0.16 m².ha-1 (1.21%), from 12.86 m².ha-1 to 13.48 m².ha-1, but the difference was not significant (df = 29, t = 0.98, p > 0.33). The variations of density and basal area were lower than those observed in a Caatinga area with evaluation period similar to that of this study; and in relation to other SDTF areas, variations observed were lower or similar (Table 1).

Table 1
Relative variations of density and basal area in seasonally dry tropical forests, including the Caatinga fragment in the municipality of Poço Verde, Sergipe, Brazil.

In both surveys, the majority of individuals are concen-trated in the three small diameter classes; thin individuals (< 14 cm) continue predominating in the community, as well as a reduced number of individuals in the larger diameter classes. However, there is a trend of diameter increase for individuals in the intermediate classes at t1 (Figure 3). An increase in the number of individuals in the first diameter class (115 or 6%) and decrease in the second (92 or 13%) and third was observed (33 or 13%), probably due to the growth of individuals of the latter two classes (Figure 3).

Figure 3
Distribution of number of individuals (a) and number of logarithmized individuals (b) by diameter classes between evaluation periods (t0 and t1) for the Caatinga fragment in the municipality of Poço Verde, Sergipe, Brazil.

The difference in the Shannon-Wiener (H’) diversity indexes between evaluation periods was not significant (df = 6463.2; t = −0.91; p > 0.35), as well as for richness (df = 29; t = 0.41; p > 0.28). These indexes, as well as the Pielou equability (J), remained stable after four years (Figure 4). In addition, no significant changes in floristic composition were observed between evaluation periods. Only four species were not found in the current survey (t1): Senegalia sp., Solanum sp., Handroanthus impetiginosus (Mart. ex DC.) Mattos and an indeterminate species. In turn, two species were unique in the current survey (t1): Vasconcellea quercifolia A.St.-Hil. and Clusia dardanoi G. Mariz & Maguire (Figure 4).

Figure 4
Venn diagram of floristic indexes evaluated for the analyzed period (t0 and t1) in the Caatinga fragment in the municipality of Poço Verde, Sergipe, Brazil, including Shannon-Wiener diversity (H’), Pielou equability (J), number of unique and common species in each period and temporal floristic similarity (Sørensen coefficient).

Fabaceae and Euphorbiaceae families remained the most abundant (17 and 10 species, respectively), with higher number of individuals (802 and 930, respectively) and, consequently, those with the highest relative IV (18.7% and 12.9% respectively) between evaluation periods. The number of families remained the same between the two evaluation periods (22). Two families (Bignoniaceae and Solanaceae) disappeared and two new ones (Caricaceae and Clusiaceae) were included. Apart from these families, the order for specific richness remained the same between the two evaluation periods. Regarding the ordering of families by number of individuals, small changes were observed and only Malvaceae, Cactaceae and Rubiaceae families increased in importance.

Alterations in the community structure were restricted to the density and absolute frequency of some species, which caused small changes in the IV parameter. Some species increased in density (Poincianella pyramidalis (Tul.) L.P.Queiroz, Psidium schenckianum Kiaersk. and Mimosa acutistipula (Mart.) Benth.), while others decreased (Myracrodruon urundeuva Allemão and Commiphora leptophloeos (Mart.) J.B.Gillett). However, because changes were small (ranging from two to eleven individuals), no major changes in the ordering of the species by IV were observed. Only for the species Guettarda angelica Mart. ex Müll.Arg. more significant changes were observed, with an increase in IV, probably due to the recruitment of individuals, since its density increased fourfold. Psidium schenckianum Kiaersk. (Myrtaceae), Cedrela odorata L. (Meliaceae) and Poincianella pyramidalis (Tul.) L.P.Queiroz (Fabaceae) continue to be the species with the highest relative IV, accounting for 28.15% of IV in the entire community. No significant differences were observed for the IV of P. schenckianum (df = 1; x2 = 2.10; p > 0.14), C. odorata (df = 1; x2 = 0.30; p > 0.58) and P. pyramidalis (df = 1; x2 = 2.10; p > 0.14) between evaluation periods.

