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Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil

Angiospermas epífitas de uma floresta montana no sul da Bahia, Brasil

Abstracts

The Atlantic Forest in southern Bahia State, Brazil, has high levels of diversity and vascular plant endemism. There have been few floristic surveys of the mountain forests there, however, especially those focusing on herbs and canopy plants, with studies of Atlantic Forest epiphytes having been limited to the southern and southeastern region of that country. The present study therefore surveyed the epiphytic angiosperms in the Serra das Lontras National Park (SLNP), the distribution of their species among genera and families, and their biogeographical patterns. Fieldwork was performed between February 2011 and March 2012. Collections were made along roads and trails and phorophytes were occasionally climbed and fallen trees and branches examined in order to sample canopy species. The study site demonstrated high epiphyte richness (256 spp.), one of the highest recorded in the Atlantic Forest. The richest families are Orchidaceae, Bromeliaceae, and Araceae, reaffirming the patterns reported in previous major reviews on epiphytes. The species exhibit high degrees of endemism (45%) to the Atlantic Forest, especially among the Bromeliaceae and Orchidaceae. One new occurrence to Brazil, four to the Atlantic Forest and 30 to Bahia state are registered. The SLNP shares more species with areas of southern and southeastern Brazil than with other northeastern states.

Atlantic forest; Araceae; Bromeliaceae; Orchidaceae; biogeographical patterns


A Floresta Atlântica do sul da Bahia apresenta elevados níveis de diversidade e endemismo de plantas vasculares. No entanto, poucos levantamentos florísticos foram realizados em áreas de floresta montana, especialmente para ervas e plantas de dossel. Estudos com epífitas estão concentrados principalmente em áreas do Sul e Sudeste do Brasil. Desta forma, o presente estudo realizou o levantamento das angiospermas epífitas do Parque Nacional da Serra das Lontras (PNSL) e verificou a distribuição das espécies pelas famílias e gêneros e seus padrões biogeográficos. As coletas foram realizadas entre fevereiro de 2011 e março de 2012 nas trilhas e estradas de acesso. Alguns forófitos foram escalados e árvores e ramos caídos foram examinados com o intuito de coletar espécies restritas ao dossel. A área de estudo apresentou alta diversidade de epífitas (256 spp.), uma das maiores já registradas para a Floresta Atlântica. As famílias com maior riqueza são Orchidaceae, Bromeliaceae e Araceae, reafirmando os padrões encontrados anteriormente em revisões do tema. As espécies apresentam alto grau de endemismo (45%) à Floresta Atlântica, especialmente em Bromeliaceae e Orchidaceae. São registradas uma nova ocorrência para o Brasil, quatro para a Floresta Atlântica e 30 para a Bahia. O PNSL possui mais espécies em comum com o Sul e o Sudeste do que com a região Nordeste.

Floresta Atlântica; Araceae; Bromeliaceae; Orchidaceae; padrões biogeográficos


Introduction

Epiphytes spend most or all of their life cycles growing on other plants (phorophytes), taking advantage of their mechanical support (Benzing 1990Benzing, D.H. 1990. Vascular epiphytes. Cambridge University Press, New York.). Epiphytes account for approximately 10% of world's vascular flora and are found in 84 angiosperm families (Gentry & Dodson 1987Gentry, A.H. & Dodson, C.H. 1987. Diversity and biogeography of neotropical vascular epiphytes. Ann. Missouri Bot. Gard. 74: 205-233. doi: 10.2307/2399395.
https://doi.org/10.2307/2399395...
, Kersten 2010Kersten, R.A. 2010. Epífitas vasculares: histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea 37(1): 9-38. doi: 10.1590/S2236-89062010000100001.
https://doi.org/10.1590/S2236-8906201000...
). Araceae, Bromeliaceae, and Orchidaceae are the most successful epiphytic angiosperm families, and the latter is responsible for over 70% of total richness.

