NEW SPECIES AND NOMENCLATURAL NOTES IN THE
PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
A. L. V. ToscAno de BriTo1, 2 And cArLyLe A. Luer3
Abstract. One new species in the genus Acianthera, A. imitator, is described and illustrated. Sixteen species are proposed as synonyms.
They are listed in alphabetical order: Acianthera gradeae as synonym of A. agathophylla; A. spilantha as synonym of A. brevilora;
A. antennata as synonym of A. gracilisepala; A. dichroa as synomym of A. strupifolia; Anathallis crebrifolia, A. montipelladensis, and
A. ourobranquensis as synonyms of A. aristulata; A. liparanges as a synomym of A. heterophylla; A. bolsanelloi, A. githaginea, and
A. nectarifera as synonyms of A. lobiserrata; A. limbata and A. marginata as synonyms of A. muscoidea; A. longiglossa as synonym of
A. paranaensis; Lankesteriana gehrtii as synonym of L. caudatipetala, and Specklinia rubidantha as synonym of Lankesteriana
imberbis. Correct provenance and habitat information are provided for the type collection of the recently described Anathallis johnsonii.
This species is herein formally recorded for Brazil. Lectotypes are selected for seven species: Anathallis heterophylla, A. nectarifera,
Lepanthes crebrifolia, Pleurothallis gracilisepala, P. microgemma, P. paranaensis, and P. spilantha, and an epitype is selected for
Pleurothallis paranaensis. Illustrations and taxonomic discussions are also provided.
Keywords: Acianthera, Anathallis, Lankesteriana, Lepanthes, Pleurothallis, Specklinia.
In the course of preparing generic treatments of
Pleurothallidinae of Brazil, material of several genera
was examined and their morphologies and nomenclature
elucidated. One species is described as new and sixteen
are proposed as synonyms. Illustrations, updated synonym
lists, typiications, and taxonomic discussions are provided.
Genera and species follow in alphabetical order.
new species
Acianthera imitator Toscano, Luer & L.Kollmann, sp. nov.
lowered peduncle 3–4 mm long, borne from the base of the
TYPE: BRAZIL. Espírito Santo: Santa Maria do Jetibá,
leaf with a spathe 2 mm long; loral bract 2–2.5 mm long;
collected and cultivated by Ludovic Kollmann, lowered
pedicel 3 mm long with a minute ilament; ovary 1.5 mm
in cultivation in Santa Teresa, 15 July 2010, A. Toscano
long; sepals leshy, glabrous, the dorsal sepal greenish with
de Brito 2866 (Holotype: MBML [43914]), C. Luer illustr.
dark purple veins, elliptical-oblong, obtuse to rounded at
21679. Fig. 1.
the tip, 9 mm long, 3 mm wide, 5-veined, the lateral sepals
Plant small, epiphytic, long-repent, the rhizome slender,
adaxially spotted with dark purple, connate into an ovate,
15–20 mm long between ramicauls; roots slender. Ramicauls
obtuse, minutely bidentate, concave synsepal, 8 mm long,
ascending, erect, 5–8 mm long, enclosed by a tubular sheath
6 mm wide expanded, 6-veined, concave below the middle;
from below the middle and another sheath at the base. Leaf
petals translucent-green, elliptical, acute, microscopically
erect, suberect to horizontal, thickly coriaceous, elliptical to
serrulate above the middle, 3.5 mm long, 1.5 mm wide,
ovate, obtuse to subacute at the apex, 20–30 mm long, 12–15
3-veined; lip dark purple with green lateral lobes, thick,
mm wide, the base rounded, sessile. Inlorescence a singleoblong-subtrilobed, 4 mm long, 2.5 mm wide, markedly
This paper is part of the project “The Pleurothallid Orchids of Brazil—Contributions to an inventory and an understanding of evolution, ecology and
conservation,” until recently partially sponsored by the National Geographic Society’s Committee for Research and Exploration, grant nr. 8906-11.
It is currently funded by the Marie Selby Botanical Gardens and a grant nr. PVE 88881.065009/2014-01 from CAPES, Brazil, under the subproject title
“Estudos Filogenéticos e Taxonômicos em Pleurothallidinae e Oncidiinae—Clado Ornithocephalus (Orchidaceae).” We acknowledge the services of
Coordenação de Proteção e Conservação Florestal, Diretoria de Uso Sustentável da Biodiversidade e Florestas, IBAMA, Brazil, and the Plant Inspection
Station at John F. Kennedy International Airport, New York, U.S.A., for issuing the CITES permits which allowed the exchange and transportation of some
specimens treated here. The authors would like to thank the following organizations, which, during recent years, have provided funds and facilities that
enabled study in the libraries and of historical collections deposited in several European and American herbaria, undertaking of ieldwork in Brazil as well
as molecular research: the Kew Latin American Research Fellowships Programme at the Royal Botanic Gardens, Kew, the Swiss Orchid Foundation at the
University of Basel, the David Rockefeller Center for Latin American Studies at Harvard University, the Harvard University Herbaria, the Missouri Botanical
Garden, the National Geographic Society, the Institute of Museum and Library Services (IMLS), and the Sarasota Orchid Society. Financial support
received from the following individuals is also acknowledged: Rudolf Jenny, Dora Gerhard, and Peter and Gail Furniss. We thank the folowing herbaria
for the privilege of visiting their collections and permission to reproduce images of herbarium specimens: AMES, BM, BR, C, HB, K, LE, P, MO, MBML,
NY, RB, SEL, SP, UPCB, and W. We are also grateful to Ronaldo Herculano Alves, Maria Rita Cabral, and Maria do Rosário de Almeida Braga for
providing material of Acianthera agathophylla for study; Luiz Filipe Varella for permission to reproduce some of his photographs and for providing
material for study; Leslie Garay for helping to interpret Reichenbach’s collections and handwriting, and also for sending literature; Thiago Meneguzzo
for assistance in locating literature and specimens at RB; Rudolf Jenny for sending literature; Eric Smidt for sending images and for helping
prepare specimens for UPCB; Nancy Karam and Wade Collier for help in assembling the illustrations; Stig Dasltröm for inking the illustrations, and the
Pleurothallidinae Alliance for making it possible.
1
Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, Florida 34236-7726 U.S.A.
2
Author for correspondence: atoscano@selby.org
3
Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110, U.S.A. Address for correspondence: 3222 Old Oak Drive, Sarasota,
Florida 34239-5019 U.S.A.; cluer@juno.com
Harvard Papers in Botany, Vol. 20, No. 1, 2015, pp. 39–68.
© President and Fellows of Harvard College, 2015
ISSN: 1938-2944, DOI: 10.3100/hpib.v20iss1.2015.n4, Published online: 30 June 2015
40
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 1. Acianthera imitator Toscano, Luer & L.Kollmann, sp. nov. Based on the holotype, A. Toscano de Brito 2866 (MBML). Drawn
by C. A. Luer.
verrucose, serrulate, narrowly rounded and thickened at
the apex, the lobes below the middle, small, erect, obtuse
to rounded, the disc narrowly channeled between a pair of
verrucose calli on the middle third, the base truncate with a
minute, pedunculated lobule at the corners; column green,
semiterete, narrowly winged toward the purple spotted
apex, 3 mm long, anther, rostellum and stigma ventral, the
foot thick, 1.5 mm long.
Etymology: From the Latin imitator, “an imitator,”
referring to its similarity to Barbosa Rodrigues’s
Pleurothallis bidentula [= Acianthera bidentula (Barb.
Rodr.) Pridgeon & M.W.Chase].
This new species belongs to a complex of longrepent, similar taxa recently referred to as the Acianthera
saundersiana (Rchbf.) Pridgeon & M.W.Chase complex
(Toscano de Brito and Luer, 2015). It resembles in habit
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
Acianthera bidentula, but it is distinguished by a markedly
verrucose lip that is narrowly rounded at the tip, instead
of broadly rounded or truncate. The creeping rhizome is
delicate and the leaves are elliptical to slightly ovate. The
lowers are relatively smaller and more delicate than those
of A. bidentula. In shape and color, the lip of A. imitator
41
resembles that of A. saundersiana, but the latter species
is a much larger one with longer ramicauls and larger
lowers. The reader should refer to Toscano de Brito and
Luer (2015) for further discussion on the identities of some
species in this complex, especially that of A. bidentula and
A. saundersiana.
nomencLATurAL noTes
Acianthera agathophylla (Rchb.f.) Pridgeon &
base and apex, may vary even in the same individual.
