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Anathallis roseopapillosa (Orchidaceae Pleurothallidinae), a New Species from

the Central Amazon Region


Author(s): Edlley Pessoa, Jefferson J. Valsko, Santelmo Vasconcelos, Ana M. Benko-Isepon, and
Marccus Alves
Source: Systematic Botany, 39(4):-. 2014.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364414X683949

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Systematic Botany (2014), 39(4)


Copyright 2014 by the American Society of Plant Taxonomists
DOI 10.1600/036364414X683949
Date of publication September 3, 2014

Anathallis roseopapillosa (Orchidaceae Pleurothallidinae), a New Species


from the Central Amazon Region
Edlley Pessoa,1,5 Jefferson J. Valsko,2 Santelmo Vasconcelos,1 Ana M. Benko-Isepon,3 and Marccus Alves4
1

Programa de Pos-Graduacao em Biologia Vegetal, Departamento de Botanica, Universidade Federal de Pernambuco,


50670-901, Recife, Pernambuco, Brazil.
2
Programa de Pos-graduacao em Diversidade Biologica, Departamento de Botanica, Universidade Federal do Amazonas,
Av. Rodrigo Otavio, 2600, Manaus, Amazonas, Brazil.
3
Departamento de Genetica, Universidade Federal de Pernambuco, 50670-901, Recife, Pernambuco, Brazil.
4
Departamento de Botanica, Universidade Federal de Pernambuco, 50670-901, Recife, Pernambuco, Brazil.
Temporary address: Senckenberg Naturmuseum. 60325, Frankfurt am Main, Germany
5
Author for correspondence (edlley_max@hotmail.com)
Communicating Editor: Min Feng
AbstractA new species of Anathallis, A. roseopapillosa, is described from the central Brazilian Amazon. It is morphologically related to
A. gracilenta, but differs by having shorter racemes arising abaxially from the leaves, with flowers opening one at a time, petals with a
papillose apex, and an oblong-elliptic to ovate-elliptic lip, which is often obscurely tri-lobed. We also provide evidence of the phylogenetic
position of the new species in the subtribe Pleurothallidinae by means of a cladistic analysis of nuclear DNA sequence data (ITS).
KeywordsBrazil, monocots, Neotropics, phylogeny, South America.

The generic affiliation of the majority of the species of the


subtribe Pleurothallidinae, originally placed in Pleurothallis
Lindl., has been changed in recent years due to the use of
molecular approaches (Pridgeon et al. 2001; Pridgeon and
Chase 2001; Barros 2002, 2003; Barros and Rodrigues 2009;
Luer 2006, 2007, 2009; Chiron et al. 2012). The studies of
Pridgeon et al. (2001) and Chiron et al. (2012), using nuclear
DNA sequences, showed that several infrageneric taxa previously proposed for Pleurothallis by Luer (1986) were not
monophyletic groups.
Pridgeon and Chase (2001) reestablished some neglected
genera, such as Acianthera Scheidw., Anathallis Barb. Rodr.,
Phloeophila Hoehne & Schltr. Pleurothallopsis Porto & Brade,
and Specklinia Lindl. Later, Chiron et al. (2012) proposed
names under Pabstiella Brieger & Senghas as suggested
by Luer (2007) for species mainly placed in Stelis Sw. and
Specklinia by Pridgeon and Chase (2001). In both studies,
Anathallis was recognized as a monophyletic group, excluding A. barbulata (Lindl.) Pridgeon & M. W. Chase (grouped
within Trichosalpinx Luer) and A. rubens (Lindl.) Pridgeon &
M. W. Chase [= Stelis neorubens Chiron, as proposed by
Chiron et al. (2012)]. According to Chiron et al. (2012), the
genus is well supported and closely related to Lepanthes
Sw., Lepanthopsis (Cogn.) Ames, Thichosalpinx Luer, and
Zootrophion Luer.
Anathallis is a Neotropical genus with about 160 species (Luer and Toscano-de-Brito 2011; Govaerts et al. 2013)
and comprises the species originally placed in Pleurothallis
subg. Acuminatia Luer and Pleurothallis subg. Speklinia sect.
Muscosae Lindl. (Pridgeon and Chase 2001). According to
Barros et al. (2014), 81 species occur in Brazil, of which
67 species are endemic to the country. Moreover, only 12 species are cited for the Brazilian portion of the Amazon forest
(Barros et al. 2014), and one of them was recently described
(Pessoa and Alves 2012).
The Amazonian species of Anathallis were mostly
included in Panmorphia by Luer (2006), which is mainly
characterized by the small size of the plants, short ramicauls,
and the presence of two basal appendages at the base

