Biota Neotropica 14(2): 1––12, 2014
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inventory
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Paula Leitman1,5, André Amorim2,3, Luiz Menini Neto4 & Rafaela C. Forzza1
1
Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão 915, Jardim Botânico, Rio de Janeiro, RJ, Brazil ,
CEP: 22460-030
2
Universidade Estadual de Santa Cruz, Departamento de Ciências Biológicas, Salobrinho, Ilhéus, BA,
Brazil, CEP: 45662900
3
Centro de Pesquisas do Cacau - Herbário CEPEC, Caixa Postal 07, Itabuna, Bahia, Brazil, CEP:
45660970
4
Centro de Ensino Superior de Juiz de Fora, Campus Arnaldo Janssen - Rua Luz Interior 345, Santa Luzia,
Juiz de Fora, MG, Brazil, CEP: 36030-776
5
Corresponding author: Paula Leitman, e-mail: paulaleitman@gmail.com
LEITMAN, P., AMORIM, A., MENINI NETO, L., FORZZA, R.C. Epiphytic angiosperms in a
–12. http://dx.doi.org/10.1590/S1676mountain forest in southern Bahia, Brazil. Biota Neotropica. 14(2): 1–
06032014001013
Abstract: The Atlantic Forest in southern Bahia State, Brazil, has high levels of diversity and vascular
plant endemism. There have been few floristic surveys of the mountain forests there, however, especially
those focusing on herbs and canopy plants, with studies of Atlantic Forest epiphytes having been limited
to the southern and southeastern region of that country. The present study therefore surveyed the
epiphytic angiosperms in the Serra das Lontras National Park (SLNP), the distribution of their species
among genera and families, and their biogeographical patterns. Fieldwork was performed between
February 2011 and March 2012. Collections were made along roads and trails and phorophytes were
occasionally climbed and fallen trees and branches examined in order to sample canopy species. The
study site demonstrated high epiphyte richness (256 spp.), one of the highest recorded in the Atlantic
Forest. The richest families are Orchidaceae, Bromeliaceae, and Araceae, reaffirming the patterns
reported in previous major reviews on epiphytes. The species exhibit high degrees of endemism (45%) to
the Atlantic Forest, especially among the Bromeliaceae and Orchidaceae. One new occurrence to Brazil,
four to the Atlantic Forest and 30 to Bahia state are registered. The SLNP shares more species with areas
of southern and southeastern Brazil than with other northeastern states.
Keywords: Atlantic forest, Araceae, Bromeliaceae, Orchidaceae, biogeographical patterns.
LEITMAN, P., AMORIM, A., MENINI NETO, L., FORZZA, R.C. Angiospermas epı́fitas de uma
floresta montana no sul da Bahia, Brasil. Biota Neotropica. 14(2): 1––12. http://dx.doi.org/10.1590/S167606032014001013
Resumo: A Floresta Atlântica do sul da Bahia apresenta elevados nı́veis de diversidade e endemismo de
plantas vasculares. No entanto, poucos levantamentos florı́sticos foram realizados em áreas de floresta
montana, especialmente para ervas e plantas de dossel. Estudos com epı́fitas estão concentrados
principalmente em áreas do Sul e Sudeste do Brasil. Desta forma, o presente estudo realizou o
levantamento das angiospermas epı́fitas do Parque Nacional da Serra das Lontras (PNSL) e verificou a
distribuição das espécies pelas famı́lias e gêneros e seus padrões biogeográficos. As coletas foram
realizadas entre fevereiro de 2011 e março de 2012 nas trilhas e estradas de acesso. Alguns forófitos foram
escalados e árvores e ramos caı́dos foram examinados com o intuito de coletar espécies restritas ao dossel.
A área de estudo apresentou alta diversidade de epı́fitas (256 spp.), uma das maiores já registradas para a
Floresta Atlântica. As famı́lias com maior riqueza são Orchidaceae, Bromeliaceae e Araceae,
reafirmando os padrões encontrados anteriormente em revisões do tema. As espécies apresentam alto
grau de endemismo (45%) à Floresta Atlântica, especialmente em Bromeliaceae e Orchidaceae. São
registradas uma nova ocorrência para o Brasil, quatro para a Floresta Atlântica e 30 para a Bahia. O
PNSL possui mais espécies em comum com o Sul e o Sudeste do que com a região Nordeste.
Palavras-chave: Floresta Atlântica, Araceae, Bromeliaceae, Orchidaceae, padrões biogeográficos.
http://dx.doi.org/10.1590/S1676-06032014001013
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Biota Neotrop., 14(2): 1–
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Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Introduction
Epiphytes spend most or all of their life cycles growing
on other plants (phorophytes), taking advantage of their
mechanical support (Benzing 1990). Epiphytes account for
approximately 10% of world’s vascular flora and are found in
84 angiosperm families (Gentry & Dodson 1987, Kersten 2010).
Araceae, Bromeliaceae, and Orchidaceae are the most successful epiphytic angiosperm families, and the latter is responsible
for over 70% of total richness.
Bromeliaceae is a typically American family, while the
largest genera of Araceae and Orchidaceae (e.g., Anthurium,
Epidendrum, Lepanthes, Philodendron and Pleurothallis s.l.)
are likewise endemic to that continent. The Neotropical
region therefore hosts the greater part of world’s epiphytic
flora (. 15.500 spp.), which are mainly concentrated in Central
America, the Andes, northwestern Amazonia and the Atlantic
Forest (Gentry & Dodson 1987, Nieder et al. 2001). Of the
14.500 vascular plants known to inhabit the Atlantic Forest,
3.000-4.000 are epiphytes (Stehmann et al. 2009, Kersten 2010).
