Angiosperm epiphytes as conservation indicators in forest fragments: A case study from southeastern Minas Gerais, Brazil

Biodivers Conserv (2009) 18:3785 3807 DOI 10.1007/s10531-009-9679-2 ORIGINAL PAPER Angiosperm epiphytes as conservation indicators in forest fragments: A case study from southeastern Minas Gerais, Brazil Luiz Menini Neto Æ Rafaela Campostrini Forzza Æ Daniela Zappi Received: 26 September 2008 / Accepted: 18 June 2009 / Published online: 1 July 2009 Ó Springer Science+Business Media B.V. 2009 Abstract Epiphytes play an important role in the flora and ecology of the tropical regions. Most floristic studies within the state of Minas Gerais focus on terrestrial, woody plant diversity, but this is a different approach, looking at epiphytic angiosperms in three highland areas in Southeastern Minas Gerais, namely Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) and Parque Estadual de Ibitipoca (PEI). Regular collections were performed in these sites between 1999 and 2007, complemented by herbarium records for PEI, and 181 species of epiphytic angiosperms were recorded in 66 genera, within 12 families. Orchidaceae, with 89 species, was by far the largest, and the most species rich genera were Peperomia (Piperaceae) and Pleurothallis s.l. (Orchidaceae), with 12 species each. Similarity analysis has shown a closer relationship between MB and RBRG, both composed by seasonal semideciduous forest, however, Jaccard (0.163) index are low. A similarity analysis including other 21 areas of southeastern and southern Brazil revealed strong influence of sazonality, vegetation type and altitude in the composition of the epiphytic flora, and a relative independence regarding the geographic proximity of the areas sampled. From the conservation standpoint, 30 species recorded for this work appear in the red list for the state of Minas Gerais, under different conservation categories. The low similarity indices obtained between the studied areas underline the importance of the conservation of each one of the remaining forest fragments in Minas L. Menini Neto (&) Centro de Ensino Superior de Juiz de Fora, Campus Arnaldo Janssen, Luz Interior 345, Juiz de Fora, MG 36030-776, Brazil e-mail: menini_neto@hotmail.com L. Menini Neto R. C. Forzza Escola Nacional de Botânica Tropical/Jardim Botânico do Rio de Janeiro, Pacheco Leão 915, Rio de Janeiro, RJ 22460-030, Brazil e-mail: rafaela@jbrj.gov.br D. Zappi HLAA, Royal Botanic Gardens, Kew TW9 3AB, UK e-mail: d.zappi@kew.org

3786 Biodivers Conserv (2009) 18:3785 3807 Gerais, as their relative geographical proximity does not necessarily mean that their epiphytic flora is similar. Keywords Atlantic forest Montane forest Seasonal semi-deciduous forest Similarity analysis Southeastern Brazil Southern Brazil Resumo Angiospermas epífitas como indicadores para conservação em fragmentos florestais. Um estudo de caso do sudeste de Minas Gerais, Brasil. Epífitas desempenham um importante papel na flora e ecologia de regiões tropicais. A maioria dos estudos florísticos no estado de Minas Gerais têm como foco a diversidade das plantas arbóreas, sendo apresentada aqui uma abordagem diferente, visando as angiospermas epífitas de três áreas de altitude no sudeste de Minas Gerais, Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) e Parque Estadual de Ibitipoca, (PEI). Coletas regulares foram realizadas nestas localidades entre 1999 e 2007, complementadas por registros de herbário para o PEI, sendo registradas 181 espécies de angiospermas epífitas, distribuídas em 66 gêneros e 12 famílias. Orchidaceae, com 89 espécies, foi a mais rica, enquanto os gêneros com o maior número de espécies foram Peperomia (Piperaceae) e Pleurothallis s.l. (Orchidaceae), com 12 espécies cada. Análises de similaridade mostraram uma relação mais próxima entre MB e RBRG, ambas compostas por floresta estacional semidecidual, no entanto, o índice de Jaccard (0,163) foi baixo. Uma análise de similaridade incluindo outras 21 áreas do sudeste e sul do Brasil revelou forte influência da sazonalidade, tipo de vegetação e altitude na composição da flora epifítica e uma relativa independência no que tange a proximidade geográfica das áreas amostradas. Em relação à conservação, 30 espécies registradas neste trabalho aparecem citadas na lista vermelha para o estado de Minas Gerais, sob diferentes categorias de conservação. Os baixos índices de similaridade obtidos entre as áreas estudadas destacam a importância da conservação de cada um dos fragmentos florestais remanescentes em Minas Gerais, bem como sua relativa proximidade geográfica não necessariamente significa que sua flora epifítica seja similar. Palavras Chave Floresta atlântica Floresta montana Floresta estacional semidecidual Análise de similaridade Sudeste do Brasil Sul do Brasil Abbreviations AA Anthropic area c. Circa CA Campo de altitude CER Cerrado CR Campo rupestre DEF Dense evergreen forest Disp Dispersal syndrome DSF Deciduous seasonal forest EC Ecological category E.E. Ecological Station GF Gallery forest HeP Primary hemiepiphyte HeS Secondary hemiepiphyte HLC Characteristic holoepiphyte HLF Facultative holoepiphyte

Biodivers Conserv (2009) 18:3785 3807 3787 MB MG P.E. PEI P.N. PR RBRG RE ReBio RJ RS SEF SSF SP Mata do Baú Minas Gerais State State Park Parque Estadual de Ibitipoca National Park Paraná State Reserva Biológica da Represa do Grama Restinga or coastal forest Biological Reserve Rio de Janeiro State Rio Grande do Sul State Subtropical evergreen forest Semideciduous Seasonal Forest São Paulo State Introduction Brazil is one of the leading countries in terms of the size of its flora (Giulietti et al. 2005) and the state of Minas Gerais is a major contributor towards Brazilian biodiversity (Mendonça and Lins 2000). Knowledge regarding this state s flora is proportionately low, and floristic surveys have tended to concentrate on woody plants (Oliveira-Filho and Fontes 2000; França and Stehmann 2004), while herbaceous species have been comprehensively surveyed only for highland areas with open vegetation, known as campo rupestre (Giulietti et al. 1987; Peron 1989; Pedralli et al. 1997; Pirani et al. 2003; Viana and Lombardi 2007; Alves and Kolbek 2009). Epiphytes represent approximately 10% of all vascular plants, and are distributed in over 80 plant families with marked richness and abundance in tropical regions, especially in the Neotropics (Gentry and Dodson 1987). Mainly herbaceous in habit, they are concentrated in the families Orchidaceae, Araceae and Bromeliaceae amongst Angiosperms, and Polypodiaceae, Aspleniaceae and Hymenophyllaceae amongst ferns (Kress 1986; Benzing 1990). In terms of species richness, around 80% of epiphytic species occur in Orchidaceae, Bromeliaceae, Polypodiaceae and Araceae (Gentry and Dodson 1987). The epiphytic community supplies the local fauna with diverse resources, such as fruits, nectar, pollen and water, and maintains fundamental specialized microhabitats for the establishment and survival of other organisms (Benzing 1990; Ingram and Nadkarni 1993). Only two previous studies involving epiphytes in Minas Gerais exist, both focused on epiphytic communities in campo rupestre vegetation (Werneck and Espírito-Santo 2002; Alves et al. 2008). Most previous research on this topic has focused on the southern states of Brazil (Waechter 1998; Dittrich et al. 1999; Kersten and Silva 2002; Borgo and Silva 2003; Gonçalves and Waechter 2003; Rogalski and Zanin 2003; Giongo and Waechter 2004; Gaiotto and Acra 2005; Kersten and Kuniyoshi 2009), with only a few in southeastern Brazil: São Paulo (Dislich and Mantovani 1998; Piliackas et al. 2000) and Rio de Janeiro (Fontoura et al. 1997). The three studied areas, Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) and Parque Estadual de Ibitipoca (PEI), feature in the atlas of priority areas for conservation in Minas Gerais (Drummond et al. 2005) with very high (MB and RBRG)