Acianthera agathophylla is a variable species, both
M.W.Chase, Lindleyana 16: 241. 2001. Fig. 2.
vegetatively and in loral morphology, but it can be
Basyonym: Pleurothallis agathophylla Rchb.f., Xenia
easily recognized by its repent rhizome, ascending
Orchid. 3: 25. 1881. TYPE: BOLIVIA. La Paz:
ramicauls, and spreading, broad elliptical to suborbicLarecaja, vinicinity of Sorata, Cerro del Iminapi,
ular leaves. The inlorescence is a short, few-lowered
alt. 2600 m, March–May 1860, G. Mandon 1132
raceme at the base of the leaf and produces 1 to 4
(Holotype: W [45938]; Isotypes: AMES [118553],
orange-yellow lowers. Most distinctive is a darker,
BM [82439], G [24887], K [584177], MPU [18064,
convex thickening above the middle of the sepals.
photo seen], NY [9220], P [489101], P [489102], P
The Brazilian collection that we studied comes from
[489103]).
the municipality of Apiacás, in the state of Mato
Synonyms: Humboldtia agathophylla (Rchb.f.) Kuntze,
Grosso, near the border with the state of Pará. It was
Revis. Gen. Pl. 2: 667. 1891.
collected by Ronaldo Herculano Alves and lowered
Pleurothallis cyclophylla Luer, Selbyana 3: 92. 1976.
in cultivation by Maria Rita Cabral in 2012. This same
TYPE: BOLIVIA. Santa Cruz: near Samaipata, alt.
collection was photographed by Luiz Filipe Varella
1500 m, lowered in cultivation at J and L Orchids,
and a couple of his photographs are reproduced
Easton, CT, 20 October 1976, F. Fuchs s.n. (Holotype:
here (Fig. 2A–B). Several others can be found at:
SEL [002781]), C. Luer illustr. 249-S.
http://www.flickr.com/photos/luizfilipevarella/sets/
Pleurothallis nakatae T.Hashim., Ann. Tsukuba Bot.
72157620101297689. Duplicates of this collection
Gard. 4: 3. 1986. TYPE: PERU. Junín: Chanchamayo,
are currently in cultivation at Rio de Janeiro Botanical
near La Merced, alt. 1700–1800 m, lowered in
Garden (Fig. 2C).
cultivation in TBG, 8 May 1985, M. Nakata 626
Additional specimen examined: BRAZIL. Mato
(Holotype: TNS [not seen]).
Grosso: Apiacás, collected by R.H. Alves s.n., lowered in
Acianthera gradeae Chiron & Benelli, Richardiana
cultivation in 25 August 2012, M.R. Cabral 6 (MBM).
13: 66. 2012, syn. nov. TYPE: BRAZIL. Mato
Grosso: Nova Bandeirante, Fazendinha da Lagoa, alt.
Acianthera brevilora (Lindl.) Luer, Monogr. Syst. Bot.
304 m, A. Petini-Benelli, A. Grade and & H.S. Lima
Missouri Bot. Gard. 95: 253. 2004. Fig. 3.
APB963 (Holotype: HERBAM [4953, not seen]);
Basionym: Pleurothallis brevilora Lindl., Edwards’s Bot.
same locality, June 2005, lowered in cultivation in
Reg. 27(Misc.): 59. 1841. TYPE: MEXICO (in
September 2005, A. Petini-Benelli, A. Grade and &
error, most likely from Brazil): imported by Messrs.
Loddiges s.n. (Holotype: K [000885713]).
H.S. Lima APB964 (Paratype: HERBAM [4954, not
Synonyms: Humboltia brevilora (Lindl.) Kuntze, Revis.
seen].
Gen. Pl. 2: 667. 1891.
Acianthera agathophylla is a common species distribPleurothallis spilantha Barb.Rodr., Gen. Spec. Orchid.
uted from central Peru through Bolivia (Váquez &
2: 32. 1881, syn. nov. TYPE: BRAZIL. Minas Gerais:
Dodson 1982; Váquez & Ibisch, 2000; Luer, 2004)
Serra das Bicas, alt. 600 m, J. Barbosa Rodrigues s.n.
and has been recently collected in the Brazilian state
[Holotype: Lost; Lectotype here designated: Barbosa
of Mato Grosso. It was redescribed as Acianthera
Rodrigues’s original illustration in Iconogr. Orchid.
gradeae, in 2012, based on two collections from the
Brésil, vol. 3, tab. 173, at the Library of Rio de Janeiro
municipality of Nova Bandeirantes, also in the state of
Botanical Garden, cited as tab. 752 (unpubl.) in Barb.
Mato Grosso, but this name is illegitimate and it is here
Rodr. loc.cit; copied and reprod. in black and white
placed in the synonym of Acianthera agathophylla.
in Cogn., Fl. Bras. (Mart.) 3(4), tab. 106, ig. 1. 1896;
This synonymy has been alluded to in some nonreprod. in color in Sprunger et al. (1996), vol. 1: 231,
scientiic websites (e.g. http://pt.wikipedia.org/wiki/
ig. A].
Acianthera) without provision of supporting evidence.
Myoxanthus spilanthus (Barb.Rodr.) Luer, Monogr.
Chiron and Petini-Benelli (2012) distinguished
Syst. Bot. Missouri Bot. Gard. 15: 38. 1986, syn. nov.
Acianthera gradeae from A. agathophylla based on
Pleurothallis paspaliformis Loefgr., Arch. Jard.
a number of characters such as rhizome length, leaf
Bot. Rio de Janeiro 2: 53. 1918, syn. nov. TYPE:
shape, leaf base and apex, spathe size, petals and lip
BRAZIL. Rio de Janeiro: Itatiaia, P. Campos Porto
shape, type of margins, lip surface, and column
s.n. (Holotype: RB [14168].
thickness. However, none of these features can be
used to distinguish these two taxa because they are
Acianthera spilantha (Barb.Rodr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 118. 2007, syn. nov.
variable characters and some of them, e.g. leaf shape,
42
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 2. Acianthera agathophylla (Rchb.f.) Pridgeon & M.W.Chase. A–B. Photographed by Luiz Filipe Varella based on a plant in
cultivation in Porto Alegre, Rio Grande do Sul, Brazil. C. Photographed by Wade Collier based on a plant in cultivation at Rio de Janeiro
Botanical Garden. All based on duplicate specimens of M.R. Cabral 6 (MBM).
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
43
Figure 3. Acianthera brevilora (Lindl.) Luer. Holotype of Pleurothallis brevilora Lindl. at K showing watercolor by John Lindley on the
type sheet. © The Board of Trustees of the Royal Botanic Gardens, Kew. Reproduced with permission.
44
HARVARD PAPERS IN BOTANY
Acianthera spilantha (Barb.Rodr.) Campacci Bol.
CAOB 69–70: 25. 2008, nom. illeg.
John Lindley described Pleurothallis brevilora [=
Acianthera brevilora] based on a specimen said to have been
imported from Mexico by the Loddiges family. No lowers
remain on the type specimen preserved at K (Fig. 3) but a
ine watercolor prepared by Lindley can be found on the
herbarium sheet. This painting and the sterile specimen at
Kew agree in every detail with Barbosa Rodrigues’s original
illustration of Pleurothallis spilantha, a frequent species in
the Atlantic forest of southeast Brazil. It also agrees with the
type specimen of Pleurothallis paspaliformis. Therefore,
these two taxa are here placed in the synonym of Acianthera
brevilora. This casts doubt on the Mexican origin of the
specimen imported by the Loddiges, which most certainly
came from Brazil. An illustration based on a collection
by Martin Martens, probably from Brazil, was published
by Luer in 2004. As in the case of Acianthera gradeae,
previously treated here, this synonymy has also been alluded
to in some non-scientiic websites (e.g. http://pt.wikipedia.
org/wiki/Acianthera) without providing further evidence.
The type specimen of Pleurothallis spilantha is lost
and the only extant original material is the illustration
that appeared in Barbosa Rodrigues’s Iconographie des
orchidées du Brésil, which is now deposited in the library
of Rio de Janeiro Botanical Garden. This illustration was
copied and reproduced in black and white in Cogniaux
(1896) and reproduced in color in Sprunger et al. (1996). It
is selected here as lectotype.