of the lip. However, it is not a monophyletic group


(Chiron et al. 2012) and the names under this genus
should be avoided. In Brazil, among the Amazonian species, A. sclerophylla (Lindl.) Pridgeon & M. W. Chase is
the only one placed out of the Panmorphia group and related
to the type species of the genus, A. fasciculata Barb. Rodr.
[= A. obovata (Lindl.) Pridgeon & M. W. Chase]. It is part
of a group of larger plants (if compared with Panmorphia
species), with elongated ramicauls (as long as, or longer
than the leaf), often with long racemes, and the sepals are
commonly acuminate.
The aim of this study is to describe and illustrate a new
species of Anathallis from the Brazilian Amazon, which
is morphologically related to the A. angustilabia (Schltr.)
Pridgeon & M. W. Chase, A. obovata complex (Luer 1981).
We also discuss its morphological affinities and phylogenetic position in Pleurothallidinae by means of a cladistic
analysis with nuclear DNA sequence data from the internal transcribed spacer (ITS) region, comprising both ITS
sequences (ITS1 and ITS 2) and the 5.8 S rDNA subunit.

Materials and Methods


Morphological ObservationsThe morphological analysis and description of the new species were based on examination of several live and
dry (rehydrated) specimens, in addition to flowers in spirit collection.
The flowers were dissected and the perianth was mounted between
glass slides and cover slips using Entellan (Merck), and it was examined
under a stereomicroscope.
A key of the Anathallis species from Brazilian Amazon is provided;
it was produced based in specimens deposited in the most representative herbaria of northern Brazil [MG, INPA, IAN, RB, and HB, acronyms according to Thiers (2013)], the opus princeps of the species, and
type images available in virtual herbaria. Anathallis fastigiata (Luer &
Toscano) F. Barros & Barberena, a species described from Para, Brazil,
was not included because there is no material deposited in the visited
herbaria, and although Luer and Toscano-de-Brito (2002) cite the holotype in the HUEFS, we could not locate it. As explained by Barros
and Barberena (2010), according to the staff of this herbarium, apparently it was never deposited there.
Taxonomic SamplingSamples were collected in a NaCl-CTAB solution as described by Rogstad (1992) to perform the DNA extraction

SYSTEMATIC BOTANY
procedures. In addition, a total of 47 ITS sequences (45 from
Pleurothallidinae species and two from Dilomilis Raf. and Neocogniauxia
Schltr. (Laeliinae) species to be used as outgroup) were obtained from
the NCBI database (Appendix 1), to access the phylogenetic placement
of the new species in the subtribe Pleurothallidinae, as well as confirm its
position in Anathallis.
Molecular MethodsDNA extraction procedures were performed
with fresh leaf tissues of the new species, following the CTAB I protocol described by Weising et al. (2005), with minor modifications. The
PCR amplification of the nuclear DNA region comprising the ITS1, the
5.8 S rDNA and the ITS2 from total genomic DNA was performed

[Volume 39

according to Starr et al. (1999), using the forward primer ITS-L (Hsiao
et al. 1994) and the reverse primer ITS-4 (White et al. 1990). The PCR
products were purified with the AxyPrepPCR cleanup kit (Axygen,
Union City, California), following the manufacturers instructions. The
bidirectional DNA sequencing was performed using an ABI PRISM
3500 genetic analyzer (Applied Biosystems, Foster City, California) at
the Sequencing Platform LABCEN/CCB of the Universidade Federal
de Pernambuco (Recife, Brazil). Bidirectional sequencing reads were
edited and assembled with Geneious 6.1 (Biomatters Ltd., Auckland).
The alignment of the sequences was performed with the ClustalW algorithm (Larkin et al. 2007), with minor posterior manual adjustments.

