Diversity high up: a cloud forest of the Serra da Mantiqueira as a vascular epiphyte hotspot

Furtado, Samyra Gomes; Menini Neto, Luiz

doi:10.1590/2175-7860201869201

Abstract

The diversity of montane environments is dictated by a variety of environmental conditions. Parque Estadual do Ibitipoca is located in the Serra da Mantiqueira, between ~1,000-1,800 m, and harbors approximately 300 ha of cloud forests. The composition of vascular epiphytes was determined by analyzing data from expeditions conducted between July 2014 and July 2015, and specimens deposited at herbaria. The 224 species were distributed into 82 genera of which Pleurothallis s.l. was the richest (13 spp.) and 23 families of which Orchidaceae was the richest (87 spp.). This richness corresponds to approximately 9.5% of the vascular epiphytic flora of the Atlantic Forest concentrated in an area that comprises 0.00085% of this phytogeographic domain, which represents one of the largest diversities ever sampled in the Brazilian Atlantic Forest. This fact is more relevant given that 13 species are threatened at the country level and 23 at the state level.

Key words:
Atlantic Forest; Dwarf-forest; epiphytic quotient; epiphytism; Parque Estadual do Ibitipoca

Resumo

A diversidade dos ambientes montanos é condicionada por inúmeras variáveis ambientais. O Parque Estadual do Ibitipoca está localizado na Serra da Mantiqueira, entre ~1.000-1.800 m, e abriga cerca de 300 ha de florestas nebulares. A composição de epífitas vasculares foi determinada pela análise dos dados de expedições, realizadas entre julho de 2014 e julho de 2015, e espécimes depositados em herbários. Foram registradas 224 espécies distribuídas em 82 gêneros dos quais Pleurothallis s.l. foi o mais rico (13 spp.) e 23 famílias das quais Orchidaceae foi a mais rica (87 spp.). Esta riqueza corresponde a aproximadamente 9,5% da flora de epífitas vasculares da Floresta Atlântica concentrados em uma área que compreende 0,00085% deste domínio fitogeográfico, representando uma das maiores diversidades já amostradas na Floresta Atlântica brasileira. Este fato é ainda mais relevante tendo em conta que 13 espécies estão ameaçadas a nível nacional e 23 em nível estadual.

Palavras-chave:
Floresta Atlântica; nanofloresta; quociente epifítico; epifitismo; Parque Estadual do Ibitipoca

Introduction

Mountainous vegetation harbors a huge diversity of plants due to a combination of factors including isolation (the mountains are comparable to islands), climatic changes that occur in little distances due to the elevational shift, and geodiversity resulting from topographically diverse terrain and differences in substrata (Körner 2004Körner C (2004) Mountain biodiversity, its causes and function. Ambio 13: 11-17. Available at <http://www.ncbi.nlm.nih.gov/pubmed/15575177>. Access on 15 September 2015.
http://www.ncbi.nlm.nih.gov/pubmed/15575... ). These factors apply to Parque Estadual do Ibitipoca, in the Serra da Mantiqueira in Minas Gerais, which harbors a remarkable plant richness despite comprising only 1,488 ha (Forzza et al. 2013Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) (2013) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. 382p.).

Epiphytes are plants that spend part or all of their life using another plant as support and are of high ecological importance because they provide shelter and resources to animals as well as capturing and storing water (Benzing 1990Benzing DH (1990) Vascular epiphytes: general biology and related biota. Cambridge University Press, Cambridge. 376p.; Zotz 2016Zotz G (2016) Plants on plants - the biology of vascular epiphytes. Springer, Cham. 282p.). This synusia also contributes to the diversity, representing 9-10% of the known vascular species (Gentry & Dodson 1987Gentry AH & Dodson CH (1987) Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden 74: 205-223.; Zotz 2013Zotz G (2013) The systematic distribution of vascular epiphytes - a critical update. Botanical Journal of the Linnean Society 171: 453-481.), with a particular diversity in Neotropical rainforests (Gentry and Dodson 1987Gentry AH & Dodson CH (1987) Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Garden 74: 205-223.). The percentage of vascular epiphytes in the Brazilian Atlantic Forest (BAF) reaches 15% of the whole domain of vascular flora, of which 78% are endemic to Brazil and around 11% are threatened with extinction. These statistics reinforce the need to acquire knowledge of the biodiversity conservation and ecosystem management practices for this epiphytic group (Freitas et al. 2016Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79.).

Studies regarding this epiphytic synusia in Minas Gerais have been intensified (Werneck & Espírito Santo 2002Werneck M & Espírito-Santo MM (2002) Species diversity and abundance of vascular epiphytes on Vellozia piresiana in Brazil. Biotropica 34: 51-57.; Alves et al. 2008Alves RJV, Kolbek J & Becker J (2008) Vascular epiphyte vegetation in rocky savannas of southeastern Brazil. Nordic Journal of Botany 26: 101-117.; Menini Neto et al. 2009aMenini Neto L, Matozinhos CN, Abreu NL, Valente ASM, Antunes K, Souza FS, Viana PL & Salimena FRG (2009b) Flora vascular não-arbórea de uma floresta de grota na Serra da Mantiqueira, Zona da Mata de Minas Gerais, Brasil. Biota Neotropica 9: 149-161.; Alves & Menini Neto 2014Alves FE & Menini Neto L (2014) Vascular epiphytes in a forest fragment of Serra da Mantiqueira and floristic relationships with Atlantic high altitude areas in Minas Gerais. Brazilian Journal of Botany 37: 187-196.; Barbosa et al. 2015Barbosa DEF, Basílio GA, Silva FR & Menini Neto L (2015) Vascular epiphytes in a remnant of seasonal semideciduous forest in Zona da Mata of Minas Gerais Brazil. Bioscience Journal 31: 623-633.; Furtado & Menini Neto 2015aFurtado SG & Menini Neto L (2015a) Diversity of vascular epiphytes in two high altitude biotopes of the Brazilian Atlantic Forest. Brazilian Journal of Botany 38: 295-310., bFurtado SG & Menini Neto L (2015b) Diversity of vascular epiphytes in urban environment: a case study in a biodiversity hotspot, the Brazilian Atlantic Forest. CES Revista 29: 82-101., 2016Furtado SG & Menini Neto L (2016) Vascular epiphytic flora of a high montane environment of Brazilian Atlantic Forest: composition and floristic relationships with other ombrophilous forests. Acta Botanica Brasilica 30: 422-436. DOI: 10.1590/0102-33062016abb0090
https://doi.org/10.1590/0102-33062016abb... ), although they can be considered scarce if taking in account the extension of this state.

A previous checklist of epiphytic angiosperms occurring in PEIB was presented by Menini Neto et al. (2009a)Menini Neto L, Matozinhos CN, Abreu NL, Valente ASM, Antunes K, Souza FS, Viana PL & Salimena FRG (2009b) Flora vascular não-arbórea de uma floresta de grota na Serra da Mantiqueira, Zona da Mata de Minas Gerais, Brasil. Biota Neotropica 9: 149-161., thus the present study aimed to complement the knowledge of vascular epiphytes in PEIB (including ferns as well as new records of angiosperms), an important conservation unit of Minas Gerais (Costa et al. 1998Costa CMR, Hermann G & Martins CS (1998) Biodiversidade em Minas Gerais: um atlas para sua conservação. Fundação Biodiversitas, Belo Horizonte. 94p.; Drummond et al. 2005Drummond GM, Martins CS, Machado ABM, Sebaio FA & Antonini Y (orgs.) (2005) Biodiversidade em Minas Gerais, um atlas para sua conservação. 2a ed. Fundação Biodiversitas, Belo Horizonte. 222p.; Forzza et al. 2013Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) (2013) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. 382p.), contributing to the biodiversity knowledge base of the Serra da Mantiqueira and Atlantic domain.