Additional specimens examined: BRAZIL. Espírito
Santo: Vargem Alta, 650 m, 3 June 1949, A.C. Brade 19966
(RB). Minas Gerais: without collection data, cultivated in
Poços de Caldas, 1 March 2004, Americo s.n. (SEL), C.
Luer illustr. 20636. São Paulo: without collection data,
cultivated in São Paulo by V.P. Castro, 28 February 2004,
C. Luer 20617 (SEL). Without collection data, cultivated in
Hort. Bot. Leuvenensis, 1837, by M. Martens s.n. (BR), C.
Luer illustr. 18913.
Acianthera gracilisepala (Brade) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 253. 2004. Fig. 4.
Basionym: Pleurothallis gracilisepala Brade, Arch. Jard.
Bot. Rio de Janeiro 11: 74. 1951. TYPE: BRAZIL.
Paraná: Campos Gerais, 1949, lowered in cultivation
4 January 1950, A. Guimarães s.n. (RB [74403, lost];
Lectotype here designated: Brade original drawings of
the holotype deposited in Guido F. J. Pabst’s reference
iles at Herbarium Bradeanum (HB), numbered Brade
20159, reproduced in Arch. Jard. Bot. Rio de Janeiro
11: plate 4, ig. 1–10. 1951.
Synonyms: Pleurothallis antennata Garay, Arch. Jard.
Bot. Rio de Janeiro 12: 170. 1953, syn. nov. TYPE:
BRAZIL. Paraná: Capão Grande, l. cult. 21 February
1915, P. Dusén 16727 (Holotype: S [S-R-4949, photo
seen]; Isotype: AMES [118552].
Acianthera antennata (Garay) Pridgeon &
M.W.Chase, Lindleyana 16: 242. 2001, syn. nov.
Study of an isotype of Pleurothallis antennata and of a
photograph of the holotype deposited at S has shown that
this species is a synonym of Pleurothallis gracilisepala
VOL. 20, NO. 1
[= Acianthera gracilisepala]. The holotype specimen of
Pleurothallis gracilisepala has not been located at RB
and is apparently lost. However, this species was nicely
illustrated in the protologue leaving no doubt that it is
conspeciic with P. antennata. Brade’s original drawings
that appeared in the protologue have also been examined
and conirm our opinion. Figure 4 in the present work is
based on the isotype of Pleurothallis antennata at AMES.
Although rehydration of the lower using ammonia was not
satisfactory and the column was still somewhat distorted,
all other loral segments match well those of Acianthera
gracilisepala. This specimen was collected by Dusén in
forests by the Guabiroba river, in Capão Grande, state of
Paraná. As far as we could ascertain, Capão Grande is a
farm in the municipality of Ponta Grossa. It was founded in
1850 and is located near what today is Vila Velha State Park.
For further information on these sites, the reader should
refer to the following website addresses: http://sigep.cprm.
gov.br/sitio099/sitio099.htm and http://www.webhotel.com.
br/parana/turismo/tc2_pontagrossa.htm
Additional specimens examined: BRAZIL. São Paulo:
Rio Claro, collected locally and cultivated by Wagner
Marques, 16 March 2007, D.H. Baptista s.n. (SEL), C.
illustr. 21131. Paraná: Ortigueira, Rio Barra Grande, 13
May 1969, G. Hatschbach 21472 (C). Without collection
data, lowered in cultivation 16 March 2007 by Wagner s.n.
(SEL), C. Luer illustr. 21131.
Acianthera strupifolia (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16: 246. 2001. Fig. 5–7.
Basyonym: Pleurothallis strupifolia Lindl., Edwards’s Bot.
Reg. 25(Misc.): 2. 1839. TYPE: BRAZIL. Without
collection data, imported by Messrs. Loddiges s.n.
(Holotype: K [000885712]).
Synonyms: Pleurothallis picta Hook., Bot. Mag. 68: t. 3897
(1841), nom. illeg., non Lindl. 1835. TYPE: MEXICO
(in error): imported by Mr. Parkinson, lowered in
cultivation at Woburn, submitted to the Botanical
Magazine in 1839 by Mr. Forbes s.n. (Holotype: K
[not located]).
Pleurothallis bicolor Lindl., Edwards’s Bot.
Reg. 28(Misc.): 76. 1842, replacement name for
Pleurothallis picta Hook.
Pleurothallis dichroa Rchb.f., Hamburger GartenBlumenzeitung 21: 356. 1865, syn. nov. TYPE:
BRAZIL. Without locality, collected by H. Blunt,
lowered in cultivation in 1863, S.H. Low s.n.
(Holotype: W [R-21887].
Pleurothallis hookeri Regel, Trudy Imp. S.
Peterburgsk. Bot. Sada 7: 545. 1881, nom. illeg.,
replacement name for Pleurothallis picta Hook.
Humboltia strupifolia (Lindl.) Kuntze, Revis. Gen. Pl.
2: 668. 1891.
Pleurothallis glaucophylla Hoehne, Arch. Bot.
São Paulo 1: 579. 1927. TYPE: BRAZIL. Minas
Gerais: Santa Bárbara do Mato Dentro, lowered in
cultivation at Horto Oswaldo Cruz, Butantan, in 30
October 1922, F.C Hoehne s.n. (Holotype: SP [8117];
Isotype: B [destroyed]).
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
45
Figure 4. Acianthera gracilisepala (Brade) Luer. Drawing based on the isotype of Pleurothallis antennata Garay, P. Dusén 16727 (AMES).
Drawn by C. A. Luer.
Acianthera dichroa (Rchb.f.) F.Barros & L.Guimarães,
Neodiversity 5: 28 (2010), syn. nov.
Study of the type specimen of Pleurothallis dichroa has
shown that this species is a synonym of the common and
showy Pleurothallis strupifolia [=Acianthera strupifolia].
Pleurothallis dichroa was described by Reichenbach
in 1865 based on a collection received from Stuart Henry
Low in 1863. Stuart was the younger son of Hugh Low
(1793–1863), founder of Hugh Low & Co., a famous
nursery at Upper Clapton, London. He received the
specimen from his plant collector in Brazil, Henry Blunt.
Three herbarium sheets labeled as Pleurothallis dichroa are
found in the Reichenbach orchid herbarium at W. One of
them, numbered 21887 (Fig. 5), contains a specimen and
a colored illustration accompanied by a short description
in Reichenbach’s almost illegible handwriting. It is labeled
“Low 63,” the number 63 meaning the year (1863) he
received it from Low. The other two sheets are numbered
41142 (Fig. 6) and 18556 (Fig. 7). The former bears a
sketch, description and notes, also in Reichenbach’s almost
illegible handwriting. On the upper right side of these notes
the number “258” can be read. The latter sheet, numbered
46
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 5. Acianthera strupifolia (Lindl.) Pridgeon & M.W.Chase. Holotype of Pleurothallis dichroa Rchb.f., herbarium sheet 21887, in
Reichenbach Orchid Herbarium at W. By permission of the Keeper, Herbarium Natural History Museum in Vienna.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
47
Figure 6. Acianthera strupifolia (Lindl.) Pridgeon & M.W.Chase. Herbarium sheet 41142 of Pleurothallis dichroa Rchb.f., Reichenbach
Orchid Herbarium at W. By permission of the Keeper, Herbarium Natural History Museum in Vienna.
48
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 7. Acianthera strupifolia (Lindl.) Pridgeon & M.W.Chase. Herbarium sheet 18556 of Pleurothallis dichroa Rchb.f., Reichenbach
Orchid Herbarium at W. By permission of the Keeper, Herbarium Natural History Museum in Vienna.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
18556, contains an actual specimen and is labeled “258
Pl. dichroa” in Reichenbach’s handwriting. The number
“258” is most certainly not Blunt’s collection number, but
an identiication number applied to each live specimen at
Low’s nursery (L. A. Garay 2013, pers. comm.). The crosslabelling on the herbarium sheets 41142 and 18556 in
Reichenbach herbarium indicates that these two sheets with
a “Blunt’s number,” one containing the drawings and the
other a specimen, refers to the same collection. Following
standard practice of his time, Reichenbach did not explicitly
cite the word “type” (or “typus”) in the protologue of
Pleurothallis dichroa. However, he explicitly cited that he
received the plant from Low in 1863 and that it had been
collected by Blunt. All three herbarium sheets mentioned
here seem to constitute original material and might
have been used by Reichenbach to base his description.
