SciELO - Scientific Electronic Library Online

 
vol.28 issue4Poaceae communities in the savannas of the Amazon Estuary in BrazilHost specificity and experimental assessment of the early establishment of the mistletoe Phoradendron crassifolium (Pohl ex DC.) Eichler (Santalaceae) in a fragment of Atlantic Forest in southeast Brazil author indexsubject indexarticles search
Home Pagealphabetic serial listing  

Services on Demand

Journal

Article

Indicators

Related links

Share


Acta Botanica Brasilica

Print version ISSN 0102-3306

Acta Bot. Bras. vol.28 no.4 Belo Horizonte Oct./Dec. 2014

http://dx.doi.org/10.1590/0102-33062014abb3432 

ARTICLES

 

Structure and floristic diversity of remnant semideciduous forest under varying levels of disturbance

 

 

Darlene GrisI, *; Lívia Godinho TemponiII; Geraldo Alves Damasceno JuniorIII

IUniversidade Estadual do Oeste do Paraná, Programa de Pós-Graduação em Conservação e Manejo de Recursos Naturais, Cascavel, PR
IIUniversidade Estadual do Oeste do Paraná, Herbário UNOP, Cascavel, PR
IIIUniversidade Federal de Mato Grosso do Sul, Departamento de Botânica, Campo Grande, MS

 

 


ABSTRACT

The perturbation of Neotropical forests generates large disturbances in biological communities. The species that suffer least from the resulting habitat fragmentation are the pioneers, because they possess greater ability to inhabit disturbed environments. Therefore, it is expected that species diversity will be greater in areas subjected to intermediate disturbance, such as the opening of gaps, because a large number of pioneer species will develop and coexist with species of more advanced successional stages. This study aimed to compare two forest remnants that differed in size and disturbance intensity, in order to determine the effects of disturbances on species diversity and the size ratios of individual trees. This was accomplished with comparative analyses of diversity, richness and diameter ratios obtained for 10 plots at two semideciduous forest sites. We recorded a total of 85 species, of which 70 were in the private nature reserve Fazenda Santa Maria, 58 were in Iguaçu National Park, and 43 were at both sites. Diversity was greater in the more disturbed remaining forest, because this area showed higher species richness, which is in accordance with some premises of the intermediate disturbance theory. There was also an increase in the number of pioneer individuals, and the less disturbed area showed individuals with larger diameters, which is likely attributable to the removal of large individuals from the more disturbed area during the anthropogenic process of forest modification.

Key words: Intermediate disturbance, species richness, Neotropical forest, tree communities


 

 

Introduction

The constant disturbance of tropical forests, whether by natural events or human activities (destruction of vegetation and conversion or loss of habitat), has led to the fragmentation of these forests into small isolated patches. Disturbances occur more frequently in the fragments, drastically altering species richness, diversity and forest structure (Tanizaki-Fonseca & Moulton 2000; Campos 2006). Tropical forests in general have high spatial and temporal heterogeneity, which strongly influence species distribution patterns. Plant species in tropical forests grow and establish in a mosaic of different successional stages, whereby the distribution and density of plant populations are influenced by disturbance dynamics (Brokaw 1985; Young & Hubbell 1991).

Upon the occurrence of a natural disturbance in a forest, such as canopy openings or border effects, pioneer species increase in density in response to increased light availability. The same phenomenon occurs in response to anthropogenic disturbances such as selective logging or changes in land use, and varying intensities and frequencies of these disturbances have different effects on a plant community (Hill & Curran 2003; Laurance et al. 2006; Bongers et al. 2009). One observed effect is the decrease in diameter of trees and the occurrence of many small individuals (Gomes et al. 2004; Villela et al. 2006), due to the increased number of young individuals and pioneer species, and reduction of the number of large individuals by selective logging, for example. Such alterations in vegetation structure as a result of disturbance have been observed in several studies on effects of fragment size, edge dynamics, and canopy gaps on woody plant composition in the Atlantic Forest (Alves & Santos 2002; Bertoncini & Rodrigues 2008; Lopes et al. 2009; Oliveira et al. 2004; Oliveira et al. 2008; Ribeiro et al. 2009; Santos et al. 2008).