Material and Methods

Study site

The Parque Estadual da Serra do Ibitipoca (PEIB) covers approximately 1,488 ha in the southeastern region of Minas Gerais, in the municipality of Lima Duarte, district of Conceição de Ibitipoca between coordinates 21º40’-21º44’S and 43º52’-43º55’W (Fig. 1). It is part of the Mantiqueira Mountains (Serra da Mantiqueira) and has an elevational range of 1,000 to 1,784 m. The climate is classified as Cwb according to the Köppen system, with dry winters and mild summers. The averages of annual precipitation and temperature are 1,532 mm and 18.9 ºC, respectively (CETEC 1983CETEC (1983) Diagnóstico ambiental do estado de Minas Gerais. Fundação Centro Tecnológico de Minas Gerais, Belo Horizonte. 158p.).

Figure 1
Location of Parque Estadual do Ibitipoca, Minas Gerais, Brazil. Adapted from Oliveira-Filho et al. (2013)Oliveira-Filho AT, Fontes MAL, Viana PL, Valente ASM, Salimena FRG & Ferreira FM (2013) O mosaico de fitofisionomias do Parque Estadual do Ibitipoca. In: Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. Pp. 53-93..

The PEIB is situated in the Atlantic domain, and its vegetation harbors a mosaic of field and forest phytophysiognomies, with the predominance of campo rupestre, which has a high floristic richness and several endemic species. Cloud forests are interspersed throughout the field vegetation (covering approximately 300 ha or 20% of the whole extension of PEIB), especially the so called Mata Grande (with a canopy of ~20 m) and the formations of dwarf-forests (with canopy of ~5 m). In this environment, the richness of vascular epiphytes is remarkable due to the moisture provided by the fog and retained by the trees (Oliveira-Filho et al. 2013Oliveira-Filho AT, Fontes MAL, Viana PL, Valente ASM, Salimena FRG & Ferreira FM (2013) O mosaico de fitofisionomias do Parque Estadual do Ibitipoca. In: Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. Pp. 53-93.) (Fig. 2).

Figure 2
Vegetation of Parque Estadual do Ibitipoca - a. patches of cloud forest interspersed with campo rupestre; b. detail of fog covering the forests and campo rupestre; c. general view of a cloud forest; d. detail of the canopy of a cloud forest. e-g. interior view of the cloud forests.

The flora of PEIB is well documented with checklists of bryophytes (hornworts, liverworts, and mosses) (Luizi-Ponzo et al. 2013Luizi-Ponzo AP, Siviero TS, Amorim ET, Henriques DK, Rocha LM, Gomes HCS, Paiva LA, Rodrigues RS, Silva IC, Silva AGD, Ribeiro GC, Gomes CQ & Campeão AS (2013) Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Krieger. In: Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. Pp. 95-122.), ferns and lycophytes (Salino et al. 2013Salino A, Almeida TE, Mynssen CM, Condack JPS & Sylvestre LS (2013) Pteridófitas do Parque Estadual do Ibitipoca. In: Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. Pp. 123-152.), and spermatophytes (angiosperms and gymnosperms) (Forzza et al. 2013Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) (2013) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. 382p.).

Data collection

We conducted field trips between July 2014 and July 2015 in order to collect fertile specimens (when needed), observe and obtain photographic recordings of epiphytic species. The resulting photographs were published as a rapid color guide of The Field Museum of Chicago (<http://fieldguides.fieldmuseum.org/sites/default/files/rapid-color-guides-dfs/712_brasil-epifitas_de_ibitipoca.pdf>). The collected specimens were deposited at Herbarium CESJ of Universidade Federal de Juiz de Fora. Specimens deposited at the herbaria BHCB, CESJ, HB, R, RB, and VIC (acronyms according to Thiers 2016Thiers B [continuously updated]. Index herbariorum: a global directory of public herbaria and associated staff. The New York Botanical Garden. Available at <http://sweetgum.nybg.org/science/ih/>. Access on 20 June 2016.
http://sweetgum.nybg.org/science/ih/... ) from previous collections in the PEIB spanning more than 40 years were also examined.

The evolutionary lineages of vascular plants are according to PPG (2016)PPG I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603. for ferns and lycophytes and APG IV (2016)APG IV - The Angiosperm Phylogeny Group (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20. for angiosperms (magnoliids, monocotyledons, and eudicotyledons).

Data about endemism in Brazil were obtained in the database compiled by BFG (2015)BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113. (available at <http://dx.doi.org/10.6084/m9.figshare.1538647>) and Prado et al. (2015)Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CGV, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.. The conservation status of species in Brazil is according to Martinelli & Moraes (2013)Martinelli G & Moraes MA (orgs.) (2013) Livro vermelho da flora do Brasil. Jardim Botânico do Rio de Janeiro, Andrea Jakobsson, Rio de Janeiro. 1100p., and similar data for Minas Gerais were obtained from Drummond et al. (2008)Drummond GM, Machado ABM, Martins CS, Mendonça MP & Stehmann JR (2008) Listas vermelhas das espécies da fauna e da flora ameaçadas de extinção em Minas Gerais. 2a ed. Fundação Biodiversitas, Belo Horizonte. CD-Rom. Available at <http://www.biodiversitas.org.br/cdlistavermelha/default.asp> Access on 20 June 2015.
http://www.biodiversitas.org.br/cdlistav... . The distribution of the genera is according to the literature for each group.

Ecological categories are according to Benzing (1990)Benzing DH (1990) Vascular epiphytes: general biology and related biota. Cambridge University Press, Cambridge. 376p., and we did not differentiate between primary and secondary hemiepiphytes.

Data analysis

We calculated the epiphytic quotient according to Hosokawa (1950)Hosokawa T (1950) Epiphyte-quotient. Botanical Magazine of Tokyo 63: 18-19., which represents the percentage of epiphytic plants identified among the vascular species that occur in the cloud forest of PEIB. One calculation set consisted of all the recorded epiphytes, and the other set excluded the accidental ones. This calculation was possible due to the existence of checklists of ferns and lycophytes (Salino et al. 2013Salino A, Almeida TE, Mynssen CM, Condack JPS & Sylvestre LS (2013) Pteridófitas do Parque Estadual do Ibitipoca. In: Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. Pp. 123-152.) and angiosperms (Forzza et al. 2013Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) (2013) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. 382p.) in the PEIB. A filtering of species occurring in PEIB cloud forests was conducted, through data obtained in herbarium sheets, literature [especially the published floras for the Park, cited by Forzza et al. (2013)Forzza RC, Menini Neto L, Salimena FRG & Zappi D (orgs.) (2013) Flora do Parque Estadual do Ibitipoca e seu entorno. Editora UFJF, Juiz de Fora. 382p.] and field observation.