Nonetheless, the herbarium sheet 21887 is the only one
which contains unambiguous information that agrees with
the protologue (Low’s name and the year Reichenbach
received the specimen), apart from agreeing with the
original description. It is therefore here interpreted as the
actual holotype of Pleurothallis dichroa. Blunt’s name is
also cited in the protologue but no number is mentioned in
the original description.
The name Restrepia liebmanniana Kraenzl. has been
usually cited in the synonymy of Acianthera strupifolia
(e.g. Barros et al. 2015; Govaerts et al. 2015) but this
Mexican species is a distinct taxon. It is actually a synonym
of Pleurothallis spectrilinguis Rchb.f. [= Muscarella
marginata (Rich.) Luer] and it is therefore here excluded
from the synonymy of Acianthera strupifolia.
Additional specimens examined: BRAZIL. Without
locality: Blunt 258 (W [R-18556 and R-41142, drawings and
notes]). Rio de Janeiro: Carmo, Rio Paquequer, date not given,
Neves-Armond 120 (BR). Espírito Santo: Serra do Castelo,
near Alto Ribeirão do Meio, alt. 1000 m, cultivated at
Fazenda Capijuma, November 2002, M. Frey 282 (SEL), C.
Luer illustr. 20253. Santa Catarina: Ilha de Santa Catarina,
Saco Grande, 24 January 1969, R.M. Klein 8131 (M).
Anathallis aristulata (Lindl.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 118. 2007. Fig. 8–10.
Basionym: Pleurothallis aristulata Lindl., Fol. Orchid. 9:
41. 1859. TYPE: BRAZIL. Rio de Janeiro, J. Miers
.n. (Holotype: K [000885711]; Isotype: BR [658536]).
Synonyms: Lepanthes crebrifolia Barb.Rodr., Gen. Spec.
Orchid. 2: 67. 1881, syn. nov. TYPE: BRAZIL. Rio
de Janeiro: near Rodeio, lowered in November, J.
Barbosa Rodrigues s.n. [Holotype: Lost; Lectotype
here designated: Barbosa Rodrigues’s original
illustration in Iconogr. Orchid. Brésil, vol. 6, tab. 298,
at the Library of Rio de Janeiro Botanical Garden,
cited as tab. 611 (unpubl.) in Barb.Rodr. loc.cit;
copied and reprod. in black and white in Cogn., Fl.
Bras. (Mart.) 3(4), tab. 98, ig. 3. 1896; reprod. in
color in Sprunger et al. (1996), vol. 1: 426, ig. E].
Pleurothallis crebrifolia (Barb.Rodr.) Cogn., Fl. Bras.
(Mart.) 3(4): 483. 1896, syn. nov.
49
Pleurothallis montipelladensis Hoehne, Arch. Inst.
Biol. Defesa Agric. 2: 36. 1929, syn. nov. TYPE:
BRAZIL. São Paulo: Morro Pellado, January 1901,
G. Edwall ex Comissão Geográica e Geológica de
São Paulo 6020 (SP [22463]).
Specklinia crebrifolia (Barb.Rodr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 259. 2004, syn. nov.
Anathallis montipelladensis (Hoehne) F.Barros,
Bradea 8: 295. 2002, syn. nov.
Specklinia aristulata (Lindl.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 258. 2004, syn. nov.
Specklinia montipelladensis (Hoehne) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 262. 2004, syn. nov.
Anathallis ourobranquensis Campacci & Menini,
Bol. Bol. CAOB 60: 123. 2005 (publ. 2006), syn.
nov. TYPE: BRAZIL. Minas Gerais, Ouro Branco,
alt. 1500 m, lowered in cultivation, 24 October 2004,
M.A. Campacci MAC-1679 (Holotype: SP [not seen]).
Anathallis crebrifolia (Barb.Rodr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 118. 2007, syn. nov.
John Lindley described Pleurothallis aristulata
[= Anathallis aristulata] based on a John Miers’s collection
from the state of Rio de Janeiro. The holotype specimen
is kept in Lindley herbarium at K. We have recently
examined it and an illustration (Fig. 8) has been prepared
by one of us (CAL). We have also examined an isotype
deposited at BR. Comparison of these collections with
illustrations, collections, and protologues of Lepanthes
crebrifolia, Pleurothallis montipelladensis, and Anathallis
ourobranquensis, shows that they are all conspeciic.
Anathallis aristulata is a widespread species in eastern
Brazil, found from Rio Grande do Sul, in the south, to the
interior of Bahia, in the northeast. It inhabits the moist
Atlantic forests and usually grows as epiphytes in shade
places at low (ca. 200 m) or moderate altitudes (700–1000
m). It also grows at higher altitudes (1000–1600 m) in
open spaces, either as lithophytes or epiphytes exposed to
cold temperatures during the night and plenty of sunshine
during the day. When growing in protected areas, plants of
A. aristulata possess elongate ramicauls and dorsiventrally
lattened, elliptical or spathulate, green leaves. When growing
in open, exposed places, however, they may have much
shorter, usually inconspicuous ramicauls and the leaves are
reduced, thicker and leshier, sometimes almost orbicular and
subcylindrical, and occasionally yellowish. Variations from
one extreme to another may occasionally be found in the
same individual. Inlorescence and loral morphology are,
however, much less variable and quite uniform throughout
its distribution range. There are usually two, occasionally
three, yellow to greenish-yellow lowers on a solitary
raceme. In specimens with thicker leaves the inlorescence
usually exceeds the leaves, but it can be much shorter in
those specimens with longer, thinner leaves. Sepals and
petals are somewhat similar and acute, and the lip, usually
minutely papillose towards the base, is slightly three-lobed
with lateral lobes below the middle. Figures 9–10 illustrate
some variants of Anathallis aristulata.
50
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 8. Anathallis aristulata (Lindl.) Luer. Based on the holotype of Pleurothallis aristulata Lindl, J. Miers s.n. (K). Drawn by C. A. Luer.
Additional specimens examined: BRAZIL. Bahia: Rio
de Contas, Pico das Almas, alt 1500 m, 25 December 1988,
R.M. Harley & D.J.N. Hind 27369 (K), C. Luer illustr. 19144.
Espírito Santo: Vargem Alta, collected in 1949, lowered in
cultivation, 20 December 1950, A.C. Brade 20560 (RB).
Minas Gerais: without collection data, lowered in cultivation
by Wladyslaw Zaslawski in 1983, A. Toscano de Brito 1584
(SEL), C. Luer illustr. 21578; Bejamim Constant, collected
in 1934, lowered in cultivation at Rio Botanical Garden, 14
January 1936, P.P. Horta s.n. (RB). Rio de Janeiro: Above
Petrópolis, alt. 1500 m, sold by Verboonen at EOC, in NJ,
21 October 1988, C. Luer 13757 (SEL). Rio Grande do Sul:
Itati, Barata, 170 m, 15 August 2010, J. Klein 057 (UPCB,
spirit). Santa Catarina: cultivated in Corupá by Orquidário
Alvim Seidel, presumably collected locally, 22 March 2007,
D.H. Baptista s.n. (SEL), C. Luer illustr. 21178. Without
collection data: W.J. Burchell 2097 (K).
Anathallis heterophylla Barb.Rodr., Gen. Spec. Orchid.
2: 74. 1881. TYPE: BRAZIL. Rio de Janeiro: Serra de
Santa Anna, lowered in May, J. Barbosa Rodrigues s.n.
(Holotype: [Lost]; Lectotype here designated: Barbosa
Rodrigues’s original illustration in Iconogr. Orchid. Brésil,
vol. 3, tab. 181, ig. B, at the Library of Rio de Janeiro
Botanical Garden, cited as tab. 658 (unpubl.) in Barb.Rodr.
loc.cit; copied and reprod. in black and white in Cogn., Fl.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
51
Figure 9. Anathallis aristulata. (Lindl.) Luer. Based on A. Toscano de Brito 1584 (SEL). Drawn by C. A. Luer.
Bras. (Mart.) 3(4), tab. 3, ig. 1. 1896; reprod. in color in
Sprunger et al. (1996), vol. 1: 239, ig. B).
Synonyms: Pleurothallis liparanges Rchb.f., Gard. Chron.,
n.s., 23: 532. 1885, syn. nov. TYPE: BRAZIL.