Fig. 1. Bayesian inference of the phylogenetic position of Anathallis roseopapillosa E. Pessoa & J. J. Valsko, sp. nov., within the genus Anathallis
and their relationships with other Pleurothallidinae groups based on nuclear ITS sequences. Only posterior probability values 0.90 are shown close
to the nodes of the groups.

2014]

PESSOA ET AL.: A NEW ORCHID SPECIES FROM THE AMAZON

The sequence data-set was analyzed with a Bayesian Markov-chain


Monte Carlo (MCMC) approach (Yang and Rannala 1997), using
MrBayes 3.2 (Ronquist et al. 2012). The search for the sequence data
relied on the GTR + G model with the number of gamma categories set

to four (Yang 1993). Bayesian runs started from independent random


starting trees and were repeated four times. Markov chain Monte Carlo
runs extended for 10 million generations, with trees sampled every
1,000th generation.

Fig. 2. Anathallis roseopapillosa sp. nov. A. Habit. B. Detail of the inflorescence. C. Flower. D. Dissected perianth. E. Detail of the petal apex. F. Lip.
G. Column. (drawn from the holotype).

SYSTEMATIC BOTANY

Taxonomic Treatment
Anathallis roseopapillosa E. Pessoa & J. J. Valsko, sp.
nov.TYPE: BRAZIL. Amazonas: Manaus, Reserva
Porto Alegre-BDFF, 02 210 1900 S, 59 570 3100 W, 118 m
elev., 22 Nov. 2012, J. J. Valsko 125 (fl.) (holotype:
INPA!; isotypes: NY!, RB!, UFP!).

Anathallis roseopapillosa is morphologically related to


A. gracilenta (Luer & R. Vasquez) Pridgeon & M. W. Chase,
but differs by having shorter racemes arising abaxially
from the leaf with flowers opening one at a time (vs. longer
racemes arising adaxially from the leaf with 23 flowers
opening at a time), petals with papillose apex (vs. apex
smooth), and lip oblong-elliptic to ovate-elliptic, often
obscurely tri-lobed (vs. oblong-ligulate, entire).
Epiphytic, cespitose herb. Roots fasciculate, about 1.0 mm
wide. Rhizome 4.017.0 0.81.0 mm. Ramicaul 18.250.0
0.71.0 mm, cylindrical, unifoliate, concealed by 35, white
to gray, close-fitting sheaths; sheath membranaceous,
6.0 17.0 mm long, the apex acute to acuminate, striate,
margin entire. Annulus 1.01.7 mm long, cylindrical. Leaf
23.046.0
3.06.5 mm, lanceolate to elliptic, pale green,
coriaceous to slightly fleshy, glabrous, the apex minutely
tri-denticulate, the base attenuate to obtuse, margin entire,
11-nerved. Inflorescence terminal, sub-congested, successive
racemes, erect, arises abaxially from the leaf; peduncle
2.05.0 mm long; rachis 1.07.0 mm long; floral bracts
about 1.01.5 mm long, amplexicaul, membranaceous, the
apex acute to acuminate, margin entire. Flowers 15, opening
successively (one at a time); pedicellate ovary 0.81.0
0.2 mm; dorsal sepal 4.5 5.5
1.3 1.5 mm, pinkishcream, free, lanceolate, membranaceous, the apex acuminate, margin entire, 3-nerved; lateral sepals 4.55.5
0.81.0 mm, pinkish-cream, connate at base, lanceolate,
membranaceous, the apex acuminate, margin entire, 3-nerved,
forming a small mentum with the column foot; petals

Cladistic AnalysisThe DNA sequencing of the nuclear


ITS region of the new species generated a sequence with
627 bp (GenBank accession KF826902). The aligned matrix
containing 48 sampled species consisted of 650 characters,
of which 369 (56.7%) were variable and 264 (40.6%) were
informative, yielding, generally, well-supported groups in
the Bayesian inference (Fig. 1). In general, Bayesian posterior probabilities (PP) were higher than 0.95.
Figure 1 shows Anathallis as monophyletic, as well as a
major group which includes the allied genera Trichosalpinx,
Lepanthes, Lepanthopsis, and Zootrophion Luer, indicated by
good values of Bayesian posterior probabilities (PP of 1). On
the other hand, some internal support values are relatively
low (PP ranging from 0.900.93). Anathallis sertularioides
remained as sister of all the other Anathallis species, as
shown by Chiron et al. (2012) and Pridgeon et al. (2001).
The species placed in Panmorphia by Luer (2006) are
arranged in three different clades, confirming its identification as a paraphyletic group, as pointed out by Chiron
et al. (2012). The group formed by the new species (referred to
as Anathallis roseopapillosa in the taxonomic treatment below),
A. citrina, A. heterophylla, and A. sclerophylla emerges with
relatively low support (PP of 0.93).

