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Articles • Braz. J. Biol. 75 (2 suppl) • May 2015 • https://doi.org/10.1590/1519-6984.0913   copy

Vascular epiphytes of the Atlantic Forest in the Sinos River basin, state of Rio Grande do Sul, Brazil: richness, floristic composition and community structure

Epífitos vasculares em Floresta Atlântica da bacia do Rio dos Sinos, estado do Rio Grande do Sul, Brasil: riqueza, composição florística e estrutura comunitária

Authorship SCIMAGO INSTITUTIONS RANKINGS

Abstracts

The Atlantic Forest, which has a vast epiphytic richness, is a priority area for preservation, listed as one of the five most important world hotspots. Vascular epiphyte richness, composition and community structure were studied in two fragments, one of the ombrophilous (29º43'42"S and 50º22'00"W) and the other of the seasonal (29º40'54"S and 51º06'56"W) forest, both belonging to the Atlantic Forest biome in the Sinos River basin, Rio Grande do Sul, Brazil. In each fragment, 40 trees, divided into four ecological zones, were analyzed. In each zone, the occurrence of the species was recorded, and the importance value of each species was calculated according to the frequency of phorophytes and intervals, and cover scores. The Shannon index was calculated for the two communities. In the fragment of the ombrophilous forest (F1), 30 epiphytic species were recorded, and in the seasonal forest (F2), 25. The highest importance value was found for Microgramma squamulosa (Kaulf.) de la Sota in both fragments. The diversity indexes for F1 (H'=2.72) and F2 (H'=2.55) were similar and reflected the subtropical location of the areas. The decrease in mean richness in both fragments in zone 3 (internal crown) to zone 4 (external crown) may be associated with time and space availability for epiphyte occupation and microclimate variations. Exclusive species were found in the areas, which suggest that a greater number of preserved fragments may result in a greater number of preserved epiphytic species in the Sinos River basin.

vertical distribution; epiphytism; phytosociology; floristic diagnostic; southern Brazil

A Floresta Atlântica apresenta uma grande riqueza de epífitos e é considerada uma área prioritária para preservação listada entre os cinco mais importantes hotspots mundiais. A riqueza, a composição e a estrutura comunitária de epífitos vasculares foram estudadas em um fragmento de floresta ombrófila (29º43'42"S e 50º22'00"O) e outro de floresta estacional (29º40'54"S e 51º06'56"O), ambos pertencentes ao Bioma Floresta Atlântica, na bacia do Rio dos Sinos, Rio Grande do Sul, Brasil. Em cada fragmento foram analisadas 40 árvores divididas em quatro zonas ecológicas. Em cada zona, foi registrada a ocorrência das espécies e o valor de importância de cada uma delas foi calculado a partir da frequência nos forófitos e nos intervalos e das notas de cobertura. O índice de Shannon foi aplicado para as duas comunidades. No fragmento de floresta ombrófila (F1) foram registradas 30 e no de floresta estacional (F2) 25 espécies epifíticas. O maior valor de importância foi observado para Microgramma squamulosa (Kaulf.) de la Sota, em ambos os fragmentos. Os índices de diversidade para o F1 (H'=2,72) e o F2 (H'=2,55) foram próximos e refletem a posição geográfica mais subtropical das áreas. A diminuição na riqueza média em ambos os fragmentos da zona 3 (copa interna) para a zona 4 (copa externa) pode estar relacionada com a disponibilidade de tempo e de espaço para a ocupação de epífitos e com variações microclimáticas. Considerando que as áreas apresentaram espécies exclusivas, ficou evidenciado que um maior número de fragmentos conservados conduzirá à manutenção de um maior número de espécies epifíticas na bacia do Rio dos Sinos.

distribuição vertical; epifitismo; fitossociologia; diagnóstico florístico; sul do Brasil

1 Introduction

Epiphytes account for a substantial part of forest diversity and up to 10% (Kress, 1986Kress, WJ., 1986. The systematic distribution of vascular epiphytes: an update. Selbyana, vol. 9, no. 1, p. 2-22.) to 50% (Kersten and Silva, 2005Kersten, RA. and Silva, SM., 2005. Florística e estrutura de comunidades de epífitas vasculares da planície litorânea. In MARQUES, MCM. and BRITEZ, RM. (Eds.). História natural e conservação da Ilha do Mel. Curitiba: Universidade Federal do Paraná. p. 125-144.) of all vascular plants in tropical forests. The Atlantic Forest, listed as one of the five most important world hotspots, has 20,000 species of vascular plants (Myers et al. 2000Myers, N., Mittermeier, RA., Mittermeier, CG., Fonseca, GA. and Kent, J., 2000. Biodiversity hotspots for conservation priorities. Nature, vol. 403, no. 6772, p. 853-858. http://dx.doi.org/10.1038/35002501. PMid:10706275.
http://dx.doi.org/10.1038/35002501... ), and 3,000 to 4,000 of them are epiphytic (Kersten, 2010Kersten, RA., 2010. Epífitas vasculares – Histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea, vol. 37, no. 1, p. 9-38. http://dx.doi.org/10.1590/S2236-89062010000100001.
http://dx.doi.org/10.1590/S2236-89062010... ). In the state of Rio Grande do Sul, Brazil, the Atlantic Forest has been reduced to only 7.48% of its original area (Fundação SOS Mata Atlântica and INPE, 2011Fundação SOS Mata Atlântica and Instituto Nacional de Pesquisas Espaciais –INPE, 2011. Atlas dos remanescentes florestais da Mata Atlântica: período 2008-2010. São Paulo: INPE. 122 p.), and ombrophilous and seasonal forests are found in some of the phytoecological regions of this biome.

Epiphytism has an important ecological function in forest communities and participates in the preservation of biological diversity by providing nutritional resources and specialized microenvironments for the fauna in the crown and by participating in nutrient cycling mechanisms; it is, therefore, considered as a local biodiversity amplifier (Nadkarni, 1984Nadkarni, NM., 1984. Epiphyte biomass and nutrient capital of a neotropical elfin forest. Biotropica, vol. 16, no. 4, p. 249-256. http://dx.doi.org/10.2307/2387932.
http://dx.doi.org/10.2307/2387932... ; Gentry and Dodson, 1987GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
http://dx.doi.org/10.2307/2399395... ; Lugo and Scatena, 1992Lugo, AE. and Scatena, FN., 1992. Epiphytes and climate change research in the Caribbean: a proposal. Selbyana, vol. 13, p. 123-130.; Waechter, 1992Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.; Rocha et al., 2004Rocha, CFD., Cogliatti-Carvalho, L., Nunes-Freitas, AF., Rocha-Pessoa, TC., Dias, AS., Ariani, CV. and Morgado, LN., 2004. Conservando uma larga proporção da diversidade biológica através da conservação de Bromeliaceae.Vidalia, vol. 2, no. 1, p. 52-68.). Knowledge on vascular epiphyte composition and community structure has been used to define quality and environmental characteristics (Krömer et al., 2007Krömer, T., Kessler, M. and Gradstein, SR., 2007. Vertical stratification of vascular ephytes in submontane and montane Forest of the Bolivian Andes: the importance of the understory. Plant Ecology, vol. 189, no. 2, p. 261-278. http://dx.doi.org/10.1007/s11258-006-9182-8.
http://dx.doi.org/10.1007/s11258-006-918... ). Epiphytic plants have been used to classify the successive regeneration stages of plant formations in the Atlantic Forest of Rio Grande do Sul, according to the CONAMA Resolution no. 33/94 (Brasil, 2012Brasil. Conselho Nacional de Meio Ambiente – CONAMA, Ministério do Meio Ambiente – MMA, 2012. Resoluções do CONAMA: Resoluções vigentes publicadas entre setembro de 1984 e janeiro de 2012. Brasília: MMA. 1126 p.).

The spatial distribution of richness and the composition of the epiphytic flora may be affected by several factors at different scales and directions, such as the regional mosaic of plant types, the vertical profile of the forests, the characteristics of phorophytes and the climate conditions inside the forest (Gentry and Dodson, 1987GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
http://dx.doi.org/10.2307/2399395... ; Nieder et al., 1997Nieder, J., Ibisch, PL. and Barthlott, W., 1997. Biodiversidad de epífitas: una cuestión de escala. Revista del Jardín Botánico Nacional, vol. 17/18, p. 59-62.; Schmitt and Windisch, 2005Schmitt, JL. and Windisch, PG., 2005. Aspectos ecológicos de . Alsophila setosa Kaulf. (Cyatheaceae, Pteridophyta) no Rio Grande do Sul, BrasilActa Botanica Brasilica, vol. 19, no. 4, p. 861-867. http://dx.doi.org/10.1590/S0102-33062005000400021.
http://dx.doi.org/10.1590/S0102-33062005... ; 2010Schmitt, JL. and Windisch, PG., 2010. Biodiversity and spatial distribution of epiphytic ferns on . Alsophila setosa Kaulf. (Cyatheaceae) caudices in Rio Grande do Sul, BrazilBrazilian Journal of Biology = Revista Brasleira de Biologia, vol. 70, no. 3, p. 521-528. http://dx.doi.org/10.1590/S1519-69842010000300008. PMid:20730338.
http://dx.doi.org/10.1590/S1519-69842010... ; Waechter, 2006Waechter, JL., 2006. Diversidade epifítica ao longo de gradientes ambientais. In MARIATH, JEA. and SANTOS, RP. (Eds.). Os Avanços da Botânica no Início do Século XXI: Conferências, Plenárias e Simpósios do 57º Congresso Nacional de Botânica. Porto Alegre, Sociedade Botânica do Brasil. p. 350-353.). Some of the microclimate factors that may affect vertical distribution are humidity, which tends to decrease from the ground to the apex, and luminosity, which increases in the same direction (Lüttge, 1989Lüttge, U., 1989. Vascular epiphytes: setting the Scene. In LÜTTGE, U. (Ed.). Vascular plants as epiphytes. New York: Springer-Verlag. p. 1-12. Ecological Studies,79:; Ter Steege and Cornelissen, 1989TER Steege, H. and Cornelissen, JHC., 1989. Distribution and ecology of vascular epiphytes in Lowland rain forest of Guiana. Biotropica, vol. 21, no. 4, p. 331-339. http://dx.doi.org/10.2307/2388283.
http://dx.doi.org/10.2307/2388283... ).

