Acessibilidade / Reportar erro

Dynamics of natural regeneration after disturbance in a remnant of Mixed Ombrophilous Forest in southern Brazil

Dinâmica da regeneração natural após distúrbio em uma remanescente de Floresta Ombrófila Mista no Sul do Brasil

Abstract

This study evaluated changes in the structure for remnant Mixed Ombrophilous Forest recovering from logging, which ceased over 40 years ago. Regarding the dynamics of the floristic composition of natural regeneration, 18 species remained (23.38% of the total) and 27 new species entered (35.06% of the total). The greatest increases were observed for Allophylus edulis, Myrsine umbellata, and Miconia cinerascens. When analyzing the dynamics of regeneration in ecological groups, it was observed that pioneer species had a similar value in both surveys (29.4% and 29.6%); secondary species decreased from 56.6% to 52.8%, and late-successional species increased from 0.2% to 6.0%. Therefore, the forest fragment studied is in a healing phase after disturbance, characterized by the increased regeneration of pioneer species as a result of sufficient light entering the interior of the forest.

Keywords:
horizontal structure; ingress; mortality; succession stages

Resumo

O presente estudo teve como objetivo avaliar as alterações ocorridas, no período de 2012 a 2016, na estrutura da regeneração natural de um remanescente de Floresta Ombrófila Mista, sem intervenções há mais de 40 anos. Em relação a dinâmica da composição florística da regeneração natural, houve a saída de 18 espécies (23,38% do total) e a entrada de 27 novas espécies (35,06% do total). Os maiores acréscimos foram observados para Allophylus edulis, Myrsine umbellata e Miconia cinerascens. Analisando a dinâmica da regeneração em grupos ecológicos, observou-se que as espécies pioneiras tiveram um valor similar em ambos os levantamentos (29,4 a 29,6%). As espécies secundárias decresceram de 56,6% para 52,8%. E as espécies de sucessão tardia aumentaram de 0,2 a 6,0%. Logo, o remanescente estudado encontra-se em fase de cicatrização de distúrbio, caracterizada pelo aumento da regeneração de espécies pioneiras, uma vez que há quantidade suficiente de luminosidade incidindo no interior da floresta.

Palavras-chave:
estágios de sucessão; estrutura horizontal; ingresso; mortalidade

1. INTRODUCTION

Mixed Ombrophilous Forest (MOF), also known as Araucaria forest, is one of the main forest types of southern Brazil (Higuchi et al., 2012aHIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F.; LINKE, C.; PAULINO, P. S. Influência de variáveis ambientais sobre o padrão estrutural e florístico do componente arbóreo, em um fragmento de Floresta Ombrófila Mista Montana em Lages, SC. Ciência Florestal, v. 22, n. 1, p. 79-90, 2012a. http://dx.doi.org/10.5902/198050985081
http://dx.doi.org/10.5902/198050985081...
). Its geographical distribution occurs predominantly in the southern plateau region and is associated with places of high altitude and low average annual temperatures ((Higuchi et al., 2012aHIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F.; LINKE, C.; PAULINO, P. S. Influência de variáveis ambientais sobre o padrão estrutural e florístico do componente arbóreo, em um fragmento de Floresta Ombrófila Mista Montana em Lages, SC. Ciência Florestal, v. 22, n. 1, p. 79-90, 2012a. http://dx.doi.org/10.5902/198050985081
http://dx.doi.org/10.5902/198050985081...
). This forest formation consists mainly of Araucaria angustifolia (Bertol.) Kuntze, in association with Ocotea porosa (Nees & Mart.) Barroso, Ilex paraguariensis A. St.-Hil., Mimosa scabrella Benth., Cedrella fissilis Vell., Roupala brasiliensis Klotzsch, in addition to numerous species belonging to the Myrtaceae family (Higuchi et al., 2012aHIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F.; LINKE, C.; PAULINO, P. S. Influência de variáveis ambientais sobre o padrão estrutural e florístico do componente arbóreo, em um fragmento de Floresta Ombrófila Mista Montana em Lages, SC. Ciência Florestal, v. 22, n. 1, p. 79-90, 2012a. http://dx.doi.org/10.5902/198050985081
http://dx.doi.org/10.5902/198050985081...
, 2012bHIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F. Floristic composition and phytogeography of the tree component of Araucaria Forest fragments in southern Brazil. Brazilian Journal of Botany, v. 35, n. 2, p. 145-157, 2012b. https://doi.org/10.1590/S0100-84042012000200004
https://doi.org/10.1590/S0100-8404201200...
, 2015HIGUCHI, P.; SILVA, A. C.; BUZZI JUNIOR, F.; NEGRINI, M.; FERREIRA, T. S.; SOUZA, S. T.; SANTOS, K. F.; VEFAGO, M. B. Fatores determinantes da regeneração natural em um fragmento de floresta com araucária no Planalto Catarinense. Scientia Forestalis, v. 43, n. 106, p. 251-259, 2015.; Dalla Rosa et al., 2016DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
).