Results

A total of 224 species of vascular epiphytes were identified (of which 213 were classified at the specific level) that belonged to 82 genera and 23 families (Tab. 1), including 152 angiosperms [10 magnoliids (4.5%), 122 (54.3%) monocotyledons and 20 (8.5%) eudicotyledons], 69 (31.4%) ferns and 3 lycophytes (1.3%).

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Table 1
Checklist of vascular epiphyte species recorded in cloud forests of the Parque Estadual do Ibitipoca, Minas Gerais, Brazil.

Orchidaceae was the richest family with 86 identified species (38.6% of the total), followed by Polypodiaceae with 27 species (12.1%), and Bromeliaceae with 25 (11.2%) (Fig. 3). The main families of vascular epiphytes recorded in the PEIB are compared with those in some Brazilian ombrophilous forests (Tab. 2). Regarding the genera, Pleurothallis R.Br. sensu lato is the richest with 13 species (5.9%), Asplenium L. with 11 species (5%), and Peperomia Ruiz & Pav. and Epidendrum L. with 10 species each.

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Table 2
Comparison of species richness in families of vascular epiphytes in some checklists of rainforests

Figure 3
Representativity of the families of vascular epiphytes and comparison with percentage of endemics and threatened with extinction in the cloud forest of Parque Estadual do Ibitipoca, Minas Gerais, Brazil.

Among the ecological categories, characteristic holoepiphytes were classified as the most prominent [127 species (57%)], followed by facultative holoepiphytes [74 (33.2%)], accidental holoepiphytes [12 (5.4%)], and hemiepiphytes [11 (4.5%)].

The values of epiphytic quotients were 30% and 28.5%, respectively, for the total and excluded accidental holoepiphytes.

Discussion

The region encompassing Ibitipoca State Park has been considered of “Extreme Biological Importance” (Drummond et al. 2005Drummond GM, Martins CS, Machado ABM, Sebaio FA & Antonini Y (orgs.) (2005) Biodiversidade em Minas Gerais, um atlas para sua conservação. 2a ed. Fundação Biodiversitas, Belo Horizonte. 222p.), a fact that is corroborated by the richness of vascular epiphytes. This area represents one of the highest richness levels ever recorded in areas of dense ombrophilous forest in Brazil (Waechter 1992Waechter JL (1992) O epifitismo vascular na planície costeira do Rio Grande do Sul. Tese de Doutorado. Universidade Federal de São Carlos, São Carlos. 163p.; Kersten & Kunyioshi 2006Kersten RA & Kunyioshi YS (2006) Epífitos vasculares na Bacia do Alto Iguaçu, Paraná, Brasil-composição florística. Estudos de Biologia 28: 55-71.; Blum et al. 2011Blum CT, Roderjan CV & Galvão F (2011) Composição florística e distribuição altitudinal de epífitas vasculares da floresta ombrófila densa na Serra da Prata, Morretes, Paraná, Brasil. Biota Neotropica 11: 141-159.; Lima et al. 2011Lima RAF, Dittrich VAO, Souza VC, Salino A, Breier TB & Aguiar OT (2011) Flora vascular do Parque Estadual Carlos Botelho, São Paulo, Brasil. Biota Neotropica 11: 173-214.). This richness is even more remarkable when accounting for the total area of this conservation unit (1,488 ha), of which only 20% presents forest vegetation (i.e., less than 300 ha are composed of this physiognomy). The richness of vascular epiphytes in the PEIB also represents approximately 9.5% of the total found in the BAF [2,256 species according to Freitas et al. (2016)Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79.], in a conservation unit that comprises only 0.001% of this phytogeographic domain (0.00085% if considering only the area covered by cloud forests).

Evolutionary lineages have a different configuration regarding the proportion identified in the BAF by Kersten (2010)Kersten RA (2010) Epífitas vasculares - histórico, participação taxonômica e aspectos relevantes com ênfase na Mata Atlântica. Hoehnea 37: 9-38., with monocotyledons presenting a slightly smaller percentage (54.3% versus 63.5%) of that attributed to the BAF. A similar result was obtained for eudicotyledons (with 8.5% in PEIB and 14.1% in BAF). On the other hand, ferns have a higher diversity (31.4%) in the PEIB than in the BAF (16.4%).

The richest families found in the PEIB are among the richest often found in similar studies in several forest physiognomies in the Neotropical Region (Hietz & Hietz-Seifert 1995Hietz P & Hietz-Seifert U (1995) Composition and ecology of vascular epiphyte communities along an altitudinal gradient in Central Veracruz, Mexico. Journal of Vegetation Science 6: 487-498.; Dittrich et al. 1999Dittrich VAO, Kozera C & Silva SM (1999) Levantamento florístico dos epífitos vasculares do Parque Barigüi, Curitiba, Paraná, Brasil. Iheringia, série Botânica 52: 11-22.; Kersten & Silva 2001Kersten RA & Silva SM (2001) Composição florística e distribuição espacial de epífitas vasculares em floresta da planície litorânea da Ilha do Mel, Paraná, Brasil. Revista Brasileira de Botânica 24: 213-226.; Arévalo & Betancur 2004Arévalo R & Betancur J (2004) Diversidad de epífitas vasculares en cuatro bosques del sector suroriental de la Serranía de Chiribiquete, Guayana Colombiana. Caldasia 26: 359-380.; Giongo & Waechter 2004Giongo C & Waechter JL (2004) Composição florística e estrutura comunitária de epífitos em uma floresta de galeria na Depressão Central do Rio Grande do Sul. Revista Brasileira de Botânica 27: 563-572.; Kersten et al. 2009Kersten RA, Kuniyoshi YS & Roderjan CV (2009) Epífitas vasculares em duas formações ribeirinhas adjacentes na bacia do rio Iguaçu - Terceiro Planalto Paranaense. Iheringia, Série Botânica 64: 33-43.; Bianchi et al. 2012Bianchi JS, Michelon C & Kersten RA (2012) Epífitas vasculares de uma área de ecótono entre as Florestas Ombrófilas Densa e Mista, no Parque Estadual do Marumbi, PR. Estudos de Biologia (Ambiente e Diversidade) 34: 37-44.; Alves & Menini Neto 2014Alves FE & Menini Neto L (2014) Vascular epiphytes in a forest fragment of Serra da Mantiqueira and floristic relationships with Atlantic high altitude areas in Minas Gerais. Brazilian Journal of Botany 37: 187-196.; Barbosa et al. 2015Barbosa DEF, Basílio GA, Silva FR & Menini Neto L (2015) Vascular epiphytes in a remnant of seasonal semideciduous forest in Zona da Mata of Minas Gerais Brazil. Bioscience Journal 31: 623-633.), although the order can be different in some cases.

These families (Tab. 2) are also the main families of vascular epiphytes in the BAF, despite the differences in representativity. According to the data of Freitas et al. (2016)Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79., Araceae have a similar proportion in the BAF and PEIB, while Orchidaceae reaches 46.5% of all vascular epiphytes of the BAF and represents 38.6% of the total families in the PEIB. A similar result was found for Bromeliaceae, which represents 26.2% of the epiphytic species of the BAF and accounts for only 11.2% of the species in the PEIB. Thus, the reduced contribution of the last two families results in a smaller contribution of the monocotyledons to the epiphytic flora.