Bromeliaceae is a typically American family, while the largest genera of Araceae and Orchidaceae (e.g., Anthurium, Epidendrum, Lepanthes, Philodendron and Pleurothallis s.l.) are likewise endemic to that continent. The Neotropical region therefore hosts the greater part of world's epiphytic flora (> 15.500 spp.), which are mainly concentrated in Central America, the Andes, northwestern Amazonia and the Atlantic Forest (Gentry & Dodson 1987Gentry, A.H. & Dodson, C.H. 1987. Diversity and biogeography of neotropical vascular epiphytes. Ann. Missouri Bot. Gard. 74: 205-233. doi: 10.2307/2399395.
https://doi.org/10.2307/2399395...
, Nieder et al. 2001Nieder, J., Prosperí, J. & Michaloud, G. 2001. Epiphytes and their contribution to canopy diversity. Plant ecol. 153: 51-63. doi: 10.1023/A:1017517119305.
https://doi.org/10.1023/A:1017517119305...
). Of the 14.500 vascular plants known to inhabit the Atlantic Forest, 3.000-4.000 are epiphytes (Stehmann et al. 2009Stehmann, J.R., Forzza, R.C., Sobral, M. & Kamino, L.H.Y. 2009. Gimnospermas e Angiospermas. In Plantas da Floresta Atlântica (J.R. Stehmann, R.C. Forzza, A. Salino, M. Sobral, D.P. Costa & L.H.Y. Kamino, eds.). Jardim Botânico do Rio de Janeiro, Rio de Janeiro. p. 27-37., Kersten 2010Kersten, R.A. 2010. Epífitas vasculares: histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea 37(1): 9-38. doi: 10.1590/S2236-89062010000100001.
https://doi.org/10.1590/S2236-8906201000...
). The Atlantic Forest has large latitudinal (from 3°S to 30°S) and altitudinal (from sea level up to 2.890 m) extensions and significant rainfall variations - resulting in high habitat heterogeneity and an extraordinary diversity of organisms (Oliveira-Filho & Fontes 2000Oliveira-Filho, A.T. & Fontes, M.A.L. 2000. Patterns of floristic differentiation among Atlantic forests in southeastern Brazil and the influence of climate. Biotropica 32(4b): 793-810. doi: 10.1111/j.1744-7429.2000.tb00619.x.
https://doi.org/10.1111/j.1744-7429.2000...
).

Southern Bahia is considered an area of great biological importance, with elevated levels of richness and endemism in its arboreal component (Mori et al. 1983Mori, S., Boom, B.M., Carvalho, A.M. & Santos, T.S. 1983. Southern Bahian moist forest. Bot. rev. 49(2): 155-232. doi: 10.1007/BF02861011.
https://doi.org/10.1007/BF02861011...
, Thomas et al. 1998Thomas, W.W., Carvalho, A.M.V., Amorim, A.M., Garrison, J. & Arbeláez, A.L. 1998. Plant endemism in two forests in southern Bahia, Brazil. Biodivers. Conserv. 7(3): 311-322. doi: 10.1023/A:1008825627656.
https://doi.org/10.1023/A:1008825627656...
, Martini et al. 2007Martini, A.M.Z., Fiaschi, P., Amorim, A.M. & Paixão, J.L. 2007. A hot-point within a hot-spot: a high diversity site in Brazil's Atlantic Forest. Biodivers. Conserv. 16(11): 3111-3128. doi: 10.1007/s10531-007-9166-6.
https://doi.org/10.1007/s10531-007-9166-...
, Thomas et al. 2009Thomas, W.W., Jardim, J.G., Fiaschi, P., Mariano-Neto, E. & Amorim, A.M. 2009. Composição florística e estrutura do componente arbóreo de uma área transicional de Floresta Atlântica no sul da Bahia, Brasil. Rev. Bras. Bot. 32(1): 65-78. doi: 10.1590/S0100-84042009000100007.
https://doi.org/10.1590/S0100-8404200900...
, Rocha & Amorim 2012Rocha, D.S.B. & Amorim, A.M.A. 2012. Heterogeneidade altitudinal na Floresta Atlântica setentrional: um estudo de caso no sul da Bahia, Brasil. Acta bot. bras. 26: 309-327.), although surveys of humid montane forests have been scarce, especially those focusing on terrestrial and epiphytic herbs (Amorim et al. 2009Amorim, A.M.A., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O. & Thomas, W.W. 2009. Angiospermas em remanescentes de Floresta Montana no sul da Bahia, Brasil. Biota Neotrop. 9(3): 313-348 http://www.biotaneotropica.org.br/v9n3/en/abstract?inventory+bn02909032009 (accessed in 15/07/2013). doi: 10.1590/S1676-06032009000300028
http://www.biotaneotropica.org.br/v9n3/e...
, Matos et al. 2010Matos, F.B., Amorim, A.M. & Labiak, P.H. 2010. The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. J. Bot. Res. Inst. Texas 4(1): 333-346.). The recently created Serra das Lontras National Park comprises a vast remnant of montane Atlantic Forest in southern Bahia (Save Brasil et al. 2009SAVE Brasil, IESB & BirdLife International 2009. Complexo Serra das Lontras e Una, Bahia: Elementos naturais e aspectos de sua conservação. SAVE Brasil, São Paulo.).

A preliminary survey in the park, covering only a small area, recorded 709 angiosperm species, of which 158 (22.3%) were epiphytes (Amorim et al. 2009Amorim, A.M.A., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O. & Thomas, W.W. 2009. Angiospermas em remanescentes de Floresta Montana no sul da Bahia, Brasil. Biota Neotrop. 9(3): 313-348 http://www.biotaneotropica.org.br/v9n3/en/abstract?inventory+bn02909032009 (accessed in 15/07/2013). doi: 10.1590/S1676-06032009000300028
http://www.biotaneotropica.org.br/v9n3/e...
). These authors highlighted the fact that many of the species encountered, including many epiphytes (e.g., Fuchsia regia [Vell.] Munz, Nematanthus lanceolatus [Poir.] Chautems, and Vriesea longicaulis Mez), were previously known only from mountainous areas in southern and southeastern Brazil.