Without locality: obtained from G.M. Binot, lowered
in cultivation at Leiden Botanical Garden, H. Witte
s.n. (Holotype: W [R-45888]; Isotype: AMES [69074,
fragment]).
Pleurothallis heterophylla (Barb.Rodr.) Cogn., Fl.
Bras. (Mart.) 3(4): 556. 1896.
Pleurothallis hoehnei Schltr., Arch. Bot. São Paulo
1: 215. 1926. TYPE: BRAZIL. São Paulo: Estação
Biológica Alto da Serra, 5 May 1921, A. Gehrt s.n.
(Holotype: B [destroyed]; Lectotype: designated
by Barros (2004): SP [5542]; Isolectotypes: AMES
[54816 and 118513]; HB [2739, not seen]).
52
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 10. Anathallis aristulata. (Lindl.) Luer. Based on C. Luer 13757 (SEL). Drawn by C. A. Luer.
Specklinia heterophylla (Barb.Rodr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 261. 2004, syn. nov.
Specklinia liparanges (Rchb.f.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 261. 2004, syn. nov.
Anathallis liparanges (Rchb.f.) Luer, Novon 18: 78.
2008, syn. nov.
Examination of the holotype of Pleurothallis liparanges
deposited at W conirms that this species is conspeciic with
Anathallis heterophylla as stated in Barros et al. (2015).
However, this name is actually a later synonym published
four years after the publication of Anathallis heterophylla.
The type specimen of Anathallis heterophylla is lost
and the only extant original material is the illustration
that appeared in Barbosa Rodrigues’s Iconographie des
orchidées du Brésil, which is now deposited in the library
of Rio de Janeiro Botanical Garden. This illustration was
copied and reproduced in black and white in Cogniaux
(1896) and reproduced in color in Sprunger et al. (1996). It
is here selected as lectotype.
Anathallis johnsonii Luer & Toscano, Harv. Pap. Bot.
16(2): 365. 2011. TYPE: ARGENTINA. Misiones: Departmento San Pedro, El Piñalito, Arroyo Manduri, 750 m,
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
collected 6 January 1991, lowered in cultivation 10 March
1991, A. Johnson 0171 (MO), C. Luer illustr. 18109.
The type specimen of Anathallis johnsonii was
discovered by the late Andrés E. Johnson (1956–2009) and
sent to one of us (CAL) for identiication in December 1996.
The specimen remained unnamed for 15 years and was only
recently recognized as new and inally published in 2011.
Johnson’s correspondence did not provide speciic locality
and the original label, which accompanied the specimen,
had been, unfortunately, lost. While studying additional
collections by Johnson, we rediscovered the original label
of the type specimen of Anathallis johnsonii. It contains
precise location and additional ecological information on
this species. The type citation of Anathallis johnsonii is,
therefore, herewith corrected and should be cited as above.
This species was found near a small stream growing as an
epiphyte on upper branches of Luehea divaricata Mart.
(Malvaceae). The bases of sepals and petals are yellowish
with the tips magenta. The tip and sides of the lip are also
magenta, and the center is yellow.
First described for Argentina, this species has recently
been discovered in the Brazilian state of Rio Grande do
Sul. It is herewith formally recorded for Brazil. A series of
photographs based on this collection can be found in the
following website: http://www.orquideasgauchas.net/P_
desc_especie.php?cod_especie=480
Additional specimens examined: BRAZIL. Rio Grande
do Sul. Cambará do Sul, 19 July 2014, A. Toscano de Brito,
L.F. Varella, J. Klein & S. Kuhn 3267 (UPCB).
Anathallis lobiserrata (Barb.Rodr.) Luer & Toscano,
Monogr. Syst. Bot. Missouri Bot. Gard. 115: 259. 2009.
Fig. 11–12.
Basionym: Lepanthes lobiserrata Barb.Rodr., Gen. Spec.
Orchid. 2: 63. 1881. TYPE: BRAZIL. Rio de Janeiro:
Rodeio, lowered in January, J. Barbosa Rodrigues
s.n. [Holotype: Lost; Lectotype selected by Barros
and Barberena (2010): Barbosa Rodrigues’s original
illustration in Iconogr. Orchid. Brésil, vol. 6, t. 298,
ig. F, at the Library of Rio de Janeiro Botanical
Garden, cited as tab. 617 (unpubl.) in Barb.Rodr. loc.
cit; copied and reprod. in black and white in Cogn.,
Fl. Bras. (Mart.) 3(4), tab. 92, ig. 4. 1896; reprod.
in color in Sprunger et al. (1996), vol. 1: 426, ig. F].
Synonyms: Anathallis nectarifera Barb.Rodr., Gen. Spec.
Orchid. 2: 74. 1881, syn. nov. TYPE: BRAZIL. Rio
de Janeiro: Serra do Mar, lowered in May, J. Barbosa
Rodrigues s.n. [Holotype: Lost; Lectotype here
designated: Barbosa Rodrigues’s original illustration
in Iconogr. Orchid. Brésil, vol. 3, tab. 145, at the
Library of Rio de Janeiro Botanical Garden, cited as
tab. 538 (unpubl.) in Barb.Rodr. loc.cit; copied and
reprod. in black and white in Cogn., Fl. Bras. (Mart.)
3(4), tab.111, ig. 4. 1896; reprod. in color in Sprunger
et al. (1996), vol. 1: 203].
Pleurothallis lobiserrata (Barb.Rodr.) Cogn., Fl.
Bras. (Mart.) 3(4): 438. 1896.
Pleurothallis nectarifera (Barb.Rodr.) Cogn., Fl.
Bras. (Mart.) 3(4): 560. 1896, syn. nov.
53
Pleurothallis githaginea Pabst & Garay, Arch. Jard.
Bot. Rio de Janeiro 14: 12. 1956, syn. nov. TYPE:
BRAZIL. Rio de Janeiro: Serra da Carioca, near
Trapicheiro, 300 m, cultivated at Rio de Janeiro
Botanical Garden, collector unknown, lowered in
July 1952, G. Pabst 1689 (Holotype: RB [81252];
Isotype: AMES [118516]; HB [1689, not seen]).
Anathallis githaginea (Pabst & Garay) Pridgeon &
M.W.Chase, Lindleyana 16: 248. 2001, syn. nov.
Specklinia lobiserrata (Barb.Rodr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 261. 2004.
Specklinia githaginea (Pabst & Garay) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 260. 2004, syn.
nov.
Specklinia nectarifera (Barb.Rodr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 262, 2004, syn.
nov.
Panmorphia githaginea (Pabst & Garay) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 105: 177.
2006, syn. nov.
Panmorphia lobiserrata (Barb.Rodr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 105: 177. 2006.
Anathallis bolsanelloi Chiron & V.P.Castro
Richardiana 9(1): 4. 2008, syn. nov. TYPE: BRAZIL.
Espírito Santo: Domingos Martins, Biriricas, 200–600
m, collected by R. Bolsanello s.n., G. Chiron 08184
(Holotype: SP [not seen]).
Anathallis lobiserrata (Barb.Rodr.) F.Barros &
Barbarena, Rodriguésia 61(1):128. 2010, nom. illeg.
Study of the protologues and original illustrations of
Lepanthes lobiserrata [= Anathallis lobiserrata], Anathallis
nectarifera and Anathallis bolsanelloi, and examination of
the type specimens of Pleurothallis githaginea have shown
that all these taxa belong within a same species concept. We
did not examine the type specimen of Anathallis bolsanelloi,
but the description, illustration, and photographs provided
in the protologue leave no doubt that it also belongs here.
Anathallis lobiserrata is frequent in the Atlantic forests
of Rio de Janeiro and Espírito Santo. It is recognized by
its oblong-elliptical leaves, which are longer than the
ramicauls, and the inlorescences which are much shorter
than the leaves and usually produced in a fascicle of several
racemes at the apex of the ramicauls. The short, fewlowered racemes produce up to ca. 6 successive lowers;
a single open lower is found near the apex of the racemes.