The Atlantic Forest has large latitudinal (from 36S to 306S) and
altitudinal (from sea level up to 2.890 m) extensions and
significant rainfall variations – resulting in high habitat
heterogeneity and an extraordinary diversity of organisms
(Oliveira-Filho & Fontes 2000).
Southern Bahia is considered an area of great biological
importance, with elevated levels of richness and endemism in its
arboreal component (Mori et al. 1983, Thomas et al. 1998,
Martini et al. 2007, Thomas et al. 2009, Rocha & Amorim
2012), although surveys of humid montane forests have been
scarce, especially those focusing on terrestrial and epiphytic
herbs (Amorim et al. 2009, Matos et al. 2010). The recently
created Serra das Lontras National Park comprises a vast
remnant of montane Atlantic Forest in southern Bahia (Save
Brasil et al. 2009).
A preliminary survey in the park, covering only a small
area, recorded 709 angiosperm species, of which 158 (22.3%)
were epiphytes (Amorim et al. 2009). These authors highlighted
the fact that many of the species encountered, including many
epiphytes (e.g., Fuchsia regia [Vell.] Munz, Nematanthus
lanceolatus [Poir.] Chautems, and Vriesea longicaulis Mez),
were previously known only from mountainous areas in
southern and southeastern Brazil.
For the preliminary survey there was no effort focused on
this habit, which presents peculiarities that hinder their
sampling, including the height they occur on the phorophyte
and the small sizes of some groups (e.g., Peperomia and
Pleurothallis s.l.). We therefore sought to inventory the
composition of epiphytic angiosperms in the Serra das
Lontras National Park to investigate their distributions into
genera and families and their biogeographical patterns to
increase our knowledge of this group in the park and in
northeastern Brazil.
Material and methods
Serra das Lontras National Park (SLNP) is located in
southern Bahia State (Figure 1), Brazil, in the municipalities of
Arataca, São José da Vitória, and Una (156079-156159S and
396159-396259W). The park comprises approximately 11.000
hectares at altitudes that vary from 400 to 1.000 m. Located
30 km from the coast, the mountains act as a first barrier to
humid air masses moving inland from the ocean, resulting in
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mean annual precipitation rates of 1.300-1.600 mm, without a
marked dry season, and a mean annual temperature of 246C
(classified as Af in the Koppen system) (Peel et al. 2007,
Amorim et al. 2009, Nacif et al. 2009).
Situated in the Atlantic Forest domain, the SLNP comprises
a vegetation mosaic of anthropogenic areas mixed with wellpreserved submontane forests (with canopies .30 m) at lower
altitudes and cloud forests above 800 m with smaller trees with
twisted trunks bearing bryophytes (Stadtmüller 1987).
A total floristic inventory of the SLNP has been underway
for the last eight years, and preliminary results have already
been published (Amorim et al. 2009; Save Brasil et al. 2009).
Visits to the area were made every two months, from February/
2011 to March/2012 to survey the epiphytic angiosperms.
Collections were made along roads and trails in order to cover
the largest possible areas, and some living phorophytes were
occasionally climbed and fallen trees examined to collect
canopy species. Accidental epiphytes were not considered in
this survey. Informations on the type of epiphytism (characteristic holoepiphyte, facultative holoepiphyte, hemiepiphyte)
were obtained from local observations and complemented with
informations from literature. This was done in order to have
more accurate data, since many facultative species might have
been seen only as epiphytes in the area or might occur as
terrestrial or rupiculous in parts of the park that were not
visited. All fertile material was deposited at the RB and CEPEC
herbaria, while sterile specimens were incorporated into the exsitu collection at the Rio de Janeiro Botanical Garden.
Specimens collected during the general floristic inventory,
as well as those from the present survey, were identified to the
species level, when possible, with the help of specialists and
specific literature (e.g., the Flora Neotropica). Nomenclature
and species concepts follow the List of Species of the Brazilian
Flora - LSBF (2013). Distribution and endemism data were
obtained from the LSBF for taxa identified to the species level.
A Rapid Color Guide, published by The Field Museum,
was prepared and images on most species can be seen at www.
fm2.fieldmuseum.org/plantguides/guideimages.asp?ID=545.
Results
The Serra das Lontras National Park shows high epiphyte
richness, with a total of 256 species identified, representing 87
genera and 18 families (Table 1) – making it one of the richest
sites for epiphytic angiosperms in the Atlantic Forest. Of the
published checklists, only Macaé de Cima in Rio de Janeiro State
reported more epiphyte species (275 spp.) (Fontoura et al. 1997).
The most important families encountered were Orchidaceae
(106 spp.), Bromeliaceae (64 spp.), Araceae (31 spp.), and
Piperaceae (14 spp.). Together, these families encompassed
more than 80% of the epiphyte flora at SLNP. The species
distributions within the families were very unequal, however,
with 11 families being represented by less than five species.
The richest genera were Vriesea (22 spp.), Peperomia and
Stelis (14 spp. each), Anthurium (13 spp.), Aechmea and
Philodendron (12 spp. each), Epidendrum (11 spp.), and
Hohenbergia (10 spp.). The species distributions within the
genera were very unequal, with half of them (43 genera) being
represented by only a single species, and 82% by less than five
species.