For the preliminary survey there was no effort focused on this habit, which presents peculiarities that hinder their sampling, including the height they occur on the phorophyte and the small sizes of some groups (e.g., Peperomia and Pleurothallis s.l.). We therefore sought to inventory the composition of epiphytic angiosperms in the Serra das Lontras National Park to investigate their distributions into genera and families and their biogeographical patterns to increase our knowledge of this group in the park and in northeastern Brazil.

Material and methods

Serra das Lontras National Park (SLNP) is located in southern Bahia State (Figure 1), Brazil, in the municipalities of Arataca, São José da Vitória, and Una (15°07′-15°15′S and 39°15′-39°25′W). The park comprises approximately 11.000 hectares at altitudes that vary from 400 to 1.000 m. Located 30 km from the coast, the mountains act as a first barrier to humid air masses moving inland from the ocean, resulting in mean annual precipitation rates of 1.300-1.600 mm, without a marked dry season, and a mean annual temperature of 24°C (classified as Af in the Koppen system) (Peel et al. 2007Peel, M.C., Finlayson, B.L. & McMahon, T.A. 2007. Updated world map of the Köppen-Geiger climate classification. Hydrol. Earth Syst. Sc. 11: 1633-1644. doi: 10.5194/hess-11-1633-2007.
https://doi.org/10.5194/hess-11-1633-200...
, Amorim et al. 2009Amorim, A.M.A., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O. & Thomas, W.W. 2009. Angiospermas em remanescentes de Floresta Montana no sul da Bahia, Brasil. Biota Neotrop. 9(3): 313-348 http://www.biotaneotropica.org.br/v9n3/en/abstract?inventory+bn02909032009 (accessed in 15/07/2013). doi: 10.1590/S1676-06032009000300028
http://www.biotaneotropica.org.br/v9n3/e...
, Nacif et al. 2009Nacif, P.G.S., Costa, O.V., Araújo, M. & Santos, P.S. 2009. Geomorfodinâmica da Região do Complexo de Serras das Lontras. In Complexo de Serras das Lontras e Una, Bahia: Elementos naturais e aspectos de sua conservação (Save Brasil, IESB e Birdlife International, eds.). SAVE Brasil, São Paulo. p. 9-14.).

Figure 1
Map showing the location of the studied area (Serra das Lontras National Park); in gray is the Atlantic Forest limits. N.P. - National Park; B.R. - Biological Reserve/Mapa de localização da área de estudo (Parque Nacional da Serra das Lontras); em cinza está demarcado o limite da Floresta Atlântica.

Situated in the Atlantic Forest domain, the SLNP comprises a vegetation mosaic of anthropogenic areas mixed with well-preserved submontane forests (with canopies >30 m) at lower altitudes and cloud forests above 800 m with smaller trees with twisted trunks bearing bryophytes (Stadtmüller 1987Stadtmüller, T. 1987. Cloud Forests in the Humid Tropics: A bibliographic review. http://archive.unu.edu/unupress/unupbooks/80670e/80670E00.htm#Contents (accessed in 08/2012).
http://archive.unu.edu/unupress/unupbook...
).

A total floristic inventory of the SLNP has been underway for the last eight years, and preliminary results have already been published (Amorim et al. 2009Amorim, A.M.A., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O. & Thomas, W.W. 2009. Angiospermas em remanescentes de Floresta Montana no sul da Bahia, Brasil. Biota Neotrop. 9(3): 313-348 http://www.biotaneotropica.org.br/v9n3/en/abstract?inventory+bn02909032009 (accessed in 15/07/2013). doi: 10.1590/S1676-06032009000300028
http://www.biotaneotropica.org.br/v9n3/e...
; Save Brasil et al. 2009SAVE Brasil, IESB & BirdLife International 2009. Complexo Serra das Lontras e Una, Bahia: Elementos naturais e aspectos de sua conservação. SAVE Brasil, São Paulo.). Visits to the area were made every two months, from February/2011 to March/2012 to survey the epiphytic angiosperms. Collections were made along roads and trails in order to cover the largest possible areas, and some living phorophytes were occasionally climbed and fallen trees examined to collect canopy species. Accidental epiphytes were not considered in this survey. Informations on the type of epiphytism (characteristic holoepiphyte, facultative holoepiphyte, hemiepiphyte) were obtained from local observations and complemented with informations from literature. This was done in order to have more accurate data, since many facultative species might have been seen only as epiphytes in the area or might occur as terrestrial or rupiculous in parts of the park that were not visited. All fertile material was deposited at the RB and CEPEC herbaria, while sterile specimens were incorporated into the ex-situ collection at the Rio de Janeiro Botanical Garden.