However, the MOF has been intensely altered by strong deforestation because of logging, the expansion of agriculture and livestock, the homogeneous reforestation of exotic species, and the advancement of urban areas, and is thus considered one of the most threatened forest types in Brazil (Ribeiro et al., 2009RIBEIRO, M. C.; METZGER, J. P.; MARTENSEN, A. C.; PONZONI, F. J.; HIROTA, M. M. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological conservation, v. 42, n. 6, p. 1141-1153, 2009. https://doi.org/10.1016/j.biocon.2009.02.021
https://doi.org/10.1016/j.biocon.2009.02...
; Vibrans et al., 2013VIBRANS, A. C.; MCROBERTS, R.; LINGNER, D. V.; NICOLETTI, A. L.; MOSER, P.; NICOLETTI, A. Extensão original e remanescentes da Floresta Ombrófila Mista em Santa Catarina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. V. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 25-31.; Souza et al., 2014SOUZA, K.; FAXINA, T. C.; SILVA, G. O.; DIAS, R. A. R.; SILVA, A. C., HIGUCHI, P. Análise fitossociológica de trilha ecológica em Floresta Ombrófila Mista. Revista de Ciências Agroveterinárias, v. 13, n. 3, p. 266-274, 2014.; Aguiar et al., 2017AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
). In fact, only 12.6% of the original MOF extension remains ((Ribeiro et al., 2009RIBEIRO, M. C.; METZGER, J. P.; MARTENSEN, A. C.; PONZONI, F. J.; HIROTA, M. M. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological conservation, v. 42, n. 6, p. 1141-1153, 2009. https://doi.org/10.1016/j.biocon.2009.02.021
https://doi.org/10.1016/j.biocon.2009.02...
). A lack of knowledge regarding native MOF management has contributed to the reduction in area and low diversity of natural remnants (Hess et al., 2010HESS, A. F.; CALGAROTTO, A. R.; PINHEIRO, R.; WANGINIAK, T. C. R. Propostas de manejo de Araucaria angustifolia utilizando o quociente de Liocourt e análise de incremento, em propriedade rural no município de Lages, SC. Pesquisa Florestal Brasileira, v. 30, n. 64, p. 337-345, 2010. https://doi.org/10.4336/2010.pfb.30.64.337
https://doi.org/10.4336/2010.pfb.30.64.3...
). Currently, in the State of Santa Catarina, only 24.4% of the original MOF remains, with forest areas distributed in fragmented remnants not exceeding 50 ha (Vibrans et al., 2013VIBRANS, A. C.; MCROBERTS, R.; LINGNER, D. V.; NICOLETTI, A. L.; MOSER, P.; NICOLETTI, A. Extensão original e remanescentes da Floresta Ombrófila Mista em Santa Catarina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. V. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 25-31.). However, even with the disturbances that have occurred, the remaining fragments can be maintained through natural ecological processes, as evidenced by the natural regeneration of tree species (Santos et al., 2018SANTOS, G. N.; HIGUCHI, P.; SILVA, A. C.; FARIAS, K. J.; MACHADO, F. D.; DUARTE, E. D.; FERNANDES, C.; VIEIRA, F.; AMARAL, R. S.; AGUIAR, V.; WALTER, F. F.; MORES, B.; REIS, M. A. Regeneração natural em uma floresta com araucária: inferências sobre o processo de construção da comunidade de espécies arbóreas. Ciência Florestal, v. 28, n. 2, p. 483-494, 2018. https://doi.org/10.5902/1980509832029
https://doi.org/10.5902/1980509832029...
).

Therefore, because of the great environmental and social importance of the MOF, the conservation and restoration of its remnant fragments is a primary objective of MOF management (Higuchi et al., 2012aHIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F.; LINKE, C.; PAULINO, P. S. Influência de variáveis ambientais sobre o padrão estrutural e florístico do componente arbóreo, em um fragmento de Floresta Ombrófila Mista Montana em Lages, SC. Ciência Florestal, v. 22, n. 1, p. 79-90, 2012a. http://dx.doi.org/10.5902/198050985081
http://dx.doi.org/10.5902/198050985081...
). In this context, studies of natural regeneration are extremely important because they generate results that enable a better understanding of the behavior of tree communities (Aguiar et al., 2017AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
). With regard to this, a greater understanding of the plant succession process in areas with different disturbance histories is essential, as it can generate data that detect the need to facilitate natural regeneration actions, as well as forest management incentives that facilitate recovery of other similar areas (Larsen et al., 2019LARSEN, J. G.; DUARTE, E.; DREYER, J. B. B.; DALLABRIDA, J. P.; MUZEKA, L. M.; SANTOS, G. N.; RODRIGUES JÚNIOR, L. C.; LIMA, C. L. L.; SILVA, A. C.; HIGUCHI, P. Variación espacio-temporal de la regeneración natural arbórea y arbustiva de bosque de alta montaña en etapa inicial de desarrollo en el sur de Brasil. Bosque, v. 40, n. 2, p. 185-193, 2019. https://doi.org/10.4067/S0717-92002019000200185
https://doi.org/10.4067/S0717-9200201900...
). Thus, these studies can support the conservation and recovery plans of forest ecosystems by enabling the identification of limiting environmental factors and inferring the future dynamics of forests (Aguiar et al., 2017AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
; Dalla Rosa et al., 2016DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
).

In this light, this study evaluated the changes that occurred from 2012 to 2016 in the structure of the natural regeneration of a remnant of Mixed Ombrophilous Forest that has remained without intervention for more than 40 years, in the state of Santa Catarina, Brazil.

2. MATERIAL AND METHODS

2.1. Characterization of the study area

The studied forest remnant is located in Fazenda Guamirim Gateados and is part of the “Emílio Einsfeld Filho” Private Reserve of Natural Heritage (PNHR), located in the municipalities of Campo Belo do Sul and Capão Alto, Santa Catarina, Brazil. The experimental area has a total of 3,365 ha, with approximately 72% forest cover, characterized by several succession stages of Mixed Ombrophilous Forest. Sampling was conducted within a radius of up to 500 m from the coordinate point 28°02'55.00” S and 50°45' 59.56” W.