The lip is slightly three-lobed, the side lobes are round and
erect with minutely pillose margins; a central, longitudinal
callosity runs from the very base up to near the middle of
the lip; at the middle or between the middle and the apical
three-quarters of the lip there is also a pair of parallel, short,
elongate callosities or elevations which are usually very faint
and dificult to see without magnifying lenses. They were
described and illustrated in the protologue of Anathallis
nectarifera, but are missing in the original descriptions
and illustrations of Lepanthes lobiserrata, Anathallis
bolsanelloi, and Pleurothallis githaginea. Examination of
a lower from an isotype at AMES has shown, however,
54
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 11. Anathallis lobiserrata (Barb.Rodr.) Luer & Toscano. Based on A. Toscano de Brito 2020 (MBML). Drawn by C. A. Luer.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
Figure 12. Anathallis lobiserrata (Barb.Rodr.) Luer & Toscano. Based on A. Toscano de Brito 1483 (SEL). Drawn by C. A. Luer.
55
56
HARVARD PAPERS IN BOTANY
that these callosities are present on the lip of Pleurothallis
githaginea. They are also present on the lip of Anathallis
bolsanelloi and can be seen in a photograph of a lower
in front view in the protologue of this species (Chiron &
Castro Neto 2008: 17, ig. b). Figures 11–12 in the present
work illustrate two collections from Espírito Santo.
The type specimen of Anathallis nectarifera is lost
and the only extant original material is the illustration
that appeared in Barbosa Rodrigues’s Iconographie des
orchidées du Brésil, which is now deposited in the library
of Rio de Janeiro Botanical Garden. This illustration, which
was copied and reproduced in black and white in Cogniaux
(1896) and reproduced in color in Sprunger et al. (1996), is
selected here as the lectotype.
Additional specimens examined: BRAZIL. Espírito
Santo: Domingos Martins, 700 m, property of Roberto
Kautsky, 15 April 1997, A. Toscano de Brito 1483 (SEL),
C. Luer illustr. 21572; same locality, lowered in cultivation
by Roberto Kautsky 1059, 24 March 1999, A. Toscano
de Brito 2020 (MBML); Santa Teresa, Estação Biológica
de Santa Lúcia, 10 Nov. 2000, L. Kollmann et al. 3121
(MBML, SEL), C. Luer illustr. 20548.
VOL. 20, NO. 1
Anathallis muscoidea (Lindl.) F.Barros & Barberena,
Rodriguésia 61(1): 129. 2010. Fig. 13–14.
Basionym: Pleurothallis muscoidea Lindl., Edwards’s Bot.
Reg. 24(Misc.): 89. 1838. TYPE: BRAZIL. Without
locality: C. Loddiges s.n. (Holotype: K [000885710]).
Synomyms: Lepanthes marginata Barb.Rodr., Gen. Spec.
Orchid. 2: 68. 1881, syn. nov. TYPE: BRAZIL. Rio de
Janeiro, Palmeiras, Pascoinha, lowered in October,
J. Barbosa Rodrigues s.n. [Holotype: Lost; Lectotype
selected by Barrros & Guimarães (2010): Barbosa
Rodrigues’s original illustration in Iconogr. Orchid.
Brésil, vol. 3, tab. 133, ig. C, at the Library of Rio de
Janeiro Botanical Garden, cited as tab. 677 (unpubl.)
in Barb.Rodr. loc.cit; copied and reprod. in black and
white in Cogn., Fl. Bras. (Mart.) 3(4), tab.98, ig.2.
1896; reprod. in color in Sprunger et al. (1996), vol.
1: 191, ig. C].
Humboldtia muscoidea (Lindl.) Kuntze, Revis. Gen.
Pl. 2: 668. 1891.
Pleurothallis limbata Cogn., Fl. Bras. (Mart.)
3(4): 485. 1896, replacement name for Lepanthes
marginata Barb.Rodr., syn. nov.
Figure 13. Anathallis muscoidea (Lindl.) F.Barros & Barberena. Holotype of Pleurothallis muscoidea Lindl. deposited in Lindley’s
herbarium at K. © The Board of Trustees of the Royal Botanic Gardens, Kew. Reproduced with permission.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
57
Figure 14. Anathallis muscoidea (Lindl.) F.Barros & Barberena. Based on A. Toscano de Brito 2575 (UPCB). Drawn C. A. Luer.
Specklinia limbata (Cogn.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 261. 2004, nom. illeg.
Specklinia muscoidea (Lindl.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 121. 2007.
Panmorphia limbata (Cogn.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 105: 177. 2006, nom. illeg.
Anathallis limbata (Cogn.) Luer & Toscano, Monogr.
Syst. Bot. Missouri Bot. Gard. 115: 258. 2009, nom.
illeg.
Anathallis marginata (Barb.Rodr.) F.Barros &
Barberena, Rodriguésia 61(1): 129. 2010 [before 15
Dec 2010], syn. nov.
Anathallis marginata (Barb.Rodr.) F.Barros &
L.R.S.Guim. Neodiversity 5(1): 30. 2010 [31 Dec
2010], nom. illeg.
Anathallis muscoidea (Lindl.) F.Barros & L.R.S.Guim.
Neodiversity 5(1): 30. 2010 [31 Dec 2010], nom.
illeg.
58
HARVARD PAPERS IN BOTANY
John Lindley described Pleurothallis muscoidea
[=Anathallis muscoidea] based on a Brazilian collection he
received from the Loddiges family. The type specimen is
deposited at K and comprises a leaf and an inlorescence
with only one tiny lower, all glued on a small piece of
paper which is kept within an envelope. We were not able,
for this reason, to dissect and properly examine that lower.
However, a detailed watercolor of a lower prepared by
Lindley is found on the herbarium sheet of Pleurothallis
muscoidea (Fig. 13) and provides enough information for
recognition of this species. It is conspeciic with Lepanthes
marginata, a later synonym irst illustrated by Barbosa
Rodrigues in his unpublished Iconographie des Orchidées
du Brésil. Figure 14 presented herein illustrates a specimen
from São Carlos in the state of São Paulo, southeast Brazil.
Photographs and an illustration of a specimen from the
same area, and most probably representing a duplicate of
this same collection, were recently published by Ferreira et
al. (2013) named as Anathallis marginata.
A lectoptiication of Lepanthes marginata was proposed
almost simultaneously by Barros and Barberena (2010) and
Barrros and Guimarães (2010). The latter reference correctly
selected Barbosa Rodrigues’s original illustration, the only
extant original material, as the lectotype of this species.
For this reason, it must be the one to be accepted. Barros
and Barberena (2010) failed to select Barbosa Rodrigues’s
actual, original illustration.
Additional specimens examined: BRAZIL. São Paulo:
São Carlos, obtained from Dalton H. Baptista, lowered
in cultivation, February 2010, A. Toscano de Brito 2575
(UPCB), C. Luer illustr. 21552.
Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 17: 100. 2002. Fig. 15–19.
Basionym: Pleurothallis paranaensis Schltr., Notizbl. Bot.
Gart. Berlin-Dahlem 7: 274. 1918. TYPE: BRAZIL.
Paraná: without locality, lowered in cultivation in
June 1918, P. Dusén s.n. (Holotype: B [destroyed];
Lectotype here designated: Schlechter’s illustration
reproduced in Feddes Repert. Spec. Nov. Regni Veg.
58, tab. 35, ig. 138. 1930. Epitype here designated:
Rio de Janeiro: Friburgo, Alto Macaé, collected by R.
Warren s.n., lowered in cultivation by Manning 940319, September 1997, C. Luer 18557 (SEL).
Synonyms: Pleurothallis microgemma Schltr. ex Hoehne,
Bol. Mus. Nac. Rio de Janeiro 12(2): 9. 1936, syn.
nov. TYPE: BRAZIL. São Paulo, Iguape, Morro
das Pedras, 20 November 1920, A.C. Brade 8055
(Holotype: B [Lost], Lectotype here designated: HB
[8232, not seen]; Brade’s original drawings: HB).
Specklinia microgemma (Schltr. ex Hoehne) F.Barros,
Hoehnea 10: 110. 1983 (publ. 1984), syn. nov.
Anathallis microgemma (Schltr. ex Hoehne) Pridgeon
& M.W.Chase, Lindleyana 16: 249. 2001, syn. nov.
Specklinia paranaensis (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 262. 2004.
Panmorphia paranaensis (Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 105: 177. 2006.
VOL. 20, NO. 1
Anathallis longiglossa Chiron & N.Sanson,
Richardiana 10: 153 (2010), syn. nov. TYPE:
BRAZIL. Espírito Santo: Conceição do Castelo, G.
Chiron 9858 (Holotype: MBML [41733]), C. Luer
illustr. 21804.