Specimens collected during the general floristic inventory, as well as those from the present survey, were identified to the species level, when possible, with the help of specialists and specific literature (e.g., the Flora Neotropica). Nomenclature and species concepts follow the List of Species of the Brazilian Flora - LSBF (2013). Distribution and endemism data were obtained from the LSBF for taxa identified to the species level.

A Rapid Color Guide, published by The Field Museum, was prepared and images on most species can be seen at www.fm2.fieldmuseum.org/plantguides/guideimages.asp?ID=545.

Results

The Serra das Lontras National Park shows high epiphyte richness, with a total of 256 species identified, representing 87 genera and 18 families (Table 1) - making it one of the richest sites for epiphytic angiosperms in the Atlantic Forest. Of the published checklists, only Macaé de Cima in Rio de Janeiro State reported more epiphyte species (275 spp.) (Fontoura et al. 1997Fontoura, T., Sylvestre, L.S., Vaz, A.M.S.F. & Vieira, C.M. 1997. Epífitas vasculares, hemiepífitas e hemiparasitas da Reserva Ecológica de Macaé de Cima. In Serra de Macaé de Cima: Diversidade florística e conservação em Mata Atlântica (H.C. Lima & R.R. Guedes-Bruni, eds.). Jardim Botânico do Rio de Janeiro, Rio de Janeiro, p. 89-101.).

Table 1
Epiphytic Angiosperms registered for Serra das Lontras National Park, Bahia, Brazil. 1 *New occurence to Bahia state; ** New occurence to the Atlantic Forest; *** New occurence to Brazil. 2 AA - André M. Amorim; AF - André P. Fontana; AJ - Alessandra B. Jardim; JJ - Jomar G. Jardim; JP - José Lima da Paixão; ML - Márdel M. Lopes; PF - Pedro Fiaschi; PL - Paula Leitman; RB - Rafael X. Borges; RP - Ricardo O. Perdiz; WT - Wm. Wayt Thomas. 3 Epi - Characteristic Holoepiphyte; Fac - Facultative Holoepiphyte; Hep - Hemiepiphyte. 4 CAF - Cerrado-Atlantic Forest; Di - Disjunct distribution Amazon-Atlantic Forest; En - Endemic to Atlantic Forest; WD - Wide distribution; O - Other; “-” - not applicable. 5 CR - Criticaly endangered; DD - Data deficient; EN - Endangered; VU - Vulnerable.

The most important families encountered were Orchidaceae (106 spp.), Bromeliaceae (64 spp.), Araceae (31 spp.), and Piperaceae (14 spp.). Together, these families encompassed more than 80% of the epiphyte flora at SLNP. The species distributions within the families were very unequal, however, with 11 families being represented by less than five species.

The richest genera were Vriesea (22 spp.), Peperomia and Stelis (14 spp. each), Anthurium (13 spp.), Aechmea and Philodendron (12 spp. each), Epidendrum (11 spp.), and Hohenbergia (10 spp.). The species distributions within the genera were very unequal, with half of them (43 genera) being represented by only a single species, and 82% by less than five species.

With many ornamental species, 10 epiphytes registered for the SLNP are cited on the Red Book of Brazilian Flora (Martinelli & Moraes 2013Martinelli, G. & Moraes, M.A. (orgs.) 2013. Livro vermelho da flora do Brasil. Andrea Jakobsson/Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1100 p.). Of this total, three are “Critically endangered”, five are “Endangered”, and two are “Vulnerable”. Four are considered to be “Data Deficient” (Table 1).