2.0 2.5
0.40.5 mm, whitish with a purple line, free,
elliptic, sub-falcate, membranaceous, the apex acuminate,
papillose, margin entire in proximal half, the distal half
ciliate, 1-nerved; lip 1.3 1.6
0.5 0.7 mm, purple, free,
oblong-elliptic to ovate-elliptic, often obscurely 3-lobed, apex
obtuse to rounded, with two basal sub-orbicular appendages
at base, margin entire, shortly papillose, 3-nerved, articulated with the column foot; column about 1.31.5 mm long,
broadly winged, clinandrium 3-dentate, erose, obtuse apex,
denticulate margin, column foot about 0.1 mm long. Fruits
were not seen. Figure 2.
EtymologyThe new species was named for the color
of the flowers and the papillae at the apex of the petals.
Distribution, Habitat and EcologyAnathallis roseopapillosa
is known from the vicinities of Manaus, central Brazilian
Amazon. The type locality (Biological Dynamics of Forest
Fragments Projects BDFF) is a stretch of dense rainforest
about 80 km north of Manaus, Amazonas, Brazil (Fig. 3).
The annual average rainfall is 1,9003,500 mm (Laurance
2001) and average temperature is 27 C (Bierregaard et al.
1992; Lovejoy and Bierregaard 1990). The soils are mostly
poor in nutrients, composed of yellow oxisols with high
clay content (Chauvel 1983). The type locality experiences
periodic flooding for a short period, usually between the
months of March to June, and forming small isolated
ponds within the forest. The understory is composed of
acaulous palms, small trees and epiphytes indicating high
local humidity. Individuals of A. roseopapillosa were found
in trees about 25 m tall, recently fallen. Specimens of
Trichosalpinx orbicularis (Lindl.) Luer, Stelis argentata Lindl.,
and Specklinia grobyi (Bateman ex Lindl.) F. Barros were
also found nearby. The flowering period is poorly known,
but based on the analyzed samples, flowers can be found
from November to May.
Morphological AffinitiesAnathallis roseopapillosa is morphologically related to species of the A. angustilabia A. obovata
complex (Luer 1981), a group of species treated in Pleurothallis
subgen. Acuminatia by Luer (1999). It is distinguished from
the group of A. sclerophylla by its longer and generally acuminate petals (vs. shorter and acute, cuspidate, obtuse or

Results and Discussion

[Volume 39

Fig. 3. Map of the type localities of Anathallis gracilenta (square) and


A. roseopapillosa (circle).

2014]

PESSOA ET AL.: A NEW ORCHID SPECIES FROM THE AMAZON

rounded petals), and erose to denticulate clinandrium margin


(vs. entire clinandrium). It has short successive racemes
which is rare among the species in this taxonomic complex,
which often have racemes as long as the leaves [such as
A. angustilabia and A. citrina (Schltr.) Pridgeon & M. W.
Chase]. Among the species with short inflorescences,
A. gracilenta, endemic to Bolivia, is the most morphologically related to A. roseopapillosa, especially because of the
narrower leaves (if compared with the other species in
this group). However, the new species is distinguished
by having shorter racemes arising abaxially from the leaf
with flowers opening one at a time (vs. longer racemes
arising adaxially in the leaf with 23 flowers opening
at a time), petals with papillose apex (vs. apex smooth),
and oblong-elliptic to ovate-elliptic, often obscurely 3-lobed
(vs. oblong-ligulate) lip. The flowers of A. gracilenta are
whitish with yellow apices and a whitish lip, while in
A. roseopapillosa they are pinkish-cream with a purple lip
(Table 1).
Two specimens collected 20 yr ago and nearby the type
locality were previously identified as A. taracuana (Schltr.)
F. Barros & Barberena. However, this species differs from
A. roseopappilacea by the sizes of the ramicaul (> 80.0 mm