3788 Biodivers Conserv (2009) 18:3785 3807 and special (PEI) status, these being the highest levels used to categorize the importance of regional flora in terms of the conservation of its species richness. The localities are found within Brazilian biodiversity hotspots, the Cerrado (MB) and the Atlantic forest (RBRG and PEI), where high biodiversity and endemism are paired with accelerated habitat loss (Myers et al. 2000), supporting the need to investigate their biological diversity as a whole and to promote their conservation. This qualitative survey of epiphytic angiosperms in three areas of montane and upper montane forest in Minas Gerais uses the epiphytic angiosperm flora to compare the similarity between the studied areas and 21 others in Southeastern and Southern Brazil, contributing towards the knowledge of the Flora of Minas Gerais and its conservation. Study areas The three study areas (Mata do Baú MB, Reserva Biológica da Represa do Grama RBRG and Parque Estadual de Ibitipoca PEI) are located in southeastern Minas Gerais, relatively close to each other (between 70 and 110 km, within an area of c. 2,900 km 2 ) (Fig. 1). The climate is classified as Cwb, mesothermic with clearly defined seasons (CETEC 1983). Mata do Baú lies in the seasonal savanna ecosystem known as Cerrado (Menini Neto et al. 2004b), while RBRG and PEI are part of Brazil s Atlantic forest (Menini Neto et al. 2004a, 2007). Mata do Baú (MB) this private reserve in the Municipality of Barroso, in the district of Campo das Vertentes (c. 21 11 0 S and 43 56 0 W), covers approximately 10 ha of seasonal semi-deciduous montane and gallery forests, at c. 900 m alt., with mean temperature of approx. 17 C and rainfall around 1,300 mm/year (Menini Neto et al. 2004a). Fig. 1 Location of the three study areas (1 3) and further 21 sites

Biodivers Conserv (2009) 18:3785 3807 3789 Reserva Biológica da Represa do Grama (RBRG) within the Zona da Mata ( forest zone ) of Minas Gerais, in the Municipality of Descoberto (c. 21 25 0 S 42 56 0 W), represents 263.8 ha of seasonal semi-deciduous montane and gallery forests, at c. 750 m alt., with mean temperature of approx. 21 C and rainfall around 1,600 mm/year (Menini Neto et al. 2004b). Parque Estadual de Ibitipoca (PEI) this State Park, also within the Zona da Mata of Minas Gerais, although considered as part of the highland area known as Serra da Mantiqueira, lies between the municipalities of Lima Duarte and Santa Rita de Ibitipoca (c. 21 40 0 21 44 0 S and 43 52 0 43 55 0 W), comprising 1923.5 ha. Altitude ranges between 1,200 and 1,784 m. It supports a complex vegetation mosaic known as campo rupestre, including mostly open highland vegetation on quartzitic/arenitic rock, but also a varied combination of gallery forest along rivers, evergreen upper montane forest and seasonal montane forest groves ( capões de mata ), between 1,200 and 1,400 m, and cloud forest above 1,500 m. The mean temperature is approx. 18.9 C and rainfall is around 2,200 mm/ year (Menini Neto et al. 2007). The most important forested areas within PEI are two fragments of evergreen upper montane forest covering 94 and 30 ha (Fontes 1997). Methods Monthly plant collections were carried out in RBRG (between August 1999 and August 2002), MB (between February 2001 and June 2003) and PEI (between October 2003 and December 2007), targeting fertile specimens of all flowering plants in selective walks in the three areas. Data were collected in the field, and herbarium specimens were databased and deposited at CESJ and RB herbaria. Former collections from PEI, dating back to the 1970s, from herbaria BHCB, CESJ, HB, RB and VIC, were also included in the database. Herbarium acronyms are according Holmgren et al. (1990). Threatened species cited in the Minas Gerais Red List (Biodiversitas 2007) were highlighted and symbols were added for different threat categories (Table 1). The ecological categories were classified according to Benzing (1990) as characteristic holoepiphyte (HLC), facultative holoepiphyte (HLF) and hemiepiphyte (divided in primary hemiepiphyte HeP and secondary hemiepiphyte HeS). The dispersal syndromes were classified in two categories, anemochory (wind dispersal) and zoochory (animal dispersal) (Table 1). Two similarity analyses (using Jaccard s coefficient with UPGMA to cluster similarities) were carried out with PAST v. 1.89 software (Hammer et al. 2001). The cophenetic values were calculated to test strength of the fit between the clusters in the dendrograms and the similarity index matrix, generating 1,000 permutations in order to stabilize the data matrix. The first analysis used a similarity matrix combining the three species lists from this survey with those from 21 other studies (Fig. 1), focusing on epiphytes or including/ highlighting epiphytic and hemiepiphytic species among their data (Table 2). A total of 24 floristic lists and 618 species were included in this analysis. The second analysis used the same 24 floristic lists but excluded the rare species (species with only one occurrence) in a matrix with 301 species. Amongst these, the lists from Parque Nacional do Itatiaia and Parque Nacional da Serra dos Órgãos (both in Rio de Janeiro state) were complemented with database data (RB database, available at www.jbrj.gov.br).