On the other hand, Polypodiaceae has greater representativity, with 12.1% of the vascular epiphytes of this conservation unit, a value substantially superior to the 4.3% found in the BAF (Freitas et al. 2016Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79.), contributing to the richness of ferns in the flora of the PEIB. The richness of this lineage was enhanced by the Hymenophyllaceae, (the fourth richest; 6.3% of total), which represents only 1.8% (in ninth position) in the BAF epiphytes, next to Aspleniaceae and Dryopteridaceae (5% each) which both have a greater relative representativity in the PEIB than in the BAF (around 1.8% each).

As reviewed by Kersten (2010)Kersten RA (2010) Epífitas vasculares - histórico, participação taxonômica e aspectos relevantes com ênfase na Mata Atlântica. Hoehnea 37: 9-38., Orchidaceae is the richest family in the dense ombrophilous forest of the BAF, followed by Bromeliaceae, Araceae, Polypodiaceae, Cactaceae, Dryopteridaceae, Gesneriaceae, Hymenophyllaceae, and Piperaceae. It is apparent that although the richest families in the PEIB (Tab. 2, Fig. 2) are among the richest of this type of forest in the BAF, there is a reduced contribution of the angiosperm families Araceae, Cactaceae, and Gesneriaceae.

A comparison with some punctual studies of vascular epiphytes in ombrophilous and montane ombrophilous forests (Tab. 2) shows a consistent greater relative richness of ferns, with Polypodiaceae as the second richest in four of six sites (all areas with elevations superior to 1,000 m). Hymenophyllaceae is well represented in all sites, and Aspleniaceae and Dryopteridaceae are also among the richest families in the majority of sites. On the other hand, there are fewer Araceae species in sites of high elevation, except in the Parque Estadual Carlos Botelho, which has a wide altitudinal gradient and lower altitudes of approximately 30 m.

Thus, ferns with only marginal importance in the BAF or globally (e. g., Aspleniaceae, Dryopteridaceae and Hymenophyllaceae) contribute less than 2% of the total richness in both cases (Madison 1977Madison M (1977) Vascular epiphytes: their systematic occurrence and salient features. Selbyana 2: 1-13.; Kersten 2010Kersten RA (2010) Epífitas vasculares - histórico, participação taxonômica e aspectos relevantes com ênfase na Mata Atlântica. Hoehnea 37: 9-38.; Zotz 2013Zotz G (2013) The systematic distribution of vascular epiphytes - a critical update. Botanical Journal of the Linnean Society 171: 453-481.; Freitas et al. 2016Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79.) in the PEIB have a greater richness to the detriment of angiosperm families of vascular epiphytes often found among the richest. For example, Araceae, Piperaceae, and Cactaceae are the fourth, fifth and sixth richest families in the BAF, respectively; however, Cactaceae did not figure among the richest of the PEIB.

Elevation could contribute to this pattern, due to the sensitivity of some families to low temperatures. For example, Araceae has a lower percentage in the physiognomy of the mixed ombrophilous forests, which are common in the southern region of Brazil (Kersten 2010Kersten RA (2010) Epífitas vasculares - histórico, participação taxonômica e aspectos relevantes com ênfase na Mata Atlântica. Hoehnea 37: 9-38.), occurring at higher latitudes (and, consequently, lower temperatures). Consistently, only one species of Araceae was observed by Furtado & Menini Neto (2016)Furtado SG & Menini Neto L (2016) Vascular epiphytic flora of a high montane environment of Brazilian Atlantic Forest: composition and floristic relationships with other ombrophilous forests. Acta Botanica Brasilica 30: 422-436. DOI: 10.1590/0102-33062016abb0090
https://doi.org/10.1590/0102-33062016abb... in a mixed ombrophilous forest in the Serra da Mantiqueira ranging from 1,600 to 1,650 m of elevation. Krömer et al. (2005)Krömer T, Kessler M, Gradstein R & Acebey A (2005) Diversity patterns of vascular epiphytes along an elevational gradient in the Andes. Journal of Biogeography 32: 1799-1809. and Cardelús et al. (2006)Cardelús CL, Colwell RK & Watkins JE (2006) Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology 94: 144-156. also recorded a reduction in the richness of this family correlating with the enhancement of elevation in the Andes. On the other hand, ferns have a greater richness that correlates with enhanced elevation (Moran 1995Moran RC (1995) The importance of mountains to pteridophytes, with emphasis on Neotropical montane forests. In: Churchill SP, Balslev H, Forero E & Luteyn JL (eds.) Biodiversity and conservation of Neotropical montane forests (Proceedings of a symposium, New York Botanical Garden, 21-26 June 1993). New York Botanical Garden, New York. Pp. 359-363.; Hietz & Hietz-Seifert 1995Hietz P & Hietz-Seifert U (1995) Composition and ecology of vascular epiphyte communities along an altitudinal gradient in Central Veracruz, Mexico. Journal of Vegetation Science 6: 487-498.; Krömer et al. 2005Krömer T, Kessler M, Gradstein R & Acebey A (2005) Diversity patterns of vascular epiphytes along an elevational gradient in the Andes. Journal of Biogeography 32: 1799-1809.; Cardelús et al. 2006Cardelús CL, Colwell RK & Watkins JE (2006) Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology 94: 144-156.). Further investigation in Brazilian mountainous areas will contribute to determining whether this is a consistent pattern associated with cloud forests.

The epiphytic quotient observed in the PEIB reinforces the relevance of epiphytes in the diversity of flora. This quotient is 15% in the BAF (Freitas et al. 2016Freitas L, Salino A, Menini Neto L, Almeida TE, Mortara SR, Stehmann JR, Amorim AM, Guimarães EF, Coelho MN, Zanin A & Forzza RC (2016) A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates. PhytoKeys 58: 65-79.), and this index varies from 13 to 32% (average of ~20%) in areas of Brazilian dense ombrophilous forests (Kersten 2010Kersten RA (2010) Epífitas vasculares - histórico, participação taxonômica e aspectos relevantes com ênfase na Mata Atlântica. Hoehnea 37: 9-38.). Thus, the quotient percentage observed in the PEIB is higher than the average in the BAF and of the majority of areas where this value was calculated. The relevance of this value must be highlighted because it is the same as that of the Equator flora (Moller-Jorgensen & León-Yánez 1999Moller-Jorgensen P & León-Yánez S (1999) Catalogue of the vascular plants of Ecuador. Missouri Botanical Garden Press, St. Louis. 1181p.; Zotz 2016Zotz G (2016) Plants on plants - the biology of vascular epiphytes. Springer, Cham. 282p.) and higher than the quotient for the flora of Panama (Foster & Hubbel 1990Foster RB & Hubbell SP (1990) The floristic composition of the Barro Colorado Island forest. In: Gentry AH (ed.). Four Neotropical rainforests. Yale University Press, New Haven. Pp. 85-98.), indicating the importance of this synusia to the cloud forest in the studied area.

The majority of the 75 recorded genera (excluding those completely accidental) are tropically distributed (34 Neotropical and 17 Pantropical), and only four genera are considered Cosmopolitan. Eight genera occur exclusively in the BAF, of which six are endemic to Brazil (Centroglossa Barb.Rodr., DyssochromaMiers, HatioraBritton & Rose, Promenaea Lindl., SchlumbergeraLem., ThysanoglossaPorto & Brade). Six genera are found in the BAF and Cerrado (CE), of which three are endemic to Brazil (GrobyaLindl., IsabeliaBarb.Rodr., NematanthusSchrad.). In addition, Nidularium is endemic to Brazil, occurring in Caatinga (CA) and the BAF, and Wittrockia Lindm. is also endemic, but occurs in CA, CE, and BAF. Griselinia J.R. Forst. & G. Forst present a disjunct distribution in South America and New Zealand, and Neoregelia L.B.Sm. is disjunct between the Andes/Amazonia (AM) and the BAF. Scuticaria Lindl. is found in AM, CE, BAF, and Racinaea M.A. Spencer & L.B.Sm. in AM and AF.