Much of the vegetation in the study area has been subjected to selective logging, which targeted high commercial value woods (Zeller, 2010ZELLER, R. H. Plano de Manejo: Reserva Particular do Patrimônio Natural Emílio Einsfeld Filho, Santa Catarina. Campo Belo do Sul: Florestal Gateados; 2010. ). According to the Köppen climate classification, the climate of the region is characterized as humid subtropical mesothermal (Cfb), with cool summers, no defined dry season, and frequent severe frosts. The average annual temperature is approximately 15.8°C, varying from 11.4°C in the coldest month to 20.3°C in the hottest month. The average annual precipitation is 1,742 mm (Alvares et al., 2013ALVARES, C. A.; STAPE, J. L.; SENTELHAS, P. C.; GONÇALVES, J. L. M.; SPAROVEK, G. Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift, v. 22, n. 6, p. 711-728, 2013. https://doi.org/10.1127/0941-2948/2013/0507
https://doi.org/10.1127/0941-2948/2013/0...
). The region that encompasses the municipality of Campo Belo do Sul is located in the southern plateau, which in the State of Santa Catarina is bordered to the east by dense ombrophilous forest in addition to deciduous forest along the banks of the Uruguay River (Vibrans et al., 2013VIBRANS, A. C.; MCROBERTS, R.; LINGNER, D. V.; NICOLETTI, A. L.; MOSER, P.; NICOLETTI, A. Extensão original e remanescentes da Floresta Ombrófila Mista em Santa Catarina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. V. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 25-31.).

2.2. Data collection

In February 2012, 20 permanent circular plots were established in a random design in the experimental area. Each plot was 5 m in diameter, and the total sample area was 392.5 m² (Figure 1). In 2016, the same sample units were remeasured. Between the two survey occasions, a gale occurred in the study area, eliminating part of the upper tree layer in four sample units. The collected data corresponded to the number, height, and identification of the regenerating individuals in the sample units. Individuals with a minimum height of 0.50 m and a diameter breast height of less than 15 cm were considered to be regenerating.

Figure 1.
Geographic location of the plots in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil.

2.3. Data analysis

Using data collected in the permanent plots from 2012 and 2016, the floristic and phytosociological parameters of natural regeneration were estimated. For this, the species found in the plots were identified by visual analysis in the field or in the expert opinions and specialized literature. For the ecological classification of species, in addition to observations in the field, the methodology described by Vibrans et al. (2013)VIBRANS, A. C.; MCROBERTS, R.; LINGNER, D. V.; NICOLETTI, A. L.; MOSER, P.; NICOLETTI, A. Extensão original e remanescentes da Floresta Ombrófila Mista em Santa Catarina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. V. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 25-31. was followed. This system considers the following categories: pioneer species (P), secondary species (SE), and late-successional species (LS).

The dynamics of regeneration were evaluated by changing the values of diversity and structure of species between the two periods surveyed. The parameters used to estimate the regeneration structure were density and frequency, based on Mueller-Dombois and Ellenberg (1974)MUELLER-DOMBOIS, D.; ELLENBERG, D. Aims and methods of vegetation ecology. New York: Wiley, 1974. 547 p., and the category of size and value of the regeneration based on Finol (1971)FINOL, U. H. Nuevos parámetros a considerarse en el analisis estructural de las selvas vírgenes tropicales. Revista Florestal Venezuelana, v. 14, p. 337-1144, 1971.. Species diversity was calculated using the Shannon’s Diversity Index (H′), Simpson’s diversity index (D), and Pielou’s equability index (J′) (Brower and Zar, 1984BROWER, J. E.; ZAR, J. H. Field and laboratory methods for general ecology. Dubuque: W. M. C. Brow, 1984. 226 p.). All analyses were done by the PAST® statistical software, Version 2.17c, according to the methodology described by Felfili and Rezende (2003)FELFILI, J. M.; REZENDE, R. P. Conceitos e métodos em fitossociologia. Brasília: Departamento de Engenharia Florestal, Universidade de Brasília, 2003. 68 p. and Magurran (2004)MAGURRAN, A. E. Measuring biological diversity. Oxford: Blackwell Science, 2004. 256 p..

The regeneration-size category parameter (absolute and relative) refers to the distribution of regenerating individuals in height classes by determining the phytosociological values of natural regeneration. The regeneration value, in absolute and relative terms, in turn expresses the importance of species in the community by averaging their relative density, frequency, and size category. The calculations covered all species sampled in the study area. Changes in species density were analyzed using the paired t-test (alpha was set at = 0.05).

3. RESULTS AND DISCUSSION

3.1. Floristic composition

In general, in both surveys, 77 species were found, represented by 51 genera and 32 botanical families. Specifically, in 2012 and 2016, 50 (12,382 ind ha-1) and 59 species (12,313 ind ha-1) were found, respectively, of which 18 were exclusive to the first survey and 28 were exclusive to the second (Table 1). The species richness found in this study was similar to that found by Aguiar et al. (2017)AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
in an MOF experiment in Lages, SC, Brazil, and by Santos et al. (2015)SANTOS, K. F.; FERREIRA, T. S.; HIGUCHI, P.; SILVA, A. C.; VANDRESEN, P. B.; COSTA, A.; SPADA, G.; SCHMITZ, V.; SOUZA, F. Regeneração natural do componente arbóreo após a mortalidade de um maciço de taquara em um fragmento de Floresta Ombrófila Mista em Lages - SC. Ciência Florestal, v. 25, n. 1, p. 107-117, 2015. https://doi.org/10.1590/1980-509820152505107
https://doi.org/10.1590/1980-50982015250...
in a fragment with the same forest typology also in Lages, SC, Brazil.