Pleurothallis paranaensis [= Anathallis paranaensis]
was described based on a specimen collected by P. Dusén
in the state of Paraná. The original description lacks an
illustration and the holotype was most certainly destroyed
by the World War II bombing of the Berlin-Dahlem
herbarium. Schlechter prepared a detailed illustration of
a loral dissection of this species, but his original drawing
remained unpublished until it was posthumously reproduced
by Mansfeld (1930). The protologue does not mention any
voucher other than the type and we were unable to locate
any original material associated with the protologue except
the illustration published in 1930. This illustration (Fig. 15)
is here selected as the lectotype. As no information on the
origin of Schlechter’s drawings was provided in the 1930
publication, it could be argued that this illustration could
have been based on a different specimen rather than on
Dusén’s collection. Here we share the opinion expressed by
Buzatto et al. (2011) who, while investigating a similar case
(Capanemia angustilabia Schltr.), found no reason to think
otherwise. We have examined several specimens which
it nicely Schlechter’s illustration and description. One of
them, C. Luer 18557 (Fig. 16) has been selected here as the
epitype as it complements the lectotype illustration which is
represented by a loral analysis.
Hoehne (1929) proposed an amendment to Schlechter’s
description of Pleurothallis paranaensis claiming that
Schlechter had not correctly diagnosed it. He provided a
Latin description and an illustration based on a different
collection from São Paulo, F. C. Hoehne s.n. (SP 17176).
Hoehne slightly altered the diagnostic features of this taxon
and cited only his collection from São Paulo, which seems
like he intended to exclude the type of P. paranaensis.
However, he kept the same speciic epithet and authority
(Schlechter) meaning that he did not want to change the
author citation of the name of the taxon. Therefore, he did not
intend and effectively publish a new name, but only proposed
an amendment to Schlechter’s description. In 1936, Hoehne
changed his opinion and concluded that the specimen he had
used for amending Schlechter’s description of Pleurothallis
paranaensis actually represented an undescribed species.
He described it as Pleurothallis microgemma, taking an
unpublished name given by Schlechter, and cited a collection
from Rio das Pedras, Iguape, in the state of São Paulo.
This collection, whose lowers possess a characteristically
emarginated lip, apparently existed in Germany and was
destroyed during the bombing of Berlin-Dahlem herbarium
(Barros, 2004). A duplicate of the specimen studied by
Hoehne and Brade’s original drawings cited and reproduced
in the protologue are preserved at HB. The former is here
selected as the lectotype of Pleurothallis microgemma.
We have studied other collections which match well the
protologue and Brade’s original illustration of Pleurothallis
microgemma, including specimens which have the
characteristic emarginated lip shown in the prologue.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
59
Figure 15. Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase. Lectotype of Pleurothallis paranaensis Schltr., selected here:
Schlechter’s illustration published by Mansfel in 1930.
60
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 16. Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase. Based on the epitype of Pleurothallis paranaensis Schltr., here
selected: C. Luer 18557 (SEL). Drawn by C. A. Luer.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
61
Figure 17. Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase. Based on a plant cultivated at Orquidário Colibri s.n., C. Luer illustr.
21106. (SEL). Drawn by C. A. Luer.
62
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 18. Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase. Based on A. Toscano de Brito 2592 (HUEFS). Drawn by C. A. Luer.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
63
Figure 19. Anathallis paranaensis (Schltr.) Pridgeon & M.W.Chase. Based on the holotype of Anathallis longiglossa Chiron & N.Sanson,
G. Chiron 9858 (MBML). Drawn by C. A. Luer.
64
HARVARD PAPERS IN BOTANY
Figure 17 in the present article illustrates one of them. This
feature has been shown to be only a variation of a spectrum
ranging from entire to emarginate lip and cannot be used to
distinguish A. microgemma from A. paranaensis. Figure 18
illustrates an intermediate variant collected in the Brazilian
state of Paraná.
Study of the holotype of Anathallis longiglossa reveals
not only that it is conspeciic with A. paranaensis, but
also shows that the illustration of the lip in the protologue
of the former (Chiron and Sanson, 2010: ig. 1, f.1–f. 2)
is inaccurate. It is so different from the lip in the type
specimen that we can only assume that the illustrator either
confused one of the lateral sepals as being the lip or based
his illustration on a distinct collection. Nevertheless, with
the exception of the lip, all other features in the protologue
and on the type specimen agree well with those of A.
paranaensis. The illustration presented herein (Fig. 19) was
made from a rehydrated lower from the type specimen of A.
longiglossa. The tiny lower was too badly lattened to allow
an adequate drawing of the lip surface and its callosities. It
is worth noticing, however, that the lip of A. longiglossa
is also emarginate just like in the type of Pleurothallis
microgemma.
Additional specimens examined: BRAZIL. Espírito
Santo: Domingos Martins, collected and cultivated by R. A.
Kautsky s.n., l. cult. 20 March 1999, A. Toscano de Brito
2002 (SEL), C. Luer illustr. 21343; Serra do Castelo, vicinity
of Fazenda Capijuma, collected by Michel Frey, lowered in
cultivation at Capijuma, 6 Mar. 2004, M. Frey s.n. (SEL),
C. Luer illustr. 20670. Paraná: Rio Tibagi, cultivated by
D. H. Baptista s.n., l. cult. 19 December 2008, A. Toscano
de Brito 2592 (HUEFS), C. Luer illustr. 21592; Curitiba,
lowered in cultivation, 21 March 2007, Marcos Klingelfus
s.n. (SEL). São Paulo: São Paulo, Parque do Estado, lowered
in cultivation, 20 April 1934, F.C. Hoehne s.n. (AMES, SP);
São Paulo, Parque Jabaquara, 2 February, 1926, F.C. Hoehne
s.n. (SP-17176); São Bernado do Campo, Serra do Mar, 800
m, M. Campacci and V.P. Castro s.n. (SEL), C. Luer illustr.
18212; without collection data, lowered in cultivation in
Embú, São Paulo, 11 March 2007, R.B. Galati s.n. (SEL),
C. Luer illustr. 21082; Caucaia, lowered in cultivation,
12 March 2007, Orquidário Colibri s.n. (SEL), C. Luer
illustr. 21106.
Lankesteriana caudatipetala (C.Schweinf.) Karremans,
Lankesteriana 13(3): 327. 2014. Fig. 20–21.
Basionym: Pleurothallis caudatipetala C.Schweinf., Bot.
Mus. Leal. 10: 175. 1942. TYPE. PERU. Junín:
Chanchamayo Valley, alt. 1500 m, March 1930, C.
Schunke 1307 (Holotype: F [622342]; Isotype: AMES
[00083282, lower from the holotype, not seen].
Synonyms: Pleurothallis gehrtii Hoehne & Schltr., Arch.
Bot. São Paulo 1: 214. 1926, syn. nov. TYPE.
BRAZIL. São Paulo: Alto da Serra, Mata da Estação
Biológica, 6 May 1921, A. Gehrt s.n. (Holotype:
destroyed at B; Lectotype selected by Barros (2004):
SP [5539, photo seen]; Isolectotype: AMES 54815).
VOL. 20, NO. 1
Pleurothallis illudens Spann., An. Prim. Reun. SulAmer. Bot. 3: 171. 1938 (publ. 1940), syn. nov. TYPE.
BRAZIL. Rio de Janeiro: Petrópolis, June 1938,
collected by Fr. Hermenegildo s.n., C. Spannagel 486
(Holotype: SP [42297, photo seen]).
Specklinia caudatipetala (C.Schweinf.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 259. 2004.
Anathallis gehrtii (Hoehne & Schltr.) F.Barros, Orchid
Memories: 10. 2004, syn. nov.
Specklinia gehrtii (Hoehne & Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot.Gard. 95: 260. 2004, syn. nov.
Panmorphia gehrtii (Hoehne & Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard.105: 177. 2006, syn.
nov.
Anathallis caudatipetala (C.Schweinf.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 115: 257. 2009.
Anathallis edmeiae F.J. de Jesus, Xim.Bols. & Chiron,
Richardiana 13: 296. 2013, syn. nov. TYPE. BRAZIL.
São Paulo: Salesópolis, at the border of Salesópolis
and Caraguatatuba, 1270 m, 9 March 2013, F. José
de Jesus & J. Rodrigues Gomes s.n. (Holotype: SP
[447912, not seen]).
Lankesteriana gehrtii (Hoehne & Schltr.) Karremans
& Luer, Orchids 84(5): 308. 2015, syn. nov.