Among the 111 endemic Brazilian species (BFG 2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113.; Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CGV, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.), four are endemic to Minas Gerais: Anthurium leoni, Neoregelia oligantha, Hoffmannseggella crispata (accidental), and Peperomia decora. Among the angiosperms, 38 represent species not recorded in the PEIB by Menini Neto et al. (2009a)Menini Neto L, Matozinhos CN, Abreu NL, Valente ASM, Antunes K, Souza FS, Viana PL & Salimena FRG (2009b) Flora vascular não-arbórea de uma floresta de grota na Serra da Mantiqueira, Zona da Mata de Minas Gerais, Brasil. Biota Neotropica 9: 149-161., of which only six are accidental.

Thirteen species are threatened with extinction at the country level, of which five are classified as Near Threatened, four as Vulnerable, three as Endangered, and one as Critically Endangered (Martinelli & Moraes 2013Martinelli G & Moraes MA (orgs.) (2013) Livro vermelho da flora do Brasil. Jardim Botânico do Rio de Janeiro, Andrea Jakobsson, Rio de Janeiro. 1100p.). There are 23 threatened species at the state level, of which seven are classified as Vulnerable, nine as Endangered, six as Critically Endangered, and one as Data Deficient (Drummond et al. 2008Drummond GM, Machado ABM, Martins CS, Mendonça MP & Stehmann JR (2008) Listas vermelhas das espécies da fauna e da flora ameaçadas de extinção em Minas Gerais. 2a ed. Fundação Biodiversitas, Belo Horizonte. CD-Rom. Available at <http://www.biodiversitas.org.br/cdlistavermelha/default.asp> Access on 20 June 2015.
http://www.biodiversitas.org.br/cdlistav... ). Orchidaceae has the greatest number of threatened species (16 at the country or state level, or both), due in particular to the ornamental appeal of several species.

The conservation of epiphytic species represents a challenge due to habitat suppression, which threatens the biodiversity as a whole or due to the sensitivity of fragmentation and collection of ornamental species. Both the richness and the complexity of epiphyte communities in the PEIB, as well as the presence of species at different levels of risk and endemism, reinforces its importance for biodiversity, revealing that relatively small areas are also relevant to the preservation, and even consolidated conservation units must improve the strategies for maintaining the biodiversity.

1
Part of Master dissertation of the first author.

Editora de área: Dra. Cassia Sakuragui

Available material also at <https://figshare.com/s/6acf229b87cdc8b07e0e>

Acknowledgements

We would like to thank the specialists who helped us with the identification of the species: ferns (Alexandre Salino, Thaís Elias Almeida, Claudine M. Mynssen, João Paulo Santos Condack, Lana da Silva Sylvestre, Vinicius Antônio Oliveira Dittrich), Araceae (Livia Godinho, Cassia Sakuragui, Marcus Nadruz Coelho), Bromeliaceae (Rafaela Campostrini Forzza, Raquel Monteiro), Cactaceae (Daniela Zappi, Diego Rafael Gonzaga), Gesneriaceae (Alain Chautems), Piperaceae (Erika Von Sohsten Medeiros, Elsie Franklin Guimarães, Daniele Monteiro); Instituto Estadual de Florestas of Minas Gerais (IEF-MG) and Programa de Pós-Graduação em Ecologia da Universidade Federal de Juiz de Fora (PGECOL/UFJF) for the logistic support; Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the study grant conceded to S.G. Furtado, and to Dr. Ana Paula Gelli de Faria and Dr. Rafaela Campostrini for the comments on the manuscript.

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Publication Dates

Publication in this collection
Apr-Jun 2018

History

Received
12 Sept 2016
Accepted
15 June 2017

This is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited

[1] 1
Part of Master dissertation of the first author.