The family with the greatest richness was Myrtaceae, followed by Lauraceae. Emphasizing the ecological importance of the Myrtaceae family, it has also been classified as highly representative of MOFs by several authors (Vibrans et al., 2011VIBRANS, A. C.; SEVEGNANI, L.; UHLMANN, A.; SCHORN, L. A.; SOBRAL, M. G.; GASPER, A. L.; LINGNER, D. V.; BROGNI, E.; KLEM, G.; GODOY, M. B.; VERDI, M. Structure of mixed ombrophyllous forests with Araucaria angustifolia (Araucariaceae) under external stress in Southern Brazil. Revista de Biologia Tropical, v. 59, n. 3, p. 1371-1387, 2011.; Meyer et al., 2013MEYER, L.; GASPER, A. L.; SEVEGNANI, L.; SCHORN, L. A.; VIBRANS, A. C.; LINGNER, D. V.; VERDI, M.; STIVAL-SANTOS, A.; DREVECK, S.; KORTE, A. Regeneração natural da Floresta Ombrófila Mista em Santa Catatina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 191-222.; Higuchi et al., 2015HIGUCHI, P.; SILVA, A. C.; BUZZI JUNIOR, F.; NEGRINI, M.; FERREIRA, T. S.; SOUZA, S. T.; SANTOS, K. F.; VEFAGO, M. B. Fatores determinantes da regeneração natural em um fragmento de floresta com araucária no Planalto Catarinense. Scientia Forestalis, v. 43, n. 106, p. 251-259, 2015.; Santos et al., 2015SANTOS, K. F.; FERREIRA, T. S.; HIGUCHI, P.; SILVA, A. C.; VANDRESEN, P. B.; COSTA, A.; SPADA, G.; SCHMITZ, V.; SOUZA, F. Regeneração natural do componente arbóreo após a mortalidade de um maciço de taquara em um fragmento de Floresta Ombrófila Mista em Lages - SC. Ciência Florestal, v. 25, n. 1, p. 107-117, 2015. https://doi.org/10.1590/1980-509820152505107
https://doi.org/10.1590/1980-50982015250...
, 2018; Dalla Rosa et al., 2016DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
; Silva et al., 2017SILVA, J. O.; SILVA, A. C.; HIGUCHI, P.; MAFRA, A. L.; GONÇALVES, D. A.; BUZZI JÚNIOR, F.; DALLA ROSA, A.; CRUZ, A. P.; FERREIRA, T. S. Floristic composition and phytogeography contextualization of the natural regeneration of an alluvial forest located in the "Planalto Sul Catarinense" Region, SC, Brazil. Revista Árvore, v. 41, 2017. https://doi.org/10.1590/1806-90882017000200003
https://doi.org/10.1590/1806-90882017000...
; Santana et al., 2018SANTANA, L. D.; RIBEIRO, J. H. C.; IVANAUSKAS, N. M.; CARVALHO, F. A. Estrutura, diversidade e heterogeneidade de uma floresta ombrófila mista altomontana em seu extremo norte de distribuição (Minas Gerais). Ciência Florestal, v. 28, n. 2, p. 567-579, 2018. https://doi.org/10.5902/1980509832039
https://doi.org/10.5902/1980509832039...
; Vefago et al., 2019VEFAGO, M. B.; SILVA, A. C.; CUCHI, T.; SANTOS, G. N.; NUNES, A. S.; RODRIGUES JUNIOR, L. C.; LIMA, C. L.; GROSS, A.; KILCA, R. V.; HIGUCHI, P. What explains the variation on the regenerative component dynamics of Araucaria Forests in southern Brazil? Scientia agrícola, v. 76, n. 5, p. 405-414, 2019. https://doi.org/10.1590/1678-992x-2017-0304
https://doi.org/10.1590/1678-992x-2017-0...
), occurring in areas of different altitudes (Dalla Rosa et al., 2016DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
).

Table 1.
Floristic and structural descriptors evaluated for natural regeneration in 2012 and 2016 in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil.

The H′ and J′ values were 3.15 and 0.81 in 2012 and 3.26 and 0.79 in 2016, respectively. The estimated value for D was 0.94 for both surveys (Table 1). The H′ value was similar to that found in an experiment by Aguiar et al. (2017)AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
in an MOF in Lages, SC, Brazil (3.34) and by Santos et al. (2018)SANTOS, G. N.; HIGUCHI, P.; SILVA, A. C.; FARIAS, K. J.; MACHADO, F. D.; DUARTE, E. D.; FERNANDES, C.; VIEIRA, F.; AMARAL, R. S.; AGUIAR, V.; WALTER, F. F.; MORES, B.; REIS, M. A. Regeneração natural em uma floresta com araucária: inferências sobre o processo de construção da comunidade de espécies arbóreas. Ciência Florestal, v. 28, n. 2, p. 483-494, 2018. https://doi.org/10.5902/1980509832029
https://doi.org/10.5902/1980509832029...
, also in an MOF in Lages, SC (3.42). Higher values were found by Dalla Rosa et al. (2016)DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
in a remnant of the same forest typology in Urubici, SC, Brazil (2.51). In contrast, lower values were found by Santos et al. (2015)SANTOS, K. F.; FERREIRA, T. S.; HIGUCHI, P.; SILVA, A. C.; VANDRESEN, P. B.; COSTA, A.; SPADA, G.; SCHMITZ, V.; SOUZA, F. Regeneração natural do componente arbóreo após a mortalidade de um maciço de taquara em um fragmento de Floresta Ombrófila Mista em Lages - SC. Ciência Florestal, v. 25, n. 1, p. 107-117, 2015. https://doi.org/10.1590/1980-509820152505107
https://doi.org/10.1590/1980-50982015250...
in an MOF fragment in Lages, SC, Brazil. According to Nascimento et al. (2001)NASCIMENTO, A. R. T.; LONGHI, S. J.; BRENA, D. A. Estrutura e padrões de distribuição espacial de espécies arbóreas em uma amostra de Floresta Ombrófila Mista em Nova Prata, RS. Ciência Florestal, v. 11, n. 1, p. 105-119, 2001. https://doi.org/10.5902/19805098499
https://doi.org/10.5902/19805098499...
, diversity values close to 3.00 characterize a medium diversity, which is expected in secondary succession stage MOF forests that have suffered anthropic influences.