Considering those disturbances to the vegetation, it is expected that, as proposed in the intermediate disturbance theory (Connell 1978), higher species diversity may be maintained at moderate levels of disturbance frequency and intensity because superior competitors, which dominate communities at more advanced successional stages, are more susceptible to disturbance. Therefore, gaps are open for the development of pioneer species, which tolerate unfavorable conditions. Conversely, when disturbance levels are very high, after intense human intervention, for example, all species are at risk of being eliminated. This study therefore aimed to evaluate the effects of disturbance on tree species richness and diversity, as well as on the proportional distribution by diameter, in forest fragments, comparing two sites: a large area of continuous forest that is less disturbed; and a nearby forest fragment that is smaller and more heavily disturbed by human activities.

 

Material and Methods

Study area

To compare effects of increased disturbance on plant community in forest remnants, we selected two areas. One was a small private nature reserve, the Fazenda Santa Maria, hereafter referred to as the FSM, which was heavily disturbed, whereas the other was less disturbed, representing a large tract of more advanced stage forest, Iguaçu National Park (INP).

The FSM is located in the municipality of Santa Terezinha do Itaipu, in the state of Paraná, and covers 242 ha. This area has been protected since 1955 and was designated a private nature reserve in 1997. The fragment had suffered a moderate degree of human intervention prior to the year 1955, when several species of economic interest were extracted and forest loss in the surrounding land was pronounced. Therefore, edge effects are evident in the remnant, as are gaps caused by logging.

The INP is an ecological reserve in the southwestern region of the state of Paraná, created in 1939 and currently encompassing 185,262 ha (Salamuni et al. 2002). It contains patches of Araucaria forest (rain forest) and semideciduous forest (Veloso 1991; IBGE 2012).

Both sampled sites are fragments of semideciduous forest and are on the Third Paraná Plateau, near Foz do Iguaçu, with elevations ranging from 120 m to 540 m (Santos et al. 2006; Maack 2012). At both sites, the soils are red Oxisols and the climate is humid subtropical (Köppen type Cfa), with an annual mean temperature of 21.5°C and annual rainfall of approximately 1800 mm, of which two thirds are distributed from October to March, mainly in December and January (Bhering & Santos 2008; SIMEPAR 2011).

Sampling and Analysis

We installed 10 permanent plots using a 20 × 20 m spacing regime, for a total of 0.4 ha in each forest remnant. Plots were demarcated at a distance of approximately 400 m from the forest edge, preventing any clearing or trail opening. Plots in the INP were located between the coordinates 25°31'56.79"S;54°17'32.14"W and 25°31'55.56"S;54°17'22.61"W, and plots in the FSM were located between the coordinates 25°29'30.70"S;54°21'30.61"W and 25°29'32.94"S;54°21'41.36"W, and the distance between the two areas is approximately 8 km. Trees whose circumference at breast height (CBH) was 15 cm were included in the sampling, as per Rodrigues & Gandolfi (2004), as were bifurcated stems, if at least one had a diameter 15 cm. The selected individuals were numbered and observed monthly from September 2010 to November 2011, in order to sample the maximum number of species in flower.

When found, fertile specimens were collected, identified and processed using usual herbarium techniques (Mori et al. 1989; Bridson & Forman 2004). Voucher specimens were registered at Western Paraná State University Herbarium (code, UNOP). Sterile individuals were also collected, identified and added to a sterile collection in the UNOP Herbarium. For identification of the plant material, we used regional checklists, as well as identification keys, specific bibliography (Lorenzi 2002a, 2002b, 2010; Ramos et al. 2008) and comparison with specimens in the UNOP Herbarium and the Herbarium of the Curitiba Municipal Botanical Museum (code, MBM). The lists were produced with the botanical families organized according to the Angiosperm Phylogeny Group III guidelines (APG III 2009) and the species names and the authors organized according to the List of Species in the Flora of Brazil (Lista de Espécies da Flora do Brasil 2013).

We analyzed the data using the programs R, version 3.0.1 (R Development Core Team 2013), PAST (Hammer et al. 2001) and FITOPAC, version 2.1.2 (Shepherd 2010). Shannon diversity and evenness indices were obtained for the remnants, as were importance values (IVs) for each species. The IV is an outcome of phytosociological analysis, obtained by summing the percentages of relative density, relative frequency and relative dominance (Shepherd 2010). To compare diversity indices, we used Hutcheson's t-test (Hutcheson 1970; Zar 1999).