With many ornamental species, 10 epiphytes registered for
the SLNP are cited on the Red Book of Brazilian Flora
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Biota Neotrop., 14(2): 1––12, 2014
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Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Figure 1. Map showing the location of the studied area (Serra das Lontras National Park); in gray is the Atlantic Forest limits. N.P. - National Park;
B.R. - Biological Reserve/Mapa de localização da área de estudo (Parque Nacional da Serra das Lontras); em cinza está demarcado o limite da
Floresta Atlântica.
(Martinelli & Moraes 2013). Of this total, three are ‘‘Critically
endangered’’, five are ‘‘Endangered’’, and two are ‘‘Vulnerable’’.
Four are considered to be ‘‘Data Deficient’’ (Table 1).
Discussion
The high numbers of species observed in Araceae,
Bromeliaceae, Orchidaceae, and Piperaceae were consonant
with global patterns of epiphytic distribution among angiosperm families, as reported by Madison (1977) and Gentry &
Dodson (1987). Similar results were reported for the
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Neotropical region (e.g., Sudgen & Robins 1979, Gentry &
Dodson 1987, Wolf & Flamenco-S 2003, Küper et al. 2004) and
the tropical and subtropical Atlantic Forest (e.g., Fontoura et
al. 1997, Borgo & Silva 2003, Giongo & Waechter 2004,
Kersten & Silva 2001, Buzatto et al. 2008, Martins et al. 2008,
Menini Neto et al. 2009, Mania & Monteiro 2010).
Table 2 compares the SLNP and other Atlantic Forest areas
with high epiphyte richness. The elevated numbers of species of
Bromeliaceae, Araceae, and Gesneriaceae is notable, representing the highest epiphyte richness ever recorded for these
families in the Atlantic Forest. Vriesea, Stelis, Anthurium,
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Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 1. Epiphytic Angiosperms registered for Serra das Lontras National Park, Bahia, Brazil. 1 *New occurence to Bahia state; ** New occurence
to the Atlantic Forest; *** New occurence to Brazil. 2 AA – André M. Amorim; AF – André P. Fontana; AJ – Alessandra B. Jardim; JJ – Jomar G.
Jardim; JP – José Lima da Paixão; ML – Márdel M. Lopes; PF – Pedro Fiaschi; PL – Paula Leitman; RB – Rafael X. Borges; RP – Ricardo O.
Perdiz; WT – Wm. Wayt Thomas. 3 Epi – Characteristic Holoepiphyte; Fac – Facultative Holoepiphyte; Hep – Hemiepiphyte. 4 CAF – CerradoAtlantic Forest; Di – Disjunct distribution Amazon-Atlantic Forest; En – Endemic to Atlantic Forest; WD – Wide distribution; O – Other; ‘‘-’’ – not
applicable. 5 CR – Criticaly endangered; DD – Data deficient; EN – Endangered; VU – Vulnerable.
Family
Acanthaceae
Araceae
Specie1
Voucher2
Habit3
Distribution4
Clistax bahiensis Profice & Leitman
Anthurium boudetii Nadruz *
Anthurium gladiifolium Schott
Anthurium gracile (Rudge) Lindl. **
Anthurium intermedium Kunth
Anthurium jilekii Schott
Anthurium pentaphyllum (Aubl.) G.Don
AA 5301
AA 6399
AA 5770
PL 211
PL 165
PL 547
AA
5770a
ML 1473
AA 6610
PL 417
JJ 4919
PL 324
JJ 4389
JJ 4337
JJ 4886
JP 956
AA 6580
JJ 4898
ML 1449
AA 5960
JJ 4896
AA 6727
AA 6599
PL 202
PL 287
AA 5296
PL 119
PL 561
JJ 4842
PL 428
AA 5298
AA
5960a
AA 6568
AA 5280
AA 5265
PL 338
AA 5275
PL 424
PL 378
PL 309
AA 6594
PL 262
PL 395
AA 7217
PL 129
ML 1484
ML 1491
PL 141
JJ 4680
JJ 5431
Hep
Fac
Epi
Epi
Fac
Hep
Hep
En
En
En
WD
En
O
WD
Hep
Fac
Epi
Epi
Epi
Epi
Epi
Hep
Hep
Hep
Hep
Hep
Hep
Fac
Hep
Hep
Hep
Hep
Hep
Hep
Hep
Hep
Hep
Epi
Hep
WD
WD
WD
WD
En
En
WD
WD
O
WD
WD
En
En
Di
Di
WD
Di
Di
WD
Hep
Hep
Hep
Hep
Hep
Fac
Fac
Fac
Fac
Fac
Fac
Fac
Fac
Epi
Epi
Epi
Epi
Epi
WD
En
En
En
O
En
En
En
En
En
CAF
Di
En
En
En
En
-
Anthurium scandens (Aubl.) Engl.
Anthurium solitarium Schott
Anthurium sp. 1
Anthurium sp. 2
Anthurium sp. 3
Anthurium sp. 4
Anthurium sp. 5
Heteropsis oblongifolia H.B.K.
Monstera adansonii Schott
Philodendron cordatum Kunth
Philodendron edmundoi G.M.Barroso *
Philodendron fragrantissimum (Hook.) G.Don
Philodendron hederaceum (Jacq.) Schott
Philodendron insigne Schott
Philodendron ornatum Schott
Philodendron pedatum (Hook.) H.B.K.
Philodendron propinquum Schott
Philodendron recurvifolium Schott
Philodendron rudgeanum Schott
Philodendron surinamense (Miq.) Engl.
Philodendron aff. williamsii Hook. f.
Rhodospatha latifolia Poepp.