Regarding J′, Dalla Rosa et al. (2016)DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
, Santos et al. (2015)SANTOS, K. F.; FERREIRA, T. S.; HIGUCHI, P.; SILVA, A. C.; VANDRESEN, P. B.; COSTA, A.; SPADA, G.; SCHMITZ, V.; SOUZA, F. Regeneração natural do componente arbóreo após a mortalidade de um maciço de taquara em um fragmento de Floresta Ombrófila Mista em Lages - SC. Ciência Florestal, v. 25, n. 1, p. 107-117, 2015. https://doi.org/10.1590/1980-509820152505107
https://doi.org/10.1590/1980-50982015250...
, Aguiar et al. (2017)AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
https://doi.org/10.1590/2179-8087.083214...
and Santos et al. (2018)SANTOS, G. N.; HIGUCHI, P.; SILVA, A. C.; FARIAS, K. J.; MACHADO, F. D.; DUARTE, E. D.; FERNANDES, C.; VIEIRA, F.; AMARAL, R. S.; AGUIAR, V.; WALTER, F. F.; MORES, B.; REIS, M. A. Regeneração natural em uma floresta com araucária: inferências sobre o processo de construção da comunidade de espécies arbóreas. Ciência Florestal, v. 28, n. 2, p. 483-494, 2018. https://doi.org/10.5902/1980509832029
https://doi.org/10.5902/1980509832029...
found values close to that of this study: 0.75, 0.76, 0.82 and 0.83, respectively. This indicates high ecological dominance. According to Brower and Zar (1984)BROWER, J. E.; ZAR, J. H. Field and laboratory methods for general ecology. Dubuque: W. M. C. Brow, 1984. 226 p., the Simpson diversity index (D) measures the probability that two individuals, selected at random from the same sample, belonging to the same species. That is, the higher the D value, the greater the diversity and the lesser the dominance of species (Klauberg et al., 2010KLAUBERG., C.; PALUDO, G. F.; BORTOLUZZI, R. L. C.; MANTOVANI, A. Florística e estrutura de um remanescente de Floresta Ombrófila Mista no Planalto Catarinense. Revista Biotemas, v. 23, n. 1, p. 35-47, 2010. https://doi.org/10.5007/2175-7925.2010v23n1p35
https://doi.org/10.5007/2175-7925.2010v2...
).

The quantitative data of the floristic descriptors evaluated (Table 1) revealed little change between the two surveys. It is observed that the diversity values found in this study reflect the pattern already verified in other studies in the same formation and confirm that the MOF in Santa Catarina is, in general, undergoing regeneration after decades of selective exploration. However, in this study, the observed changes were small, which indicates the existence of disturbances such as gusts and tree falls in the area, which can increase environmental filters and facilitate the entry of pioneer species.

3.2. Phytosociological parameters of natural regeneration

In the 2012 survey, the species Eugenia sp., Allophylus edulis (A.St.-Hil., Cambess. & A. Juss.) Radlk., Casearia decandra Jacq., Matayba elaeagnoides Radlk., and Myrsine umbelata Mart. were the most important in terms of density, which together accounted for 46.7% of the total stems (Table 2). These species were also the most representative in terms of frequency. In 2016, these same species remained the most important; however, there was a change in their order: Allophylus edulis, Myrsine umbellata, Casearia decandra, Matayba elaeagnoides and Eugenia sp. The mentioned species added up to 43.4% of the total density (Table 2). According to Higuchi et al. (2015)HIGUCHI, P.; SILVA, A. C.; BUZZI JUNIOR, F.; NEGRINI, M.; FERREIRA, T. S.; SOUZA, S. T.; SANTOS, K. F.; VEFAGO, M. B. Fatores determinantes da regeneração natural em um fragmento de floresta com araucária no Planalto Catarinense. Scientia Forestalis, v. 43, n. 106, p. 251-259, 2015., the variation of the most abundant species can be explained due to their different life strategies and respective capacities to develop in the forest understory. Some species may produce more propagules; however, they are inefficient competitors. Other species may produce few propagules, but with good competitive capacity.

Approximately half of the species (51.67%) were found in only 5% of the sample units, demonstrating the great diversity of species present in the regeneration of this forest type. In a study by Vefago et al. (2019)VEFAGO, M. B.; SILVA, A. C.; CUCHI, T.; SANTOS, G. N.; NUNES, A. S.; RODRIGUES JUNIOR, L. C.; LIMA, C. L.; GROSS, A.; KILCA, R. V.; HIGUCHI, P. What explains the variation on the regenerative component dynamics of Araucaria Forests in southern Brazil? Scientia agrícola, v. 76, n. 5, p. 405-414, 2019. https://doi.org/10.1590/1678-992x-2017-0304
https://doi.org/10.1590/1678-992x-2017-0...
, Allophylus edulis and Matayba elaeagnoides were also among the most representative species. Araucaria angustifolia (Bertol.) Kuntze, despite being considered the species that most characterizes MOF, went from 7th to 13th rank between the 2012 and 2016 survey; that is, it represented 4.3% of the density in 2012 (535.0 ind ha-1), and only 2.7% in 2016 (331.2 ind ha-1). However, despite the decrease in density in the plots, the species were more concentrated spatially in 2012 (FR: 3.2%) than in 2016 (FR: 4.1%). Other studies have also shown that this species had low representativeness (Dalla Rosa et al., 2016DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
; Vefago et al., 2019VEFAGO, M. B.; SILVA, A. C.; CUCHI, T.; SANTOS, G. N.; NUNES, A. S.; RODRIGUES JUNIOR, L. C.; LIMA, C. L.; GROSS, A.; KILCA, R. V.; HIGUCHI, P. What explains the variation on the regenerative component dynamics of Araucaria Forests in southern Brazil? Scientia agrícola, v. 76, n. 5, p. 405-414, 2019. https://doi.org/10.1590/1678-992x-2017-0304
https://doi.org/10.1590/1678-992x-2017-0...
). According to Dalla Rosa et al. (2016)DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
https://doi.org/10.1590/0100-67622016000...
, A. angustifolia has a limited number of regenerating individuals in the understory of the MOF and this limitation is even more extreme at high altitudes (>1,500 m).