Rarefaction curves, with their respective 95% confidence intervals, were built according to the number of individuals, in order to compare the species richness estimated for both areas. Results of rarefaction curves constitute an unbiased comparison with other studies, because they are not influenced by variations in the density of individuals and can simulate small sample sizes (Colwell & Coddington 1994; Gotelli & Colwell 2001). Therefore, the curves were constructed to compare the estimated richness between sites, in case the sampling number were equal between them.

On the basis of data in the literature, were classified the species, by successional stage, as pioneer or non-pioneer species. As is well known, sites that are more disturbed, have canopy gaps or are under greater edge effects are colonized by greater numbers of species and individuals. Therefore, we made two calculations to determine the differences in patterns between the two sites: the chi-square test for total non-pioneer and pioneer species per site; and the chi-square test for proportions of individuals belonging to the pioneer and non-pioneer groups per site.

To examine the difference between the mean diameters in the studied fragments, we used the nonparametric Wilcoxon test at a 5% significance level to identify differences in distributions of tree diameters in the FSM and INP (Zar 1999). In addition, to determine the differences between the two sites, in terms of the diameter ratios, the number of individuals per class at intervals of 5 cm was calculated, as proposed by Soares et al. (2006), starting from a minimum inclusion diameter of 4.77 cm, corresponding to the minimum CBH of 15 cm.

 

Results

In the study area as a whole, we found a total of 1032 individuals belonging to 85 species (Tab. 1): 70 species were present in the FSM, 58 were present in the INP, and 43 were present at both sites. We sampled only one exotic species, Citrus X aurantium, which occurred in the INP.

The first 10 species, in order of IV, accounted for 50% of the total, reflecting a few species with high IVs. The species that were the most well represented, in terms of the number of individuals, were Euterpe edulis, Sorocea bonplandii and Guarea kunthiana (in both areas); Chrysophyllum gonocarpum and Balfourodendron riedelianum (in the FSM only); and Citrus X aurantium and Trichilia catigua (in the INP only).

The Shannon-Wiener diversity values were 2.71 for the IPN and 3.37 for the FSM (Tab. 2), and the difference between the two was statistically significant (t=6.48, p=1.3311). The evenness values were 0.67 for the INP and 0.79 for the FSM. The higher richness in the FSM was observed in the rarefaction curves (Fig. 1).

 

 

As can be seen in Tab. 3, there was no significant difference between the two sites with respect to the number of species belonging to the analyzed successional groups, pioneer (χ2=1.50, p=0.22) and non-pioneer (χ2=0.35, p=0.56). Furthermore, the number of individuals belonging to the non-pioneer group did not differ between the two areas (χ2=0.34, p=0.56). However, the FSM had a significantly higher number of individuals of pioneer species than did the INP (χ2=19.63, p=0.935) (Tab. 3).

The diameter class distribution was significantly different between the two sites (W=110215.5, p=4.266). Despite the fact that individuals with a diameter < 15 cm accounted for 80% of the individuals in the FSM (n = 412) and 86% of the individuals in the INP (n = 447), the INP presented some specimens with a diameter > 70 cm, whereas the FSM did not (Fig. 2).

 

 

Discussion

The two analyzed areas shared important species for the Atlantic Forest Biome, such as Aspidosperma polyneuron, Cedrela fissilis and Balfourodendron riedelianum. These species are rare and endangered because they are attractive targets for illegal logging, which underscores the importance of preserving the remaining fragments of semideciduous forest (Hatschbach & Ziller 1995; Ribas et al. 2003; IUCN 2009).

Ten species collectively accounted for 50% of the total IV, and seven of those species occurred at both study sites. These high IVs may be related to large diameters, combined with a reasonable number of individuals, reflecting a higher basal area, as noted for Aspidosperma polyneuron, Alchornea triplinervia, Cabralea canjerana and Chrysophyllum gonocarpum, or to small diameters, combined with high numbers of individuals, as noted for Euterpe edulis; Guarea kunthiana and Sorocea bonplandii. In addition, excepting G. kunthiana and S. bonplandii, the five species with the highest IVs were also the most representative for basal area at both sites.

The Shannon-Wiener diversity indices differed between the INP and the FSM. Although other studies of semideciduous forest (Tab. 2) have yielded results consistent with those we obtained for the FSM, the index we obtained for the INP was lower than those reported in all of the studies we reviewed. The same pattern was observed for the evenness index for the INP, which was lower in the present study than in other studies, whereas the evenness index for the FSM was consistent with those of the same studies (Ivanauskas et al. 1999; Bianchini et al. 2003; Jurinitz & Jarenkov 2003; Silva et al. 2004; Costa-Filho et al. 2006; Prado-Júnior et al. 2011).