In general, epiphytes are extremely diversified and profuse in dense ombrophilous forests when compared with deciduous and semi-deciduous seasonal forests (Rambo, 1954Rambo, B., 1954. História da flora do litoral rio-grandense. Sellowia, vol. 6, no. 6, p. 113-172.; Klein, 1975Klein, RM., 1975. Southern Brazilian phytogeographic features and the probable influence of upper quaternary climatic changes in the floristic distribution. Boletim Paranaense de Geociências, vol. 33, p. 67-88.; Roderjan et al., 2002Roderjan, CV., Galvão, F., Kuniyoshi, YS. and Hatschbach, GG., 2002. As unidades fitogeográficas do estado do Paraná. Ciência e Ambiente, vol. 24, no. 1, p. 75-92.), a similar distribution to that of fern richness (Sehnem, 1977Sehnem, A., 1977. As filicíneas do Sul do Brasil, sua distribuição geográfica, sua ecologia e suas rotas de migração. Pesquisas, vol. 31, p. 1-108.; 1979Sehnem, A., 1979. Semelhanças e diferenças nas formações florestais do sul do Brasil. Acta Biologica Leopoldensia, vol. 1, no. 1, p. 111-135.). In the southern region of Brazil, studies have been conducted with different vegetation types, including coastal plains (Waechter, 1986Waechter, JL., 1986. Epífitos vasculares da mata paludosa do Faxinal, Torres, Rio Grande do Sul, Brasil. Iheringia, vol. 34, p. 39-49.; 1992Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.; 1998Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.; Kersten and Silva, 2001Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... ; Gonçalves and Waechter, 2003Gonçalves, CN. and Waechter, JL., 2003. Aspectos florísticos e ecológicos de epífitos vasculares sobre figueiras isoladas no norte da planície costeira do Rio Grande do Sul. Acta Botanica Brasilica, vol. 17, no. 1, p. 89-100. http://dx.doi.org/10.1590/S0102-33062003000100007.
http://dx.doi.org/10.1590/S0102-33062003... ; Staudt et al., 2012Staudt, MG., Lippert, APU., Cunha, S., Becker, DFP., Marchioretto, MS. and Schmitt, JL., 2012. Composição florística de epífitos vasculares do Parque Natural Municipal Tupancy, Arroio do Sal, RS – Brasil. Pesquisas, vol. 63, p. 177-188.; Becker et al., 2013Becker, DFP., Cunha, S., Marchioretto, MS. and Schmitt, JL., 2013. Riqueza, estrutura comunitária e distribuição vertical de epífitos vasculares no Parque Natural Municipal Tupancy, Arroio do Sal, RS, Brasil. Pesquisas, vol. 64, p. 127-139.), semi-deciduous (Aguiar et al., 1981Aguiar, LW., Citadini-Zanette, V., Martau, L. and Backes, A., 1981. Composição florística de epífitos vasculares numa área localizada nos municípios de Montenegro e Triunfo, Rio Grande do Sul, Brasil. Iheringia Série Botânica, vol. 28, no. 1, p. 55-93.; Borgo et al., 2002Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130.; Dettke et al., 2008Dettke, GA., Orfrini, AC. and Milanese-Gutierre, MA., 2008. Composição florística e distribuição de epífitas vasculares em um remanescente alterado da Floresta Estacional Semidecidual do Paraná, Brasil. Rodriguésia, vol. 59, no. 4, p. 859-872.) and deciduous (Rogalski and Zanin, 2003Rogalski, JM. and Zanin, EM., 2003. Composição florística de epífitos vasculares no estreito de Augusto César, Floresta Estacional Decidual do Rio Uruguai, RS, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 4, p. 551-556. http://dx.doi.org/10.1590/S0100-84042003000400014.
http://dx.doi.org/10.1590/S0100-84042003... ) seasonal forests, dense (Blum et al., 2011Blum, CT., Roderjan, CV. and Galvão, F., 2011. Composição florística e distribuição altitudinal de epífitas vasculares da Floresta Ombrófila Densa na Serra da Prata, Morretes, Paraná, Brasil. Biota Neotropica, vol. 11, no. 4, p. 141-159. http://dx.doi.org/10.1590/S1676-06032011000400015.
http://dx.doi.org/10.1590/S1676-06032011... ) and mixed (Borgo and Silva, 2003Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
http://dx.doi.org/10.1590/S0100-84042003... ; Hefler and Faustioni, 2004Hefler, SM. and Faustioni, P., 2004. Levantamento florístico de epífitos vasculares do bosque São Cristovão –Curitiba – Paraná – Brasil. Revista Estudos de Biologia, vol. 26, no. 54, p. 11-19.; Schmitt et al., 2005Schmitt, JL., Budke, JC. and Windisch, PG., 2005. Aspecto florísticos e ecológicos de pteridófitas epifíticas em cáudices de Hook. (Pteridophyta, Dicksoniaceae), São Francisco de Paula, RS, Brasil. Dicksonia sellowianaPesquisas, vol. 56, p. 161-172.; Brustulin and Schmitt, 2008Brustulin, J. and Schmitt, JL., 2008. Composição florística, distribuição vertical e floração de orquídeas epifíticas em três parques municipais do estado do Rio Grande do Sul, Brasil. Pesquisas, vol. 59, p. 143-158.; Buzatto et al., 2008Buzatto, C.R., SEVERO, B M.A. and WAECHTER, J.L., 2008. Composição florística e distribuição ecológica de epífitos vasculares na Floresta Nacional de Passo Fundo, Rio Grande do Sul. Iheringia Serie Botânica, vol. 63, no. 2, p. 231-239.; Bonnet et al., 2009Bonnet, A., Lavoranti, OJ. and Curcio, GR., 2009. Epífitos vasculares no Corredor de Biodiversidade Araucária, bacia do rio Iguaçu, Paraná, Brasil. Cadernos da Biodiversidade, vol. 6, p. 49-70.; Kersten and Kuniyoshi, 2009Kersten, RA. and Kuniyoshi, YS., 2009. Conservação das florestas na Bacia do alto Iguaçú, Paraná – Avaliação da comunidade de epífitas vasculares em diferentes estágios serais. Revista Floresta, vol. 39, no. 1, p. 51-66.; Kersten et al., 2009Kersten, RA., Kuniyidhi, YS. and Roderjan, CV., 2009. Epífitas vasculares em duas formações ribeirinhas adjacentes na bacia do rio Iguaçu – Terceiro Planalto Paranaense. Iheringia Série Botânica, vol. 64, no. 1, p. 33-43.) ombrophilous forests, riparian forests (Giongo and Waechter, 2004Giongo, C. and Waechter, JL., 2004. Composição florística e estrutura comunitária de epífitos vasculares em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 27, no. 3, p. 563-572. http://dx.doi.org/10.1590/S0100-84042004000300015.
http://dx.doi.org/10.1590/S0100-84042004... ) and areas of transition from semi-deciduous seasonal to mixed ombrophilous forests (Bonnet et al., 2010Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2010. Relações de epífitos vasculares com fatores ambientais nas florestas do Rio Tibagi, Paraná, Brasil. Biotemas, vol. 23, no. 3, p. 37-47., 2011Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2011. Flora epifítica vascular em três unidades vegetacionais do Rio Tibagi, Paraná, Brasil.Rodriguésia, vol. 62, no. 3, p. 491-498.).

Among the studies that compared the epiphytism in different areas, Barthlott et al. (2001)BarthloTt, W., Schmit-Neuerburg, V., Nieder, J. and Engwald, S., 2001. Diversity and abundance of vascular epiphytes: a composition of secondary vegetation and primary montane rain forest in the Venezuelan Andes. Plant Ecology, vol. 152, no. 2, p. 145-156. http://dx.doi.org/10.1023/A:1011483901452.
http://dx.doi.org/10.1023/A:101148390145... analysed secondary and primary vegetations in a humid Montana forest in Venezuela. Flores-Palacios and García-Franco (2008)Flores-Palacios, A. and García-Franco, JG., 2008. Habitat isolation changes the beta diversity of vascular epiphyte community in lower montane forest, Veracruz, Mexico. Biodiversity and Conservation, vol. 17, no. 1, p. 191-207. http://dx.doi.org/10.1007/s10531-007-9239-6.
http://dx.doi.org/10.1007/s10531-007-923... compared the relationship between tree size and species richness in forest and pastureland trees. Krömer et al. (2007)Krömer, T., Kessler, M. and Gradstein, SR., 2007. Vertical stratification of vascular ephytes in submontane and montane Forest of the Bolivian Andes: the importance of the understory. Plant Ecology, vol. 189, no. 2, p. 261-278. http://dx.doi.org/10.1007/s11258-006-9182-8.
http://dx.doi.org/10.1007/s11258-006-918... studied the vertical stratification in a submontane and montane forest of the Bolivian Andes. In Brazil, Menini-Neto et al. (2009)MeniniNeto, L., Forzza, RC. and Zappi, D., 2009. Angiosperm epiphytes as conservation indicators in forest fragments: A case study from southeastern Minas Gerais, Brasil. Biodiversity and Conservation, vol. 18, no. 14, p. 3785-3807. http://dx.doi.org/10.1007/s10531-009-9679-2.
http://dx.doi.org/10.1007/s10531-009-967... analysed the angiosperm epiphytes in forest fragments in southeastern Minas Gerais. Bataghin et al. (2010)Bataghin, FA., Barros, F. and Pires, JSR., 2010. Distribuição da comunidade de epífitos vasculares em sítios sob diferentes graus de perturbação na Floresta Nacional do Ipanema, São Paulo, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 33, no. 3, p. 501-512. http://dx.doi.org/10.1590/S0100-84042010000300012.
http://dx.doi.org/10.1590/S0100-84042010... compared the distribution of epiphytes in three sites with different degrees of disturbance in a semi-deciduous seasonal forest in the Floresta Nacional de Ipanema, in the state of São Paulo. In the state of Paraná, Kersten and Kuniyoshi (2009)Kersten, RA. and Kuniyoshi, YS., 2009. Conservação das florestas na Bacia do alto Iguaçú, Paraná – Avaliação da comunidade de epífitas vasculares em diferentes estágios serais. Revista Floresta, vol. 39, no. 1, p. 51-66. evaluated epiphyte composition and distribution in eight areas of the mixed ombrophilous forest of the Alto Iguaçu river basin. In the same forest type, Bonnet et al. (2009)Bonnet, A., Lavoranti, OJ. and Curcio, GR., 2009. Epífitos vasculares no Corredor de Biodiversidade Araucária, bacia do rio Iguaçu, Paraná, Brasil. Cadernos da Biodiversidade, vol. 6, p. 49-70. evaluated the vascular epiphyte diversity in four sites from the Corredor Araucária, relating it to factors of environmental degradation and forest characteristics. Bonnet et al. (2010)Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2010. Relações de epífitos vasculares com fatores ambientais nas florestas do Rio Tibagi, Paraná, Brasil. Biotemas, vol. 23, no. 3, p. 37-47. analysed three sections of riparian vegetation in the Tibagi River to evaluate the composition, distribution and association with environmental factors in each area. The present study evaluated the richness, composition and community structure of vascular epiphytes in two fragments in the upper and lower sections of the Sinos River basin, belonging to ombrophilous and semi-deciduous forests, respectively.