Rhodospatha oblongata Poepp.
Stenospermation spruceanum Schott
Syngonium vellozianum Schott
Araliaceae
Begoniaceae
Bromeliaceae
Oreopanax capitatus (Jacq.) Decne. & Planch.
Begonia convolvulacea A.DC.
Begonia fruticosa (Klotzsch) A.DC.
Begonia aff. itaguassuensis Brade
Begonia radicans Vell.
Begonia grisea A.DC. **
Aechmea burlemarxii E. Pereira
Aechmea conifera L.B.Sm. *
Aechmea froesii (L.B.Sm.) Leme & J.A.Siqueira
Aechmea gustavoi J.A.Siqueira & Leme
Aechmea multiflora L.B.Sm.
Aechmea nudicaulis Griseb.
Aechmea patentissima (Mart. ex Schult. & Schult. f.) Baker
Aechmea subintegerrima (Philcox) Leme
Aechmea tentaculifera Leme et al.
Aechmea turbinocalyx Mez
Aechmea viridostigma Leme & H.Luther
Aechmea sp.
Threat
category5
CR
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Biota Neotrop., 14(2): 1––12, 2014
5
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 1. Continued.
Specie1
Voucher2
Habit3
Distribution4
Billbergia euphemiae E.Morren
Billbergia saundersii Bull
Canistrum camacaense Martinelli & Leme
Canistrum montanum Leme
Guzmania lingulata (L.) Mez
Hohenbergia augusta (Vell.) E.Morren *
Hohenbergia brachycephala L.B.Sm.
Hohenbergia capitata Schult. & Schult.f.
Hohenbergia edmundoi L.B.Sm. & Read
Hohenbergia minor L.B.Sm.
Hohenbergia sp. 1
Hohenbergia sp. 2
Hohenbergia sp. 3
Hohenbergia sp. 4
Hohenbergia sp. 5
Lymania azurea Leme
Neoregelia azevedoi Leme
Neoregelia crispata Leme
Neoregelia kerryi Leme
Neoregelia pauciflora L.B.Sm.
Neoregelia wilsoniana M.B.Foster
Nidularium innocentii Lem.
Nidularium procerum Lindm.
AA 5752
JP 976
AA 5751
AA 5269
RB 591
PL 149
PL 439
AA 6602
JJ 4920
PL 158
PL 398
PL 134
PL 307
PL 391
PL 305
JJ 5316
AA 6624
AA 5991
AA 5807
AA 6590
PL 267
AA 6696
WT
14093b
PL 435
AA 6597
AA 5351
AA 5820
AA 6391
JJ 4895
AA 6086
PL 208
AA 5303
PL 380
JP 965
JP 975
RB 590
PL 489
AA 5797
AA 6758
AA 6646
PL 368
AA 6759
PL 301
PL 205
PL 150
PL 345
AA 6097
AA 6688
PL 289
PL 114
PL 181
PL 442
JJ 4401
PL 230
PL 200
Fac
Epi
Fac
Epi
Fac
Fac
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Fac
Epi
Epi
Fac
Fac
En
En
En
En
WD
En
En
En
En
En
En
En
En
En
En
En
En
En
Epi
Epi
Fac
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
En
En
En
WD
CAF
CAF
CAF
En
En
En
En
En
En
En
En
En
Di
O
En
En
En
En
O
En
En
WD
O
En
Family
Cactaceae
Portea filifera L.B.Sm.
Portea nana Leme
Quesnelia clavata Amorim & Leme
Racinaea spiculosa (Griseb.) M.A.Spencer & L.B.Sm.
Tillandsia geminiflora Brong.
Tillandsia sprengeliana Klotzsch ex Mez
Tillandsia stricta Sol.
Vriesea breviscapa (E.Pereira & I.A.Penna) Leme
Vriesea dictyographa Leme
Vriesea drepanocarpa (Baker) Mez
Vriesea duvaliana E.Morren
Vriesea ensiformis (Vell.) Beer
Vriesea flammea L.B.Sm.
Vriesea guttata Linden & André *
Vriesea longiscapa Ule
Vriesea longisepala A.F.Costa
Vriesea paratiensis E.Pereira
Vriesea procera (Mart. ex Schult. & Schult.f.) Wittm.
Vriesea regnelli Mez
Vriesea rhodostachys L.B.Sm.
Vriesea rodigasiana E.Morren
Vriesea ruschii L.B.Sm.
Vriesea simplex (Vell.) Beer
Vriesea tijucana E. Pereira
Vriesea vagans (L.B.Sm.) L.B.Sm. *
Vriesea sp. 1
Vriesea sp. 2
Vriesea sp. 3
Vriesea sp. 4
Epiphyllum phyllanthus (L.) Haw.
Lepismium cruciforme (Vell.) Miq.
Rhipsalis floccosa Salm-Dyck ex Pfeiff.
Threat
category5
EN
EN
EN
EN
DD
DD
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Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 1. Continued.
Family
Clusiaceae
Cyclanthaceae
Gesneriaceae
Lentibulariaceae
Marcgraviaceae
Melastomataceae
Voucher2
Habit3
Distribution4
Threat
category5
Rhipsalis hileiabaiana (N.P.Taylor & Barthlott) N.
Korotkova & Barthlott
Rhipsalis lindbergiana K.Schum
Rhipsalis neves-armondii K.Schum
Rhipsalis oblonga Loefgr.