Connell (1978) proposed that higher species diversity is maintained under intermediate levels of disturbance. Such disturbances occurring at moderate frequency and intensity should promote increased environmental heterogeneity, because pioneer species coexist with species of more advanced successional stages (shade-tolerant species), thus increasing diversity. Therefore, it was not surprising that the more heavily disturbed FSM site, with a history of more intense human activity, had higher diversity and more individuals of pioneer species, which also directly affect diversity, than did the INP site, given that degradation and loss of natural habitat at intermediate levels enable a wider range of species to become established (Hunter 1996; Wilson 1997; Hernandez-Stefanoni 2005). Nunes et al. (2003) found similar results in a study conducted in a semideciduous forest, where the borders of fragments were more diverse than were the interiors, due to the emergence of light-demanding pioneer species. In dry areas of a tropical forest, similar to our study area, Bongers et al. (2009) showed that increases in pioneer species and diversity were interconnected with increased disturbance, supporting the theory of intermediate disturbance. However, the authors found that the occurrence of disorders did not significantly alter the diversity in wetlands.

We can conclude that the alterations caused by disturbances in the FSM did not lead to a significant increase in the richness of pioneer species, as would have been expected, but rather in the number of pioneer individuals compared to the pattern observed in the INP. Disturbed areas are expected to show a pattern in which individuals in the smaller diameter classes are more numerous and there are fewer large trees. Individuals with a diameter < 15 cm predominated in the FSM and INP (respectively accounting for 80% and 86% of the total), this pattern (many individuals in the smaller diameter classes) being commonly observed in tropical forests (Meyer 1952). However, we observed a significant difference in the diameter distribution between the two sites, individuals with a CBH > 70 cm being found only in the INP. Although the FSM has a history of human disturbance, no timber extraction or other type of human activities has taken place there in the last 60 years. Nevertheless, this lack of large individuals may be due to mortality of older, larger individuals after intense disturbance or by timber extraction, as observed in remnants of Amazon rain forest (Laurance et al. 2000) and submontane Atlantic Forest (Carvalho & Nascimento 2009). Therefore, the exclusion of individuals of the canopy could favor an increase in the number of small individuals, suggesting that there was a change in the successional processes and in the current structural conformation in this forest fragment.

The results obtained in the present study seem to agree with some premises of the intermediate disturbance theory, showing, as expected, an increase in diversity in the area with a known history of disturbance (the FSM) in relation to a more conserved one (INP). It would be useful to investigate a third area, with very high disturbance, to have a comparative that could confirm the hypothesis, because other potential determinants were not evaluated. In the FSM, the increased diversity is related to the increase in species richness. The FSM also showed a pattern of individuals in the smaller diameter classes, which is likely attributable to the extraction of large individuals during the anthropogenic process of forest modification.

 

Acknowledgments

We are grateful to the researchers at the Curitiba Municipal Botanical Museum, for their assistance in the identification of specimens, to Adaíldo Policena, for the assistance in the fieldwork, and to Dr. Arnildo Pott, for reviewing the English. This study received financial support from the Parque Tecnológico Itaipu Ciência, Tecnologia e Inovação da Fundação Parque Tecnológico Itaipu - Brasil (PTI C&T/FPTI-BR, Itaipu Technology Park for Science, Technology and Innovation via the Itaipu Technology Park Foundation of Brazil; master's scholarship to DG); the Araucaria Foundation (Grant no. 223/2010); the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, São Paulo Research Foundation; Grant no. 2010/17400-3); the Brazilian Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, National Council for Scientific and Technological Development; Grant no. 562240/2010-1); and the Centro de Ciências Biológicas e da Saúde-Universidade Estadual do Oeste do Paraná (CCBS-Unioeste, Western Paraná State University Center for Biological and Health Sciences).