2 Material and Methods

2.1 Study area

This study was conducted in two secondary forest fragments located in the Sinos River basin in the state of Rio Grande do Sul, Brazil: (a) fragment 1 (F1) was in the upper section of the basin, in a rural area, next to the source of the Sinos River, in the municipality of Caraá (29º43'42"S and 50º22'00"W; 408.1 m alt.); 60 ha of predominantly dense ombrophilous forest with some remaining mixed ombrophilous forest; (b) fragment 2 (F2) was in the lower section of the basin, in the municipality of Novo Hamburgo (29º40'54"S and 51º06'56"W; 16.4 m alt.), in the Henrique Luís Roessler park, formally declared a Preservation Area in 2009, in the category of Relevant Area of Ecological Interest. This park is in the urban area of the town and has an area of 54.4 ha of fields and forests (Rosa, 2010Rosa, NA., 2010. Plano de Manejo – Parque Municipal Henrique Luís Roessler – Parcão. Novo Hamburgo: Prefeitura Municipal de Novo Hamburgo. 846 p.) classified as lowlands semi-deciduous seasonal forest (Teixeira et al., 1986Teixeira, MB., Coura-Neto, AB., Pastore, U. and Rangel-Filho, ALR., 1986. Vegetação: as regiões fitoecológicas, sua natureza e seus recursos econômicos – estudo fitogeográfico. In Levantamento de Recursos Naturais.Rio de Janeiro, IBGE, vol. 33, p. 541-620.). The climate in the region of the two fragments is Cfa according to the Köppen classification: temperate with rainfall regularly distributed over the year; and temperature of the hottest month higher than 22 ºC (Moreno, 1961Moreno, JA., 1961. Clima do Rio Grande do Sul.Porto Alegre: Secretaria da Agricultura. 42 p. Secção de Geografia.). In Caraá (F1), rainfall is about 1,600 mm per year (Brigada Militar de Caraá, 2012Brigada Militar de Caraá, 2012. Boletim pluviométrico. Caraá: Brigada Militar de Caraá.); in the weather station close to F2, in the city of Campo Bom (29º41'S; 51º03'W; 25.8 m alt.), rainfall is about 1,500 mm per year.

2.2 Sampling

In both fragments, at least 40 phorophytes of arboreal angiosperms, measuring at least 10 cm in diameter at breast height (CBH) were selected for the study of the epiphytic community using the centered quadrant method (Cottam and Curtis, 1956Cottam, G. and CURTIS, JT., 1956. The use of distance measures in phytosociological sampling. Ecology, vol. 37, no. 3, p. 451-460. http://dx.doi.org/10.2307/1930167.
http://dx.doi.org/10.2307/1930167... ). Sampling points were defined at 20 m intervals along a 200 m transect. The trees were divided according to Braun-Blanquet (1979)Braun-Blanquet, J., 1979. Fitossociologia: bases para el estudio de las comunidades vegetales. Madrid: Blume. 820 p. into four ecological zones: (1) lower bole, (2) upper bole or transition from upper bole to internal crown, (3) internal crown, and (4) external crown.

2.3 Floristic inventory

The floristic inventory was made during bimonthly visits in 2010 and 2011 by means of direct observation of the plants in the epiphytic environment or by using a combination of phorophyte climbing and observation from a distance with binoculars (BUSHNELL® - 96m AT 1000M). Representative fertile specimens were collected, labelled and classified according to the technique described by Guedes-Bruni et al. (2002)Guedes-Bruni, RR., Morim, MPM., LIMA, HC. and SYLVESTRE, L., 2002. Inventário florístico. In SYLVESTRE, LS. and ROSA, MMT. (Eds.). Manual metodológico para estudos botânicos na Mata Atlântica.Rio de Janeiro: Universidade Rural. p. 24-49.. Species were identified according to the taxonomic literature, comparisons with botanical collections in herbaria, or consultations with specialists. Angiosperms were classified according to the APG III system (APG, 2009The Angiosperm Phylogeny Group – APG, 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, vol. 161, no. 2, p. 105-121. http://dx.doi.org/10.1111/j.1095-8339.2009.00996.x.
http://dx.doi.org/10.1111/j.1095-8339.20... ), and the ferns, according to Smith et al. (2006Smith, AR., Kathleen, MP., Schuettpelz, E., Korall, P., Schneider, H. and Wolf, PG., 2006. A classification for extant ferns. Taxon, vol. 55, no. 3, p. 705-731. http://dx.doi.org/10.2307/25065646.
http://dx.doi.org/10.2307/25065646... ; 2008Smith, AR., Pryer, KM., Schuettpelz, E., Korall, P., Schneider, H. and Wolf, PG., 2008. Fern Classification. In RANKER, TA. and HAUFLER, CH. The Biology and Evolution of Ferns and Lycophytes. Cambridge: Cambridge University Press. p. 417-467.). Voucher material was deposited in the Herbarium Anchieta (PACA) in São Leopoldo, Brazil. Species were classified according to the type of association with phorophytes in the ecological categories described by Benzing (1995)Benzing, DH., 1995. The physical mosaic and plant variety in forest canopies. Selbyana, vol. 16, no. 2, p. 159-168.: holoepiphytes do not have any relation with the ground, as they complete all their life cycle in the epiphytic environment, and may be habitual, accidental or facultative; hemiepiphytes have a relationship with the ground, which may be primary, when their growth begins in the phorophyte, or secondary, when their growth begins in the ground. Floristic similarity between the two epiphyte communities was calculated using the Jaccard index based on a matrix of presence and absence of species.

2.4 Richness

For both fragments, a rarefaction curve was obtained (Gotelli and Colwell, 2001Gotelli, NJ. and Colwell, RK., 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness.Ecology Letters, vol. 4, no. 4, p. 379-391. http://dx.doi.org/10.1046/j.1461-0248.2001.00230.x.
http://dx.doi.org/10.1046/j.1461-0248.20... ) to evaluate the association between the increase of specific richness and the number of phorophyte samples, at 95% confidence intervals and using the EstimateS 7.5 software. The curve stabilisation was considered an asymptote. The number of species for the total sample was estimated with the same software and 50 random repeated sampling with the Jackknife 1 nonparametric estimator, which uses data about species presence or absence.

The normal distribution of data about richness was evaluated using the Shapiro-Wilk test using the SPSS 20.0 program. Differences between the richness of the zones in each fragment were evaluated using the Kruskal-Wallis test followed by the Student-Newman-Keuls test, at 5% probability. Mean richness values between fragments were compared using the Mann-Whitney test, and the level of significance was set at 5%.

2.5 Phytosociological parameters

Absolute and relative frequencies of species per phorophyte and zone were calculated using the method described by Waechter (1998)Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.. For each species, a cover score was assigned to the corresponding zone according to the scale described by Kersten and Kunyioshi (2006)Kersten, RA. and Kunyioshi, YS., 2006. Epífitos vasculares na Bacia do Alto Iguaçu, Paraná, Brasil – composição florística. Estudos de Biologia, vol. 28, p. 55-71.: (1) very small and isolated individuals; (3) few small individuals or one larger individual; (5) medium-sized individuals or many small individuals; (7) large individuals or many medium-sized individuals; and (10) very large individuals or many large individuals. The sum of cover scores per zone was used to calculate relative cover for each species. The importance value was the mean of the sum of relative frequencies in the phorophytes and in the zones, and of the relative cover. Specific diversity was determined by the Shannon index (H'), using the natural logarithm of frequency data; equability was determined according to the Pielou index (J’) (Magurran, 1988Magurran, AE., 1988. Ecological diversity and its measurement. Cambridge: British Library.).

3 Results

3.1 Floristic inventory

In F1, there were 30 species distributed into 15 genera and four families, and in F2, 25 species in 19 genera and eight families (Table 1). The three richest families in both fragments were Polypodiaceae, Orchidaceae and Bromeliaceae. Aspleniaceae, Blechnaceae and Pteridaceae had only one species each. Besides Pleopeltis Humb. & Bonpl. ex Willd. in F1, Tillandsia L. was the richest genus in both fragments. A total of eight genera had only one species. The analysis of ecological category revealed that habitual holoepiphytes were predominant, with 97.56% (39 species) of the total number of species. Asplenium claussenii and Blechnum binervatum were the only species classified as accidental holoepiphytes and secondary hemiepiphytes, both found only in F2. Of all the species found in the two fragments, 15 were common to both fragments, 15 were exclusive of F1, and 10, of F2. The similarity between F1 and F2 was low, and the Jaccard index was 0.38.

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Table 1
Families and species of vascular epiphytes recorded in fragments 1 and 2, located in the upper and lower sections of the Sinos River basin. (HAB – habitual holoepiphyte; SHE = secondary hemiepiphyte; ACC = accidental holoepiphyte).

3.2 Richness

The rarefaction curve based on the number of sampling units in both areas was not an asymptote (Figure 1). The richness estimator (Jackknife 1) showed that there might be even more species, and that 32.9 were estimated for F1 and 29.9 for F2. Mean total richness in F1 was 6.5 ± 2.7 species per tree, and a minimum of two and maximum of 13 species per phorophyte were recorded. In F2, mean total richness was 6.0 ± 2.1 species per tree, and a minimum of three and maximum of 11 species per phorophyte were recorded. Statistical analyses showed that means were not significantly different (U = 710.00; P = 0.386). In F1, phorophyte zone 2 had the highest mean richness without, however, being significantly different from zone 1. The highest richness means in F2 were recorded in the intermediate phorophyte zones (2 and 3). The comparison of the two areas revealed that mean richness recorded in zone 1 of the trees in F1 was higher than in the same zone of the phorophytes in F2, and the opposite was found in zones 3 and 4. In F1 and F2, zone 2 had statistically equivalent mean values of epiphytic species (Table 2).