3.3. Dynamics of floristic composition of natural regeneration

There were considerable changes in the floristic composition of natural regeneration between the two surveys (Table 3). For example, 18 species remained (23.38% of the total) and 27 new species entered (35.06% of the total). From this, we can see the dynamism that occurred in natural regeneration, which shows that the occurrence of species in the regenerative stratum will not guarantee their presence in the tree stratum in the future.

Table 2.
Phytosociological parameters of 20 species of greatest of values relative natural regeneration in 2012 and 2016 in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil.
Table 3.
Dynamics of the structure of the regenerative stratum in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil.

The greatest increases were observed for Allophylus edulis (+3.4%), Myrsine umbellata (+2.7%), and Miconia cinerascens Miq. (+1.9%), of which only M. cinerascens was classified as a pioneer, while the others were classified as secondary species. This fact indicates that this remnant forest is in a reestablishment phase after disturbance, since there is sufficient light shining into the forest to initiate the development/ recruitment of both pioneer and secondary species. Regarding the decreases, the species Eugenia sp., Ocotea pulchella (Nees & Mart.) Mez, and Cupania vernalis Cambess. stood out (-5.5%, -4.1%, and -2.7%) and the latter two were classified as pioneers. A study by Gross et al. (2018)GROSS, A.; SILVA, A. C.; CRUZ, A. P.; KILKA, R. V.; NUNES, A. S.; DUARTE, E.; VEFAGO, M. B.; SANTOS, G. S.; LIMA, C. L.; SALAMI, B.; HIGUCHI, P. Fragmentation as a key driver of tree community dynamics in mixed subtropical evergreen forests in Southern Brazil. Forest Ecology and Management, v. 411, p. 20-26, 2018. https://doi.org/10.1016/j.foreco.2018.01.013
https://doi.org/10.1016/j.foreco.2018.01...
reported that the dynamics of the Araucaria forest varied mainly in terms of tree mortality and rates of basal area loss.

3.4. Dynamics of the regeneration of ecological groups

Analyzing the dynamics of regeneration in ecological groups, it was observed that the pioneer species had similar values in both surveys (29.4% and 29.6%, respectively) (Figure 2). In contrast, secondary species decreased from 56.6% to 52.8%, while late-successional species increased from 0.2% to 6.0% between the first and the second survey, respectively. Meyer et al. (2013)MEYER, L.; GASPER, A. L.; SEVEGNANI, L.; SCHORN, L. A.; VIBRANS, A. C.; LINGNER, D. V.; VERDI, M.; STIVAL-SANTOS, A.; DREVECK, S.; KORTE, A. Regeneração natural da Floresta Ombrófila Mista em Santa Catatina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 191-222., when evaluating the natural regeneration of the MOF in Santa Catarina, Brazil, observed that the secondary, pioneer, and late-successional species corresponded to 54.37%, 29.66% and 15.97% of the total, respectively, which partially reflects the history of use of this forest.

In general, pioneer species showed the highest rates of positive change (189.8%), while secondary species showed the highest rates of negative change (-573.25%) (Figure 3). In this case, it becomes evident that the availability of light in the interior of the forest is caused by the fall of individuals from the tree layer due to the actions of climatic events, which is reflected in the increase of pioneer individuals, also resulting in the mortality of secondary species in natural regeneration. In this sense, the changes observed in the floristic composition and distribution of ecological groups can be attributed to disturbances that occur in the forest.

Chazdon (2016)CHAZDON, R. L. Renascimento de florestas: regeneração na era do desmatamento. São Paulo: Oficina de Textos, 2016. 430 p. mentioned that disturbances in natural forests can be caused by either natural or anthropogenic activities. Considering that selective logging ceased more than 40 years ago in the study area, natural disturbances, such as gales and the fall of mature trees, are undoubtedly causing the regeneration gaps. This, consequently, provides opportunities for changes in the distribution of ecological groups in the forest, especially with the increased representativeness of pioneer species that are extremely important for the forest succession, creating conditions for the species of more advanced successional stages to establish themselves. Thus, the colonization of clearings by pioneer species detected in this study shows the occurrence of forest reestablishment after disturbances (Narvaes et al., 2005NARVAES, I. S.; BRENA, D. A.; LONGHI, S. J. Estrutura da regeneração natural em Floresta Ombrófila Mista na Floresta Nacional de São Francisco de Paula, RS. Ciência Florestal, v. 15, n. 4, p. 331-342, 2005. https://doi.org/10.5902/198050981871
https://doi.org/10.5902/198050981871...
).

Figure 2.
Relative natural regeneration by ecological groups in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil. P: Pioneers species; SE: Secondary species; LS: late-successional species; ND: Ecological group not defined.

Figure 3.

Average rates of change in the regenerative stratum by ecological groups in a remnant of Mixed Ombrophilous Forest in Santa Catarina, Brazil.

P: Pioneers species; SE: Secondary species; LS: late-successional species; ND: Ecological group not defined.


The changes that occurred in this fragment of MOF in the period of four years between the two vegetation censuses show that there was an increase in pioneer species and decreases in secondary and late-successional species. Therefore, the results show that the dynamics of natural regeneration of the forest may not follow the path theoretically expected in such a short period of time, as internal and external disturbances can affect the succession processes.

4. CONCLUSIONS

Considerable changes occurred in the structure and floristic composition dynamics of natural regeneration between the two data surveys.

Pioneer species had the highest rates of positive change, while secondary species had the highest rates of negative change.

This studied remnant forest fragment is in the healing phase after disturbance, since there is a sufficient amount of light falling inside the forest.