 

References

Alves, L.F. & Santos, F.A.M. 2002. Tree allometry and crown shape of four tree species in Atlantic rain forest, south-east Brazil. Journal of Tropical Ecology 18: 245-260.         [ Links ]

APG III. 2009 An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105-121.         [ Links ]

Bertoncini, A.P. & Rodrigues, R.R. 2008. Forest restoration in an indigenous land considering a forest remnant influence (Avai, São Paulo State, Brazil). Forest Ecology and Management 255: 513-521.         [ Links ]

Bhering, S.B. & Santos, H.G. (Orgs.). 2008. Mapa de Solos do Estado do Paraná - Legenda Atualizada. Rio de Janeiro, Embrapa Florestas, Embrapa Solos, IAPAR.         [ Links ]

Bianchini, E.; Popolo, R.S.; Dias, M.C. & Pimenta, J.A. 2003. Diversidade e estrutura de espécies arbóreas em área alagável do município de Londrina, Sul do Brasil. Acta Botanica Brasilica 17(3): 405-419.         [ Links ]

Bongers, F.; Poorter, L.; Hawthorne, W.D. & Sheil, D. 2009. The Intermediate Disturbance Hypothesis applies to tropical forests, but disturbance contributes little to tree diversity. Ecology Letters 12(8): 798-805.         [ Links ]

Bridson, D.M. & Forman, L. 2004. The Herbarium Handbook. 3rd ed. Kew, Royal Botanic Gardens.         [ Links ]

Brokaw, N. 1985. Gap-phase regeneration in a tropical forest. Ecology 66: 682-687.         [ Links ]

Campos, J.B. 2006. A fragmentação de ecossistemas, efeitos decorrentes e corredores de biodiversidade. Pp. 165-174. In: Campos, J.B., Tossulino, M.G.P. & Muller, C.R.C (Eds.). Unidades de Conservação: ações para a valorização da biodiversidade. Curitiba, Instituto Ambiental do Paraná         [ Links ].

Carvalho, F.A. & Nascimento, M.T. 2009. Estrutura diamétrica da comunidade e das principais populações arbóreas de um remanescente de Floresta Atlântica Submontana (Silva Jardim-RJ, Brasil). Revista Árvore 33(2): 327-337.         [ Links ]

Colwell, R.K. & Coddington, J.A. 1994. Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions of the Royal Society (Series B) 345: 101-118.         [ Links ]

Connell, J.H. 1978. Diversity in Tropical Rain Forests and Coral Reefs. Science 199(4335): 1302-1310.         [ Links ]

Costa-Filho, L.V.; Nanni, M.R. & Campos, J.B. 2006. Floristic and Phytosociological Description of a Riparian Forest and the Relationship with the Edaphic Environment in Caiuá Ecological Station - Paraná - Brazil. Brazilian Archives of Biology and Technology 49(5): 785-798.         [ Links ]

Gomes, A.P.C.; Souza, A.L. & Meira Neto, J.A.A. 2004. Alteração estrutural de uma área florestal explorada convencionalmente na bacia do Paraíba do Sul, Minas Gerais, nos domínios de Floresta Atlântica. Revista Árvore 28(3): 407-417.         [ Links ]

Gotelli, N.J. & Colwell, R.K. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters 4: 379-391.         [ Links ]

Hammer, O.; Harper, D.A.T. & Ryan, P.D. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1): 1-9.         [ Links ]

Hatschbach, G.G. & Ziller, S.R. 1995. Lista vermelha de plantas ameaçadas de extinção no estado do Paraná. Curitiba, Secretária de Estado do Meio Ambiente.         [ Links ]

Hernandez-Stefanoni, J.L. 2005. Relationships between landscape patterns and species richness of trees, shrubs and vines in a tropical forest. Plant Ecology 179: 53-65.         [ Links ]

Hill, J.L. & Curran, P.J. 2003. Area, shape and isolation of tropical forest fragments: effects on tree species diversity and implications for conservation. Journal of Biogeography 30: 1391-1403.         [ Links ]

Hunter, M.L. 1996. Fundamentals of conservation biology. Cambridge, Blackwell Science.         [ Links ]

Hutcheson, K. 1970. A test for comparing diversities based on the Shannon formula. Journal of Theoretical Biology 29: 151-154.         [ Links ]

IBGE - Instituto Brasileiro de Geografia e Estatística. 1992. Manual Técnico da Vegetação Brasileira. Rio de Janeiro, IBGE.         [ Links ]

IUCN - International Union for Conservation of Nature. 2009. IUCN Red List of Threatened Species. Available from: http://www.iucnredlist.org. Cited 2013 Aug 01.         [ Links ]