Table 1. Diagnostic differences between Anathallis gracilenta and


A. roseopapillosa.
Character

A. gracilenta

A. roseopapillosa

Inflorescence size
Inflorescence position
No. of flowers
open at a time
Flower color

10.015.0 mm long
Adaxial from the leaf
24

3.012.0 mm long
Abaxial from the leaf
1

Whitish with
yellow apices
Not papillose
Oblong-ligulate

Pinkish-cream

Petal apex
Lip morphology

Papillose
Oblong-elliptic to
ovate-elliptic,
often obscurely
tri-lobed

long vs. < 50.0 mm long) and the peduncle of the inflorescence (ca. 23.0 mm long vs. < 5.0 mm long), shape and size
of the leaves (oblanceolate to oblong and > 65.0 mm long
vs. lanceolate to elliptic and < 50.0 mm long), and shape of
the sepals (oblong vs. lanceolate).
Representative Specimens ExaminedBRAZIL. Amazonas: Manaus,
Reserva Ducke, 02 530 S, 59 580 W, 3 Nov. 1994, J.E.L.Ribeiro et al. 1485,
fl. (INPA); ibid., 39 May 1996, J.E.L.Ribeiro 1832, fl. (INPA).

Key to Species of ANATHALLIS FROM THE BRAZILIAN AMAZON


1.

1.

Lateral sepals completely connate or up to the 4/5 of the length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2


2. Apex of the petals and lip glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. imberbis (Luer & Hirtz) Luer
2. Apex of the petals and lip ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. barbulata
Lateral sepals free or connate only at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Ramicauls > 80.0 mm long; leafs > 65.0 20.0 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Rachis > 50 mm long; flowers creamy, sepals > 15.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sclerophylla
4. Rachis < 15.0 mm long; flowers purple, sepals < 10.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. taracuana
3. Ramicauls 50.0 mm long, leafs < 50.0 15.0 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Ramicauls as long as or longer than the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Sepals apex acuminate; apex of petals and lip papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. roseopapillosa
6. Sepals apex acute; apex of petals and lip smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. brevipes (H.Focke) Pridgeon & M. W.Chase
5. Ramicauls shorter than the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Petals broadly elliptic or obovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Cespitose herb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. funerea (Barb.Rodr.) F. Barros & Barberena
8. Sub-repent herb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Petals with acute apex, lip 3.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. polygonoides (Griseb.) Pridgeon & M. W.Chase
9. Petals with obtuse apex, lip 2.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nanifolia (Foldats) F. Barros & Barberena
7. Petals narrowly elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Inflorescence with inconspicuous peduncle, lip entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. amazonica E. Pessoa & M. Alves
10. Inflorescence long pedunculate (10.040.0 mm long), lip minutely trilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Leaves with purple punctuations; flowers orangish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. humilis (C. Schweinf.) Luer
11. Leaves without punctuations; flowers whitish . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sertularioides (Sw.) Pridgeon & M. W.Chase

Acknowledgments. This work was supported by the following


agencies: CNPq (Conselho Nacional de Desenvolvimento Cientfico e
Tecnologico) and CAPES (Coordenacao de Aperfeicoamento de Pessoal
de Nvel Superior). The authors thank the Sequencing Platform LABCEN/
CCB of the Universidade Federal de Pernambuco for the use of its facilities, the PDBFF (Projeto Dinamica Biologica de Fragmentos Florestais) for
the support for fieldwork, Regina Carvalho for the line drawing of the
new species and Scott Heald for the English review of the manuscript.