3790 Biodivers Conserv (2009) 18:3785 3807 Table 1 Angiosperm species list of epiphytes and hemiepiphytes recorded at Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) and Parque Estadual de Ibitipoca (PEI), Minas Gerais, Brazil Species Area Habitat EC Disp Voucher specimens MB RBRG PEI ARACEAE Anthurium boudetii Nadruz X GF HeS Zo Temponi 397 (@) A. contum Schott. X X SSF/GF HeS Zo Forzza 1750 (?), Temponi 400 (@) A. leonii E.G.Gonç. X GF HLF Zo Forzza 4270 (@) A. minarum Sakuragui & X GF HeS Zo Temponi 390 (@) Mayo A. pentaphyllum (Aubl.) G. X SSF HeS Zo Forzza 1729 (?) Don A. scandens (Aubl.) Engl. X X SSF/GF HLC Zo Almeida 14 (?), Forzza 2663 (@) A. solitarium Schott X SSF HLF Zo Almeida 9 (?) Anthurium sp. X SSF HLC Zo Castro 539 (?) Heteropsis salicifolia Kunth X SSF HeS Zo Forzza 1697 (?) Monstera adansonii Schott X SSF HeS Zo Faria s.n. (CESJ 31035) Philodendron appendiculatum Nadruz & Mayo X X SSF/DEF/GF HeS Zo Almeida 13 (?), Forzza 3638 (@) P. bipinnatifidum Schott X GF HeS Zo Temponi 410 (@) P. cordatum Kunth ex Schott X SSF HeS Zo Assis 418 (?) P. curvilobum Schott X SSF HeS Zo Forzza 1698 (?) P. hastatum K.Koch & Sellow? X SSF HeS Zo Forzza 1699 (?) P. minarum Engl. X GF HeS Zo Forzza 2653 (@) P. ornatum Schott X SSF HeS Zo Almeida 32 (?) P. propinquum Schott X X X SSF/DEF/ GF HeS Zo Forzza 1935 (?), Sakuragui 1638 (@), Temponi 398 (@) P. speciosum Schott ex X SSF HeS Zo Almeida 18 (?) Endl. Rhodospatha latifolia Poepp. X SSF HeS Zo Almeida 17 (?) BEGONIACEAE Begonia angulata Vell. X DEF HeS An Temponi 393 (@) B. fruticosa A. DC. X SSF HeS An Faria s.n. (CESJ 34482) Begonia sp. X DEF HLF An Forzza 4287 (@) BROMELIACEAE Aechmea bromeliifolia X SSF HLC Zo Assis 180 (?) (Rudge) Baker A. nudicaulis (L.) Griseb. X SSF HLF Zo Monteiro 13 (@) A. ramosa Mart. ex Schult. F. X SSF HLF Zo Almeida 5 (?) Billbergia alfonsijoanis Reitz X SSF HLC Zo Martinelli 15283 (@) B. euphemiae E. Morren X SSF HLC Zo Almeida 11 (?) B. tweedieana Baker? X SSF HLC Zo Almeida 21 (?)

Biodivers Conserv (2009) 18:3785 3807 3791 Table 1 continued Species Area Habitat EC Disp Voucher specimens MB RBRG PEI B. zebrina (Herb.) Lindl. X X SSF HLC Zo Assis 19 (?), Forzza 2052 (?) Neoregelia farinosa (Ule) L.B.Sm. X SSF HLC Zo Almeida 22 (?) N. ibitipocensis (Leme) Leme X? HLF Zo Forzza 3338 @ N. lymaniana R. Braga & Sucre X? HLC Zo Leme 1478 (#) N. oligantha L.B.Sm. X GF HLF Zo Monteiro 38 (@) Nidularium ferdinandocoburgii Wawra X GF HLF Zo Martinelli 15290 (@) N. longiflorum Ule? X SSF HLC Zo Almeida 27 (?) N. marigoi Leme? X GF HLF Zo Forzza 3232 @ Portea petropolitana (Wawra) Mez X SSF HLC An Castro 563 (?) Quesnelia arvensis (Vell.) X SSF HLF Zo Almeida 8 (?) Mez Q. augusto-coburgii Wawra X SSF HLF Zo Almeida 20 (?) Racinaea aerisincola (Mez) Spencer & L.B.Sm. X GF HLC An Sucre 7147 (@) Tillandsia gardneri (Vell.) Mez X X SSF HLF An Assis 118 (?), Sucre 7184 (@) T. geminiflora Brongn. X X X SSF/GF HLC An Assis 20 (?), Forzza 1695 (?), Krieger s.n. (CESJ 8595) T. tenuifolia L. X X SSF/DEF/GF HLF An Assis 11 (?), Andrade 1131 (*) T. recurvata (L.) L. X X SSF HLC An Assis 28 (?), Sucre 7234 (@) T. stricta Sol. ex Sims X X SSF/GF/CR HLC An Faria s.n. (CESJ 34138), Câmara s.n. (CESJ 9405) T. usneoides (L.) L. X X X SSF HLC An Assis 397 (?), Almeida 34 (?) Forzza 3022 (@) Vriesea bituminosa L.B.Sm. X SSF/GF/DEF HLF An Monteiro 28 (@) V. carinata Wawra X? HLC An Paula s.n. (VIC 26470) V. friburgensis Mez X SSF/GF/CR HLF An Martinelli 15281 (@) V. gigantea Mart. ex Schult. X SSF HLC An Castro 463 (?) F. V. gradata (Baker) Mez X SSF HLC An Castro 146 (?) V. guttata Linden & André X GF HLC An Monteiro 25 (@) V. heterostachys (Baker) X DEF/GF HLF An Martinelli 15301 (@) L.B.Sm. V. longicaulis (Baker) Mez X DEF/GF HLF An Martinelli 15314 (@) V. pauperrima E. Pereira X SSF HLC An Forzza 2205 (?)

3792 Biodivers Conserv (2009) 18:3785 3807 Table 1 continued Species Area Habitat EC Disp Voucher specimens MB RBRG PEI V. penduliflora L.B.Sm.? X GF/CR HLF An Leme 1476 (#) V. scalaris E. Morren X SSF HLC An Forzza 2188 (?) Wittrockia gigantea (Baker) X DEF/GF HLC Zo Martinelli 15313 (@) Leme CACTACEAE Epiphyllum phyllanthus (L.) Haw. X SSF HLC Zo Forzza 2957 (?) Hatiora salicornioides (Haw.) Britton & Rose Hylocereus setaceus (Salm- Dyck) Ralf Bauer Lepismium cruciforme (Vell.) Miq. L. houlletianum (Lem.) Barthlott Rhipsalis elliptica G. Lindb. ex K. Schum. R. floccosa Salm-Dyck ex Pfeiff. R. juengeri Barthlott & N.P. Taylor X DEF/CR HLF Zo Brügger s.n. (CESJ 21541) X X SSF HLF Zo Assis 300 (?); Forzza 2062 (?) X SSF HLC Zo Forzza 2044 (?) X? HLC Zo Oliveira s.n. (CESJ 24194) X SSF HLC Zo Forzza 2057 (?) X GF/CR HLC Zo Krieger s.n. (CESJ 8589) X DEF/GF HLC Zo Krieger s.n. (CESJ 8594) R. lindbergiana K. Schum. X SSF HLC Zo Forzza 2920 (?) R. pulchra Loefgr. X GF HLC Zo Krieger s.n. (9269) R. russellii Britton & Rose? X GF HLC Zo Forzza 3226 (@) Schlumbergera opuntioides (Loefgr. & Dusén) D.R. Hunt? X GF HLF Zo Forzza 3180 (@) CYCLANTHACEAE Asplundia brachypus (Drude) X SSF HeS Zo Forzza 1743 (?) Harling GESNERIACEAE Nematanthus crassifolius (Schott) Wiehler? X X SSF/DEF HLC Zo Forzza 1738 (?), Forzza 4274 (@) N. lanceolatus (Poir.) Chautems X SSF HLF Zo Castro 67 (?) Sinningia magnifica (Otto & X GF HLF An Forzza s.n. (CESJ 27323) A.Dietr.) Wiehler GRISELINIACEAE Griselinia ruscifolia (Goy) X? HeP Zo Krieger s.n. (14340) Taub. MARCGRAVIACEAE Marcgravia polyantha Delp. X SSF HeS Zo Castro 100 (?) Marcgravia sp. X SSF HeS Zo Franco 64 (?) MELASTOMATACEAE Clidemia blepharodes DC. X SSF HeP Zo Forzza 1721 (?)