Ivanauskas, N.M.; Rodrigues, R.R. & Nave, A.G. 1999. Fitossociologia de um trecho de Floresta Estacional Semidecidual em Itatinga, SP, Brasil. Scientia Forestalis 56: 83-99.         [ Links ]

Jurinitz, C.F. & Jarenkow, J.A. 2003. Estrutura do componente arbóreo de uma floresta estacional na Serra do Sudeste, Rio Grande do Sul, Brasil. Revista Brasileira de Botânica 26(4): 475-487.         [ Links ]

Laurance, W.F.; Delamônica, P.; Laurance, S.G.; Vasconcelos, H.L. & Lovejoy, T.E. 2000. Rainforest fragmentation kills big trees. Nature 404: 836.         [ Links ]

Laurance, W.F.; Fernside, P.M.; Nascimento, H.E.M.; Laurance, G., Andrade, A.C.; Ribeiro, E.L. & Capretz, R.L. 2006. Rain forest fragmentation and the proliferation of successional trees. Ecology 87(2): 469-482.         [ Links ]

Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. 2013. Available from: http://floradobrasil.jbrj.gov.br. Cited 2013 Apr 03.         [ Links ]

Lopes, A.V.; Girão, L.C.; Santos, B.A.; Peres, C.A. & Tabarelli, M. 2009. Long-term erosion of tree reproductive trait diversity in edge-dominated Atlantic forest fragments. Biological Conservation 142: 1154-1165.         [ Links ]

Lorenzi, H. 2002a. Árvores Brasileiras: manual de identificação e cultivo de plantas arbóreas do Brasil, v. 1. São Paulo, Nova Odessa.         [ Links ]

Lorenzi, H. 2002b. Árvores Brasileiras: manual de identificação e cultivo de plantas arbóreas do Brasil, v. 2. São Paulo, Nova Odessa.         [ Links ]

Lorenzi, H. 2010. Árvores Brasileiras: manual de identificação e cultivo de plantas arbóreas do Brasil, v. 3. São Paulo, Nova Odessa.         [ Links ]

Maack, R. 2012. Geografia física do estado do Paraná. 4th ed. Ponta Grossa, Editora UEPG.         [ Links ]

Meyer, H.A. 1952. Structure, growth and drain in balanced uneven-aged forest. Journal of Forestry 50: 85-92.         [ Links ]

Mori, S.A.; Mattos-Silva, L.A.; Lisboa, G. & Coradin, L. 1989. Manual de manejo do herbário fanerogâmico. 2nd ed. Ilhéus, CEPLAC.         [ Links ]

Nunes, Y.R.F.; Mendonça, A.V.R.; Botezelli, L.; Machado, E.L.M. & Oliveira-Filho, A.T. 2003. Variações da fisionomia, diversidade e composição de guildas da comunidade arbórea em um fragmento de Floresta Semidecidual em Lavras, MG. Acta Botanica Brasilica 17(2): 213-229.         [ Links ]

Oliveira, M.A.; Grillo, A.S. & Tabarelli, M. 2004. Forest edge in the Brazilian Atlantic forest: drastic changes in tree species assemblages. Oryx 38: 389-394.         [ Links ]

Oliveira, M.A.; Santos, A.M.M. & Tabarelli, M. 2008. Profound impoverishment of the large-tree stand in a hyper-fragmented landscape of the Atlantic forest. Forest Ecology and Management 256: 1910-1917.         [ Links ]

Prado-Júnior, J.A.; Vale, V.S.; Oliveira, A.P.; Gusson, A.E.; Neto, O.C.D.; Lopes, S.F. & Schiavini, I. 2011. Estrutura da comunidade arbórea em um fragmento de Floresta Estacional Semidecidual, localizada na reserva legal da Fazenda Irara, Uberlândia, MG. Bioscience Journal 26(4): 638-647.         [ Links ]

R Core Team. 2013. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: http://www.R-project.org/. Cited 2013 Aug 02.         [ Links ]

Ramos, V.S.; Durigan, G.; Franco, G.A.D.C.; Siqueira, M.F. & Rodrigues, R.R. 2008. Árvores da Floresta Estacional Semidecidual. Guia de identificação de espécies. São Paulo, EDUSP - FAPESP.         [ Links ]

Ribas, L.L.F.; Zanette, F.; Kulchetscki, L. & Guerra, M.P. 2003. Estabelecimento de culturas assépticas de Aspidosperma polyneuron. Ciência Florestal 13(1): 115-122.         [ Links ]