Discussion

The high numbers of species observed in Araceae, Bromeliaceae, Orchidaceae, and Piperaceae were consonant with global patterns of epiphytic distribution among angiosperm families, as reported by Madison (1977)Madison, M. 1977. Vascular epiphytes: their systematic occurrence and salient features. Selbyana 2: 1-13. and Gentry & Dodson (1987)Gentry, A.H. & Dodson, C.H. 1987. Diversity and biogeography of neotropical vascular epiphytes. Ann. Missouri Bot. Gard. 74: 205-233. doi: 10.2307/2399395.
https://doi.org/10.2307/2399395...
. Similar results were reported for the Neotropical region (e.g., Sudgen & Robins 1979Sudgen, A. & Robins, R. 1979. Aspects of the Ecology of Vascular Epiphytes in Colombian Cloud Forests, I. The Distribution of the epiphytic flora. Biotropica 11(3): 173-188. doi: 10.2307/2388037.
https://doi.org/10.2307/2388037...
, Gentry & Dodson 1987Gentry, A.H. & Dodson, C.H. 1987. Diversity and biogeography of neotropical vascular epiphytes. Ann. Missouri Bot. Gard. 74: 205-233. doi: 10.2307/2399395.
https://doi.org/10.2307/2399395...
, Wolf & Flamenco-S 2003Wolf, J.H.D. & Flamenco-S, A. 2003. Patterns in species richness and distribution of vascular epiphytes in Chiapas, Mexico. J. biogeogr. 30: 1689-1707. doi: 10.1046/j.1365-2699.2003.00902.x.
https://doi.org/10.1046/j.1365-2699.2003...
, Küper et al. 2004Küper, W., Kreft, H., Nieder, J., Köster, N. & Barthlott, W. 2004. Large-scale diversity patterns of vascular epiphytes in Neotropical montane rain forests. J. biogeogr. 31: 1477-1487. doi: 10.1111/j.1365-2699.2004.01093.x.
https://doi.org/10.1111/j.1365-2699.2004...
) and the tropical and subtropical Atlantic Forest (e.g., Fontoura et al. 1997Fontoura, T., Sylvestre, L.S., Vaz, A.M.S.F. & Vieira, C.M. 1997. Epífitas vasculares, hemiepífitas e hemiparasitas da Reserva Ecológica de Macaé de Cima. In Serra de Macaé de Cima: Diversidade florística e conservação em Mata Atlântica (H.C. Lima & R.R. Guedes-Bruni, eds.). Jardim Botânico do Rio de Janeiro, Rio de Janeiro, p. 89-101., Borgo & Silva 2003Borgo, M. & Silva, S.M. 2003. Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Rev. Bras. Bot. 26(3): 391-401. doi: 10.1590/S0100-84042003000300012.
https://doi.org/10.1590/S0100-8404200300...
, Giongo & Waechter 2004Giongo, C. & Waechter, J.L. 2004. Composição florística e estrutura comunitária de epífitos vasculares em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Rev. Bras. Bot. 27(3): 563-572. doi: 10.1590/S0100-84042004000300015.
https://doi.org/10.1590/S0100-8404200400...
, Kersten & Silva 2001Kersten, R.A. & Silva, S.M. 2001. Composição florística e estrutura do componente epifítico vascular em floresta da planície litorânea na Ilha do Mel, Paraná, Brasil. Rev. Bras. Bot. 24(2): 213-226. doi: 10.1590/S0100-84042001000200012.
https://doi.org/10.1590/S0100-8404200100...
, Buzatto et al. 2008Buzatto, C.R., Severo, B.M.A. & Waechter, J.L. 2008. Composição florística e distribuição ecológica de epífitos vasculares na Floresta Nacional de Passo Fundo, Rio Grande do Sul. Iheringia, Bot. 63(2): 231-239., Martins et al. 2008Martins, S.E., Rossi, L., Sampaio, P.S.P. & Magenta, M.A.G. 2008. Caracterização florística de comunidades vegetais de restinga em Bertioga, SP, Brasil. Acta bot. bras. 22(1): 249-274. doi: 10.1590/S0102-33062008000100024.
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, Menini Neto et al. 2009Menini Neto, L., Forzza, R.C., & Zappi, D. 2009. Angiosperm epiphytes as conservation indicators in forest fragments: a case study from southeastern Minas Gerais, Brazil. Biodivers. Conserv. 18: 3785-3807. doi: 10.1007/s10531-009-9679-2.
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, Mania & Monteiro 2010Mania, L. & Monteiro, R. 2010. Florística e ecologia de epífitas vasculares em um fragmento de floresta de restinga, Ubatuba, SP, Brasil. Rodriguésia 61(4): 705-713.).

Table 2 compares the SLNP and other Atlantic Forest areas with high epiphyte richness. The elevated numbers of species of Bromeliaceae, Araceae, and Gesneriaceae is notable, representing the highest epiphyte richness ever recorded for these families in the Atlantic Forest. Vriesea, Stelis, Anthurium, Aechmea, and Hohenbergia also exhibited their greatest epiphytic species diversity in the SLNP. It is interesting that Hohenbergia was among the richest genera in the study area, as this genus does not commonly figure among the most species rich taxa in epiphyte surveys. The greatest diversity of this genus is encountered in southern Bahia, where 21 of its 25 Atlantic Forest species occur (Martinelli et al. 2008Martinelli, G., Vieira, C.M., Gonzalez, M., Leitman, P., Piratininga, A., Costa, A.F. & Forzza, R.C. 2008. Bromeliaceae da Mata Atlântica Brasileira: lista de espécies, distribuição e conservação. Rodriguésia 59(1): 209-258., Forzza et al. 2013Forzza, R.C., Costa, A., Siqueira Filho, J.A., Martinelli, G., Monteiro, R.F., Santos-Silva, F., Saraiva, D. P. & Paixão-Souza, B. 2013. Bromeliaceae. In Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB66 (accessed in 15/05/2013).
http://floradobrasil.jbrj.gov.br/jabot/f...
).

Table 2
Comparison of species richness for the main families and genera with epiphytic habit between Serra das Lontras National Park and other surveys carried at the Atlantic Forest. 1 N.P. - National Park; S.P. - State Park; N.R. - Natural Reserve. “-” - Information not available.