Clusia panapanari (Aubl.) Choisy
Asplundia gardneri (Hook,) Harling
PL 521
Epi
CAF
DD
PL 367
PL 513
ML 1019
AA 7205
PL 213
Epi
Epi
Epi
Hep
Hep
O
En
En
O
WD
Asplundia maximiliani Harling
Evodianthus funifer (Poit.) Lindm.
Thoracocarpus bissectus (Vell.) Harling
Codonanthe cordifolia Chautems
Codonanthe gracilis (Mart.) Hanst.
Codonanthe uleana Fritsch
Columnea sanguinea (Pers.) Hanst.
Nematanthus albus Chautems
Nematanthus corticola Schrad.
Nematanthus lanceolatus (Poir.) Chautems
Utricularia jamesoniana Oliv. **
PL 506
ML 1464
PL 427
JJ 4892
PL 411
AA 7221
RB 622
JJ 4907
JP 973
AA 5276
JJ 4908
Hep
Hep
Hep
Epi
Epi
Epi
Epi
Epi
Epi
Fac
Epi
En
WD
WD
CAF
En
WD
WD
En
En
CAF
O
Marcgravia coriacea Vahl
AA 6679
Hep
WD
Marcgravia polyantha Delpino
Bertolonia alternifolia Baumgratz et al.
JJ 4697
AJ 170
Hep
Epi
CAF
En
Bertolonia bullata Baumgratz et al.
Bertolonia ovata DC.
AA 5979
WT
14581
JJ 4921
WT
14112
PF 2811
FF 1447
AA 5806
PL 412
PL 361
Epi
Epi
En
En
Epi
Hep
En
WD
Hep
Hep
Hep
Epi
Epi
WD
O
CAF
En
En
AF 2682
Epi
Di
PL 251
AA 5809
PL 376
Fac
Epi
Epi
WD
WD
En
RP 536
JJ 4677
PL 225
JJ 5383
AF 2666
AF 2688
JJ 4385
PL 244
PL 355
JJ 4922
AA 6626
PL 354
AA 5816
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
En
En
En
CAF
O
En
Di
En
En
CAF
Pleiochiton blepharodes (DC.) Reginato & Goldenberg
Ficus arpazusa Casar.
Moraceae
Onagraceae
Orchidaceae
Specie1
Ficus castellviana Dugand
Ficus hirsuta Schott
Fuchsia regia (Vand. ex Vell.) Munz
Acianthera auriculata (Lindl.) Pridgeon & M.W.Chase
Acianthera capanemae (Barb.Rodr.) Pridgeon &
M.W.Chase *
Acianthera hygrophila (Barb.Rodr.) Pridgeon &
M.W.Chase *
Anathallis rubens (Lindl.) Pridgeon & M.W.Chase
Anathallis sclerophylla (Lindl.) Pridgeon & M.W.Chase
Baptistonia gutfreundiana (Chiron & V.P.Castro) Chiron &
V.P.Castro
Baptistonia truncata (Pabst) Chiron & V.P.Castro *
Brachionidium restrepioides Hoehne & Pabst
Bulbophyllum micropetaliforme J.E.Leite
Bulbophyllum napellii Lindl.
Camaridium carinatum (Barb.Rodr.) Hoehne
Camaridium cf. micranthum M.A. Blanco
Catasetum cf. hookeri Lindl.
Catasetum purum Nees & Sinnings
Cirrhaea cf. seidelii Pabst
Coppensia flexuosa (Sims) Campacci
Coppensia hookeri (Rolfe) F.Barros & L.Guimarães
Dichaea brevicaulis Cogn.
Dichaea cogniauxiana Schltr.
CR
VU
Continued on next page
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Biota Neotrop., 14(2): 1––12, 2014
7
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 1. Continued.
Specie1
Voucher2
Habit3
Distribution4
Dichaea pendula (Aubl.) Cogn.
Elleanthus brasiliensis (Lindl.) Rchb.f.
Elleanthus crinipes Rchb.f.
Elleanthus hymenophorus (Rchb.f.) Rchb.f. ***
Elleanthus linifolius C.Presl
Encyclia fimbriata A.C.Bastos et al.
PL 353
PL 549
AF 2673
AA 4960
AA 4983
WT
14604
PL 326
AF 2659
AA 6581
PL 463
PL 432
AA 6413
ML 539
JP 971
AA 6412
AA 5810
AF 2685
JJ 4901
JJ 4904
AA 4448
PL 413
PL 191
WT
14594
AF 2675
PL 555
AF 2684
AF 2654
WT
14097
JJ 4912
ML 1023
AA 5329
JJ 4379
PL 154
JJ 4681
PL 374
PL 314
PL 316
Epi
Epi
Epi
Epi
Epi
Epi
WD
O
CAF
WD
WD
En
Fac
Epi
Epi
Fac
Epi
Epi
Epi
Epi
Fac
Epi
Epi
Epi
Epi
Fac
Epi
Epi
Epi
CAF
En
O
WD
Di
O
WD
WD
WD
CAF
En
WD
CAF
En
Di
En
Epi
Epi
Epi
Epi
Epi
WD
O
En
Di
CAF
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
En
Di
Di
WD
En
En
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Di
WD
O
O
En
En
WD
En
Di
En
CAF
WD
-
Family
Encyclia patens Hook.
Encyclia unaensis Fowlie
Epidendrum armeniacum Lindl.
Epidendrum densiflorum Lindl.
Epidendrum latilabre Lindl. *
Epidendrum macrocarpum Rich.
Epidendrum nocturnum Jacq.
Epidendrum ochrochlorum Barb.Rodr.
Epidendrum paranaense Barb.Rodr.