Literature Cited
Barros, F. 2002. Notas nomenclaturais em Pleurothallidinae (Orchidaceae),
principalmente Brasileiras. Bradea 8: 293297.
Barros, F. 2003. Notas taxonomicas sobre especies brasileiras dos generos
Catasetum, Isabelia, Veyretia, Acianthera e Anathallis (Orchidaceae).
Hoehnea 30: 181191.
Barros, F., F. Vinhos, V. T. Rodrigues, F. F. V. A. Barberena, C. N. Fraga,
E. M. Pessoa, and W. Forster. 2014. Orchidaceae in Lista de especies
da flora do Brasil. Jardim Botanico do Rio de Janeiro. Available in

http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB179. Accessed
on: 12 May 2014.
Barros, F. and F. F. V. A. Barberena. 2010. Nomenclatural notes and new
combinations on Anathallis and Specklinia (Orchidaceae). Rodriguesia
61: 127131.
Barros, F. and V. T. Rodrigues. 2009. Nomenclatural notes and new combinations on Acianthera Scheidw. and Specklinia Lindl. (Orchidaceae).
Bradea 14: 2126.
Bierregaard, R. O., T. Lovejoy, V. Kapos, A. dos Santos, and R. Hutchings.
1992. The biological dynamics of tropical rain forest fragments.
Bioscience 42: 859866.
Chauvel, A. 1983. Os latossolos amarelos, alicos, argilosos dentro dos
ecossistemas das bacias experimentais do INPA e da regiao vizinha.
Acta Amazonica 12: 4760.
Chiron, G. R., J. Guiard, and C. van den Berg. 2012. Phylogenetic relationships in Brazilian Pleurothallis sensu lato (Pleurothallidinae,
Orchidaceae): evidence from nuclear ITS rDNA sequences. Phytotaxa
46: 3458.
Govaerts, R., J. Pfahl, M. A. Campacci, D. H. Baptista, H. Tigges, J.
Shaw, P. Cribb, A. George, K. Kreuz, and J. Wood. 2013. World

SYSTEMATIC BOTANY
checklist of Orchidaceae. Kew: Royal Botanic Gardens. Available
in http://apps.kew.org/wcsp.
Hsiao, C., N. J. Chatterton, K. H. Asay, and K. B. Jensen. 1994.
Phylogenetic relationships of 10 grass species: an assessment of
phylogenetic utility of the internal transcribed spacer region in
nuclear ribosomal DNA in Monocots. Genome 37: 112120.
Karremans, A. P., F. T. Bakker, F. Pupulin, R. Solano-Gomez, and
M. J. M. Smulders. 2013. Phylogenetics of Stelis and closely
related genera (Orchidaceae: Pleurothallidinae). Plant Systematics
and Evolution 299: 151176.
Larkin, M. A., G. Blackshields, N. P. Brown, R. Chenna, P. A. McGettigan,
H. McWilliam, F. Valentin, I. M. Wallace, A. Wilm, R. Lopez, J. D.
Thompson, T. J. Gibson, and D. Higgins. 2007. Clustal W and
Clustal X version 2.0. Bioinformatics 23: 29472948.
Laurance, W. F. 2001. The hyper-diverse flora of the central Amazon:
an overview. Pp 4753 in Lessons from Amazonia: Ecology and conservation of a fragmented forest. eds. R. O. Bierregaard, C. Gascon,
T. E. Lovejoy, R. Mesquita. New Haven: Yale University Press.
Lovejoy, T. E. and R. O. Bierregaard. 1990. Central Amazonian forest
and the minimum critical size of ecosystems project. Pp. 6071
in Four Neotropical rainforests. ed A. H. Gentry. YNew Haven: ale