Figure 1
Rarefaction curves and vascular epiphytic richness estimate (Jackknife 1) in fragments of the Atlantic Forest in the (a) upper (F1) and (b) lower (F2) sections of the Sinos River basin, in the state of Rio Grande do Sul, Brazil. Bars indicate confidence interval.
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Table 2
Mean vascular epiphytic richness in the different phorophyte zones in a fragment of the Atlantic Forest of the upper (F1) and lower (F2) sections of the Sinos River basin, in the state of Rio Grande do Sul, Brazil.

3.3 Phytosociological parameters

Microgramma squamulosa had the highest importance value (IV) in both fragments and accounted for 87.5% of the phorophytes in F1 and 80.0% in F2. In F1, the second most important species was Pleopeltis pleopeltifolia (11.65%), and in F2, Campyloneurum nitidum (15.74%), which accounted for 62.5% and 82.5% of the phorophytes, respectively (Tables 3 and 4). In F1, Microgramma vacciniifolia, Cattleya intermedia and Lophiaris pumila were recorded in only one phorophyte and had, consequently, the lowest importance value (Table 3). In F2, Billbergia nutans, Vriesea rodigasiana, Blechnum binervatum and Pleopeltis hirsutissima were also found in only one phorophyte and had the lowest importance value (Table 4). The Shannon diversity indices (H’) estimated for F1 and F2 were 2.72 and 2.55, and the equability indices (J) were 0.80 and 0.79.

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Table 3
Community structure of vascular epiphytes in fragment 1 in decreasing order of importance value (IV). (np= number of phorophytes with species; nz= number of zones with species; AFf = absolute frequency of species on phorophytes; RFf = relative frequency of species on phorophytes; AFz = absolute frequency of species on zones; RFz = relative frequency of species on zones; ADo = absolute dominance of species (sum of coverage grades); RDo = relative dominance of species).
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Table 4
Community structure of vascular epiphytes in fragment 2 in decreasing order of importance value (IV). (np= number of phorophytes with species; nz= number of zones with species; AFf = absolute frequency of species on phorophytes; RFf = relative frequency of species on phorophytes; AFz = absolute frequency of species on zones; RFz = relative frequency of species on zones; Ado = absolute dominance of species (sum of coverage grades); RDo = relative dominance of species).

4 Discussion

The area where the dense ombrophilous forest (F1) was predominant had a higher specific richness than the semi-deciduous seasonal forest (F2). The lower epiphytic richness of seasonal forests when compared with ombrophilous forests has already been described by other authors in the southern region of Brazil (Aguiar et al., 1981Aguiar, LW., Citadini-Zanette, V., Martau, L. and Backes, A., 1981. Composição florística de epífitos vasculares numa área localizada nos municípios de Montenegro e Triunfo, Rio Grande do Sul, Brasil. Iheringia Série Botânica, vol. 28, no. 1, p. 55-93.; Waechter, 1998Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.; Borgo et al., 2002Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130.; Borgo and Silva, 2003Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
http://dx.doi.org/10.1590/S0100-84042003... ). The specific richness in F1 was lower than that found by Blum et al. (2011)Blum, CT., Roderjan, CV. and Galvão, F., 2011. Composição florística e distribuição altitudinal de epífitas vasculares da Floresta Ombrófila Densa na Serra da Prata, Morretes, Paraná, Brasil. Biota Neotropica, vol. 11, no. 4, p. 141-159. http://dx.doi.org/10.1590/S1676-06032011000400015.
http://dx.doi.org/10.1590/S1676-06032011... who recorded 278 species in 6.3 ha of ombrophilous forest which characterize a primary forest with little disturbance (Blum, 2006Blum, CT., 2006. A Floresta Ombrófila Densa na Serra da Prata, Parque Nacional Saint-Hilaire/Lange, PR - Caracterização florística, fitossociológica e ambiental de um gradiente altitudinal. Curitiba: Universidade Federal do Paraná. 195 p. Dissertação de Mestrado em Ciências Florestais.). The specific richness in F2 was greater than that found in other studies also conducted in the semi-deciduous seasonal forest. Aguiar et al. (1981)Aguiar, LW., Citadini-Zanette, V., Martau, L. and Backes, A., 1981. Composição florística de epífitos vasculares numa área localizada nos municípios de Montenegro e Triunfo, Rio Grande do Sul, Brasil. Iheringia Série Botânica, vol. 28, no. 1, p. 55-93. recorded 19 species in 37 phorophytes; Dettke et al. (2008)Dettke, GA., Orfrini, AC. and Milanese-Gutierre, MA., 2008. Composição florística e distribuição de epífitas vasculares em um remanescente alterado da Floresta Estacional Semidecidual do Paraná, Brasil. Rodriguésia, vol. 59, no. 4, p. 859-872., 22 species in 90 phorophytes; and Bataghin et al. (2010)Bataghin, FA., Barros, F. and Pires, JSR., 2010. Distribuição da comunidade de epífitos vasculares em sítios sob diferentes graus de perturbação na Floresta Nacional do Ipanema, São Paulo, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 33, no. 3, p. 501-512. http://dx.doi.org/10.1590/S0100-84042010000300012.
http://dx.doi.org/10.1590/S0100-84042010... , 21 species in 270 phorophytes. Only Borgo et al. (2002)Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130. recorded a richer epiphytic flora (32 species), but based on the observation of 2,895 trees.

The families with the highest number of species in this study (Orchidaceae, Bromeliaceae and Polypodiaceae) are also the richest worldwide (Madison, 1977Madison, M., 1977. Vascular epiphytes: their systematic occurrence and salient features. Selbyana, vol. 2, no. 1, p. 1-13.; Kress, 1986Kress, WJ., 1986. The systematic distribution of vascular epiphytes: an update. Selbyana, vol. 9, no. 1, p. 2-22.; Benzing, 1990Benzing, DH., 1990. Vascular epiphytes. Cambridge: Cambridge University Press. 354 p.), in neotropical regions (Gentry and Dodson, 1987GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
http://dx.doi.org/10.2307/2399395... ) and in Brazil (Kersten, 2010Kersten, RA., 2010. Epífitas vasculares – Histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea, vol. 37, no. 1, p. 9-38. http://dx.doi.org/10.1590/S2236-89062010000100001.
http://dx.doi.org/10.1590/S2236-89062010... ). Other inventories of vascular epiphytes conducted in the Brazilian states of Rio Grande do Sul (Waechter, 1986Waechter, JL., 1986. Epífitos vasculares da mata paludosa do Faxinal, Torres, Rio Grande do Sul, Brasil. Iheringia, vol. 34, p. 39-49., 1998Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.; Gonçalves and Waechter, 2003Gonçalves, CN. and Waechter, JL., 2003. Aspectos florísticos e ecológicos de epífitos vasculares sobre figueiras isoladas no norte da planície costeira do Rio Grande do Sul. Acta Botanica Brasilica, vol. 17, no. 1, p. 89-100. http://dx.doi.org/10.1590/S0102-33062003000100007.
http://dx.doi.org/10.1590/S0102-33062003... ; Rogalski and Zanin, 2003Rogalski, JM. and Zanin, EM., 2003. Composição florística de epífitos vasculares no estreito de Augusto César, Floresta Estacional Decidual do Rio Uruguai, RS, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 4, p. 551-556. http://dx.doi.org/10.1590/S0100-84042003000400014.
http://dx.doi.org/10.1590/S0100-84042003... ; Buzatto et al., 2008Buzatto, C.R., SEVERO, B M.A. and WAECHTER, J.L., 2008. Composição florística e distribuição ecológica de epífitos vasculares na Floresta Nacional de Passo Fundo, Rio Grande do Sul. Iheringia Serie Botânica, vol. 63, no. 2, p. 231-239.; Staudt et al., 2012Staudt, MG., Lippert, APU., Cunha, S., Becker, DFP., Marchioretto, MS. and Schmitt, JL., 2012. Composição florística de epífitos vasculares do Parque Natural Municipal Tupancy, Arroio do Sal, RS – Brasil. Pesquisas, vol. 63, p. 177-188.) and of Paraná (Kersten and Silva, 2001Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... ; Borgo and Silva, 2003Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
http://dx.doi.org/10.1590/S0100-84042003... ; Hefler and Faustioni, 2004Hefler, SM. and Faustioni, P., 2004. Levantamento florístico de epífitos vasculares do bosque São Cristovão –Curitiba – Paraná – Brasil. Revista Estudos de Biologia, vol. 26, no. 54, p. 11-19.; Dettke et al., 2008Dettke, GA., Orfrini, AC. and Milanese-Gutierre, MA., 2008. Composição florística e distribuição de epífitas vasculares em um remanescente alterado da Floresta Estacional Semidecidual do Paraná, Brasil. Rodriguésia, vol. 59, no. 4, p. 859-872.; Kersten et al., 2009Kersten, RA., Kuniyidhi, YS. and Roderjan, CV., 2009. Epífitas vasculares em duas formações ribeirinhas adjacentes na bacia do rio Iguaçu – Terceiro Planalto Paranaense. Iheringia Série Botânica, vol. 64, no. 1, p. 33-43.; Bonnet et al., 2010Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2010. Relações de epífitos vasculares com fatores ambientais nas florestas do Rio Tibagi, Paraná, Brasil. Biotemas, vol. 23, no. 3, p. 37-47.; 2011Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2011. Flora epifítica vascular em três unidades vegetacionais do Rio Tibagi, Paraná, Brasil.Rodriguésia, vol. 62, no. 3, p. 491-498.; Blum et al., 2011Blum, CT., Roderjan, CV. and Galvão, F., 2011. Composição florística e distribuição altitudinal de epífitas vasculares da Floresta Ombrófila Densa na Serra da Prata, Morretes, Paraná, Brasil. Biota Neotropica, vol. 11, no. 4, p. 141-159. http://dx.doi.org/10.1590/S1676-06032011000400015.
http://dx.doi.org/10.1590/S1676-06032011... ) showed that these families have a great specific richness.