Epidendrum ramosum Jacq.
Epidendrum saximontanum Pabst
Epidendrum secundum Jacq.
Epidendrum sp.
Gomesa recurva Barb.Rodr.
Grandiphyllum pohlianum (Cogn.) Docha Neto *
Heterotaxis brasiliensis (Brieger & Illg) F.Barros
Houlletia brocklehurstiana Lindl.
Jacquiniella globosa (Jacq.) Schltr.
Koellensteinia graminea Rchb.f.
Lankesterella longicollis (Cogn.) Hoehne *
Lockhartia lunifera Rchb.f.
Masdevallia infracta Lindl.
Maxillaria bradei Schltr. ex Hoehne *
Maxilaria candida Lodd. ex Lindl. *
Maxillaria leucaimata Barb.Rodr.
Maxillaria ochroleuca Lodd. ex Lindl.
Maxilaria aff. ringens Rchb.f.
Maxillaria spiritusanctensis Pabst
Maxilaria aff. splendens Poepp. & Endl.
Maxilaria sp.
Maxillariella robusta (Barb. Rodr.) M.A.Blanco &
Carnevali *
Mormolyca rufescens (Lindl.) M.A.Blanco
Nitidobulbon nasutum (Rchb.f.) Ojeda & Carnevali
Octomeria concolor Barb.Rodr. *
Octomeria crassifolia Lindl.
Octomeria decumbens Cogn.
Octomeria geraensis Barb.Rodr. *
Octomeria grandiflora Lindl.
Octomeria tricolor Rchb.f.
Octomeria sp.
Ornithidium rigidum (Barb.Rodr.) M.A.Blanco & Ojeda *
Pabstiella bradei (Schltr.) Luer *
Pabstiella ramphastorhyncha (Barb. Rodr.) L. Kollmann
Pabstiella aff. trifida (Lindl.) Luer
Pleurothallis ruscifolia (Jacq.) R.Br.
Pleurothallis sp.
JJ 4683
AA 4970
PL 464
AF 2657
PL 400
PL 265
PL 226
JJ 4665
PL 493
RP 301
PL 377
PL 360
AF 2658
PF 2821
PL 222
Threat
category5
EN
DD
VU
Continued on next page
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8
Biota Neotrop., 14(2): 1–
–12, 2014
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 1. Continued.
Family
Specie1
Voucher2
Habit3
Distribution4
Polycycnis silvana F.Barros
Polystachya estrellensis Rchb.f.
Promenaea silvana F.Barros & Cath.
Promenaea xanthina Lindl.
Prosthechea bueraremensis (Campacci) Campacci
Prosthechea calamaria (Lindl.) W.E.Higgins *
Prosthechea fragans (Sw.) W.E.Higgins
Prosthechea pachysepala (Klotzsch) Chiron & V.P.Castro
Prosthechea pygmaea (Hook.) W.E.Higgins
AA 5805
RP 294
JJ 4815
RP 330
AF 2662
AF 2674
AA 5361
RP 335
WT
14101
ML 549
AA 5355
PL 399
AF 2672
WT
14096
AF 2651
PL 494
AA 6588
PL 532
WT
14102
PL 256
AA 5746
PL 135
PL 430
PL 175
PL 359
PL 431
PL 519
JJ 4713
WT
14103
ML 1187
PL 221
RP 69
PL 310
AA 7264
AF 2665
JJ 4346
PL 100
AA 6649
WT
14541
ML 1020
JJ 4798
AA 5952
AA 5946
MJ 840
AA 6357
PL 548
PL 421
AA 6681
AA 5287
AA 5800
AA 5741
JJ 4851
AA 4962
Epi
Fac
Epi
Epi
Epi
Epi
Epi
Epi
Epi
En
WD
En
En
En
En
WD
CAF
WD
Fac
Epi
Epi
Fac
Epi
WD
Di
WD
WD
O
Epi
Epi
Epi
Epi
Epi
WD
WD
O
En
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
Epi
O
En
Di
Di
En
-
Epi
Epi
Epi
Fac
Epi
Epi
Fac
Fac
Fac
Fac
En
WD
WD
En
WD
O
WD
O
Fac
Epi
Fac
Fac
Fac
Fac
Fac
Fac
Fac
Fac
Epi
Epi
Hep
Hep
WD
WD
WD
WD
WD
En
WD
CAF
WD
WD
WD
En
Di
En
Prosthechea vespa (Vell.) W.E.Higgins *
Rhetinantha notylioglossa (Rchb.f.) M.A.Blanco
Scaphyglottis fusiformis (Griseb.) Schultes *
Scaphyglottis modesta (Rchb.f.) Schltr.
Scaphyglottis reflexa Lindl.
Sobralia cf. sessilis Lindl.
Specklinia grobyi Batem. ex Lindl.
Stelis aprica Lindl.
Stelis caespitosa Lindl. **
Stelis deregularis Barb.Rodr.
Stelis aff. fraterna Lindl.
Stelis intermedia Poepp. & Endl. *
Stelis megantha Barb.Rodr. *
Stelis papaquerensis Rchb.f. *
Stelis pauciflora Lindl. *
Stelis ruprechtiana Rchb.f. *
Stelis sp. 1
Stelis sp. 2
Stelis sp. 3
Stelis sp. 4
Piperaceae
Rubiaceae
Solanaceae
http://www.scielo.br/bn
Stelis sp. 5
Trichopilia santoslimae Brade *
Xylobium colleyi (Bateman ex Lindl.) Rolfe *
Xylobium variegatum (Ruiz & Pav.) Mansf. *
Zygopetalum crinitum Lodd. *
Zygopetalum cf. maxillare Lodd.