4. DISCUSSION

In studies carried out in Brazilian SDTF, temporal changes in vegetation composition and structure are mainly related to the history of the evaluated area. Changes in quantitative parameters such as richness, H’, J, basal area and/or coverage, density and phytosociological indexes by species, and qualitative, such as species and family composition, were influenced by factors such as presence and anthropization level and the regeneration time (Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
; Fernandes et al., 2015Fernandes MRM, Matricardi EAT, Almeida AQ, Fernandes MM. Mudanças do uso e de cobertura da terra na região semiárida de Sergipe. Floresta e Ambiente 2015; 22(4): 472-482. 10.1590/2179-8087.121514
https://doi.org/10.1590/2179-8087.121514...
; Silva et al., 2009Silva MR, Araujo GM. Dinâmica da comunidade arbórea de uma floresta semidecidual em Uberlândia, MG, Brasil. Acta Botanica Brasilica 2009; 23(1): 49-56. 10.1590/S0102-33062009000100006
https://doi.org/10.1590/S0102-3306200900...
; Werneck & Franceschinelli, 2004Werneck MS, Franceschinelli EV. Dynamics of a dry forest fragment after the exclusion of human disturbance in southeastern Brazil. Plant Ecology 2004; 174(2): 337-346. 10.1023/B:VEGE.0000049112.06259.4c
https://doi.org/10.1023/B:VEGE.000004911...
; Werneck et al., 2000Werneck MS, Franceschinelli EV, Tameirão-Neto E. Changes in the floristic and structure of a dry deciduous forest during a period of 4-years (1994-1998), southeastern Brazil. Brazilian Journal of Botany 2000; 23(4): 401-413. 10.1590/S0100-84042000000400006
https://doi.org/10.1590/S0100-8404200000...
).

Considering that temporal variations in vegetation are associated with the intensity of the anthropic action, it is believed that the results found in the fragment indicate little anthropization (Bhat et al., 2000Bhat DM, Naik MB, Patagar SG, Hegde GT, Kanade YG, Hegde GN et al. Forest dynamics in tropical rain forests of Uttara Kannada district in Western Ghats, India. Current Science 2000; 79(7): 975-985.; Cavalcanti & Rodal, 2010Cavalcanti ADC, Rodal MJN. Efeito de borda e dinâmica de plantas lenhosas em áreas de Caatinga em Carnaubais, RN. Revista Caatinga 2010; 23(2): 41-50.; Cavalcanti et al., 2009Cavalcanti ADC, Rodal MJN, Sampaio EVSB, Costa KCC. Floristic and structural changes after five years in a Caatinga community in Pernambuco state, Brazil. Acta Botanica Brasilica 2009; 23(4): 1210-1212. 10.1590/S0102-33062009000400032
https://doi.org/10.1590/S0102-3306200900...
; Condit et al., 1992Condit R, Hubbell SP, Foster RB. Short-term dynamics of a neotropical forest. BioScience 1992; 42(11): 822-828. 10.2307/1312081
https://doi.org/10.2307/1312081...
; Pereira et al., 2001Pereira IM, Andrade LA, Costa JRM, Dias JM. Natural regeneration in a Caatinga fragment under different disturbance levels. Acta Botanica Brasilica 2001; 15(3): 413-426. 10.1590/S0102-33062001000300010
https://doi.org/10.1590/S0102-3306200100...
; Sampaio et al., 1993Sampaio EVSB, Salcedo IH, Kauffman JB. Effect of different fire severities on coppicing of Caatinga vegetation in Serra Talhada, PE, Brazil. Biotropica 1993; 25(4): 452-460. 10.2307/2388868
https://doi.org/10.2307/2388868...
; Silva et al., 2004Silva CT, Reis GG, Reis MGF, Silva E, Chaves RA. Avaliação temporal da florística arbórea de uma floresta secundária no município de Viçosa, Minas Gerais. Revista Árvore 2004; 28(3): 429-441. 10.1590/S0100-67622004000300014
https://doi.org/10.1590/S0100-6762200400...
). The temporal maintenance of all quantitative parameters analyzed and the minimal changes in the structure of some species and in the floristic composition allow inferring that the vegetation remained stable after four years.