Lineages, families and species	Voucher	CS	EC
Lycophytes (1/3)
Lycopodiaceae (1/3)
Phlegmariurus biformis (Hook.) B.Øllg. *	L.Krieger (CESJ 2665)		CHL
Phlegmariurus fontinaloides(Spring) B.Øllg. *	L.Krieger (CESJ 9347)		CHL
Phlegmariurus heterocarpon(Fée) B.Øllg.	S.G.Furtado 316		CHL
Ferns (35/71)
Aspleniaceae (1/10)
Asplenium auriculatumSw.	S.G.Furtado 308		FHL
Asplenium auritumSw.	R.F.Novelino et al. 842		FHL
Asplenium clausseniiHieron.	C.M.Mynssen 825		FHL
Asplenium feeiKunze ex Fee	C.M.Mynssen 824		FHL
Asplenium geraense (C.Chr.) Sylvestre	S.G.Furtado 309		FHL
Asplenium harpeodes Kunze	R.F.Novelino et al. 919		FHL
Asplenium oligophyllumKaulf.	P.B.Pitta 281		FHL
Asplenium praemorsumSw.	S.G.Furtado 319		FHL
Asplenium pseudonitidumRaddi *	R.C.Forzza 3588		FHL
Asplenium raddianumGaudich.	R.C.Forzza 3149		FHL
Asplenium serra Langsd. & Fisch.	L.Krieger (CESJ 8383)		FHL
Blechnaceae (1/1)
Lomaridium acutum(Desv.) Gasper & V.A.O. Dittrich	R.F.Novelino 860		FHL
Dryopteridaceae (4/11)
Arachniodes denticulata (Sw.) Ching	B.R.Silva 1367		AHL
Elaphoglossum gayanum (Fée) T.Moore	S.G.Furtado 314		FHL
Elaphoglossum glabellum J.Sm.	C.M.Mynssen 786		FHL
Elaphoglossum lingua (C.Presl) Brack. *	R.F.Novelino 1017		FHL
Elaphoglossum lisboaeRosenst. *	R.F.Novelino 1029		CHL
Elaphoglossum luridum(Fee) Christ	R.F.Novelino 1014		FHL
Elaphoglossum pachydermum (Fée) T.Moore *	J.E.Z.Oliveira 217		AHL
Elaphoglossum strictum (Raddi) T.Moore	J.E.Z.Oliveira 398		CHL
Elaphoglossum vagans (Mett.) Hieron. *	R.F.Novelino 1188		FHL
Polybotrya speciosa Schott *	C.M.Mynssen 828		Hem
Rumohra adiantiformis (G.Forst.) Ching	L.Menini Neto 1354		FHL
Hymenophylaceae (4/14)
Didymoglossum hymenoides (Hedw.) Desv.	L.Krieger (BHCB 4220)	DD MG	CHL
Didymoglossum krausii(Hook. & Grev.) C.Presl	L.Krieger (CESJ 18823)		CHL
Hymenophyllum crispum Kunth	T.E.Almeida 1171		CHL
Hymenophyllum fragile(Hedw.) C.V.Morton	T.E.Almeida 1186		CHL
Hymenophyllum fucoides (Sw.) Sw.	T.E.Almeida 1241		CHL
Hymenophyllum hirsutum(L.) Sw.	L.Krieger (CESJ 8375)		FHL
Hymenophyllum polyanthos(Sw.) Sw.	L.Krieger (CESJ 3549)		CHL
Hymenophyllum pulchellumSchltdl. & Cham.	T.E.Almeida 1167		CHL
Hymenophyllum sp.	S.G.Furtado 289		CHL
Polyphlebium angustatum (Carmich.) Ebihara & Dubuisson	C.M.Mynssen 831		CHL
Polyphlebium diaphanum(Kunth) Ebihara & Dubuisson *	L.Krieger (CESJ 16218)		FHL
Polyphlebium hymenophylloides(Bosch) Ebihara & Dubuisson	T.E.Almeida 1181		CHL
Trichomanes pilosum Raddi	R.F.Novelino 870		AHL
Trichomanes polypodioidesRaddi	C.M.Mynssen 779		CHL
Nephrolepidaceae (1/1)
Nephrolepis sp.	Not collected		AHL
Polypodiaceae (12/27)
Campyloneurum angustifolium(Sw.) Fee	R.C.Forzza 3152		CHL
Campyloneurum nitidum(Kaulf.) C.Presl	S.G.Furtado 310		FHL
Campyloneurum phyllitidis(L.) C.Presl	J.E.Z.Oliveira 228		CHL
Cochlidium punctatum(Raddi) L.E.Bishop *	L.Krieger (CESJ 11674)		CHL
Cochlidium serrulatum(Sw.) L.E.Bishop	S.G.Furtado 294		FHL
Lellingeria apiculata(Kunze ex Klotzsch) A.R.Sm. & R.C.Moran	L.Krieger (CESJ 64862)		CHL
Leucotrichum sp.	Not collected		CHL
Melpomene flabelliformis(Poir.) A.R.Sm. & R.C.Moran *	L. Krieger (CESJ 15133)		CHL
Melpomene peruviana(Desv.) A.R.Sm. & R.C.Moran	L.Krieger (CESJ 8380)	CR MG	FHL
Melpomene pilosissima (M.Martens & Galeotti) A.R.Sm. & R.C.Moran	S.G.Furtado 288		CHL
Microgramma percussa(Cav.) de la Sota	L.Krieger (CESJ 21276)		FHL
Microgramma squamulosa (Kaulf.) de la Sota	R.F.Novelino 933		FHL
Moranopteris achilleifolia(Kaulf.) R.Y. Hirai & J. Prado *	S.G.Furtado 287		CHL
Moranopteris gradata (Baker) R.Y. Hirai & J. Prado *	R.C.Forzza 4436		FHL
Niphidium crassifolium(L.) Lellinger	S.G.Furtado 323		FHL
Pecluma pectinatiformis (Lindm.) M.G.Price	C.M.Mynssen 788		CHL
Pecluma recurvata(Kaulf.) M.G.Price *	R.C.Forzza 3158		CHL
Pecluma truncorum (Lindm.) M.G.Price *	C.M.Mynssen 833		CHL
Phlebodium pseudoaureum(Cav.) Lellinger	R.C.Forzza 3587		FHL
Pleopeltis astrolepis (Liebm.) E.Fourn.	S.G.Furtado 285		CHL
Pleopeltis hirsutissima(Raddi) de la Sota	L.Krieger (CESJ 27419)		CHL
Pleopeltis macrocarpa(Bory ex Willd.) Kaulf.	R.F.Novelino 1263		CHL
Pleopeltis minarum(Weath.) Salino *	R.C.Forzza 3715		CHL
Serpocaulon catharinae (Langsd. & Fisch.) A.R.Sm. *	D.Sucre 6737		FHL
Serpocaulon fraxinifolium (Jacq.) A.R.Sm.	J.E.Z.Oliveira 375		Hem
Serpocaulon latipes(Langsd. & Fisch.) A.R.Sm. *	L.Krieger (CESJ 8378)		AHL
Polypodiaceae indet.	S.G.Furtado 292		CHL
Pteridaceae (2/4)
Radiovittaria gardneriana (Fée) E.H.Crane	C.M.Mynssen 813		CHL
Radiovittaria stipitata(Kunze) E.H.Crane	J.E.Z.Oliveira 397		FHL
Vittaria graminifolia Kaulf.	S.G.Furtado 315		FHL
Vittaria lineata (L.) Sm.	R.F.Novelino 59		FHL
Angiosperms
Magnoliids (1/10)
Piperaceae (1/10)
Peperomia augescensMiq. #*	F.Salimena (CESJ 27411)		AHL
Peperomia corcovadensis Gardn. #	D.Monteiro 625		CHL
Peperomia crinicaulis C.DC. #*	S.G.Furtado 291		CHL
Peperomia decoraDahlst. * (MG)	L.Krieger (CESJ 8541)		FHL
Peperomia diaphanoides Dahlst. #	S.G.Furtado 306		FHL
Peperomia galioidesKunth #	S.G.Furtado 303		FHL
Peperomia loxensisKunth #	D.Sucre 7246		CHL
Peperomia mandioccanaMiq. *	S.G.Furtado 302		FHL
Peperomia rotundifolia (L.) Kunth #	H.Magalhães (R 86560)		CHL
Peperomia tetraphylla(G.Forst.) Hook. & Arn.	L.Krieger (CESJ 16238)		CHL
Monocotyledons (42/120)
Amaryllidaceae (1/1)
Hippeastrum aulicum (Ker Gawl.) Herb. #*	P.M.Andrade (BHCB 92378)		FHL
Araceae (2/9)
Anthurium boudetii Nadruz *	L.Temponi 397		FHL
Anthurium comtum Schott *	L.Temponi 400		FHL
Anthurium leonii E.G.Gonç. * (MG)	R.C.Forzza 4270	CR MG	FHL
Anthurium minarumSakur. & Mayo *	L.Temponi 390		FHL
Anthurium scandens (Aubl.) Engl.	R.C.Forzza 2663		CHL
Philodendron appendiculatum Nadruz & Mayo *	R.C.Forzza 3638		Hem
Philodendron bipinnatifidum Schott	L.Temponi 410		Hem
Philodendron minarumEngl. *	R.C.Forzza 2653		Hem
Philodendron propinquumSchott *	L.Temponi 398		Hem
Bromeliaceae (8/25)
Aechmea bromeliifolia(Rudge) Baker #	E.M.C.Leme 1474		CHL
Aechmea nudicaulis(L.) Griseb. *	R.Monteiro 13		CHL
Billbergia alfonsijoannisReitz *	E.M.C.Leme 1475		CHL
Serpocaulon fraxinifolium (Jacq.) A.R.Sm.	J.E.Z.Oliveira 375		Hem
Serpocaulon latipes(Langsd. & Fisch.) A.R.Sm. *	L.Krieger (CESJ 8378)		AHL
Polypodiaceae indet.	S.G.Furtado 292		CHL
Pteridaceae (2/4)
Radiovittaria gardneriana (Fée) E.H.Crane	C.M.Mynssen 813		CHL
Radiovittaria stipitata(Kunze) E.H.Crane	J.E.Z.Oliveira 397		FHL
Vittaria graminifolia Kaulf.	S.G.Furtado 315		FHL
Vittaria lineata (L.) Sm.	R.F.Novelino 59		FHL
Angiosperms
Magnoliids (1/10)
Piperaceae (1/10)
Peperomia augescensMiq. #*	F.Salimena (CESJ 27411)		AHL
Peperomia corcovadensis Gardn. #	D.Monteiro 625		CHL
Peperomia crinicaulis C.DC. #*	S.G.Furtado 291		CHL
Peperomia decoraDahlst. * (MG)	L.Krieger (CESJ 8541)		FHL
Peperomia diaphanoides Dahlst. #	S.G.Furtado 306		FHL
Peperomia galioidesKunth #	S.G.Furtado 303		FHL