There was a concentration of vascular epiphyte species in few families, as found by Madison (1977)Madison, M., 1977. Vascular epiphytes: their systematic occurrence and salient features. Selbyana, vol. 2, no. 1, p. 1-13., Kress (1986)Kress, WJ., 1986. The systematic distribution of vascular epiphytes: an update. Selbyana, vol. 9, no. 1, p. 2-22., Gentry and Dodson (1987)GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
http://dx.doi.org/10.2307/2399395... and Kersten (2010)Kersten, RA., 2010. Epífitas vasculares – Histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea, vol. 37, no. 1, p. 9-38. http://dx.doi.org/10.1590/S2236-89062010000100001.
http://dx.doi.org/10.1590/S2236-89062010... . According to Gentry and Dodson (1987)GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
http://dx.doi.org/10.2307/2399395... , a small number of families may also reflect the specialization of some of these families in the environments under study. This trend in the distribution of species was also found in other inventories of epiphytic communities conducted in southern and southeastern Brazil (Waechter, 1986Waechter, JL., 1986. Epífitos vasculares da mata paludosa do Faxinal, Torres, Rio Grande do Sul, Brasil. Iheringia, vol. 34, p. 39-49.; 1992Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.; 1998Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.; Kersten and Silva, 2001Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... , Borgo et al., 2002Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130.; Borgo and Silva, 2003Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
http://dx.doi.org/10.1590/S0100-84042003... ; Gonçalves and Waechter, 2003Gonçalves, CN. and Waechter, JL., 2003. Aspectos florísticos e ecológicos de epífitos vasculares sobre figueiras isoladas no norte da planície costeira do Rio Grande do Sul. Acta Botanica Brasilica, vol. 17, no. 1, p. 89-100. http://dx.doi.org/10.1590/S0102-33062003000100007.
http://dx.doi.org/10.1590/S0102-33062003... ; Rogalski and Zanin, 2003Rogalski, JM. and Zanin, EM., 2003. Composição florística de epífitos vasculares no estreito de Augusto César, Floresta Estacional Decidual do Rio Uruguai, RS, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 4, p. 551-556. http://dx.doi.org/10.1590/S0100-84042003000400014.
http://dx.doi.org/10.1590/S0100-84042003... ; Hefler and Faustioni, 2004Hefler, SM. and Faustioni, P., 2004. Levantamento florístico de epífitos vasculares do bosque São Cristovão –Curitiba – Paraná – Brasil. Revista Estudos de Biologia, vol. 26, no. 54, p. 11-19.; Buzatto et al., 2008Buzatto, C.R., SEVERO, B M.A. and WAECHTER, J.L., 2008. Composição florística e distribuição ecológica de epífitos vasculares na Floresta Nacional de Passo Fundo, Rio Grande do Sul. Iheringia Serie Botânica, vol. 63, no. 2, p. 231-239.; Dettke et al., 2008Dettke, GA., Orfrini, AC. and Milanese-Gutierre, MA., 2008. Composição florística e distribuição de epífitas vasculares em um remanescente alterado da Floresta Estacional Semidecidual do Paraná, Brasil. Rodriguésia, vol. 59, no. 4, p. 859-872.; Menini-Neto et al., 2009MeniniNeto, L., Forzza, RC. and Zappi, D., 2009. Angiosperm epiphytes as conservation indicators in forest fragments: A case study from southeastern Minas Gerais, Brasil. Biodiversity and Conservation, vol. 18, no. 14, p. 3785-3807. http://dx.doi.org/10.1007/s10531-009-9679-2.
http://dx.doi.org/10.1007/s10531-009-967... ; Mania and Monteiro, 2010Mania, LF. and Monteiro, R., 2010. Florística e ecologia de epífitas vasculares em um fragmento de floresta de restinga, Ubatuba, SP, Brasil. Rodriguésia, vol. 61, no. 4, p. 705-713.; Staudt et al., 2012Staudt, MG., Lippert, APU., Cunha, S., Becker, DFP., Marchioretto, MS. and Schmitt, JL., 2012. Composição florística de epífitos vasculares do Parque Natural Municipal Tupancy, Arroio do Sal, RS – Brasil. Pesquisas, vol. 63, p. 177-188.). The three richest families accounted for 82.5% of the recorded species. According to Kersten (2010)Kersten, RA., 2010. Epífitas vasculares – Histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea, vol. 37, no. 1, p. 9-38. http://dx.doi.org/10.1590/S2236-89062010000100001.
http://dx.doi.org/10.1590/S2236-89062010... , 91% of the species in Brazil are part of the 10 richest families, and in the case of the 20 richest families, this percentage may reach 98% of the total species recorded. The success of these three families in the epiphytic environment is associated with several adaptations to drought resistance and water saving (Staudt et al., 2012Staudt, MG., Lippert, APU., Cunha, S., Becker, DFP., Marchioretto, MS. and Schmitt, JL., 2012. Composição florística de epífitos vasculares do Parque Natural Municipal Tupancy, Arroio do Sal, RS – Brasil. Pesquisas, vol. 63, p. 177-188.).

The predominance of habitual holoepiphytes over the other categories has also been observed by Aguiar et al. (1981)Aguiar, LW., Citadini-Zanette, V., Martau, L. and Backes, A., 1981. Composição florística de epífitos vasculares numa área localizada nos municípios de Montenegro e Triunfo, Rio Grande do Sul, Brasil. Iheringia Série Botânica, vol. 28, no. 1, p. 55-93., Waechter (1986Waechter, JL., 1986. Epífitos vasculares da mata paludosa do Faxinal, Torres, Rio Grande do Sul, Brasil. Iheringia, vol. 34, p. 39-49.; 1998Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.); Kersten and Silva (2001Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... ; 2002Kersten, RA. and Silva, SM., 2002. Florística e estrutura do componente epifítico vascular em Floresta Ombrófila Mista aluvial do Rio Barigüi, Paraná, Brasil. Revista Brasileira de Botânica, São Paulo, vol. 25, no. 3, p. 259-267.); Borgo et al. (2002)Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130.; Borgo and Silva (2003)Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
http://dx.doi.org/10.1590/S0100-84042003... ; Gonçalves and Waechter (2003)Gonçalves, CN. and Waechter, JL., 2003. Aspectos florísticos e ecológicos de epífitos vasculares sobre figueiras isoladas no norte da planície costeira do Rio Grande do Sul. Acta Botanica Brasilica, vol. 17, no. 1, p. 89-100. http://dx.doi.org/10.1590/S0102-33062003000100007.
http://dx.doi.org/10.1590/S0102-33062003... ; Rogalski and Zanin (2003)Rogalski, JM. and Zanin, EM., 2003. Composição florística de epífitos vasculares no estreito de Augusto César, Floresta Estacional Decidual do Rio Uruguai, RS, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 4, p. 551-556. http://dx.doi.org/10.1590/S0100-84042003000400014.
http://dx.doi.org/10.1590/S0100-84042003... , Giongo and Waechter (2004)Giongo, C. and Waechter, JL., 2004. Composição florística e estrutura comunitária de epífitos vasculares em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 27, no. 3, p. 563-572. http://dx.doi.org/10.1590/S0100-84042004000300015.
http://dx.doi.org/10.1590/S0100-84042004... ; Kersten and Kunyioshi (2006)Kersten, RA. and Kunyioshi, YS., 2006. Epífitos vasculares na Bacia do Alto Iguaçu, Paraná, Brasil – composição florística. Estudos de Biologia, vol. 28, p. 55-71. and Schmitt and Windisch (2010)Schmitt, JL. and Windisch, PG., 2010. Biodiversity and spatial distribution of epiphytic ferns on . Alsophila setosa Kaulf. (Cyatheaceae) caudices in Rio Grande do Sul, BrazilBrazilian Journal of Biology = Revista Brasleira de Biologia, vol. 70, no. 3, p. 521-528. http://dx.doi.org/10.1590/S1519-69842010000300008. PMid:20730338.
http://dx.doi.org/10.1590/S1519-69842010... . Habitual holoepiphytes had more specialized and diversified vegetative adaptations, which favoured a more generalized distribution of forest formations (Fraga et al., 2008Fraga, LL., Silva, LB. and Schmitt, JL., 2008. Composição e distribuição vertical de pteridófitas epifíticas sobre Hook. (Dicksoniaceae), em Floresta Ombrófila Mista no sul do Brasil. Dicksonia sellowianaBiota Neotropica, vol. 8, no. 4, p. 123-129. http://dx.doi.org/10.1590/S1676-06032008000400011.
http://dx.doi.org/10.1590/S1676-06032008... ).

The inventory of all species in an area is virtually impossible (Santos, 2004Santos, AJ., 2004. Estimativas de riqueza em espécies. In CULLEN-JÚNIOR, L., VALLADARES-PADUA, C. and RUDY, R. (Eds.). Métodos de estudos em biologia da conservação e manejo da vida silvestre. Curitiba: UFPR. p. 19-41.). The richness estimator indicated that 91% and 83% of the total richness was sampled in F1 and F2. These estimates are not precise previews of the actual number of species, but they indicate minimum expected values (Colwell et al., 2004Colwell, RK., Mao, CX. and Chang, J., 2004. Interpolatin, extrapolatin, and comparing incidence-based species accumulation curves.Ecology, vol. 85, no. 10, p. 2717-2727. http://dx.doi.org/10.1890/03-0557.
http://dx.doi.org/10.1890/03-0557... ) in the epiphytic communities in both fragments.

Microgramma squamulosa (Polypodiaceae) had the highest importance value in both fragments, in the same way as found in studies conducted by Kersten and Silva (2001)Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... , Kersten and Kuniyoshi (2009)Kersten, RA. and Kuniyoshi, YS., 2009. Conservação das florestas na Bacia do alto Iguaçú, Paraná – Avaliação da comunidade de epífitas vasculares em diferentes estágios serais. Revista Floresta, vol. 39, no. 1, p. 51-66. and Geraldino et al. (2010)Geraldino, HCL., Caxambú, MG. and SOUZA, DC., 2010. Composição florística e estrutura da comunidade de epífitas vasculares em uma área de ecótono em Campo Mourão, PR, Brasil. Acta Botanica Brasilica, vol. 24, no. 2, p. 469-482. http://dx.doi.org/10.1590/S0102-33062010000200018.
http://dx.doi.org/10.1590/S0102-33062010... . This species has been described as a pioneer (Kersten and Silva, 2001Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... ; Geraldino et al., 2010Geraldino, HCL., Caxambú, MG. and SOUZA, DC., 2010. Composição florística e estrutura da comunidade de epífitas vasculares em uma área de ecótono em Campo Mourão, PR, Brasil. Acta Botanica Brasilica, vol. 24, no. 2, p. 469-482. http://dx.doi.org/10.1590/S0102-33062010000200018.
http://dx.doi.org/10.1590/S0102-33062010... ), which confirms the secondary character of the fragments under analysis. Microgramma squamulosa is found in not very dense primary and secondary forests, over isolated trees, even in parks, as well as on stone walls (Sehnem, 1970Sehnem, A.,1970. Polipodiáceas. In REITZ, R. (Ed.). Flora ilustrada catarinense. Itajaí: Herbário Barbosa Rodrigues. p. 1-173.), which confirms its high plasticity and adaptation to different environments. It extends over boles and branches and takes up large areas, which is the reason why it has small individuals and reptant growth.