In Caatinga anthropization may not exceed the resilience capacity of the vegetation; low anthropization levels allow for rapid recovery of plants, maintaining vegetation stability against periodic disturbances (Holanda et al., 2015Holanda AC, Lima FTD, Silva BM, Dourado RG, Alves AR. Estrutura da vegetação em remanescentes de Caatinga com diferentes históricos de perturbação em Cajazeirinhas (PB). Revista Caatinga 2015; 28(4): 142-150. 10.1590/1983-21252015v28n416rc
https://doi.org/10.1590/1983-21252015v28...
; Sampaio et al., 1993Sampaio EVSB, Salcedo IH, Kauffman JB. Effect of different fire severities on coppicing of Caatinga vegetation in Serra Talhada, PE, Brazil. Biotropica 1993; 25(4): 452-460. 10.2307/2388868
https://doi.org/10.2307/2388868...
). Knowing the factors that contribute to the stability of the Caatinga vegetation provides subsidies for rational exploration models (Maia et al., 2003Maia ACD, Iannuzzi L, Nobre CEB, Albuquerque CMR. Padrões locais de diversidade de Cerambycidae (Insecta, Coleoptera) em vegetação de caatinga. In: Leal IR, Tabarelli M, Silva JMC, editors. Ecologia e conservação da caatinga. Recife: Editora UFPE; 2003. p. 391-434.; Pereira et al., 2001Pereira IM, Andrade LA, Costa JRM, Dias JM. Natural regeneration in a Caatinga fragment under different disturbance levels. Acta Botanica Brasilica 2001; 15(3): 413-426. 10.1590/S0102-33062001000300010
https://doi.org/10.1590/S0102-3306200100...
). Thus, the sustainable use of natural resources of the study fragment is possible through planned forest management, guaranteeing source of income for the local population. The direct use of natural resources by local populations without affecting ecosystem support capacity constitutes an excellent conservation strategy (Brasil, 2008Brasil, Ministério do Meio Ambiente. Uso sustentável e conservação dos recursos florestais da caatinga. Brasília, DF: Serviço Florestal Brasileiro; 2008.; Diegues, 2001Diegues ACS. O mito moderno da natureza intocada. São Paulo: Hucitec; 2001.).

The stability observed in this study reflects differences between evaluation periods and does not mean that the community remained static over time, since changes constantly occur in time and space (Pinto & Hay, 2005Pinto JRR, Hay JDV. Mudanças florísticas e estruturais na comunidade arbórea de uma floresta de vale no Parque Nacional da Chapada dos Guimarães, Mato Grosso, Brasil. Revista Brasileira de Botânica 2005; 28(3): 523-539. 10.1590/S0100-84042005000300010
https://doi.org/10.1590/S0100-8404200500...
). For example, fluctuations in community structure and composition may have occurred in response to climatic variability observed for the period. The alternation of annual precipitation exerts a strong influence on the forest dynamics, causing fluctuations in the balance between mortality and recruitment (Carvalho & Felfili, 2011Carvalho FA, Felfili JM. Variações temporais na comunidade arbórea de uma floresta decidual sobre afloramentos calcários no Brasil Central: composição, estrutura e diversidade florística. Acta Botanica Brasilica 2011; 25(1): 203-214. 10.1590/S0102-33062011000100024
https://doi.org/10.1590/S0102-3306201100...
).

The distribution of individuals per diameter class in both evaluation periods follows the inverted “J” pattern, considered characteristic of native forests (Oliveira et al., 2013Oliveira DG, Ferreira RA, Mello AA, Almeida ES. Estrutura diamétrica da vegetação arbustivo-arbórea no entorno de nascentes da bacia hidrográfica do rio Piauitinga, Salgado-SE, Brasil. Biotemas 2013; 26(2): 19-31. 10.5007/2175-7925.2013v26n2p19
https://doi.org/10.5007/2175-7925.2013v2...
). Although a large number of small and thin individuals may indicate the occurrence of past disturbances (Nunes et al., 2003Nunes YRF, Mendonça AVR, Botezelli L, Machado ELM, Oliveira-Filho AT. Variações da fisionomia, diversidade e composição de guildas da comunidade arbórea em um fragmento de floresta semidecidual em Lavras, MG. Acta Botanica Brasilica 2003; 17(2): 213-229. S0102-33062003000200005
https://doi.org/S0102-33062003000200005...
), the observed tendency in the growth of the individuals’ diameter in intermediate classes in t1 may indicate successional evolution (Chagas et al., 2001Chagas RK, Oliveira-Filho AT, Van Den Berg E, Scolforo JRS. Dinâmica de populações arbóreas em um fragmento de floresta estacional semidecidual montana em Lavras, Minas Gerais. Revista Árvore 2001; 25(1): 39-57., Paula et al., 2002Paula A, Silva AF, Souza AL, Santos FAM. Alterações florísticas ocorridas num período de quatorze anos na vegetação arbórea de uma Floresta Estacional Semidecidual em Viçosa-MG. Revista Árvore 2002; 26(6): 743-749. 10.1590/S0100-67622002000600010
https://doi.org/10.1590/S0100-6762200200...
).