Peperomia loxensisKunth #	D.Sucre 7246		CHL
Peperomia mandioccanaMiq. *	S.G.Furtado 302		FHL
Peperomia rotundifolia (L.) Kunth #	H.Magalhães (R 86560)		CHL
Peperomia tetraphylla(G.Forst.) Hook. & Arn.	L.Krieger (CESJ 16238)		CHL
Monocotyledons (42/120)
Amaryllidaceae (1/1)
Hippeastrum aulicum (Ker Gawl.) Herb. #*	P.M.Andrade (BHCB 92378)		FHL
Araceae (2/9)
Anthurium boudetii Nadruz *	L.Temponi 397		FHL
Anthurium comtum Schott *	L.Temponi 400		FHL
Anthurium leonii E.G.Gonç. * (MG)	R.C.Forzza 4270	CR MG	FHL
Anthurium minarumSakur. & Mayo *	L.Temponi 390		FHL
Anthurium scandens (Aubl.) Engl.	R.C.Forzza 2663		CHL
Philodendron appendiculatum Nadruz & Mayo *	R.C.Forzza 3638		Hem
Philodendron bipinnatifidum Schott	L.Temponi 410		Hem
Philodendron minarumEngl. *	R.C.Forzza 2653		Hem
Philodendron propinquumSchott *	L.Temponi 398		Hem
Bromeliaceae (8/25)
Aechmea bromeliifolia(Rudge) Baker #	E.M.C.Leme 1474		CHL
Aechmea nudicaulis(L.) Griseb. *	R.Monteiro 13		CHL
Billbergia alfonsijoannisReitz *	E.M.C.Leme 1475		CHL
Cattleya loddigesiiLindl. *	Without collector (CESJ 27534)	EN MG	CHL
Centroglossa macrocerasRchb.f. *	R.C.Forzza 54		CHL
Dichaea cogniauxiana Schltr. *	L.Menini Neto 142		CHL
Elleanthus brasiliensis(Lindl.) Rchb.f. #	L.Menini Neto 126		FHL
Encyclia patens Hook. *	S.G.Furtado 299		CHL
Epidendrum armeniacumLindl.	L.Menini Neto 175		CHL
Epidendrum avicula Lindl. #	M.C.Brügger (CESJ 24693)		CHL
Epidendrum chlorinumBarb.Rodr. *	L.Menini Neto 171		CHL
Epidendrum cf. filicaule Lindl. #	L.Menini Neto 1350		CHL
Epidendrum pseudodifformeHoehne & Schltr. *	L.Menini Neto 97		CHL
Epidendrum ochrochlorumBarb.Rodr. *	S.G.Furtado 297	EN MG	CHL
Epidendrum paranaenseBarb.Rodr. *	L.Menini Neto 131		FHL
Epidendrum ramosumJacq. #	R.C.Forzza 16		FHL
Epidendrum rigidum Jacq.	L.Menini Neto 71		CHL
Epidendrum secundum Jacq. #	L. Menini Neto 46		FHL
Eurystyles actinosophila (Barb.Rodr.) Schltr. #	S.G. Furtado 326		CHL
Eurystyles cogniauxii (Kraenzl.) Pabst *	L.Menini Neto 77		CHL
Gomesa glaziovii Cogn. *	L.Menini Neto 76		FHL
Gomesa gomezoides(Barb.Rodr.) Pabst *	L.Menini Neto 30		CHL
Gomesa recurvaR.Br.	L.Menini Neto 154		CHL
Grobya amherstiaeLindl. *	R.C.Forzza 26		CHL
Hadrolaelia coccinea (Lindl.) Chiron & V.P.Castro	L.Menini Neto 161	EN MG	CHL
Hoffmannseggella crispata(Thunb.) H.G.Jones #* (MG)	Not collected	NT BR EN MG	AHL
Isabelia violacea (Lindl.) van den Berg & M.W.Chase *	S.G.Furtado 282		CHL
Isabelia virginalisBarb.Rodr.	L.Menini Neto 47	VU BR	CHL
Isochilus linearis(Jacq.) R.Br	L.Menini Neto 44		CHL
Lankesterella gnoma (Kraenzl.) Hoehne *	L.Menini Neto 139		CHL
Masdevallia infracta Lindl.	L.Menini Neto 173		CHL
Maxillaria brasiliensisBrieger & Illg #*	L.Menini Neto 88		FHL
Maxillaria gracilisLodd. #*	L.C.S.Assis 1080		FHL
Maxillaria notylioglossa Rchb.f.	L.Menini Neto 119		CHL
Maxillaria ochroleuca Lodd. ex Lindl.	L.Menini Neto 87		CHL
Maxillaria picta Hook. #	R.C.Forzza 92		FHL
Maxillaria subulataLindl. *	L.Menini Neto 48		FHL
Octomeria crassifolia Lindl.	L.Menini Neto 138		CHL
Octomeria diaphanaLindl. *	L.Menini Neto 111		CHL
Octomeria grandifloraLindl.	S.G.Furtado 300		CHL
Octomeria rubrifolia Barb.Rodr. *	L.Menini Neto 40		CHL
Octomeria wawrae Rchb.f. *	L.Menini Neto 168	EN BR	CHL
Oncidium divaricatumLindl. *	R.C.Forzza 2190	VU BR	CHL
Oncidium gravesianumRolfe *	L.Menini Neto 112		CHL
Oncidium hookeriRolfe *	L.Menini Neto 96		CHL
Oncidium longipesLindl.	L.Menini Neto 163		CHL
Oncidium truncatumPabst *	L.Menini Neto 95	CR BR	CHL
Oncidium warmingiiRchb.f. #	G.Martinelli 15300	VU MG	AHL
Pleurothallis cryptophoranthoides Loefgr. *	L.Menini Neto 176	EN MG	CHL
Pleurothallis heliconiscapa Hoehne #*	H.C.Sousa (BHCB 9833)		CHL
Pleurothallis hypnicola Lindl.	L.Menini Neto 134		CHL
Pleurothallis liparanges Rchb.f. *	L.Menini Neto 177	CR MG	CHL
Pleurothallis luteola Lindl.	L.Menini Neto 158		CHL
Pleurothallis malachantha Rchb.f. *	L.Menini Neto 90	VU MG	CHL
Pleurothallis marginalis Rchb.f.	L.Menini Neto 162		CHL
Pleurothallis quartzicola (Barb.Rodr.) Cogn. *	S.G.Furtado 283		CHL
Pleurothallis recurva Lindl.	L.Menini Neto 236		CHL
Pleurothallis rubens Lindl.	L.Menini Neto 31		CHL
Pleurothallis saundersiana Rchb.f. *	L.Menini Neto 37		CHL
Pleurothallis cf. saurocephala Lodd. #*	Not collected		CHL
Pleurothallis tricarinata Poepp. & Endl.	L.Menini Neto 118		CHL
Polystachya hoehneanaKraenzl. *	L.Menini Neto 91	VU MG	CHL
Polystachya estrellensisRchb.f. #*	L.Menini Neto 1348		CHL
Prescottia stachyodes (Sw.) Lindl. #	D.R.Gonzaga 43		AHL
Promenaea xanthina(Lindl.) Lindl. *	L.Menini Neto 130		CHL
Prosthechea allemanoides (Hoehne) W.E.Higgins *	L.Menini Neto 26		FHL
Prosthechea calamaria (Lindl.) W.E.Higgins *	L.Menini Neto 180		CHL
Prosthechea pachysepala(Klotzsch) Chiron & V.P.Castro *	L.Menini Neto 36		FHL
Prosthechea aff. pachysepala (Klotzsch) Chiron & V.P.Castro #	D.R.Gonzaga 44		CHL
Scaphyglottis modesta(Rchb.f.) Schltr.	L.Menini Neto 52		FHL
Scuticaria hadwenii (Lindl.) Planch. *	R.C.Forzza 15	EN MG	CHL
Stelis apricaLindl.	L.Menini Neto 127		CHL
Stelis intermedia Poepp. & Endl.	L.Menini Neto 159		CHL
Stelis megantha Barb.Rodr. *	L.Menini Neto 148		CHL
Stelis papaquerensis Rchb.f.	L.Menini Neto 157		CHL
Stelis aff. caespitosa Lindl.	L.Menini Neto 25		CHL
Thysanoglossa organensisBrade *	L.Menini Neto 89		CHL
Eudicotiledôneas (13/19)
Begoniaceae (1/2)
Begonia angulata Vell. *	S.G.Furtado 321		FHL
Begonia sp.	R.C.Forzza 4287		FHL
Cactaceae (5/9)
Arthrocereus melanurus (K.Schum.) Diers et al. subsp. magnus N.P.Taylor & Zappi #* (MG)	Not collected	EN BR CR MG	AHL
Hatiora salicornioides (Haw.) Britton & Rose *	M.C.Brügger (CESJ 21541)		FHL
Lepismium cruciforme (Vell.) Miq. #	L.Menini Neto 1351		CHL
Lepismium houlletianum (Lem.) Barthlott	S.G.Furtado 313		CHL
Rhipsalis ellipticaG.Lindb. ex K.Schum. *	R.C.Forzza 3226		CHL
Rhipsalis floccosaSalm-Dyck ex Pfeiff.	L.Krieger (CESJ 8589)		CHL
Rhipsalis juengeriBarthlott & N.P.Taylor *	L.Krieger (CESJ 8594)		CHL
Rhipsalis pulchra Loefgr. *	L.Krieger (CESJ 9296)		CHL
Schlumbergera opuntioides (Loefgr. & Dusén) D.R.Hunt *	D.C.Zappi 258	VU BR VU MG	FHL
Gesneriaceae (2/3)
Nematanthus crassifolius (Schott) Wiehler *	R.C.Forzza 4274	VU MG	CHL
Nematanthus strigillosus (Mart.) H.E.Moore #*	L.Krieger (CESJ 13168)	NT BR	FHL
Sinningia magnifica (Otto & A.Dietr.) Wiehler *	R.C.Forzza (CESJ 27323)		FHL
Griseliniaceae (1/1)
Griselinia ruscifolia (Clos) Taub.	S.G.Furtado 322	NT BR	Hem
Lentibulariaceae (1/1)
Utricularia reniformis A.St.-Hil. #*	R.C.Forzza 3095		CHL
Moraceae (1/1)
Ficus cf. mexiae Standl. #*	Not collected		Hem
Rubiaceae (1/1)
Hillia parasitica Jacq.	L.Menini Neto 1353		FHL
Solanaceae (1/1)
Dyssochroma viridiflorum(Sims) Miers #*	S.G.Furtado 301		Hem
Urticaceae (1/1)
Coussapoa microcarpa (Schott) Rizzini #	not collected		Hem