Biodivers Conserv (2009) 18:3785 3807 3793 Table 1 continued Species Area Habitat EC Disp Voucher specimens MB RBRG PEI ORCHIDACEAE Bifrenaria vitellina Lindl. X GF HLC An Forzza 8 (?) Bulbophyllum cribbianum Toscano X CR HLF An Forzza 10 (?) B. glutinosum (Barb. Rodr.) X SSF HLC An Menini Neto 125 (?) Cogn. B. granulosum Barb. Rodr. X GF HLC An Menini Neto 107 (?) B. pabstii Garay X SSF HLC An Assis 788 (?) B. regnellii Rchb. f. X SSF HLC An Menini Neto 124 (?) Campylocentrum aromaticum Barb. Rodr. X SSF HLC An Assis 350 (?) C. linearifolium Schltr. ex Mansf. X X SSF HLC An Assis 470 (?), Assis 878 (?) C. micranthum (Lindl.) Rolfe X SSF HLC An Assis 815 (?) C. neglectum (Rchb. f. & X GF HLC An Menini Neto 28 (?) Warm.) Cogn. C. robustum Cogn. X X SSF HLC An Assis 257 (?), Menini Neto 193 (?) Capanemia thereziae Barb. Rodr. X SSF HLC An Assis 819 (?) Catasetum cernuum (Lindl.) Rchb. f. X SSF HLF An Almeida 23 (?) Cattleya bicolor Lindl.? X SSF HLC An Menini Neto 178 (?) C. loddigesii Lindl. X X SSF HLC An Assis 269 (?), sem coletor (CESJ 27534) Centroglossa macroceras Barb. Rodr. X DEF HLC An Forzza 54 (?) Comparettia coccinea Lindl. X X SSF HLC An Assis 469 (?), Faria s.n. (CESJ 31100) Dichaea cogniauxiana Schltr. X GF HLC An Forzza 21 (?) Encyclia patens Hook. X X X SSF/DEF HLF An Assis 691 (?), Salimena s.n. (CESJ 31246), Menini Neto 165 (?) Epidendrum armeniacum X SSF HLC An Menini Neto 175 (?) Lindl. E. chlorinum Barb. Rodr. X GF HLC An Menini Neto 171 (?) E. cristatum Ruiz & Pav. X SSF HLF An Assis 839 (?) E. densiflorum Hook. X X SSF HLF An Assis 849 (?), Menini Neto 2 (?) E. difforme Lindl. X SSF HLC An Menini Neto 97 (?) E. ochrochlorum Barb. Rodr. X DEF HLC An Forzza 36 (?) E. paranaense Barb. Rodr. X DEF/GF HLF An Forzza 83 (?) E. rigidum Jacq. X X SSF/GF HLF An Assis 847 (?), Menini Neto 71 (?)

3794 Biodivers Conserv (2009) 18:3785 3807 Table 1 continued Species Area Habitat EC Disp Voucher specimens Eurystyles actinosophila (Barb. Rodr.) Schltr. E. cogniauxii (Kraenzl.) Pabst MB RBRG PEI X X SSF HLC An Assis 808 (?), Castro 125 (?) X GF HLC An Forzza 61 (?) Gomesa glaziovii Cogn. X GF HLC An Forzza 51 (?) G. recurva Lodd. X X X SSF/DEF HLF An Menini Neto 17 (?), Faria s.n. (CESJ 31067), Forzza 38 (?) G. sessilis Barb. Rodr. X SSF HLC An Menini Neto 7 (?) Grobya amherstiae Lindl. X SSF HLC An Forzza 26 (?) Hadrolaelia coccinea (Lindl.) Chiron & V.P. Castro X SSF/CR HLF An Forzza 17 (?) Huntleya meleagris Lindl. X SSF HLC An Almeida 26 (?) Isabelia violacea (Lindl.) Van den Berg & M.W.Chase X CR HLF An Salimena-Pires s.n. (CESJ 24458) I. virginalis Barb. Rodr. X SSF HLC An Menini Neto 47 (?) Isochilus linearis (Jacq.) R. Br. Lankesterella gnomus (Kraenzl.) Hoehne X X SSF HLC An Assis 647 (?), Forzza 40 (?) X GF HLC An Menini Neto 139 (?) Masdevallia infracta Lindl. X DEF HLC An Menini Neto 173 (?) Maxillaria acicularis Herb. ex Lindl. X SSF HLC An Forzza 35 (?) M. madida Lindl. X CR HLF An Forzza 47 (?) M. notylioglossa Rchb. f. X SSF HLC An Menini Neto 119 (?) M. ochroleuca Lodd. ex Lindl. X SSF HLC An Forzza 71 (?) M. valenzuelana (A. Rich.) Nash X SSF HLF An Assis 848 (?) Notylia aff. sagittifera (Kunth) Link, Klotzsch & Otto X SSF HLC An Menini Neto 16 (?) Octomeria alpina Barb. Rodr. X CR/GF HLF An Forzza 77 (?) O. aff. diaphana Lindl. X DEF HLC An Forzza 89 (?) O. grandiflora Lindl. X GF HLC An Forzza 63 (?) O. aff. rubrifolia Barb. Rodr. X SSF/CR HLC An Sousa s.n. (BHCB 14766) O. wawrae Rchb. f. ex Wawra X SSF/CR HLC An Forzza 41 (?) Oncidium ciliatum Lindl. X SSF HLC An Assis 31 (?) O. divaricatum Lindl. X DEF HLC An Forzza 2190 (@) O. gardneri Lindl. X SSF HLC An Assis 455 (?) O. gravesianum Rolfe X SSF/DEF HLC An Forzza 23 (?) O. hookeri Rolfe X SSF/DEF HLC An Forzza 39 (?)

Biodivers Conserv (2009) 18:3785 3807 3795 Table 1 continued Species Area Habitat EC Disp Voucher specimens MB RBRG PEI O. longipes Lindl. X SSF HLC An Menini Neto 163 (?) O. pubes Lindl. X SSF HLC An Assis 266 (?) O. pumilum Lindl. X SSF HLC An Assis 353 (?) O. truncatum Pabst X SSF HLC An Menini Neto 95 (?) Pleurothallis cryptophoranthoides Loefgr. X DEF HLC An Sousa s.n. (BHCB 16647) P. heterophylla (Barb. X SSF HLC An Andrade 1160 (*) Rodr.) Cogn. P. hypnicola Lindl. X X SSF/DEF HLC An Forzza 2187 (?), Menini Neto 134 (?) P. luteola Lindl. X DEF HLC An Menini Neto 158 (?) P. malachantha Rchb. f.? X SSF/DEF HLC An Sousa s.n. (BHCB 9832) P. marginalis Rchb. f. X SSF HLC An Menini Neto 162 (?) P. pubescens Lindl. X SSF HLC An Menini Neto 18 (?) P. quartzicola (Barb. Rodr.) X DEF HLC An Menini Neto 114 (?) Cogn. P. recurva Lindl. X DEF HLC An Menini Neto 236 (?) P. rubens Lindl. X SSF/DEF/CR HLF An Forzza 78 (?) P. saundersiana Rchb. f. X SSF HLC An Forzza 32 (?) P. tricarinata Poepp. & Endl. X DEF HLC An Menini Neto 118 (?) Polystachya estrellensis Rchb. f. X X SSF HLF An Assis 188 (?), Almeida 6 (?) P. hoehneana Kraenzl.? X SSF HLC An Forzza 37 (?) P. micrantha Schltr.? X SSF HLC An Assis 193 (?) Promenaea xanthina Lindl. X DEF HLC An Eiterer s.n. (CESJ 25549) Prosthechea allemanoides X SSF/CR HLF An Forzza 55 (?) (Hoehne) W.E.Higgins P. vespa (Vell.) W.E.Higgins X SSF/CR HLF An Forzza 67 (?) Rodrigueziella gomezoides (Barb. Rodr.) Berman X GF HLC An Sousa s.n. (BHCB 16117) Scaphyglottis modesta Schltr. X SSF HLF An Forzza 4 (?) Scuticaria aff. kautskyi Pabst X GF HLC An Forzza 15 (?) Stelis aprica Lindl. X GF HLC An Menini Neto 127 (?) S. intermedia Poepp. & Endl. X DEF HLC An Menini Neto 159 (?) S. megantha Barb. Rodr. X DEF HLC An Forzza 60 (?) S. papaquerensis Rchb. f. X GF HLC An Sousa s.n. (BHCB 14764) S. parvula Lindl. X GF HLC An Krieger s.n. (CESJ 8593) Thysanoglossa organensis Brade X SSF HLC An Menini Neto 89 (?) Vanilla cf. gardneri Rolfe X SSF HeS An Forzza 2115 (?) Xylobium variegatum (Ruiz X SSF HLF An Almeida 24 (?) & Pav.) Garay & Dunst. PIPERACEAE