University Press.
Luer, C. A. 1981. Miscellaneous new species and combinations in the
Pleurothallidinae (Orchidaceae). Phytologia 49: 197240.
Luer, C. A. 1986. Icones Pleurothallidinarum III: Systematics of the
Pleurothallis (Orchidaceae). Monographs in Systematic Botany from the
Missouri Botanical Garden 20: 1109.
Luer, C. A. 1999. Icones Pleurothallidinarum XVIII. Systematics of
Pleurothallis Subgenus Pleurothallis Section Pleurothallis, Subsection
Antenniferae, Subsection Longiracemosae, Subsection MacrophyllaeRacemosae, Subsection Perplexae, Subgenus Pseudostelis, Subgenus
Acuminatia (Orchidaceae). Monographs in Systematic Botany from the
Missouri Botanical Garden 76: 1182.
Luer, C. A. 2006. Icones Pleurothallidinarum XXVIII. Reconsideration
of Masdevallia and the systematics of Specklinia and vegetatively
similar taxa. Monographs in Systematic Botany from the Missouri
Botanical Garden 105: 1300.
Luer, C. A. 2007. Icones Pleurothallidinarum XXIX: a third century of
Stelis of Ecuador. Systematics of Apoda-Prorepentia. Systematics
of miscellaneous small genera addenda: New genera, species, and
combinations (Orchidaceae). Monographs in Systematic Botany from
the Missouri Botanical Garden 112: 119121.
Luer, C. A. 2009. Icones Pleurothallidinarum XXX. Lepanthes of Jamaica
and systematics of Stelis. Stelis of Ecuador, part four and addenda:
systematic of Masdevallia, new species of Lepanthes from Ecuador,
and miscellaneous new combinations. Monographs in Systematic
Botany from the Missouri Botanical Garden 115: 1265.
Luer, C. A. and A. L. V. Toscano de Brito. 2002. Miscellaneous new
species of Pleurothallidinae from Brazil. Selbyana 23: 181195.
Luer, C. A. and A. L. V. Toscano de Brito. 2011. Miscellaneous new
species and combinations in the Pleurothallidinae (Orchidaceae)
from Brazil and Argentina. Harvard Papers in Botany 16: 361382.
Pessoa, E. and M. Alves. 2012. A new Anathallis (Orchidaceae:
Pleurothallidinae) from the Brazilian Amazon. Phytotaxa 73: 1316.
Pridgeon, A. M. and M. W. Chase. 2001. A phylogenetic reclassification
of Pleurothallidinae (Orchidaceae). Lindleyana 16: 235271.
Pridgeon, A. M., R. Solano, and M. W. Chase. 2001. Phylogenetic relationships in Pleurothallidinae (Orchidaceae): combined evidence
from nuclear and plastid DNA sequences. American Journal of Botany
88: 22862308.
Rogstad, S. H. 1992. Saturated NaCl-CTAB solution as a means of field
preservation of leaves for DNA analyses. Taxon 41: 701708.
Ronquist, F., M. Teslenko, P. van-der-Mark, D. L. Ayres, A. Darling,
S. Hohna, B. Larget, L. Liu, M. A. Suchard, and J. P. Huelsenbeck.
2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and
model choice across a large model space. Systematic Biology 61:
539542.
Starr, J. R., R. J. Bayer, and B. A. Ford. 1999. The phylogenetic position
of Carex section Phyllostachys and its implications for phylogeny