Parque Estadual do Ibitipoca		Serra do Cruz (Minas Gerais)		Serra Negra (Minas Gerais)		Parque Estadual da Serra do Papagaio (Minas Gerais)		Parque Estadual Carlos Botelho (São Paulo)		Parque Estadual Marumbi (Paraná)
Area (ha)/ elevation (m) 1488,5 ha /1100-1784		-/1300-1600		10000/900-1700		22000/1600-1950		37644/30-1000		8745/~1000
Orch	86	Orch	50	Orch	66	Orch	51	Orch	75	Orch	29
Poly	27	Poly	23	Brom	29	Poly	23	Brom	43	Brom	23
Brom	25	Brom	19	Poly	29	Brom	10	Poly	23	Poly	20
Hyme	14	Hyme	8	Pipe	10	Pipe	10	Arac	19	Hyme	12
Aspl	11	Pipe	8	Dryo	9	Aspl	5	Hyme	16	Arac	5
Dryo	11	Arac	5	Hyme	6	Hyme	4	Cact	15	Aspl	5
Pipe	10	Aspl	4	Arac	3	Lyco	4	Pipe	13	Cact	5
Arac	9	Cact	4	Gesn	3	Dryo	3	Dryo	12	Gesn	5
Other families	31	Other families	33	Other families	29	Other families	28	Other families	44	Other families	23
Total	224	Total	154	Total	184	Total	138	Total	260	Total	127
Present study		Alves & Menini Neto (2014)Alves FE & Menini Neto L (2014) Vascular epiphytes in a forest fragment of Serra da Mantiqueira and floristic relationships with Atlantic high altitude areas in Minas Gerais. Brazilian Journal of Botany 37: 187-196.		Menini Neto (2009b)Menini Neto L, Matozinhos CN, Abreu NL, Valente ASM, Antunes K, Souza FS, Viana PL & Salimena FRG (2009b) Flora vascular não-arbórea de uma floresta de grota na Serra da Mantiqueira, Zona da Mata de Minas Gerais, Brasil. Biota Neotropica 9: 149-161.; *Souza et al. (2012)Souza FS, Salino A, Viana PL & Salimena FRG (2012) Pteridófitas da Serra Negra, Minas Gerais, Brasil. Acta Botanica Brasilica 26: 378-390.; Salimena et al. (2013)*Salimena FRG, Matozinhos CN, Abreu NL, Ribeiro JH, Souza FS & Menini Neto L (2013) Flora fanerogâmica da Serra Negra, Minas Gerais, Brasil. Rodriguésia 64: 311-320.		Furtado & Menini Neto (2016)Furtado SG & Menini Neto L (2016) Vascular epiphytic flora of a high montane environment of Brazilian Atlantic Forest: composition and floristic relationships with other ombrophilous forests. Acta Botanica Brasilica 30: 422-436. DOI: 10.1590/0102-33062016abb0090 https://doi.org/10.1590/0102-33062016abb...		Breier (2005)Breier TB (2005) O epifitismo vascular em florestas do sudeste do Brasil. Tese de Doutorado. Universidade Estadual de Campinas, São Paulo. 139p.; *Lima et al. (2011)*Lima RAF, Dittrich VAO, Souza VC, Salino A, Breier TB & Aguiar OT (2011) Flora vascular do Parque Estadual Carlos Botelho, São Paulo, Brasil. Biota Neotropica 11: 173-214.		Bianchi et al. (2014)

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