3796 Biodivers Conserv (2009) 18:3785 3807 Table 1 continued Species Area Habitat EC Disp Voucher specimens MB RBRG PEI Peperomia alata Ruiz & Pav. X X SSF HLF Zo Assis 105 (?), Castro 55 (?) P. blanda (Jacq.) Kunth X SSF HLF Zo Forzza 2101 (?) P. corcovadensis Gardner X SSF HLF Zo Assis 265 (?) P. decora Dahlst. X HLF Zo Krieger s.n. (CESJ 13193) P. mandioccana Miq. X SSF/DEF HLF Zo Krieger s.n. (CESJ 8578) P. martiana Miq. X SSF HLF Zo Assis 104 (?) P. polystachyoides Dahlst. X SSF HLF Zo Assis 456 (?) P. quadrifolia (L.) Kunth X SSF HLF Zo Assis 103 (?) P. stenocarpa Regel X SSF HLF Zo Castro 498 (?) P. subpilosa Yunck. X SSF HLF Zo Forzza 2099 (?) P. tetraphylla (Forst.) Hook. X SSF/DEF HLF Zo Krieger s.n. (CESJ 8523) P. urocarpa Fisch. & C.A. X X SSF HLF Zo Assis 593 (?) Mey RUBIACEAE Hillia parasitica Jacq. X DEF HeP An Pereira s.n. (CESJ 28425) EC ecological category (Benzing 1990), HLC characteristic holoepiphyte, HLF facultative holoepiphyte, HeP primary hemiepiphyte, HeS secondary hemiepiphyte, Disp dispersal syndromes, An Anemochory, Zo Zoochory. Voucher specimens are found at BHCB (*), CESJ (?), HB (#), RB (@) and VIC ( ) herbaria. Habitat: SSF seasonal semideciduous forest (at PEI, these are the groves at 1,200 m elev.), DEF dense evergreen forest, GF gallery forest or forests near depressions or grutas (highlands of PEI, above 1,500 m elev.), CR campo rupestre Species in bold are cited in the Minas Gerais Red List (Biodiversitas 2007) as: Critically Endangered ( ); Endangered ( ); Vulnerable (?); Near Threatened ( ); Data Deficient ( ) The map was prepared using Arc View GIS 3.2 Ò on the Americas Base Map (Bletter et al. 2004). Results One hundred and eighty-one species of epiphytes (MB 41 spp., RBRG 59 spp. and PEI 113 spp.), representing 65 genera and 12 families of epiphytes were recorded. Most species (102, 56.3%) were characteristic holoepiphytes, while 54 (29.8%) were facultative holoepiphytes and 25 (13.8%) were hemiepiphytes (3 primary hemiepiphytes and 22 secondary hemipiphytes). Of these species, 113 showed dispersion by anemochory while 68 had evidence of zoochory (Table 1). A comparison between the number of epiphytic species found and the total angiosperm species listed for each area shows that the percentage of the flora represented by epiphytes is around 9 11%, being slightly lower in MB, the area influenced by Cerrado vegetation, which is drier than other two areas. Araceae, Bromeliaceae, Cactaceae, Orchidaceae and Piperaceae are the only flowering plant families common to all three areas. Exclusive occurrences at family level were

Biodivers Conserv (2009) 18:3785 3807 3797 Table 2 Lists used in floristic similarity analysis Acronym Locality Altitude N spp Area (ha) Coordinates Vegetation types Reference AC Estreito Augusto César, Rio 650 63 27 24 0 S 51 27 0 W DSF Rogalski and Zanin (2003) Uruguai (RS) Ber Bertioga (SP) 0 96 23 45 0 S 45 58 0 W RE Martins et al. (2008) Ctba Fragmentos Curitiba (PR) 900 80 96,9 25 25 0 S 49 16 0 W MEF Borgo and Silva (2003) CUASO Reserva da Cidade Universitária Armando Salles de Oliveira (SP) DC Estação Experimental Agronômica (RS) 735 770 29 10 23 33 0 S 46 43 0 W SSF, DEF Dislich and Mantovani (1998) Groppo and Pirani (2005) 40 40 30 04 0 S 51 40 0 W GF Giongo and Waechter (2004) FGA Fazenda Gralha Azul (PR) 870 920 30 475 25 37 0 S 49 17 0 W MEF Gaiotto and Acra (2005) Fig Figueiras isoladas planície 20 70 29 35 0 S 50 04 0 W AA Gonçalves and Waechter (2003) costeira (RS) IM Ilha do Mel (PR) 0 61 2,760 25 30 0 S 48 23 0 W DEF, RE Kersten and Silva (2001) Jur-Ita E. E. Juréia-Itatins (SP) 0 1,400 96 79,830* 24 18 0 S 47 30 0 W DEF, RE Mamede et al. (2004) MB Mata do Baú (MG) 900 41 10 21 11 0 S 43 56 0 W SSF, GF Present work NC Núcleo Curucutu, P. E. da 750 850 45 12,090 23 59 0 S 46 44 0 W DEF, CA Garcia and Pirani (2005) Serra do Mar (SP) PEC P. E. do Cerrado de 750 900 16 426,62 24 09 0 S 50 18 0 W SSF, GF, CER Linsingen et al. (2006) Jaguariaíva (PR) PEI-MG P. E. Ibitipoca (MG) 1,200 1,784 113 1923,5 21 40 0 S 43 52 0 W SSF, GF, DEF, CR Present work PEI-RS P.E. Itapuã (RS) 50 265 48 5,566 30 24 0 S 50 58 0 W RE Musskopf (2006) PEI-SP P. E. Intervales, Base 70 250 55 48,000* 24 14 0 S 48 04 0 W SSF Ziparro et al. (2005) Saibadela (SP) PESB P. E. da Serra do Brigadeiro 1,000 2,000 111 13,210 20 43 0 S 42 29 0 W SSF, CA Leoni and Tinte (2004) (MG) PI Parque do Ingá (PR) 550 25 47,3 23 25 0 S 51 25 0 W SSF Dettke et al. (2008) PNI P. N. do Itatiaia (RJ) 500 2,789 106 30,000* 22 30 0 S 44 35 0 W DEF, CA JABOT** PNSO P. N. Serra dos Órgãos (RJ) 1,200 2,263 87 10,653* 22 26 0 S 43 06 0 W DEF, CA JABOT**