5. REFERENCES

  • AGUIAR, M. D.; SILVA, A. C.; HIGUCHI, P.; NEGRINI, M.; SCHOLLEMBERG, A. L. Similaridade entre adultos e regenerantes do componente arbóreo em floresta com araucária. Floresta e Ambiente, v. 24, 2017. https://doi.org/10.1590/2179-8087.083214
    » https://doi.org/10.1590/2179-8087.083214
  • ALVARES, C. A.; STAPE, J. L.; SENTELHAS, P. C.; GONÇALVES, J. L. M.; SPAROVEK, G. Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift, v. 22, n. 6, p. 711-728, 2013. https://doi.org/10.1127/0941-2948/2013/0507
    » https://doi.org/10.1127/0941-2948/2013/0507
  • BROWER, J. E.; ZAR, J. H. Field and laboratory methods for general ecology. Dubuque: W. M. C. Brow, 1984. 226 p.
  • CHAZDON, R. L. Renascimento de florestas: regeneração na era do desmatamento. São Paulo: Oficina de Textos, 2016. 430 p.
  • DALLA ROSA, A.; SILVA, A. C.; HIGUCHI, P.; MARCON, A. K.; MISSIO, F. F.; BENTO, M. A.; SILVA, J. O.; GONÇALVES, D. A.; RODRIGUES JÚNIOR, L. C. Natural regeneration of tree species in a cloud forest in Santa Catarina, Brazil. Revista Árvore, v. 40, n. 6, p. 1073-1082, 2016. https://doi.org/10.1590/0100-67622016000600013
    » https://doi.org/10.1590/0100-67622016000600013
  • FELFILI, J. M.; REZENDE, R. P. Conceitos e métodos em fitossociologia. Brasília: Departamento de Engenharia Florestal, Universidade de Brasília, 2003. 68 p.
  • FINOL, U. H. Nuevos parámetros a considerarse en el analisis estructural de las selvas vírgenes tropicales. Revista Florestal Venezuelana, v. 14, p. 337-1144, 1971.
  • GROSS, A.; SILVA, A. C.; CRUZ, A. P.; KILKA, R. V.; NUNES, A. S.; DUARTE, E.; VEFAGO, M. B.; SANTOS, G. S.; LIMA, C. L.; SALAMI, B.; HIGUCHI, P. Fragmentation as a key driver of tree community dynamics in mixed subtropical evergreen forests in Southern Brazil. Forest Ecology and Management, v. 411, p. 20-26, 2018. https://doi.org/10.1016/j.foreco.2018.01.013
    » https://doi.org/10.1016/j.foreco.2018.01.013
  • HESS, A. F.; CALGAROTTO, A. R.; PINHEIRO, R.; WANGINIAK, T. C. R. Propostas de manejo de Araucaria angustifolia utilizando o quociente de Liocourt e análise de incremento, em propriedade rural no município de Lages, SC. Pesquisa Florestal Brasileira, v. 30, n. 64, p. 337-345, 2010. https://doi.org/10.4336/2010.pfb.30.64.337
    » https://doi.org/10.4336/2010.pfb.30.64.337
  • HIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F.; LINKE, C.; PAULINO, P. S. Influência de variáveis ambientais sobre o padrão estrutural e florístico do componente arbóreo, em um fragmento de Floresta Ombrófila Mista Montana em Lages, SC. Ciência Florestal, v. 22, n. 1, p. 79-90, 2012a. http://dx.doi.org/10.5902/198050985081
    » http://dx.doi.org/10.5902/198050985081
  • HIGUCHI, P.; SILVA, A. C.; FERREIRA, T. S.; SOUZA, S. T.; GOMES, J. P.; SILVA, K. M.; SANTOS, K. F. Floristic composition and phytogeography of the tree component of Araucaria Forest fragments in southern Brazil. Brazilian Journal of Botany, v. 35, n. 2, p. 145-157, 2012b. https://doi.org/10.1590/S0100-84042012000200004
    » https://doi.org/10.1590/S0100-84042012000200004
  • HIGUCHI, P.; SILVA, A. C.; BUZZI JUNIOR, F.; NEGRINI, M.; FERREIRA, T. S.; SOUZA, S. T.; SANTOS, K. F.; VEFAGO, M. B. Fatores determinantes da regeneração natural em um fragmento de floresta com araucária no Planalto Catarinense. Scientia Forestalis, v. 43, n. 106, p. 251-259, 2015.
  • KLAUBERG., C.; PALUDO, G. F.; BORTOLUZZI, R. L. C.; MANTOVANI, A. Florística e estrutura de um remanescente de Floresta Ombrófila Mista no Planalto Catarinense. Revista Biotemas, v. 23, n. 1, p. 35-47, 2010. https://doi.org/10.5007/2175-7925.2010v23n1p35
    » https://doi.org/10.5007/2175-7925.2010v23n1p35
  • LARSEN, J. G.; DUARTE, E.; DREYER, J. B. B.; DALLABRIDA, J. P.; MUZEKA, L. M.; SANTOS, G. N.; RODRIGUES JÚNIOR, L. C.; LIMA, C. L. L.; SILVA, A. C.; HIGUCHI, P. Variación espacio-temporal de la regeneración natural arbórea y arbustiva de bosque de alta montaña en etapa inicial de desarrollo en el sur de Brasil. Bosque, v. 40, n. 2, p. 185-193, 2019. https://doi.org/10.4067/S0717-92002019000200185
    » https://doi.org/10.4067/S0717-92002019000200185
  • MAGURRAN, A. E. Measuring biological diversity. Oxford: Blackwell Science, 2004. 256 p.
  • MEYER, L.; GASPER, A. L.; SEVEGNANI, L.; SCHORN, L. A.; VIBRANS, A. C.; LINGNER, D. V.; VERDI, M.; STIVAL-SANTOS, A.; DREVECK, S.; KORTE, A. Regeneração natural da Floresta Ombrófila Mista em Santa Catatina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 191-222.