In the study fragment, records indicate that the vegetation cutting is selective, prioritizing trees of better quality wood, such as Myracrodruon urundeuva Allemão and Handroanthus impetiginosus (Mart. Ex DC) Mattos (Carvalho, 2003Carvalho PER. Aroeira verdadeira. Colombo: Embrapa Florestas; 2003.; Peixoto et al., 2002Peixoto AM, Souza JSI, Toledo FF, Reichardt K, Molina-Filho J. Enciclopédia agrícola brasileira: I-M. São Paulo: Edusp; 2002.; Silva et al., 2014Silva MAS, Lopes SF, Vitorio LAP, Santiago RR, Mattos EA, Trovão DMBM. Plant functional groups of species in semiarid ecosystems in Brazil: wood basic density and SLA as an ecological indicator. Brazilian Journal of Botany 2014; 37(3): 229-237. 10.1007/s40415-014-0063-4
https://doi.org/10.1007/s40415-014-0063-...
) and those easy to handle, such as Commiphora leptophloeos (Mart.) J.B.Gillett (Carvalho, 2009Carvalho PER. Imburana-de-Espinho: Commiphora leptophloeos. Colombo: Embrapa Florestas; 2009.). It is believed that the extraction of individuals of these three species was the main reason for the temporal decrease in the density of M. urundeuva and C. leptophloeos and the disappearance of H. impetiginosus. The latter could be considered locally rare due to the low population density (Rambaldi & Oliveira, 2003Rambaldi D, Oliveira DAS. Fragmentação de ecossistemas: causas, efeitos sobre a biodiversidade e recomendações de políticas públicas. Brasília, DF: Ministério do Meio Ambiente; 2003.), which made it very inclined to local extinction (Primack & Rodrigues, 2001Primack R, Rodrigues E. Biologia da conservação. Londrina: Planta; 2001.).

In the study area, Guettarda angelica Mart. ex Müll.Arg was not found in the sub-forest, only developing in edges and/or clearings, probably because it preferred to colonize these environments. Therefore, it is believed that the extraction of wood from canopy species may have allowed a better development of individuals due to the greater light input, which would justify their temporal increases in density and IV.

Also, Guettarda angelica can be considered an example of a “winning species”, that is, the one that benefits from the anthropization of landscapes, rapidly proliferating in these environments. In contrast, Myracrodruon urundeuva and Handroanthus impetiginosus can be considered as examples of “losing species”, due to the selective cutting of wood by the local population, causing population reduction, consequently leading to their disappearance (Tabarelli et al., 2012Tabarelli M, Aguiar AV, Ribeiro CM, Metzger JP. A conversão da Floresta Atlântica em paisagens antrópicas: lições para a conservação da diversidade biológica das florestas tropicais. Interciência 2012; 37(2): 88-92.).

The current state of knowledge does not yet allow understanding whether the Caatinga biome is more stable over time than other plant formations. In addition to factors such as the time scale of investigation and presence and intensity of disturbances, the time of the disorder in relation to the investigation should also be considered as determinant for the detection of stability in temporal evaluation studies. Continuous vegetation monitoring over the evaluation period, i.e., four years, is advisable because it improves the ability to infer about the dynamic processes of the vegetation (Braga & Rezende, 2007Braga FMS, Rezende AV. Dinâmica da vegetação arbórea da Mata de Galeria do Catetinho, Brasília-DF. Cerne 2007; 13(2): 138-148.; Mognon et al., 2012Mognon F, Dallagnol F, Sanquetta C, Corte AP, Maas G. Uma década de dinâmica florística e fitossociológica em floresta ombrófila mista montana no sul do Paraná. Revista de Estudos Ambientais 2012; 14(1): 43-59. 10.7867/1983-1501.2012v14n1p43-59
https://doi.org/10.7867/1983-1501.2012v1...
).

5. CONCLUSIONS

Although the occurrence of possible fluctuations in the community in response to climatic variability observed during the evaluation period is not ruled out, the vegetation under study remained stable over time, tolerating current anthropization levels. Public policies that provide planned management of the study area by the local population should be established, thus becoming an alternative for the conservation of this remnant. Continued monitoring of the area is necessary, and further studies focusing on population dynamics are recommended.

ACKNOWLEDGEMENTS

The authors are grateful to the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Capes) for concession of the masters scholarship to the first author; to the Postgraduate Support Program (Proap-Capes) for financial support; to Eladio dos Santos, Mauricio Larios Ulloa and Irandir Félix de Jesus for fieldwork assistance; to José Bráz de Jesus (Mr. Mizé) for hospitality and aid in the field research; to the Universidade Federal do Sergipe (UFS) for providing transport and laboratory facilities; finally, to the ASE Herbarium staff for their help.

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  • FINANCIAL SUPPORT This study was partly financed by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Capes) - Finance Code 001. E.V.S.O. was supported by CAPES/FAPITEC (#88881.157451/2017-01)

Publication Dates

  • Publication in this collection
    08 May 2020
  • Date of issue
    2020

History

  • Received
    16 Aug 2017
  • Accepted
    26 Aug 2018
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