3798 Biodivers Conserv (2009) 18:3785 3807 Table 2 continued Acronym Locality Altitude N spp Area (ha) Coordinates Vegetation types Reference RB Rio Barigüi (PR) 900 33 8,6 25 34 0 S 49 20 0 W MEF Kersten and Silva (2002) RBRG ReBio da Represa do Grama (MG) 750 59 263,8 21 25 0 S 42 56 0 W SSF, GF Present work RI Bacia do Rio Iguaçu (PR) 865 950 83 53,9 25 40 0 S 54 26 0 W MEF Kersten and Kuniyoshi (2009) SSJ Serra de São José (MG) 900 1,430 41 21 05 0 S 44 09 0 W SSF, GF, CR, CER Alves and Kolbek (2009) Tor Torres (RS) 0 103 29 18 0 S 49 41 0 W DEF Waechter (1986) Number of species refers only to epiphytic Angiosperms, excluding fern species. ReBio Biological Reserve, P.E. State Park, P.N. National Park, E.E. Ecological Station, SSF semideciduous seasonal forest, DSF deciduous seasonal forest, DEF dense evergreen forest, MEF mixed evergreen forest, RE restinga or coastal forest, GF gallery forest, CR campo rupestre, CA campo de altitude, CER Cerrado, AA anthropic area Citations signalled with * refer to the whole conservation area as the authors did not specify the area sampled. JABOT ** is the database of the Jardim Botânico do Rio de Janeiro (RB), available at www.jbrj.gov.br

Biodivers Conserv (2009) 18:3785 3807 3799 Fig. 2 Species richness of the five most representative families in the three study areas: Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) and Parque Estadual de Ibitipoca (PEI) Minas Gerais, Brazil recorded for RBRG (Cyclanthaceae 1 sp., Marcgraviaceae 2 spp., Melastomataceae 1 sp.) and PEI (Griseliniaceae 1 spp., Rubiaceae 1 spp.). Highest species richness was found in Orchidaceae (89 spp.), Bromeliaceae (36 spp.), Araceae (20 spp.), Cactaceae and Piperaceae (12 spp. each), these families representing 93% of the total species recorded. However, such families occupied different positions within the three areas (Fig. 2). While Orchidaceae was the largest family at MB and PEI (24 and 66 spp. respectively), followed by Bromeliaceae (7 and 22 spp.), Piperaceae at MB (6 spp.) and Araceae at PEI (9 spp.), it appears only in third position at RBRG (13 sp.), after Bromeliaceae (16 spp.) and Araceae (14 spp.). Of the 65 genera found, only 12 were common to all three areas, while 25 were exclusive to PEI, 11 were found only at RBRG and two were exclusive to MB. Of the 181 species recorded, approximately 58% belong to 13 genera (i.e. less than one third of all recorded genera), of which eight were Orchidaceae. The most diverse genera were Peperomia and Pleurothallis s.l. (12 spp.), followed by Epidendrum, Oncidium and Philodendron (9 spp.), Anthurium (8 spp.), Rhipsalis and Tillandsia (6 spp.), Bulbophyllum, Campylocentrum, Maxillaria, Octomeria and Stelis (5 spp.). Amongst the top five species-rich genera, Tillandsia was the only one common to all areas. The floristic analysis of similarity between the three study areas, calculated using Jaccard s coefficient (MB/RBRG = 0.163, MB/PEI = 0.085 and RBRG/PEI = 0.067), and also represented in the Venn diagram (Fig. 3), shows three rather distinct floras. A small overlap exists between the epiphytic floras of the areas, with only five species common to all three: Encyclia patens, Gomesa recurva Philodendron propinquum, Tillandsia geminiflora and T. usneoides. Even in the two areas with more similar vegetation formations, RBRG and MB, the overlap covers only nine species: Billbergia zebrina, Hylocereus setaceus, Campylocentrum linearifolium, Comparettia coccinea, Epidendrum densiflorum, Eurystyles actinosophila, Polystachya estrellensis, Peperomia alata and P. urocarpa: all commonly found in seasonal semi-deciduous forests throughout South America. Results of the similarity analyses performed between the three study areas and another 21 surveys from the states of Minas Gerais, São Paulo, Rio de Janeiro, Paraná and Rio Grande do Sul show slighthly different results in the dendrograms (Figs. 4, 5).

3800 Biodivers Conserv (2009) 18:3785 3807 Fig. 3 Venn diagram showing exclusive and shared species between the three study areas: Mata do Baú (MB), Reserva Biológica da Represa do Grama (RBRG) and Parque Estadual de Ibitipoca (PEI) Minas Gerais, Brazil The similarity indices presented in the analyses are relatively low. The dendrogram generated in the first analysis (all species included) obtained a cophenetic value of 0.8339 (Fig. 4). Two major groups, A and B, can be recognized in both dendrograms. Group A includes primarily areas of Southeastern Brazil (Minas Gerais, Rio de Janeiro and São Paulo), including PEI, while group B shows a predominance of Southern Brazilian areas (Paraná and Rio Grande do Sul), but includes the study areas MB and RBRG. In the second analysis (excluding rare species, Fig. 5), a reconfiguration of the subgroups is seen, and all studied groups are included within group A, the Jaccard s coefficient is slightly higher than before, with only one subgroup at more than 0.5 (Curitiba and Rio Barigui). The cophenetic value is 0.8289. Discussion The 12 angiosperm families registered in this study correspond to approx. 30% of those with epiphytes in the Neotropics, according to the estimates provided by Gentry and Dodson (1987). The families common to all studied areas, Araceae, Bromeliaceae, Cactaceae, Orchidaceae and Piperaceae, coincided with the largest proportion of epiphytic species generally found (Benzing 1990), repeating the pattern found in other epiphytic plant surveys (Ingram et al. 1996; Kersten and Silva 2002; Borgo and Silva 2003; Gonçalves and Waechter 2003; Rogalski and Zanin 2003; Giongo and Waechter 2004; Gaiotto and Acra 2005). Regarding ecological categories, the characteristc holoepiphytes are the dominant category, almost two times higher than facultative holoepiphytes, as found in other studies (e.g., Nieder et al. 2000; Borgo and Silva 2003; Rogalski and Zanin 2003; Gaiotto and Acra 2005). However, Alves and Kolbek (2009) recorded a different proportion for campo rupestre vegetation, where they found facultative holoepiphytes to be far more numerous than characteristic holoepiphytes. This opposes what was found during the present study in PEI where campo rupestre is also predominant. Reproductive strategies are an important factor in the success of epiphytes (Gentry and Dodson 1987) and the occurrence of anemochory as the predominant dispersal syndrome among species was highlighted by Benzing (1987). The results of this study corroborate this hypothesis, as 62.4% are dispersed through anemochory and the remaining 47.6% are zoochoric species. This percentage of anemochory in species reflects the high number of orchid and bromeliads species registered in the surveys, which is also supported by our results.