In addition to H. brachycephala and H. minor, both endemic to Bahia, H. edmundoi, a poorly-known species from Bahia for which no habitat information was available (Smith & Downs 1979Smith, L.B. & Downs, R.J. 1979. Bromelioideae (Bromeliaceae). Flora neotrop. 14(3): 1-649., Forzza et al. 2013Forzza, R.C., Costa, A., Siqueira Filho, J.A., Martinelli, G., Monteiro, R.F., Santos-Silva, F., Saraiva, D. P. & Paixão-Souza, B. 2013. Bromeliaceae. In Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB66 (accessed in 15/05/2013).
http://floradobrasil.jbrj.gov.br/jabot/f...
), was found in a submontane forest in the SLNP; H. augusta, previously considered to be restricted to southeastern Brazil, was also collected. These results reaffirm the importance of southern Bahia State in the distribution and taxonomy of this genus.

Vriesea (Tillandsioideae) and Aechmea (Bromelioideae) represent 12% and 7%, respectively, of the total number of species in the Atlantic Forest (Forzza et al. 2013Forzza, R.C., Costa, A., Siqueira Filho, J.A., Martinelli, G., Monteiro, R.F., Santos-Silva, F., Saraiva, D. P. & Paixão-Souza, B. 2013. Bromeliaceae. In Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB66 (accessed in 15/05/2013).
http://floradobrasil.jbrj.gov.br/jabot/f...
). Fontoura et al. (2012)Fontoura, T., Scudeller, V.V. & Costa, A.F. 2012. Floristics and environmental factors determining the geographic distribution of epiphytic bromeliads in the Brazilian Atlantic Rain Forest. Flora (Jena), 9(): 662-672. doi: 10.1016/j.flora.2012.05.003.
https://doi.org/10.1016/j.flora.2012.05....
reported that Bromelioideae taxa were encountered 1.6 times more frequently in local surveys of epiphytic species in the Atlantic Forest than representatives of Tillandsioideae. This proportion was lower in the SLNP, where Vriesea is very diverse, with almost the same numbers of species being encountered in each subfamily (34 spp. Bromelioideae x 30 spp. Tillandsioideae).

Another family that demonstrated elevated richness in the SLNP was Araceae, with one third of the 65 species known to the Atlantic Forest being encountered there (Coelho et al. 2013Coelho, M.A.N., Sakuragui, C.M., Mayo, S., Soares, M.L., Temponi, L.G., Calazans, L.S.B., Gonçalves, E.G., & Andrade, I.M. 2013. Araceae. In Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB51 (accessed in 15/05/2013).
http://floradobrasil.jbrj.gov.br/jabot/f...
). The most diversified genera of this family were likewise well represented in the park, with more than half of the species of Anthurium and Philodendron known from Bahia being encountered in the SLNP. Additionally, two new occurrence records for Bahia State (A. boudetii and P. edmundoi), one for the Atlantic Forest domain (A. gracile), and five morphotypes of Anthurium reinforce the importance of additional studies in mountainous areas there to improve our knowledge of the distribution of this family within the state. In contrast to the high endemism levels of Bromeliaceae and Orchidaceae, representatives of Araceae exhibit wide (and sometimes disjunct) distributions (Coelho et al. 2013Coelho, M.A.N., Sakuragui, C.M., Mayo, S., Soares, M.L., Temponi, L.G., Calazans, L.S.B., Gonçalves, E.G., & Andrade, I.M. 2013. Araceae. In Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB51 (accessed in 15/05/2013).
http://floradobrasil.jbrj.gov.br/jabot/f...
).

Orchidaceae is responsible for approximately 70% of the global epiphytic flora, and is always among the top families in every floristic study of that habit (Benzing 1990Benzing, D.H. 1990. Vascular epiphytes. Cambridge University Press, New York., Kersten 2010Kersten, R.A. 2010. Epífitas vasculares: histórico, participação taxonômica e aspectos relevantes, com ênfase na Mata Atlântica. Hoehnea 37(1): 9-38. doi: 10.1590/S2236-89062010000100001.
https://doi.org/10.1590/S2236-8906201000...
, Zotz 2013Zotz, G. 2013. The systematic distribution of vascular epiphytes - a critical update. Bot. j. Linn. Soc. 171: 453-481. doi: 10.1111/boj.12010.
https://doi.org/10.1111/boj.12010...
). Most orchid diversity is found in the tropics, especially in mountainous areas (Dressler 1993Dressler, R.L. 1993. Phylogeny and classification of the orchid family. Dioscorides Press, Portland.). The majority of the orchid taxa encountered in the SLNP is relatively small (e.g., Acianthera, Anathallis, Brachionidium, Jacquiniella, Lankesterella, Octomeria, Pabstiella, Pleurothallis, Promenaea, Specklinia, and Stelis) and commonly neglected in floristic surveys - and therefore poorly represented in scientific collections (Dressler 2005Dressler, R.L. 2005. How many orchid species? Selbyana 26(1,2): 155-158.). As the present study focused on epiphytes (and ex-situ cultivation was undertaken), high numbers of species were included on the final list - with 22 new occurrences for Bahia State (Table 1).