Peperomia alata Ruiz & Pav.
Peperomia corcovadensis Gardner *
Peperomia elongata Kunth
Peperomia emarginella (Sw.) C.DC.
Peperomia hernandiifolia (Vahl) A.Dietr.
Peperomia macrostachya (Vahl) A.Dietr.
Peperomia magnoliifolia (Jacq.) A.Dietr.
Peperomia obtusifolia (L.) A.Dietr.
Peperomia pilicaulis C.DC.
Peperomia pseudoestrelensis C.DC.
Peperomia serpens (Sw.) Loud
Peperomia tenella (Sw.) A.Dietr.
Peperomia tetraphylla (G.Forst.) Hook. & Arn.
Peperomia urocarpa Fisch. & C.A.Mey.
Hillia parasitica Jacq.
Notopleura bahiensis C.M.Taylor
Schradera polycephala DC.
Dyssochroma viridiflorum (Sims) Miers
Threat
category5
CR
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Biota Neotrop., 14(2): 1––12, 2014
9
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
Table 2. Comparison of species richness for the main families and genera with epiphytic habit between Serra das Lontras National Park and other
surveys carried at the Atlantic Forest. 1 N.P. – National Park; S.P. – State Park; N.R. – Natural Reserve. ‘‘-’’ – Information not available.
1
Serra das
Lontras
N.P., BA
Macaé de
Cima, RJ
Upper
Iguaçu
River
Basin, PR
Prata
Moutain
Range,
PR
Ibitipoca
S.P., MG
Carlos
Botelho
S.P., SP
Ilha do
Cardoso
S.P., SP
Vale do Rio
Doce N.R.,
ES
Taxon
Present
study
Kersten &
Kunyioshi
2006
Blum
et al. 2011
Menini
Neto et al.
2009
Breier
et al. 2005
Breier
et al. 2005
Coelho
2010
Total area
Total epiphytic
Angiosperms
Orchidaceae
Bromeliaceae
Araceae
Piperaceae
Cactaceae
Gesneriaceae
Vriesea
Peperomia
Stelis
Anthurium
Philodendron
Aechmea
Epidendrum
Hohenbergia
11,000ha
256
Fontoura
et al. 1997;
Costa &
Wendt
2007
7,000ha
275
254
6.3ha
204
1,923.5ha
181
10.24ha
121
10.24ha
136
22.000ha
-
103
62
29
15
7
7
22
15
14
13
12
11
11
9
184
45
5
5
7
6
18
5
4
10
4
10
0
147
39
9
24
12
3
11
22
6
2
7
9
4
0
103
38
14
14
9
5
18
14
7
6
6
5
12
0
66
22
9
3
7
2
7
3
5
5
4
1
6
0
42
27
13
8
11
6
12
7
1
3
8
5
4
0
72
33
11
6
1
3
15
5
1
3
7
3
8
0
22
4
12
-
Study area
Aechmea, and Hohenbergia also exhibited their greatest
epiphytic species diversity in the SLNP. It is interesting that
Hohenbergia was among the richest genera in the study area, as
this genus does not commonly figure among the most species
rich taxa in epiphyte surveys. The greatest diversity of this
genus is encountered in southern Bahia, where 21 of its 25
Atlantic Forest species occur (Martinelli et al. 2008, Forzza
et al. 2013).
In addition to H. brachycephala and H. minor, both
endemic to Bahia, H. edmundoi, a poorly-known species from
Bahia for which no habitat information was available (Smith &
Downs 1979, Forzza et al. 2013), was found in a submontane
forest in the SLNP; H. augusta, previously considered to be
restricted to southeastern Brazil, was also collected. These
results reaffirm the importance of southern Bahia State in the
distribution and taxonomy of this genus.
Vriesea (Tillandsioideae) and Aechmea (Bromelioideae)
represent 12% and 7%, respectively, of the total number of
species in the Atlantic Forest (Forzza et al. 2013). Fontoura
et al. (2012) reported that Bromelioideae taxa were encountered
1.6 times more frequently in local surveys of epiphytic species in
the Atlantic Forest than representatives of Tillandsioideae. This
proportion was lower in the SLNP, where Vriesea is very
diverse, with almost the same numbers of species being
encountered in each subfamily (34 spp. Bromelioideae x 30
spp. Tillandsioideae).
Another family that demonstrated elevated richness in the
SLNP was Araceae, with one third of the 65 species known to
the Atlantic Forest being encountered there (Coelho et al.
2013). The most diversified genera of this family were likewise
http://dx.doi.org/10.1590/S1676-06032014001013
well represented in the park, with more than half of the species
of Anthurium and Philodendron known from Bahia being
encountered in the SLNP. Additionally, two new occurrence
records for Bahia State (A. boudetii and P. edmundoi), one for
the Atlantic Forest domain (A. gracile), and five morphotypes
of Anthurium reinforce the importance of additional studies in
mountainous areas there to improve our knowledge of the
distribution of this family within the state. In contrast to the
high endemism levels of Bromeliaceae and Orchidaceae,
representatives of Araceae exhibit wide (and sometimes
disjunct) distributions (Coelho et al. 2013).