Biodivers Conserv (2009) 18:3785 3807 3801 Fig. 4 Similarity tree (Jaccard s coefficient) obtained by the analysis of all species found within the floristic lists of the 24 areas The lower epiphyte proportion in MB was an expected result, because the area is drier than PEI or RBRG. Dry environments are generally poorer in epiphytic species, as highlighted by Gentry and Dodson (1987). Orchidaceae present the highest species richness in most studies of epiphytic flora in Neotropical vegetations, generally followed by Bromeliaceae (Freiberg 1996; Ingram et al. 1996; Freiberg and Freiberg 2000; Bussmann 2001; Kersten and Silva 2002; Borgo and Silva 2003; Rogalski and Zanin 2003; Giongo and Waechter 2004; Hefler and Faustioni 2004; Gaiotto and Acra 2005; Krömer et al. 2005; Alves et al. 2008; Kersten and Kuniyoshi 2009). Other flowering plant families occasionally occupy the first position, as was found by Benavides et al. (2006) in the Colombian Amazon, with Araceae. Arévalo and Betancur (2004) also obtained an unexpected result in four vegetation types in the Colombian Guayana, with two areas where Orchidaceae was the richest family, but another two areas where Araceae was the most species rich. The present study shows that Bromeliaceae and Araceae were richer than Orchidaceae in RBRG, an unusual result when compared with the studies cited above for the Neotropics. However, most of the research where Orchidaceae appears as the top epiphytic family was carried out either in evergreen lowland or cloud forests, while the present results suggest that family dominance may be more variable in seasonal forests. On the other hand, Ziparro et al. (2005) found more species of Araceae than of Orchidaceae and Bromeliaceae in lowland dense evergreen

3802 Biodivers Conserv (2009) 18:3785 3807 Fig. 5 Similarity tree (Jaccard s coefficient) obtained by the analysis of floristic lists excluding rare species of the 24 areas forest in São Paulo, but that area was the subject of specific Araceae studies which may have caused a bias in their results. The greater similarity of RBRG and PEI (8 genera in common, c. 12%), despite their different altitudes and vegetation types may reflect their position within the Atlantic forest biome, while MB is surrounded by Cerrado. The results suggest that the distance between the sites, which is greater between PEI and RBRG than MB-RBRG or PEI-MB, is not correlated to generic composition. Given that MB and RBRG share similar altitudes (900 and 750 m) and forest types (seasonal semi-deciduous and gallery forest), it was expected that they would present more commonality than was actually found (only two shared genera, c. 3%). Each study area presented one genus as its most diverse, highlighting the uniqueness of the epiphytic flora in each particular location: Peperomia, with 6 species, was the largest genus in MB, while 7 Philodendron species were recorded at RBRG and 11 species of Pleurothallis s.l. at PEI. Diverse factors may have contributed towards the species dominance of these genera in the different areas: for example, the succulence in species of Peperomia allows for wider tolerance of more marked seasonality in forests associated with the Cerrado biome, such as MB, and the fact that most species of this genus were recorded as facultative epiphytes also points towards a wider ecological spectrum, also

Biodivers Conserv (2009) 18:3785 3807 3803 supported by its pantropical distribution. On the other hand, the predominance of species of Philodendron and other Araceae at RBRG may be linked to this family s affinities with the Atlantic forest biome, particularly at lower altitudes. In the case of PEI and Pleurothallis,a genus particularly rich in species from cloud forests, it may indicate floristic links between this site and other highlands in Eastern Brazil. The position of Tillandsia must be emphasized, as it was the only genus amongst the top five species-rich genera common to all three areas of this study. Versieux and Wendt (2007) remark on the wide distribution of Tillandsia species in the Neotropics, justifying their presence in all three areas, despite the floristic, altitude and/or vegetation type discrepancies between them. Also, anemochory could help explain this wide distribution. Martinelli et al. (2008) also remark on the richness and wide distribution of Tillandsia in the Atlantic forest domain. The genus also appears amongst the richest within the epiphytic flora in other surveys in Southern Brazil (Kersten and Silva 2002; Borgo and Silva 2003; Gonçalves and Waechter 2003; Rogalski and Zanin 2003; Giongo and Waechter 2004; Gaiotto and Acra 2005; Linsingen et al. 2006). However, surveys performed within Southeastern Brazil, in Minas Gerais and São Paulo states, do not show Tillandsia amongst the most representative elements of the epiphytic flora: Leoni and Tinte (2004) cite only two species for the Parque Estadual da Serra do Brigadeiro Minas Gerais while Mamede et al. (2004) (Juréia-Itatins), Garcia and Pirani (2005) (Parque Estadual da Serra do Mar Núcleo Curucutu) and Ziparro et al. (2005) (Parque Estadual Intervales Base Saibadela), all in São Paulo, recorded only four, two and one species, respectively, thus highlighting the variation of epiphytic species composition between forests in the Southeastern and Southern regions of Brazil. The only five species common to all three areas (Encyclia patens, Gomesa recurva, Philodendron propinquum, Tillandsia geminiflora and T. usneoides) are widely distributed species occurring in a variety of habitats, therefore not contradicting the indication that each area has its own particular floristic composition. The low coefficient of similarity between the two areas with the more similar vegetation formations (seasonal semideciduous forest) can be explained by the isolation caused by the Cerrado vegetation surrounding MB, while RBRG is found within the Atlantic forest biome. The low similarity coefficients found between three areas within a total of only 2,900 km 2 highlights the need for further study of the epiphytes in the forest fragments of Minas Gerais. The cophenetic values obtained in the analyses was of 0.8339 (including all species) and 0.8289 (excluding the rare species), supporting the results seen in the dendrograms. The low Jaccard s index found indicates that the epiphytic floras in Southern and Southeastern Brazil can be considered distinct. In both analyses, Parque Estadual de Ibitipoca is more similar to Parque Nacional do Itatiaia: a result expected as both areas located in the Serra da Mantiqueira are also geographically close. The association between Parque Estadual da Serra do Brigadeiro and the Parque Nacional da Serra dos Órgãos may be explained by their similar altitudes rather than by their vegetation, as the forest in Parque Estadual da Serra do Brigadeiro is markedly more seasonal than the Parque Nacional da Serra dos Órgãos. However, all four areas are relatively close geographically and the similarity of their epiphytic flora is more likely to be influenced by species adaptation to high altitude (i.e., their tolerance to low or near freezing temperatures and the particular role played by mist in species establishment) than by their affinities in terms of vegetation associations. According to Gentry and Dodson (1987), altitude plays an important role in epiphyte richness, with the peak of epiphyte diversity in the Andes being found between 1,000 and