  • MUELLER-DOMBOIS, D.; ELLENBERG, D. Aims and methods of vegetation ecology. New York: Wiley, 1974. 547 p.
  • NARVAES, I. S.; BRENA, D. A.; LONGHI, S. J. Estrutura da regeneração natural em Floresta Ombrófila Mista na Floresta Nacional de São Francisco de Paula, RS. Ciência Florestal, v. 15, n. 4, p. 331-342, 2005. https://doi.org/10.5902/198050981871
    » https://doi.org/10.5902/198050981871
  • NASCIMENTO, A. R. T.; LONGHI, S. J.; BRENA, D. A. Estrutura e padrões de distribuição espacial de espécies arbóreas em uma amostra de Floresta Ombrófila Mista em Nova Prata, RS. Ciência Florestal, v. 11, n. 1, p. 105-119, 2001. https://doi.org/10.5902/19805098499
    » https://doi.org/10.5902/19805098499
  • RIBEIRO, M. C.; METZGER, J. P.; MARTENSEN, A. C.; PONZONI, F. J.; HIROTA, M. M. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological conservation, v. 42, n. 6, p. 1141-1153, 2009. https://doi.org/10.1016/j.biocon.2009.02.021
    » https://doi.org/10.1016/j.biocon.2009.02.021
  • SANTANA, L. D.; RIBEIRO, J. H. C.; IVANAUSKAS, N. M.; CARVALHO, F. A. Estrutura, diversidade e heterogeneidade de uma floresta ombrófila mista altomontana em seu extremo norte de distribuição (Minas Gerais). Ciência Florestal, v. 28, n. 2, p. 567-579, 2018. https://doi.org/10.5902/1980509832039
    » https://doi.org/10.5902/1980509832039
  • SANTOS, K. F.; FERREIRA, T. S.; HIGUCHI, P.; SILVA, A. C.; VANDRESEN, P. B.; COSTA, A.; SPADA, G.; SCHMITZ, V.; SOUZA, F. Regeneração natural do componente arbóreo após a mortalidade de um maciço de taquara em um fragmento de Floresta Ombrófila Mista em Lages - SC. Ciência Florestal, v. 25, n. 1, p. 107-117, 2015. https://doi.org/10.1590/1980-509820152505107
    » https://doi.org/10.1590/1980-509820152505107
  • SANTOS, G. N.; HIGUCHI, P.; SILVA, A. C.; FARIAS, K. J.; MACHADO, F. D.; DUARTE, E. D.; FERNANDES, C.; VIEIRA, F.; AMARAL, R. S.; AGUIAR, V.; WALTER, F. F.; MORES, B.; REIS, M. A. Regeneração natural em uma floresta com araucária: inferências sobre o processo de construção da comunidade de espécies arbóreas. Ciência Florestal, v. 28, n. 2, p. 483-494, 2018. https://doi.org/10.5902/1980509832029
    » https://doi.org/10.5902/1980509832029
  • SILVA, J. O.; SILVA, A. C.; HIGUCHI, P.; MAFRA, A. L.; GONÇALVES, D. A.; BUZZI JÚNIOR, F.; DALLA ROSA, A.; CRUZ, A. P.; FERREIRA, T. S. Floristic composition and phytogeography contextualization of the natural regeneration of an alluvial forest located in the "Planalto Sul Catarinense" Region, SC, Brazil. Revista Árvore, v. 41, 2017. https://doi.org/10.1590/1806-90882017000200003
    » https://doi.org/10.1590/1806-90882017000200003
  • SOUZA, K.; FAXINA, T. C.; SILVA, G. O.; DIAS, R. A. R.; SILVA, A. C., HIGUCHI, P. Análise fitossociológica de trilha ecológica em Floresta Ombrófila Mista. Revista de Ciências Agroveterinárias, v. 13, n. 3, p. 266-274, 2014.
  • VEFAGO, M. B.; SILVA, A. C.; CUCHI, T.; SANTOS, G. N.; NUNES, A. S.; RODRIGUES JUNIOR, L. C.; LIMA, C. L.; GROSS, A.; KILCA, R. V.; HIGUCHI, P. What explains the variation on the regenerative component dynamics of Araucaria Forests in southern Brazil? Scientia agrícola, v. 76, n. 5, p. 405-414, 2019. https://doi.org/10.1590/1678-992x-2017-0304
    » https://doi.org/10.1590/1678-992x-2017-0304
  • VIBRANS, A. C.; SEVEGNANI, L.; UHLMANN, A.; SCHORN, L. A.; SOBRAL, M. G.; GASPER, A. L.; LINGNER, D. V.; BROGNI, E.; KLEM, G.; GODOY, M. B.; VERDI, M. Structure of mixed ombrophyllous forests with Araucaria angustifolia (Araucariaceae) under external stress in Southern Brazil. Revista de Biologia Tropical, v. 59, n. 3, p. 1371-1387, 2011.
  • VIBRANS, A. C.; MCROBERTS, R.; LINGNER, D. V.; NICOLETTI, A. L.; MOSER, P.; NICOLETTI, A. Extensão original e remanescentes da Floresta Ombrófila Mista em Santa Catarina. In: VIBRANS, A. C.; SEVEGNANI, L.; GASPER, A. L.; LINGNER, D. V. Inventário Florístico Florestal de Santa Catarina: Floresta Ombrófila Mista. Blumenau: Edifurb, 2013. p. 25-31.
  • ZELLER, R. H. Plano de Manejo: Reserva Particular do Patrimônio Natural Emílio Einsfeld Filho, Santa Catarina. Campo Belo do Sul: Florestal Gateados; 2010.

Publication Dates

  • Publication in this collection
    14 June 2021
  • Date of issue
    2021

History

  • Received
    18 Nov 2020
  • Accepted
    19 Apr 2021
Instituto de Pesquisas Ambientais em Bacias Hidrográficas Instituto de Pesquisas Ambientais em Bacias Hidrográficas (IPABHi), Estrada Mun. Dr. José Luis Cembranelli, 5000, Taubaté, SP, Brasil, CEP 12081-010 - Taubaté - SP - Brazil
E-mail: ambi.agua@gmail.com