Orchidaceae is responsible for approximately 70% of the
global epiphytic flora, and is always among the top families in
every floristic study of that habit (Benzing 1990, Kersten 2010,
Zotz 2013). Most orchid diversity is found in the tropics,
especially in mountainous areas (Dressler 1993). The majority
of the orchid taxa encountered in the SLNP is relatively small
(e.g., Acianthera, Anathallis, Brachionidium, Jacquiniella,
Lankesterella, Octomeria, Pabstiella, Pleurothallis, Promenaea,
Specklinia, and Stelis) and commonly neglected in floristic
surveys – and therefore poorly represented in scientific
collections (Dressler 2005). As the present study focused on
epiphytes (and ex-situ cultivation was undertaken), high
numbers of species were included on the final list – with 22
new occurrences for Bahia State (Table 1).
Northeastern Brazil represents one of the 10 areas in the
world considered most likely to have undescribed species, and
where models predict a 30% increase in known diversity (Joppa
et al. 2011). Since initiating preliminary studies of the flora of
the SLNP (Amorim et al. 2009), six new epiphyte species have
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10
Biota Neotrop., 14(2): 1–
–12, 2014
Epiphytic angiosperms in a mountain forest in southern Bahia, Brazil
been described for the area: Bertolonia alternifolia, B. bullata,
Clistax bahiensis, Encyclia fimbriata, Quesnelia clavata, and
Vriesea longisepala, and one is about to be published. The
present survey found other possible taxonomic novelties, and
recent studies in neighboring areas encountered several new
species with epiphytic habits (e.g., Amorim & Leme 2009, Leme
& Kollmann 2011, Costa et al. 2012, Reginato et al. 2013) –
emphasizing the need of further investigations of this group in
southern Bahia.
The present study expands the distribution of many taxa,
with one new occurrence for Brazil (Elleanthus hymenophorus
(Rchb.f.) Rchb.f.), four for the Atlantic Forest biome, and 30
for Bahia State (Table 1). Utricularia jamsoniana Oliv., a species
previously known only from altitudinal regions of Central
America and northern South America (Taylor 1989), was
encountered in a cloud forest in the highest area of the park
(Peito de Moça and Javi mountain ranges). Over half (16 spp.)
of these new records were known only from collections in
strictly southern and southeastern regions. The epiphytic flora
of the SLNP shares more species with southern and southeastern Brazil (71 spp.) than with northeastern states (five
species – four Aechmea and one Hohenbergia). This same
pattern was found for ferns and lycophytes on a nearby
mountainous area in the Serra Bonita Private Reserve, with
almost a quarter of the surveyed species being previously
restricted to southeastern Brazil (Matos et al. 2010).
Almost half of the 221 taxa identified to species level in
the present study are endemic to the Atlantic Forest (100
spp. - 45%) (Table 1), and most of them belong to
Bromeliaceae and Orchidaceae (44 and 32 species respectively). These two families exhibit high degrees of endemism
in the Atlantic Forest, with 80% of Bromeliaceae and 63% of
Orchidaceae being restricted to this domain (Martinelli et al.
2008, Stehmann et al. 2009). These high levels of endemism
may be related to limited connections with the other major
South American forest formation in the Amazon (Mori et al.
1981). These two forests are separated by an open area
known as the ‘‘dry diagonal’’ (Prado & Gibbs 1993) that acts
as a barrier to species dispersion, especially less droughttolerant taxa.
On the other hand, 60 species showed broad distributions in
Brazil or the Neotropical region, and 24 had disjunct
distributions between the Atlantic and Amazon forests (Table
1); 19 species also occur in the ‘‘Cerrado’’ (Brazilian savanna)
region, suggesting that dry formations do not act as barriers to
all epiphyte taxa, especially those with broader ecophysiological tolerances (Oliveira-Filho & Ratter 1995, Ibisch et al. 1996).
Deciduous and semi-deciduous seasonal forest patches and
gallery forests that crisscross the Cerrado may offer dispersal
pathways for many species and establish tenuous connections
between these two major forest formations (Leitão Filho 1987,
Oliveira-Filho & Ratter 1995).
All of the 10 species cited on the Red Book of Brazilian
Flora (Martinelli & Moraes 2013) as threatened belong to
Bromeliaceae and Orchidaceae, two very ornamental families.
The five bromeliads are restricted to Northeast Brazil, and four
of them are endemic to Bahia. On the other hand, the orchids
indicated as threatened are new occurrences to the state. These
results reinforce the importance of the park as a conservation
unit and the relevance of floristic surveys, especially for often
neglected groups such as epiphytes. Recent floristic studies in
mountainous areas in southern Bahia have revealed large
http://www.scielo.br/bn
numbers of new species and new occurrences of epiphytes –
pointing to the need for further investigations of this group.
Acknowledgments
We thank FAPESB, FAPERJ and CAPES for fieldwork
financial support. Paula Leitman thank CNPq and CAPES for
master’s scholarship. Rafaela C. Forzza and André Amorim
are CNPq research fellows. We also thank the specialists who
helped identifying the species - Alain Chautems, Andrea Costa,
Charlotte Taylor, Claudio Nicoletti Fraga, Daniela Zappi,
Daniele Monteiro, Eliane de Lima Jacques, Elton Leme, Elsie
Franklin Guimarães, João Renato Stehmann, Ludovic
Kollmann, Marcus Nadruz, Nigel Taylor, Ricardo Perdiz,
Sheila Profice & Simon Mayo – and fieldwork helpers – José
Lima Paixão, Carlinhos, Lukas Daneu and Ricardo Perdiz.
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Received 01/08/2013
Accepted 14/02/2014
http://www.scielo.br/bn
http://dx.doi.org/10.1590/S1676-06032014001013