Ribeiro, M.C.; Metzger, J.P.; Martensen, A.C.; Ponzoni, F.J. & Hirota, M.M. 2009. The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 142: 1141-1153.         [ Links ]

Rodrigues, R.R. & Gandolfi, S. 2004. Conceitos, Tendências e Ações para a Recuperação de Florestas Ciliares. Pp. 235-247. In: Rodrigues, R.R. & Leitão-Filho, H.F. (Eds.). Matas Ciliares: Conservação e Recuperação. São Paulo, EDUSP - FAPESP.         [ Links ]

Salamuni, R.; Salamuni, E.; Rocha, L.A. & Rocha, A.L. 2002. Parque Nacional do Iguaçu, PR - Cataratas de fama mundial. Pp. 313-312. In: Schobbenhaus, C.; Campos, D.A.; Queiroz, E.T.; Berbert-Born, M.L.C. (Eds.). Sítios Geológicos e Paleontológicos do Brasil. Brasília, DNPM/CPRM - Comissão Brasileira de Sítios Geológicos e Paleobiológicos (SIGEP).         [ Links ]

Santos, L.J.C.; Oka-Fiori, C.; Canali, N.E.; Fiori, A.P.; Silveira, C.T.; Silva, J.M.F. & Ross, J.L.S. 2006. Mapeamento geomorfológico do Estado do Paraná. Revista Brasileira de Geomorfologia 7: 3-11.         [ Links ]

Santos, B.A.; Peres, C.A.; Oliveira, M.A.; Grillo, A.; Alves-Costa; C.P. & Tabarelli, M. 2008. Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil. Biological Conservation 141: 249-260.         [ Links ]

Shepherd, G.J. 2010. FITOPAC 2.1.2: Manual do usuário. Campinas, UNICAMP.         [ Links ]

Silva, N.R.S.; Martins, S.V.; Meira-Neto, J.A.A. & Souza, A.L. 2004. Composição florística e estrutura de uma floresta estacional semidecidual montana em Viçosa, MG. Revista Árvore 28(3): 397-405.         [ Links ]

SIMEPAR- Sistema Meteorológico do Paraná, 2011. Daily data of rainfall and temperature for the town of Foz do Iguaçu for the period of 01/01/2010 to 01/01/2011.         [ Links ]

Soares, C.P.B.; Paula-Neto, F.; Souza, A.L. 2006. Dendrometria e inventário florestal. Viçosa, Editora UFV.         [ Links ]

Tanizaki-Fonseca, K. & Moulton, T.P. 2000. A fragmentação da Mata Atlântica no Estado do Rio de Janeiro e a perda de biodiversidade. Pp. 23-35. In: Bergallo, H. G., Rocha, C. F. D., Alves, M. A. S. & Sluys M. V. (Eds.). A fauna ameaçada de extinção do Estado do Rio de Janeiro. Rio de Janeiro, Editora UERJ.         [ Links ]

Veloso P.H.P.; Rangel Filho, A.L.R.R. & Lima, J.C.A. 1991. Classificação da Vegetação Brasileira, Adaptada a um Sistema Universal. Rio de Janeiro, IBGE - Departamento de Recursos Naturais e Estudos Ambientais.         [ Links ]

Villela, D.M.; Nascimento, M.T.; Aragão, L.E.O.C. & Gama, D.M. 2006. Effect of selective logging on forest structure and nutrient cycling in a seasonally dry Brazilian Atlantic forest. Journal of Biogeography 33: 506-516.         [ Links ]

Wilson, E.O. 1997. A situação atual da diversidade biológica. Pp. 3-24. In: Wilson E.O. & Peter, F.M. (Eds.). Biodiversidade. Rio de Janeiro, Nova Fronteira.         [ Links ]

Young, T.P. & Hubbell, S.P. 1991. Crown asymmetry, treefalls, and repeat disturbance of broad-leaved forest gaps. Ecology 72(4): 1464-1471.         [ Links ]

Zar, J.H. 1999. Biostatistical analysis. 4th ed. New Jersey, Prentice-Hall.         [ Links ]

 

 

Received: 21 November, 2013
Accepted: 2 June, 2014

 

 

* Author for correspondence: darlenegris@hotmail.com

Creative Commons License All the contents of this journal, except where otherwise noted, is licensed under a Creative Commons Attribution License