3804 Biodivers Conserv (2009) 18:3785 3807 2,000 m. Krömer et al. (2005) pointed at a peak of richness at c. 1,500 m and a diminution above 2,200 m in Bolivia. This reduction was also observed by Bussmann (2001) in Ecuador but the altitudinal ranges in his study are higher: 1,800 3,150 m. Gentry (1988) also found a decrease in species richness at higher altitudes (between 1,500 3,100 m) in the Andes. The data found for PEI (1,200 1,780 m) shows that the species number of epiphytes in highland forest was higher. These trends contrast with those found for woody species in the same region, generally higher in lowland forests and decreasing at higher altitudes (Oliveira-Filho and Fontes 2000). There aren t studies about the influence of altitude in epiphyte diversity in Brazil, but the numbers presented in Table 2 appear to confirm higher epiphyte richness in highaltitude environments, bearing in mind that the altitude range considered here is lower than 2,000 m, where cold nights and the occurrence of frost would certainly affect the distribution of many tropical species. The position of MB and RBRG differs between the two analyses. In the analysis that takes into account all species, both areas cluster within branch B, together with areas from southern Brazil, while in the analysis without the rare species MB and RBRG appear in branch A, together with southeastern areas that are closer geographically. This can be explained by the fact that analysis 1 has a trend to group richer floras together in branch A, and MB and RBRG, which are in drier locations and have comparatively poorer epiphytic flora, are excluded from branch A and fall into branch B. Overall, the comparison of epiphytes from 24 areas from five different states (Table 2) showed remarkably low floristic similarity, except in Paraná, where areas of mixed evergreen forest at altitudes between 865 950 m (RB, FGA, Ctba and RI) have shown similarity higher than 0.5. Nevertheless, some subgroups based on altitudinal quotas could be recognized, such as the subgroup formed by areas within São Paulo (PEI-SP, Jur-Ita, Ber, IM, all between 0 250 m except Jur-Ita, 0 1,400 m) and the subgroup from Rio Grande do Sul (Fig, PEI-RS, Tor, DC, between 0 265 m). Elevation is an important factor in composition of both rain and semideciduous forest areas of Atlantic domain, based on analysis of arboreal stratum as presented by Oliveira-Filho and Fontes (2000). Low similarity coefficients between different areas were also found by Safford (2007) when working on the phytogeography of the Southeastern Brazilian highlands, or Brazilian Páramos. Also, comparative studies developed with woody strata in montane seasonal semideciduous forests in Minas Gerais and São Paulo show relatively low similarity (Oliveira-Filho and Machado 1993; Meira-Neto and Martins 2002). More surveys focusing on non-woody plant groups are needed, as most work developed in Brazil is centered on tree studies. Different study approaches would provide a deeper insight both into the actual diversity and the biogeography of the distinct areas. Conservation The low similarity coefficients found between the epiphytic floras of the three study areas in Minas Gerais and between 24 forest fragment sites within Southeastern and Southern Brazil underline the importance of conserving all possible fragments, independent of the size of the area or biome where they occur. In comparison with woody plant surveys (Oliveira-Filho and Fontes 2000), it can also be suggested that epiphytes appear to be more species rich than trees in high altitude forests, therefore representing a better biodiversity indicator for this type of vegetation.

Biodivers Conserv (2009) 18:3785 3807 3805 In terms of species conservation, the 30 recorded epiphytic species (c. 16.5%) listed within the Minas Gerais Red List (Biodiversitas 2007; Table 1) are found in the following families: Orchidaceae (12 spp.), Bromeliaceae (8 spp.), Araceae (4 spp.), Gesneriaceae (3 spp.), Cactaceae (2 spp.) and Piperaceae (1 sp.). The breakdown in conservation status is: Critically Endangered (6 spp.); Endangered (7 spp.); Vulnerable (12 spp.); Near Threatened (2 spp.); Data Deficient (4 spp.). PEI presents the largest number of threatened species (20 spp.), followed by RBRG (9 spp.) and MB (4 spp.). Due to public awareness of epiphytic orchids, bromeliads, aroids and cacti as ornamentals, epiphytic families are strongly emphasized within the Minas Gerais Red List (Biodiversitas 2007), making this list particularly useful to benchmark the findings of this work. Acknowledgments The authors would like to thank Eve Lucas, William Milliken and Nigel Taylor (all RBG Kew) for critically reading and suggesting improvements to the manuscript, two anonymous reviewers for their valuable suggestions, and to Marco Antônio Manhães (UFJF) for his help with the statistical analysis. References Alves RJV, Kolbek J (2009) Summit vascular flora of the Serra de São José, Minas Gerais, Brasil. Check List 5(1):35 73 Alves RJV, Kolbek J, Becker J (2008) Vascular epiphyte vegetation in rocky savannas of southeastern Brazil. Nord J Bot 26:101 117 Arévalo R, Betancur J (2004) Diversidad de epífitas vasculares en cuatro bosques del sector suoriental de la Serrania de Chiribiquete, Guayana Colombiana. Caldasia 26(2):359 380 Benavides AM, Wolf JHD, Duivenvoorden JF (2006) Recovery and succession of epiphytes in upper Amazonian fallows. J Trop Ecol 22:705 717 Benzing DH (1987) Vascular epiphytism: taxonomic participation and adaptive diversity. Ann Mo Bot Gard 74(2):183 204 Benzing DH (1990) Vascular epiphytes. Cambridge University Press, New York Biodiversitas (2007) Revisão das listas das espécies da flora e da fauna ameaçadas de extinção do estado de Minas Gerais. Fundação Biodiversitas, Belo Horizonte. www.biodiversitas.org.br Bletter N, Janovec J, Brosi B, Douglas CD (2004) A digital base map for studying the Neotropical flora. Taxon 53(2):46 477 Borgo M, Silva SM (2003) Epífitos vasculares em fragmentos de Floresta Ombrófila Mista, Curitiba, Paraná, Brasil. Rev Bras Bot 26(3):391 401 Bussmann RW (2001) Epiphyte diversity in a tropical Andean forest Reserva Biológica San Francisco, Zamora-Chinchipe, Ecuador. Ecotropica 7:43 59 CETEC (1983) Diagnóstico ambiental de Minas Gerais. Fundação Centro Tecnológico de Minas Gerais, Belo Horizonte Dettke GA, Orfrini AC, Milaneze-Gutierre MA (2008) Composição florística e distribuição de epífitas vasculares em remanescente alterado de Floresta Estacional Semidecidual no Paraná, Brasil. Rodriguésia 59(4):859 872 Dislich R, Mantovani W (1998) Flora de epífitas vasculares da Reserva da Cidade Universitária Armando de Salles Oliveira (São Paulo, Brasil). Boletim de Botânica da Universidade de São Paulo 17:61 83 Dittrich VAO, Kozera C, Silva SM (1999) Levantamento florístico de epífitos vasculares no Parque Barigüi, Paraná, Brasil. Iheringia (Série Botânica) 52:11 22 Drummond GM, Martins CS, Machado ABM, Sebaio FA, Antonini Y (orgs) (2005) Biodiversidade em Minas Gerais, um atlas para sua conservação, 2nd edn. Fundação Biodiversitas, Belo Horizonte Fontes MAL (1997) Análise da composição florística das florestas nebulares do Parque Estadual de Ibitipoca, Minas Gerais. Dissertation, Universidade Federal de Lavras-MG Fontoura T, Sylvestre LS, Vaz MAS, Vieira CM (1997) Epífitas vasculares, hemiepífitas e hemiparasitas da Reserva Ecológica de Macaé de Cima. In: Lima HC, Guedes-Bruni RR (eds) Serra de Macaé de Cima: diversidade florística e conservação da Mata Atlântica. Editora do Jardim Botânico do Rio de Janeiro, Rio de Janeiro