The species with the second highest importance value in F1, Pleopeltis pleopeltifolia (Polypodiaceae), was also the second in the mixed ombrophilous forest (Kersten and Kuniyoshi, 2009Kersten, RA. and Kuniyoshi, YS., 2009. Conservação das florestas na Bacia do alto Iguaçú, Paraná – Avaliação da comunidade de epífitas vasculares em diferentes estágios serais. Revista Floresta, vol. 39, no. 1, p. 51-66.) in the state of Paraná, as well as in the ecotone between the semi-deciduous seasonal and the mixed ombrophilous forests (Geraldino et al., 2010Geraldino, HCL., Caxambú, MG. and SOUZA, DC., 2010. Composição florística e estrutura da comunidade de epífitas vasculares em uma área de ecótono em Campo Mourão, PR, Brasil. Acta Botanica Brasilica, vol. 24, no. 2, p. 469-482. http://dx.doi.org/10.1590/S0102-33062010000200018.
http://dx.doi.org/10.1590/S0102-33062010... ) in the same state. In the same way as in F2, Campyloneurum nitidum (Polypodiaceae) had the second highest importance value in the inventory conducted by Schneider and Schmitt (2011)Schneider, PH. and Schmitt, JL., 2011. Composition, community structure and vertical distribution of epiphytic ferns on . Alsophila setosa Kaulf., in a Semideciduous Seasonal Forest, Morro Reuter, RS, BrasilActa Botanica Brasilica, vol. 25, no. 3, p. 557-562. http://dx.doi.org/10.1590/S0102-33062011000300007.
http://dx.doi.org/10.1590/S0102-33062011... in a semi-deciduous seasonal forest in the state of Rio Grande do Sul, but they considered only phorophytes of Alsophila setosa Kaulf. In these three Polypodiaceae species, adaptations such as frond trichomes (Müller et al., 1981Müller, L., Starnecker, G. and Winkler, S., 1981. Zur Ökologie epiphytischer Farne in Südbrasilien. I. Saugschuppen. Flora, vol. 171, p. 55-65.), rhizome succulence (Waechter, 1992Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.), poikilohydry (Benzing, 1990Benzing, DH., 1990. Vascular epiphytes. Cambridge: Cambridge University Press. 354 p.) and the higher sclerophylly index and stomatal density prevent dehydration (Rocha et al., 2013Rocha, LD., Droste, A., Gehlen, G. and Schmitt, JL., 2013. Leaf dimorphism of (Polypodiaceae): a qualitative and quantitative analysis focusing on adaptations to epiphytism. Microgramma squamulosaRevista de Biología Tropical, vol. 61, no. 1, p. 291-299. PMid:23894982.) and facilitate a greater occupation of phorophytes, increasing their importance in the organisation of the epiphytic community.

Diversity indices found in F1 and F2 were very similar, and mean total richness recorded in phorophytes was statistically equivalent. The values of the Shannon index in the fragments were close to those recorded by Waechter (1998)Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66. in Emboaba, in the city of Osório (2.99), and by Waechter (1992)Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais. in Taim, in the city of Rio Grande (2.89), in the coastal plains of Rio Grande do Sul, as well as by Kersten and Silva (2002)Kersten, RA. and Silva, SM., 2002. Florística e estrutura do componente epifítico vascular em Floresta Ombrófila Mista aluvial do Rio Barigüi, Paraná, Brasil. Revista Brasileira de Botânica, São Paulo, vol. 25, no. 3, p. 259-267. in a mixed ombrophilous forest (2.71) in the state of Paraná, Brazil. They were lower than the values found by Giongo and Waechter (2004)Giongo, C. and Waechter, JL., 2004. Composição florística e estrutura comunitária de epífitos vasculares em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 27, no. 3, p. 563-572. http://dx.doi.org/10.1590/S0100-84042004000300015.
http://dx.doi.org/10.1590/S0100-84042004... in riparian forests in Rio Grande do Sul (3.43), Kersten and Silva (2001)Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
http://dx.doi.org/10.1590/S0100-84042001... in Ilha do Mel (3.61), Kersten and Kunyioshi (2006)Kersten, RA. and Kunyioshi, YS., 2006. Epífitos vasculares na Bacia do Alto Iguaçu, Paraná, Brasil – composição florística. Estudos de Biologia, vol. 28, p. 55-71. in a dense ombrophilous forest (4.07) and Geraldino et al. (2010)Geraldino, HCL., Caxambú, MG. and SOUZA, DC., 2010. Composição florística e estrutura da comunidade de epífitas vasculares em uma área de ecótono em Campo Mourão, PR, Brasil. Acta Botanica Brasilica, vol. 24, no. 2, p. 469-482. http://dx.doi.org/10.1590/S0102-33062010000200018.
http://dx.doi.org/10.1590/S0102-33062010... in a transition area between a mixed ombrophilous forest and a semi-deciduous seasonal forest (3.17) in the state of Paraná. The lower values found in this study, particularly in relation to other places in Paraná, reflect their geographic location, which was characteristically more subtropical in the areas under analysis. The high diversity and the richness of vascular epiphytes in the tropical regions visually decrease towards the south because of the lower temperatures and greater rainfall, as well as the variations in environmental conditions due to the topography (Waechter, 1992Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.).

The trend towards an increase in mean richness from zone 1 to zone 2 in both fragments is associated with the fact that boles are areas that do not usually promote epiphyte colonisation because of their verticality (Bøgh, 1992Bøgh, A., 1992. The composition and distribution of the vascular epiphyte flora of an Ecuadorian montane rain forest. Selbyana, vol. 13, p. 25-34.), less luminosity (Parker, 1995Parker, GG., 1995. Structure and microclimate of forest canopies. In LOWMAN, MD. and NADKARNI, NM. (Eds.). Forest canopies. San Diego: Academic Press. p. 73-106.) and exposure to the action of herbivorous animals (bovine cattle) that move around the trees (Gonçalves and Waechter, 2002Gonçalves, CN. and Waechter, JL., 2002. Epífitos vasculares sobre espécimes de .Ficus organensis isolados no norte da planície costeira do Rio Grande do Sul: padrões de abundância e distribuiçãoActa Botanica Brasilica, vol. 16, no. 4, p. 429-441. http://dx.doi.org/10.1590/S0102-33062002000400006.
http://dx.doi.org/10.1590/S0102-33062002... ), as well as to the extraction of ornamental epiphytic plants. Human action may be one of the determinant factors of differences in richness found in zone 1 in F1 when compared with F2. In F2, an urban park used for leisure, public visitation is intense, and epiphytes with any ornamental value may have been removed illegally from the tree boles. Epiphytes are usually more diversified in intermediate zones, where phorophyte structures provide the necessary space and support for epiphytic flora (Nieder et al., 1999Nieder, J., Engwald, S. and Barthlott, W., 1999. Patterns of neotropical epiphyte diversity. Selbyana, vol. 20, no. 1, p. 66-75.), as seen in both areas under analysis, but more evident in F2. The decrease in mean richness in both fragments from zone 3 to zone 4 is associated with the fact that the latter is the most recent part of the tree and, consequently, it had the shortest time to receive epiphytes. Yeaton and Gladstone (1982)Yeaton, RI. and Gladstone, DE., 1982. The pattern of colonization of epiphytes on Calabash Trees (. Crescentia alata HBK.) in Guanacaste Province, Costa RicaBiotropica, vol. 14, no. 2, p. 137-140. http://dx.doi.org/10.2307/2387743.
http://dx.doi.org/10.2307/2387743... reported that substrate availability time is an important factor in the development of epiphytic plants in phorophytes. The external crown branches are thinner with smaller areas and less diversified structurally, which may result in a smaller number of species. In addition, in the external crown there is great microclimate variability characterised by long periods of low humidity, high temperatures and strong winds (Krömer et al., 2007Krömer, T., Kessler, M. and Gradstein, SR., 2007. Vertical stratification of vascular ephytes in submontane and montane Forest of the Bolivian Andes: the importance of the understory. Plant Ecology, vol. 189, no. 2, p. 261-278. http://dx.doi.org/10.1007/s11258-006-9182-8.
http://dx.doi.org/10.1007/s11258-006-918... ).

Although both F1 and F2 belong to the Atlantic Forest Biome and had similar richness means and diversity indices, there were exclusive species in each area, which resulted in low similarity and revealed a characteristic floristic identity in each area. This low coefficient of similarity between epiphytic floras of the two study areas was expected due to environmental differences such as altitude and forest type. Regardless of the nature of the matrix, whether urban or rural, in which the fragments are located, all should be taken into consideration when defining handling and preservation procedures. Therefore, if more fragments are protected, a larger number of epiphytic species may be preserved in the Sinos River basin.

Acknowledgements

The authors wish to thank: the Universidade Feevale for the use of the laboratory and financial support; the Coordenação de Aperfeiçoamento de Pessoal de Nivel Superior (CAPES) for the Master´s grant awarded to M.D. Barbosa; the Fundação de Amparo à Pesquisa no Rio Grande do Sul (FAPERGS) for the scholarship awarded to S. Cunha; the land owner and the Henrique Luís Roessler park staff for research permission and logistic assistance.