Northeastern Brazil represents one of the 10 areas in the world considered most likely to have undescribed species, and where models predict a 30% increase in known diversity (Joppa et al. 2011Joppa, L.N., Roberts, D.L., Myers, N. & Pimm, S.L. 2011. Biodiversity hotspots house most undiscovered plant species. P. Natl. Acad. Sci. USA 108(32): 13171-13176. doi: 10.1073/pnas.1109389108.
https://doi.org/10.1073/pnas.1109389108...
). Since initiating preliminary studies of the flora of the SLNP (Amorim et al. 2009Amorim, A.M.A., Jardim, J.G., Lopes, M.M.M., Fiaschi, P., Borges, R.A.X., Perdiz, R.O. & Thomas, W.W. 2009. Angiospermas em remanescentes de Floresta Montana no sul da Bahia, Brasil. Biota Neotrop. 9(3): 313-348 http://www.biotaneotropica.org.br/v9n3/en/abstract?inventory+bn02909032009 (accessed in 15/07/2013). doi: 10.1590/S1676-06032009000300028
http://www.biotaneotropica.org.br/v9n3/e...
), six new epiphyte species have been described for the area: Bertolonia alternifolia, B. bullata, Clistax bahiensis, Encyclia fimbriata, Quesnelia clavata, and Vriesea longisepala, and one is about to be published. The present survey found other possible taxonomic novelties, and recent studies in neighboring areas encountered several new species with epiphytic habits (e.g., Amorim & Leme 2009Amorim, A.M.A. & Leme, E.M.C. 2009. Two new species of Quesnelia (Bromeliaceae: Bromelioideae) from the Atlantic Rainforest of Bahia, Brazil. Brittonia 61(1): 14-21. doi: 10.1007/s12228-008-9049-5.
https://doi.org/10.1007/s12228-008-9049-...
, Leme & Kollmann 2011Leme, E.M.C. & Kollmann, L.J.C. 2011. New species and a new combination of Brazilian Bromeliaceae. Phytotaxa 16: 1-36., Costa et al. 2012Costa, A.F., Fontoura, T. & Amorim, A.M. 2012. Novelties in Bromeliaceae from the northeastern Brazilian Atlantic Rainforest. J. Torrey Bot. Soc. 139(1): 34-45. doi: 10.3159/TORREY-D-11-00061.1.
https://doi.org/10.3159/TORREY-D-11-0006...
, Reginato et al. 2013Reginato, M., Baumgratz, J.F.A. & Goldenberg, R. 2013. A taxonomic revision of Pleiochiton (Melastomataceae, Miconieae). Brittonia 65(1): 16-41. doi: 10.1007/s12228-012-9258-9.
https://doi.org/10.1007/s12228-012-9258-...
) - emphasizing the need of further investigations of this group in southern Bahia.

The present study expands the distribution of many taxa, with one new occurrence for Brazil (Elleanthus hymenophorus (Rchb.f.) Rchb.f.), four for the Atlantic Forest biome, and 30 for Bahia State (Table 1). Utricularia jamsoniana Oliv., a species previously known only from altitudinal regions of Central America and northern South America (Taylor 1989Taylor, P. 1989. The genus Utricularia - a taxonomic monograph. Kew Bulletin, Additional Series XIV, London.), was encountered in a cloud forest in the highest area of the park (Peito de Moça and Javi mountain ranges). Over half (16 spp.) of these new records were known only from collections in strictly southern and southeastern regions. The epiphytic flora of the SLNP shares more species with southern and southeastern Brazil (71 spp.) than with northeastern states (five species - four Aechmea and one Hohenbergia). This same pattern was found for ferns and lycophytes on a nearby mountainous area in the Serra Bonita Private Reserve, with almost a quarter of the surveyed species being previously restricted to southeastern Brazil (Matos et al. 2010Matos, F.B., Amorim, A.M. & Labiak, P.H. 2010. The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. J. Bot. Res. Inst. Texas 4(1): 333-346.).