[Volume 39

and subgeneric circumscription in Carex (Cyperaceae). American


Journal of Botany 86: 563577.
Thiers, B. 2013. [continuously updated] Index herbariorum: A global
directory of public herbaria and associated staff. New York Botanical
Garden. http://sweetgum.nybg.org/ih.
Weising, K., H. Nybom, K. Wolff, and G. Kahl. 2005. DNA fingerprinting
in plants: Principles, methods, and applications, 2nd ed. Boca Raton:
CRC Press.
White, T. J., T. Bruns, S. Lee, and J. Taylor. 1990. Amplification and
direct sequencing of fungal ribosomal RNA genes for phylogenetics. Pp 315322 in PCR protocols: A guide to methods and application, eds. M. Innis, D. Gelfand, J. Sninsky, and T. White. San
Diego: Academic Press.
Yang, Z. 1993. Maximum-likelihood estimation of phylogeny from
DNA sequences when substitution rates differ over sites. Molecular
Biology and Evolution 10: 13961401.
Yang, Z. and B. Rannala. 1997. Bayesian phylogenetic inference using
DNA sequences: a Markov chain Monte Carlo method. Molecular
Biology and Evolution 14: 717724.
Appendix 1. Voucher information is listed as follows: taxon name,
synonym (marked with asterisks and cited only for species that had
been previously registered in GenBank with that name), collector name
and number (herbarium) or related publication [Pridgeon et al. 2001
(as P), Chiron et al. 2012 (as C), Karremans et al. 2013 (as K)],
GenBank accessions (ITS).
Acianthera papillosa (Lindl.) Pridgeon & M. W. Chase. (C) JQ306501.
Acianthera prolifera (Herb. ex Lindl.) Pridgeon & M. W. Chase.
*Pleurothallis prolifera, (P) AF275688. Acianthera saundersiana (Rchb. f.)
Pridgeon & M. W. Chase. (C) JQ306452. Acianthera sicaria (Lindl.)
Pridgeon & M. W. Chase. *Pleurothallis sicaria, (P) AF262848. Anathallis
adenochila (Loefgr.) F. Barros. (C) JQ306490. Anathallis aristulata
(Lindl.) Luer. (C) JQ306338. Anathallis citrina (Schltr.) Pridgeon &
M. W. Chase. (C) JQ306498. Anathallis corticicola (Schltr. ex Hoehne)
Pridgeon & M. W. Chase. *Pleurothallis corticicola, (P) AF262870. Anathallis
heterophylla Barb. Rodr. (C) JQ306339. Anathallis linearifolia (Cogn.)
Pridgeon & M. W. Chase. *Pleurothallis linearifolia, (P) AF262869. Anathallis
petropolitana (Hoehne) Luer & Toscano. (C) JQ306491. Anathallis
roseopapillosa E. Pessoa & J. J. Valsko. J. J. Valsko 125 (INPA) KF826902.
Anathallis sclerophylla (Lindl.) Pridgeon & M. W. Chase. (C) JQ306348.
Anathallis sertularioides (Sw.) Pridgeon & M. W. Chase. *Pleurothallis
sertularioides, (P) AF262871. Barbosella cogniauxiana (Speg. & Kraenzl.)
Schltr. (C) JQ306503. Barbosella orbicularis Luer. (P) AF262814.
Brachionidium valerioi Ames & C. Schweinf. (P) AF262913. Dilomilis
montana (Sw.) Summerh. (P) AF262915. Dracula bella (Rchb. f.) Luer.
(P) AF262760. Dryadella edwallii (Cogn.) Luer. (P) AF262824. Lepanthes
felis Luer & R. Escobar. (P) AF262891. Lepanthopsis astrophora
(Rchb. f. ex Kraenzl.) Garay. (P) AF262893. Lepanthopsis floripecten
(Rchb. f.) Ames. (C) JQ306336. Masdevallia venezuelana H. R. Sweet.
(P) AF262782. Myoxanthus exasperatus (Lindl.) Luer. (P) AF262882.
Myoxanthus lonchophyllus (Barb. Rodr.) Luer. (P) AF262884. Neocogniauxia
hexaptera (Cogn.) Schltr. (P) AY429385. Octomeria gracilis Lodd. ex
Lindl. (P) AF262911. Octomeria lithophila (Barb. Rodr.) Barb. Rodr.
(P) AF262912. Pabstiella lingua (Lindl.) Luer. (C) JQ306386. Pabstiella
uniflora (Lindl.) Luer. (C) JQ306469. Pabstiella yauaperyensis (Barb.
Rodr.) F. Barros. *Pleurothallis mentosa, (P) AF262864. Phloeophila
peperomioides (Ames) Garay *Pleurothallis peperomioides, (P) AF275690.
Platystele stenostachya (Rchb. f.) Garay. (P) AF262821. Pleurothallis
cardiantha Rchb. f. (P) AF262832. Pleurothallis ruscifolia (Jacq.) R. Br.
(P) AF262836. Porroglossum amethystinum (Rchb. f.) Garay. (P) AF262804.
Scaphosepalum grande Kraenzl. (P) AF262819. Specklinia grobyi (Bateman
ex Lindl.) F. Barros (P) JQ306388. Stelis argentata Lindl. (P) JQ306421.
Stelis deregularis Barb. Rodr. (K) JF934771. Stelis gelida (Lindl.)
Pridgeon & M. W. Chase. (K) JF934780. Stelis velaticaulis (Rchb. f.)
Pridgeon & M. W. Chase. *Plerothallis velaticaulis, (P) AF262847.
Trichosalpinx egleri (Pabst) Luer. (C) JQ306347. Trichosalpinx orbicularis
(Lindl.) Luer. (P) AF262886. Trisetella triglochin (Rchb. f.) Luer. (P)
AF262807. Zootrophion atropurpureum (Lindl.) Luer. (P) JQ306415.
Zootrophion hirtzii Luer. (P) AF262897.

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