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References

  • Aguiar, LW., Citadini-Zanette, V., Martau, L. and Backes, A., 1981. Composição florística de epífitos vasculares numa área localizada nos municípios de Montenegro e Triunfo, Rio Grande do Sul, Brasil. Iheringia Série Botânica, vol. 28, no. 1, p. 55-93.
  • BarthloTt, W., Schmit-Neuerburg, V., Nieder, J. and Engwald, S., 2001. Diversity and abundance of vascular epiphytes: a composition of secondary vegetation and primary montane rain forest in the Venezuelan Andes. Plant Ecology, vol. 152, no. 2, p. 145-156. http://dx.doi.org/10.1023/A:1011483901452.
    » http://dx.doi.org/10.1023/A:1011483901452
  • Bataghin, FA., Barros, F. and Pires, JSR., 2010. Distribuição da comunidade de epífitos vasculares em sítios sob diferentes graus de perturbação na Floresta Nacional do Ipanema, São Paulo, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 33, no. 3, p. 501-512. http://dx.doi.org/10.1590/S0100-84042010000300012.
    » http://dx.doi.org/10.1590/S0100-84042010000300012
  • Becker, DFP., Cunha, S., Marchioretto, MS. and Schmitt, JL., 2013. Riqueza, estrutura comunitária e distribuição vertical de epífitos vasculares no Parque Natural Municipal Tupancy, Arroio do Sal, RS, Brasil. Pesquisas, vol. 64, p. 127-139.
  • Benzing, DH., 1990. Vascular epiphytes. Cambridge: Cambridge University Press. 354 p.
  • Benzing, DH., 1995. The physical mosaic and plant variety in forest canopies. Selbyana, vol. 16, no. 2, p. 159-168.
  • Blum, CT., 2006. A Floresta Ombrófila Densa na Serra da Prata, Parque Nacional Saint-Hilaire/Lange, PR - Caracterização florística, fitossociológica e ambiental de um gradiente altitudinal. Curitiba: Universidade Federal do Paraná. 195 p. Dissertação de Mestrado em Ciências Florestais.
  • Blum, CT., Roderjan, CV. and Galvão, F., 2011. Composição florística e distribuição altitudinal de epífitas vasculares da Floresta Ombrófila Densa na Serra da Prata, Morretes, Paraná, Brasil. Biota Neotropica, vol. 11, no. 4, p. 141-159. http://dx.doi.org/10.1590/S1676-06032011000400015.
    » http://dx.doi.org/10.1590/S1676-06032011000400015
  • Bøgh, A., 1992. The composition and distribution of the vascular epiphyte flora of an Ecuadorian montane rain forest. Selbyana, vol. 13, p. 25-34.
  • Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2010. Relações de epífitos vasculares com fatores ambientais nas florestas do Rio Tibagi, Paraná, Brasil. Biotemas, vol. 23, no. 3, p. 37-47.
  • Bonnet, A., Curcio, GR., Lavorant, OJ. and GalvÃo, F., 2011. Flora epifítica vascular em três unidades vegetacionais do Rio Tibagi, Paraná, Brasil.Rodriguésia, vol. 62, no. 3, p. 491-498.
  • Bonnet, A., Lavoranti, OJ. and Curcio, GR., 2009. Epífitos vasculares no Corredor de Biodiversidade Araucária, bacia do rio Iguaçu, Paraná, Brasil. Cadernos da Biodiversidade, vol. 6, p. 49-70.
  • Borgo, M. and Silva, SM., 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 3, p. 391-401. http://dx.doi.org/10.1590/S0100-84042003000300012.
    » http://dx.doi.org/10.1590/S0100-84042003000300012
  • Borgo, M., Petean, M. and Silva, SM., 2002. Epífitos vasculares em um remanescente de floresta estacional semidecidual, município de Fênix, PR, Brasil. Acta Biologica Leopoldensia, vol. 24, no. 2, p. 121-130.
  • Brasil. Conselho Nacional de Meio Ambiente – CONAMA, Ministério do Meio Ambiente – MMA, 2012. Resoluções do CONAMA: Resoluções vigentes publicadas entre setembro de 1984 e janeiro de 2012. Brasília: MMA. 1126 p.
  • Braun-Blanquet, J., 1979. Fitossociologia: bases para el estudio de las comunidades vegetales. Madrid: Blume. 820 p.
  • Brigada Militar de Caraá, 2012. Boletim pluviométrico. Caraá: Brigada Militar de Caraá.
  • Brustulin, J. and Schmitt, JL., 2008. Composição florística, distribuição vertical e floração de orquídeas epifíticas em três parques municipais do estado do Rio Grande do Sul, Brasil. Pesquisas, vol. 59, p. 143-158.
  • Buzatto, C.R., SEVERO, B M.A. and WAECHTER, J.L., 2008. Composição florística e distribuição ecológica de epífitos vasculares na Floresta Nacional de Passo Fundo, Rio Grande do Sul. Iheringia Serie Botânica, vol. 63, no. 2, p. 231-239.
  • Colwell, RK., Mao, CX. and Chang, J., 2004. Interpolatin, extrapolatin, and comparing incidence-based species accumulation curves.Ecology, vol. 85, no. 10, p. 2717-2727. http://dx.doi.org/10.1890/03-0557.
    » http://dx.doi.org/10.1890/03-0557
  • Cottam, G. and CURTIS, JT., 1956. The use of distance measures in phytosociological sampling. Ecology, vol. 37, no. 3, p. 451-460. http://dx.doi.org/10.2307/1930167.
    » http://dx.doi.org/10.2307/1930167
  • Dettke, GA., Orfrini, AC. and Milanese-Gutierre, MA., 2008. Composição florística e distribuição de epífitas vasculares em um remanescente alterado da Floresta Estacional Semidecidual do Paraná, Brasil. Rodriguésia, vol. 59, no. 4, p. 859-872.
  • Flores-Palacios, A. and García-Franco, JG., 2008. Habitat isolation changes the beta diversity of vascular epiphyte community in lower montane forest, Veracruz, Mexico. Biodiversity and Conservation, vol. 17, no. 1, p. 191-207. http://dx.doi.org/10.1007/s10531-007-9239-6.
    » http://dx.doi.org/10.1007/s10531-007-9239-6
  • Fraga, LL., Silva, LB. and Schmitt, JL., 2008. Composição e distribuição vertical de pteridófitas epifíticas sobre Hook. (Dicksoniaceae), em Floresta Ombrófila Mista no sul do Brasil. Dicksonia sellowianaBiota Neotropica, vol. 8, no. 4, p. 123-129. http://dx.doi.org/10.1590/S1676-06032008000400011.
    » http://dx.doi.org/10.1590/S1676-06032008000400011
  • Fundação SOS Mata Atlântica and Instituto Nacional de Pesquisas Espaciais –INPE, 2011. Atlas dos remanescentes florestais da Mata Atlântica: período 2008-2010. São Paulo: INPE. 122 p.
  • GENTRY, AH. and DODSON, CH., 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden, vol. 74, no. 2, p. 205-233. http://dx.doi.org/10.2307/2399395.
    » http://dx.doi.org/10.2307/2399395
  • Geraldino, HCL., Caxambú, MG. and SOUZA, DC., 2010. Composição florística e estrutura da comunidade de epífitas vasculares em uma área de ecótono em Campo Mourão, PR, Brasil. Acta Botanica Brasilica, vol. 24, no. 2, p. 469-482. http://dx.doi.org/10.1590/S0102-33062010000200018.
    » http://dx.doi.org/10.1590/S0102-33062010000200018
  • Giongo, C. and Waechter, JL., 2004. Composição florística e estrutura comunitária de epífitos vasculares em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 27, no. 3, p. 563-572. http://dx.doi.org/10.1590/S0100-84042004000300015.
    » http://dx.doi.org/10.1590/S0100-84042004000300015
  • Gonçalves, CN. and Waechter, JL., 2002. Epífitos vasculares sobre espécimes de .Ficus organensis isolados no norte da planície costeira do Rio Grande do Sul: padrões de abundância e distribuiçãoActa Botanica Brasilica, vol. 16, no. 4, p. 429-441. http://dx.doi.org/10.1590/S0102-33062002000400006.
    » http://dx.doi.org/10.1590/S0102-33062002000400006
  • Gonçalves, CN. and Waechter, JL., 2003. Aspectos florísticos e ecológicos de epífitos vasculares sobre figueiras isoladas no norte da planície costeira do Rio Grande do Sul. Acta Botanica Brasilica, vol. 17, no. 1, p. 89-100. http://dx.doi.org/10.1590/S0102-33062003000100007.
    » http://dx.doi.org/10.1590/S0102-33062003000100007
  • Gotelli, NJ. and Colwell, RK., 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness.Ecology Letters, vol. 4, no. 4, p. 379-391. http://dx.doi.org/10.1046/j.1461-0248.2001.00230.x.
    » http://dx.doi.org/10.1046/j.1461-0248.2001.00230.x
  • Guedes-Bruni, RR., Morim, MPM., LIMA, HC. and SYLVESTRE, L., 2002. Inventário florístico. In SYLVESTRE, LS. and ROSA, MMT. (Eds.). Manual metodológico para estudos botânicos na Mata Atlântica.Rio de Janeiro: Universidade Rural. p. 24-49.
  • Hefler, SM. and Faustioni, P., 2004. Levantamento florístico de epífitos vasculares do bosque São Cristovão –Curitiba – Paraná – Brasil. Revista Estudos de Biologia, vol. 26, no. 54, p. 11-19.
  • Kersten, RA., 2010. Epífitas vasculares – Histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea, vol. 37, no. 1, p. 9-38. http://dx.doi.org/10.1590/S2236-89062010000100001.
    » http://dx.doi.org/10.1590/S2236-89062010000100001
  • Kersten, RA. and Kunyioshi, YS., 2006. Epífitos vasculares na Bacia do Alto Iguaçu, Paraná, Brasil – composição florística. Estudos de Biologia, vol. 28, p. 55-71.
  • Kersten, RA. and Kuniyoshi, YS., 2009. Conservação das florestas na Bacia do alto Iguaçú, Paraná – Avaliação da comunidade de epífitas vasculares em diferentes estágios serais. Revista Floresta, vol. 39, no. 1, p. 51-66.
  • Kersten, RA., Kuniyidhi, YS. and Roderjan, CV., 2009. Epífitas vasculares em duas formações ribeirinhas adjacentes na bacia do rio Iguaçu – Terceiro Planalto Paranaense. Iheringia Série Botânica, vol. 64, no. 1, p. 33-43.
  • Kersten, RA. and Silva, SM., 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 24, no. 2, p. 213-226. http://dx.doi.org/10.1590/S0100-84042001000200012.
    » http://dx.doi.org/10.1590/S0100-84042001000200012
  • Kersten, RA. and Silva, SM., 2002. Florística e estrutura do componente epifítico vascular em Floresta Ombrófila Mista aluvial do Rio Barigüi, Paraná, Brasil. Revista Brasileira de Botânica, São Paulo, vol. 25, no. 3, p. 259-267.