Almost half of the 221 taxa identified to species level in the present study are endemic to the Atlantic Forest (100 spp. - 45%) (Table 1), and most of them belong to Bromeliaceae and Orchidaceae (44 and 32 species respectively). These two families exhibit high degrees of endemism in the Atlantic Forest, with 80% of Bromeliaceae and 63% of Orchidaceae being restricted to this domain (Martinelli et al. 2008Martinelli, G., Vieira, C.M., Gonzalez, M., Leitman, P., Piratininga, A., Costa, A.F. & Forzza, R.C. 2008. Bromeliaceae da Mata Atlântica Brasileira: lista de espécies, distribuição e conservação. Rodriguésia 59(1): 209-258., Stehmann et al. 2009Stehmann, J.R., Forzza, R.C., Sobral, M. & Kamino, L.H.Y. 2009. Gimnospermas e Angiospermas. In Plantas da Floresta Atlântica (J.R. Stehmann, R.C. Forzza, A. Salino, M. Sobral, D.P. Costa & L.H.Y. Kamino, eds.). Jardim Botânico do Rio de Janeiro, Rio de Janeiro. p. 27-37.). These high levels of endemism may be related to limited connections with the other major South American forest formation in the Amazon (Mori et al. 1981Mori, S., Boom, B.M. & Prance, G.T. 1981. Distribution patterns and conservation of eastern Brazilian coastal forest tree species. Brittonia 33(2): 233-245. doi: 10.2307/2806330.
https://doi.org/10.2307/2806330...
). These two forests are separated by an open area known as the “dry diagonal” (Prado & Gibbs 1993Prado, D. & Gibbs, P. 1993. Patterns of species distributions in the dry seasonal forests of South America. Ann. Missouri Bot. Gard. 80(4): 902-927. doi: 10.2307/2399937.
https://doi.org/10.2307/2399937...
) that acts as a barrier to species dispersion, especially less drought-tolerant taxa.

On the other hand, 60 species showed broad distributions in Brazil or the Neotropical region, and 24 had disjunct distributions between the Atlantic and Amazon forests (Table 1); 19 species also occur in the “Cerrado” (Brazilian savanna) region, suggesting that dry formations do not act as barriers to all epiphyte taxa, especially those with broader ecophysiological tolerances (Oliveira-Filho & Ratter 1995Oliveira-Filho, A.T. & Ratter, J.A. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinb. j. bot. 52(2): 141-194. doi: 10.1017/S0960428600000949.
https://doi.org/10.1017/S096042860000094...
, Ibisch et al. 1996Ibisch, P.L., Boegner, A., Nieder, J. & Barthlott, W. 1996. How diverse are neotropical epiphytes? An analysis based on the “Catalogue of the flowering plants and gymnosperms of Peru”. Ecotropica 2: 13-28.). Deciduous and semi-deciduous seasonal forest patches and gallery forests that crisscross the Cerrado may offer dispersal pathways for many species and establish tenuous connections between these two major forest formations (Leitão Filho 1987Leitão Filho, H.F. 1987. Considerações sobre a florística de florestas tropicais e subtropicais do Brasil. IPEF 35: 41-46., Oliveira-Filho & Ratter 1995Oliveira-Filho, A.T. & Ratter, J.A. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinb. j. bot. 52(2): 141-194. doi: 10.1017/S0960428600000949.
https://doi.org/10.1017/S096042860000094...
).

All of the 10 species cited on the Red Book of Brazilian Flora (Martinelli & Moraes 2013Martinelli, G. & Moraes, M.A. (orgs.) 2013. Livro vermelho da flora do Brasil. Andrea Jakobsson/Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1100 p.) as threatened belong to Bromeliaceae and Orchidaceae, two very ornamental families. The five bromeliads are restricted to Northeast Brazil, and four of them are endemic to Bahia. On the other hand, the orchids indicated as threatened are new occurrences to the state. These results reinforce the importance of the park as a conservation unit and the relevance of floristic surveys, especially for often neglected groups such as epiphytes. Recent floristic studies in mountainous areas in southern Bahia have revealed large numbers of new species and new occurrences of epiphytes - pointing to the need for further investigations of this group.

We thank FAPESB, FAPERJ and CAPES for fieldwork financial support. Paula Leitman thank CNPq and CAPES for master's scholarship. Rafaela C. Forzza and André Amorim are CNPq research fellows. We also thank the specialists who helped identifying the species - Alain Chautems, Andrea Costa, Charlotte Taylor, Claudio Nicoletti Fraga, Daniela Zappi, Daniele Monteiro, Eliane de Lima Jacques, Elton Leme, Elsie Franklin Guimarães, João Renato Stehmann, Ludovic Kollmann, Marcus Nadruz, Nigel Taylor, Ricardo Perdiz, Sheila Profice & Simon Mayo - and fieldwork helpers - José Lima Paixão, Carlinhos, Lukas Daneu and Ricardo Perdiz.

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Publication Dates

  • Publication in this collection
    14 Feb 2014
  • Date of issue
    April-June 2014

History

  • Received
    01 Aug 2013
  • Accepted
    14 Feb 2014
Instituto Virtual da Biodiversidade | BIOTA - FAPESP Departamento de Biologia Vegetal - Instituto de Biologia, UNICAMP CP 6109, 13083-970 - Campinas/SP, Tel.: (+55 19) 3521-6166, Fax: (+55 19) 3521-6168 - Campinas - SP - Brazil
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