  • Kersten, RA. and Silva, SM., 2005. Florística e estrutura de comunidades de epífitas vasculares da planície litorânea. In MARQUES, MCM. and BRITEZ, RM. (Eds.). História natural e conservação da Ilha do Mel. Curitiba: Universidade Federal do Paraná. p. 125-144.
  • Klein, RM., 1975. Southern Brazilian phytogeographic features and the probable influence of upper quaternary climatic changes in the floristic distribution. Boletim Paranaense de Geociências, vol. 33, p. 67-88.
  • Kress, WJ., 1986. The systematic distribution of vascular epiphytes: an update. Selbyana, vol. 9, no. 1, p. 2-22.
  • Krömer, T., Kessler, M. and Gradstein, SR., 2007. Vertical stratification of vascular ephytes in submontane and montane Forest of the Bolivian Andes: the importance of the understory. Plant Ecology, vol. 189, no. 2, p. 261-278. http://dx.doi.org/10.1007/s11258-006-9182-8.
    » http://dx.doi.org/10.1007/s11258-006-9182-8
  • Lugo, AE. and Scatena, FN., 1992. Epiphytes and climate change research in the Caribbean: a proposal. Selbyana, vol. 13, p. 123-130.
  • Lüttge, U., 1989. Vascular epiphytes: setting the Scene. In LÜTTGE, U. (Ed.). Vascular plants as epiphytes. New York: Springer-Verlag. p. 1-12. Ecological Studies,79:
  • Madison, M., 1977. Vascular epiphytes: their systematic occurrence and salient features. Selbyana, vol. 2, no. 1, p. 1-13.
  • Magurran, AE., 1988. Ecological diversity and its measurement. Cambridge: British Library.
  • Mania, LF. and Monteiro, R., 2010. Florística e ecologia de epífitas vasculares em um fragmento de floresta de restinga, Ubatuba, SP, Brasil. Rodriguésia, vol. 61, no. 4, p. 705-713.
  • MeniniNeto, L., Forzza, RC. and Zappi, D., 2009. Angiosperm epiphytes as conservation indicators in forest fragments: A case study from southeastern Minas Gerais, Brasil. Biodiversity and Conservation, vol. 18, no. 14, p. 3785-3807. http://dx.doi.org/10.1007/s10531-009-9679-2.
    » http://dx.doi.org/10.1007/s10531-009-9679-2
  • Moreno, JA., 1961. Clima do Rio Grande do Sul.Porto Alegre: Secretaria da Agricultura. 42 p. Secção de Geografia.
  • Müller, L., Starnecker, G. and Winkler, S., 1981. Zur Ökologie epiphytischer Farne in Südbrasilien. I. Saugschuppen. Flora, vol. 171, p. 55-65.
  • Myers, N., Mittermeier, RA., Mittermeier, CG., Fonseca, GA. and Kent, J., 2000. Biodiversity hotspots for conservation priorities. Nature, vol. 403, no. 6772, p. 853-858. http://dx.doi.org/10.1038/35002501. PMid:10706275.
    » http://dx.doi.org/10.1038/35002501
  • Nadkarni, NM., 1984. Epiphyte biomass and nutrient capital of a neotropical elfin forest. Biotropica, vol. 16, no. 4, p. 249-256. http://dx.doi.org/10.2307/2387932.
    » http://dx.doi.org/10.2307/2387932
  • Nieder, J., Engwald, S. and Barthlott, W., 1999. Patterns of neotropical epiphyte diversity. Selbyana, vol. 20, no. 1, p. 66-75.
  • Nieder, J., Ibisch, PL. and Barthlott, W., 1997. Biodiversidad de epífitas: una cuestión de escala. Revista del Jardín Botánico Nacional, vol. 17/18, p. 59-62.
  • Parker, GG., 1995. Structure and microclimate of forest canopies. In LOWMAN, MD. and NADKARNI, NM. (Eds.). Forest canopies. San Diego: Academic Press. p. 73-106.
  • Rambo, B., 1954. História da flora do litoral rio-grandense. Sellowia, vol. 6, no. 6, p. 113-172.
  • Rocha, CFD., Cogliatti-Carvalho, L., Nunes-Freitas, AF., Rocha-Pessoa, TC., Dias, AS., Ariani, CV. and Morgado, LN., 2004. Conservando uma larga proporção da diversidade biológica através da conservação de Bromeliaceae.Vidalia, vol. 2, no. 1, p. 52-68.
  • Rocha, LD., Droste, A., Gehlen, G. and Schmitt, JL., 2013. Leaf dimorphism of (Polypodiaceae): a qualitative and quantitative analysis focusing on adaptations to epiphytism. Microgramma squamulosaRevista de Biología Tropical, vol. 61, no. 1, p. 291-299. PMid:23894982.
  • Roderjan, CV., Galvão, F., Kuniyoshi, YS. and Hatschbach, GG., 2002. As unidades fitogeográficas do estado do Paraná. Ciência e Ambiente, vol. 24, no. 1, p. 75-92.
  • Rogalski, JM. and Zanin, EM., 2003. Composição florística de epífitos vasculares no estreito de Augusto César, Floresta Estacional Decidual do Rio Uruguai, RS, Brasil. Revista Brasileira de Botânica. Brazilian Journal of Botany, vol. 26, no. 4, p. 551-556. http://dx.doi.org/10.1590/S0100-84042003000400014.
    » http://dx.doi.org/10.1590/S0100-84042003000400014
  • Rosa, NA., 2010. Plano de Manejo – Parque Municipal Henrique Luís Roessler – Parcão. Novo Hamburgo: Prefeitura Municipal de Novo Hamburgo. 846 p.
  • Santos, AJ., 2004. Estimativas de riqueza em espécies. In CULLEN-JÚNIOR, L., VALLADARES-PADUA, C. and RUDY, R. (Eds.). Métodos de estudos em biologia da conservação e manejo da vida silvestre. Curitiba: UFPR. p. 19-41.
  • Schmitt, JL. and Windisch, PG., 2005. Aspectos ecológicos de . Alsophila setosa Kaulf. (Cyatheaceae, Pteridophyta) no Rio Grande do Sul, BrasilActa Botanica Brasilica, vol. 19, no. 4, p. 861-867. http://dx.doi.org/10.1590/S0102-33062005000400021.
    » http://dx.doi.org/10.1590/S0102-33062005000400021
  • Schmitt, JL. and Windisch, PG., 2010. Biodiversity and spatial distribution of epiphytic ferns on . Alsophila setosa Kaulf. (Cyatheaceae) caudices in Rio Grande do Sul, BrazilBrazilian Journal of Biology = Revista Brasleira de Biologia, vol. 70, no. 3, p. 521-528. http://dx.doi.org/10.1590/S1519-69842010000300008. PMid:20730338.
    » http://dx.doi.org/10.1590/S1519-69842010000300008
  • Schmitt, JL., Budke, JC. and Windisch, PG., 2005. Aspecto florísticos e ecológicos de pteridófitas epifíticas em cáudices de Hook. (Pteridophyta, Dicksoniaceae), São Francisco de Paula, RS, Brasil. Dicksonia sellowianaPesquisas, vol. 56, p. 161-172.
  • Schneider, PH. and Schmitt, JL., 2011. Composition, community structure and vertical distribution of epiphytic ferns on . Alsophila setosa Kaulf., in a Semideciduous Seasonal Forest, Morro Reuter, RS, BrasilActa Botanica Brasilica, vol. 25, no. 3, p. 557-562. http://dx.doi.org/10.1590/S0102-33062011000300007.
    » http://dx.doi.org/10.1590/S0102-33062011000300007
  • Sehnem, A.,1970. Polipodiáceas. In REITZ, R. (Ed.). Flora ilustrada catarinense. Itajaí: Herbário Barbosa Rodrigues. p. 1-173.
  • Sehnem, A., 1977. As filicíneas do Sul do Brasil, sua distribuição geográfica, sua ecologia e suas rotas de migração. Pesquisas, vol. 31, p. 1-108.
  • Sehnem, A., 1979. Semelhanças e diferenças nas formações florestais do sul do Brasil. Acta Biologica Leopoldensia, vol. 1, no. 1, p. 111-135.
  • Smith, AR., Kathleen, MP., Schuettpelz, E., Korall, P., Schneider, H. and Wolf, PG., 2006. A classification for extant ferns. Taxon, vol. 55, no. 3, p. 705-731. http://dx.doi.org/10.2307/25065646.
    » http://dx.doi.org/10.2307/25065646
  • Smith, AR., Pryer, KM., Schuettpelz, E., Korall, P., Schneider, H. and Wolf, PG., 2008. Fern Classification. In RANKER, TA. and HAUFLER, CH. The Biology and Evolution of Ferns and Lycophytes. Cambridge: Cambridge University Press. p. 417-467.
  • Staudt, MG., Lippert, APU., Cunha, S., Becker, DFP., Marchioretto, MS. and Schmitt, JL., 2012. Composição florística de epífitos vasculares do Parque Natural Municipal Tupancy, Arroio do Sal, RS – Brasil. Pesquisas, vol. 63, p. 177-188.
  • Teixeira, MB., Coura-Neto, AB., Pastore, U. and Rangel-Filho, ALR., 1986. Vegetação: as regiões fitoecológicas, sua natureza e seus recursos econômicos – estudo fitogeográfico. In Levantamento de Recursos Naturais.Rio de Janeiro, IBGE, vol. 33, p. 541-620.
  • TER Steege, H. and Cornelissen, JHC., 1989. Distribution and ecology of vascular epiphytes in Lowland rain forest of Guiana. Biotropica, vol. 21, no. 4, p. 331-339. http://dx.doi.org/10.2307/2388283.
    » http://dx.doi.org/10.2307/2388283
  • The Angiosperm Phylogeny Group – APG, 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, vol. 161, no. 2, p. 105-121. http://dx.doi.org/10.1111/j.1095-8339.2009.00996.x.
    » http://dx.doi.org/10.1111/j.1095-8339.2009.00996.x
  • Waechter, JL., 1986. Epífitos vasculares da mata paludosa do Faxinal, Torres, Rio Grande do Sul, Brasil. Iheringia, vol. 34, p. 39-49.
  • Waechter, JL., 1992. O epifistismo vascular na Planície Costeira do Rio Grande do Sul.São Carlos: Universidade Federal de São Carlos. 163 p. Tese de Doutorado em Ecologia e Recursos Naturais.
  • Waechter, JL., 1998. O epifitismo vascular em uma floresta de restinga do Brasil subtropical. Revista Ciência e Natura, vol. 20, no. 4, p. 43-66.
  • Waechter, JL., 2006. Diversidade epifítica ao longo de gradientes ambientais. In MARIATH, JEA. and SANTOS, RP. (Eds.). Os Avanços da Botânica no Início do Século XXI: Conferências, Plenárias e Simpósios do 57º Congresso Nacional de Botânica. Porto Alegre, Sociedade Botânica do Brasil. p. 350-353.
  • Yeaton, RI. and Gladstone, DE., 1982. The pattern of colonization of epiphytes on Calabash Trees (. Crescentia alata HBK.) in Guanacaste Province, Costa RicaBiotropica, vol. 14, no. 2, p. 137-140. http://dx.doi.org/10.2307/2387743.
    » http://dx.doi.org/10.2307/2387743

Publication Dates

  • Publication in this collection
    May 2015

History

  • Received
    24 May 2013
  • Accepted
    26 Aug 2013
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