Lankesteriana caudatipetala is widely distributed from
Central America into the Andes to southern Brazil. It has been
described for Brazil on three occasions: irst as Pleurothallis
gehrtii, second as Pleurothallis illudens, and more recently
as Anathallis edmeiae. We concur with Karremans (2015),
who recently recognised Anathallis edmeiae as conspeciic
with Lankesteriana gerthii. However, these two species are
only forms of a same taxon and are therefore here placed in
the synonymy of L. caudatipetala.
This species is frequent in the Brazilian Atlantic forest
ocurring in Santa Catarina (Spannagel, 1940), Rio de
Janeiro, São Paulo, and in Espírito Santo. It is characterized
by a lax, lexuous, successively lowered raceme that usually
surpasses the leaves. Flowers are very variable in color;
sepals, petals and lip run from dark purple, through redpurple to greenish-yellow and intermediate combinations
within this spectrum. The sepals are acute, vary from
glabrous to ciliate or pubescent, and are connate more or
less at or above the middle into a biid, ovate lamina. The
petals are glabrous or pubescent, usually contracted near
the middle into a narrowly caudate or linear, apical portion.
The lip is oblong to subpandurate and ciliate. A pair of
longitudinal calli begins tall above the middle, diverges
near the middle, and in apposition at the base. At base of the
lip there are a couple of minute lobules, a common feature
found in many other Lankesteriana and Anathallis species.
These lobules are missing in the illustration of the lip in
the protologue of A. edmeiae. The drawings in the present
work illustrate two collections from Rio de Janeiro showing
variation in tepals pubescence. The specimen in Fig. 20 has
glabrous sepal margins, the synsepal is minutely pubescent
in the basal, central portion, and petals are glabrous.
The specimen in Fig. 21 has ciliate sepals and minutely
ciliate petals.
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
65
Figure 20. Lankesteriana caudatipetala (C.Schweinf.) Karremans. Based on A. Toscano de Brito s.n., C. A. Luer illustr. 18306 (SEL).
Drawn by C. A. Luer.
66
HARVARD PAPERS IN BOTANY
VOL. 20, NO. 1
Figure 21. Lankesteriana caudatipetala (C.Schweinf.) Karremans. Based on C. Luer 14512 (SEL). Drawn by C. A. Luer.
Additional specimens examined: BRAZIL. Rio de
Janeiro: Teresópolis: Picada do Rancho Frio, 1500 m,
lowered in cultivation 20 Nov. 1940, A.C. Brade 16746
(RB); Petrópolis: cultivated at Orquidário Binot by Jorge
and Maurício Verboonen, 27 Jan. 1997, A. Toscano de Brito
s.n. (SEL), C. Luer illustr. 18306; near Petrópolis, 1000
m, cultivated at Orquidário Binot by Jorge and Maurício
Verboonen, 30 Nov. 1989, C. Luer 14512 (SEL). São Paulo:
São Bernardo do Campo, 800 m, 19 Jan. 1997, M. Campacci
& V.P. Castro s.n. (SEL), C. Luer illustr. 18204. Espírito Santo:
Alfredo Chaves, São Bento de Urânia, 1200 m, 20 March 1999,
A. Toscano de Brito & R. Kautsky 1954 (MBML). PERU:
Cuzco: Huancabamba, Machu Picchu, 2040 m, 16 April
1943, C. Vargas 3367 (AMES); Urubamba, Machu Picchu,
on rocks, 2040 m, ca. 16 April 1948, C. Vargas 3367 (AMES).
BOLIVIA: La Paz: Nor Yungas, Yierbani, alt. 2750 m,
collected by M. Manon, 5 Feb. 1980, C. Luer 5132 (SEL).
Cochabamba: Chapare, between Cochabamba and Villa
Tunari, alt. 1200 m, 17 Jan. 1988, C. Luer, J. Luer & R. Vásquez
12854 (MO). ECUADOR: Pichincha: old road between
Quito and Santo Domingo, collected by C.H. Dodson,
cultivated at SEL, 1975, C. Luer 565 (SEL). COSTA RICA:
San José: near Cascajal, 1700 m, collected and cultivated by
W. Ballestero, 21 Mar. 1986, C. Luer & B. Luer 12137 (MO).
2015
TOSCANO DE BRITO AND LUER, PLEUROTHALLIDINAE (ORCHIDACEAE) FROM BRAZIL
67
Figure 22. Lankesteriana imberbis (Luer & Hirtz) Karremans. Based on A. Toscano de Brito 1573 (MBML). Drawn by C. A. Luer.
Lankesteriana imberbis (Luer & Hirtz) Karremans,
Lankesteriana 13(3): 327. 2014. Fig. 22.
Basionym: Pleurothallis imberbis Luer & Hirtz, Lindleyana
11: 163. 1996. TYPE: ECUADOR. Morona-Santiago:
Cordillera de Cuticú, epiphytic in forest along the new
road between Mendez and Morona, alt. ca. 900 m, 19
January 1989, A. Hirtz, A. Andreetta & S. Ortega 4136
(Holotype: MO [5259628]), C. Luer illustr. 14917.
Synonyms: Pleurothallis aondae Carnevali & G.A.Romero
in G.A.Romero & G.Carnevali, Orchids Venez. (ed.
2): 1141. 2000. TYPE: VENEZUELA. Bolívar: forest
of Río Aonda, Auyantepui, alt. 600 m, September
1972, C.G.K. Dunsterville 1239 (Holotype: AMES
[00287231, drawing]).
Specklinia imberbis (Luer & Hirtz) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 261. 2004.
Panmorphia imberbis (Luer & Hirtz) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 105: 160. 2006.
Specklinia rubidantha Chiron & Xim.Bols.
Richardiana 9(3): 125. 2009, syn. nov. TYPE:
BRAZIL. Espírito Santo, Domingos Martins,
Biriricas, 200–600 m, collected by R.X. Bolsanello
90511, G. Chiron 9381 (MBML [not located]; Isotype:
LY [not seen]).
68
HARVARD PAPERS IN BOTANY
Anathallis imberbis (Luer & Hirtz) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 115: 258. 2009.
Lankesteriana rubidantha (Chiron & Xim.Bols.)
Karremans, Lankesteriana 13(3): 327. 2014, syn. nov.
Lankesteriana imberbis was originally described
in the genus Pleurothallis R.Br. based on a collection
from lowland Ecuador. Two other collections, one from
Amazonian Bolivia and one from Amazonian Brazil, were
also cited in the protologue. Comparison of the collections
cited in the protologue and two additional collections from
Domingos Martins, in Espírito Santo, southeast Brazil,
with the protologue of the recently described Specklinia
rubidantha, whose type specimen also comes from
Domingos Martins, leaves no doubt that S. rubidantha is a
synonym of Lankesteriana imberbis and corroborates the
view of Karremans (2015) who alluded to this synonymy.
The illustration of the lip in the protologue of Specklinia
rubidantha (page 126, ig. 1–2) is crude, seems to be
inaccurate and therefore misleading: the lip is shown with disc
free of callosities and the truncate base lacks basal lobules,
features always present on the lip of L. imberbis. We have
been unable to locate the holotype of S. rubidantha at MBML
VOL. 20, NO. 1
during a recent visit by one of us (ALVTB). Apparently
a specimen has not been deposited in this herbarium. The
two collections from Domingos Martins cited in the present
article, A. Toscano de Brito 1449 and A. Toscano de Brito
1573, conirm the presence of Lankesteriana imberbis in
southeast Brazil. One of them, A. Toscano de Brito 1573,
is illustrated in Fig. 22. This species seems to rival in
distribution Lankesteriana barbulata (Lindl.) Karremans, a
frequent and apparently closely related species. We believe
that other collections of L. imberbis might exist in Brazilian
herbaria misidentiied as L. barbulata.
Additional specimens examined: ECUADOR. Napo:
between Puerto Napo and Misahualli, alt. 430 m, 23
June 1987, C.H. Dodson and M.W. Chase 17230 (MO).
BRAZIL. Espírito Santo: Domingos Martins, collected by
R.A. Kautsky s.n., 1996, lowered in cultivation in February
1997, A. Toscano de Brito 1573 (MBML), C. Luer illustr.
21589; same locality, collected by R.A. Kautsky s.n., 1996,
lowered in cultivation 14 April 1997, A. Toscano de
Brito 1449 (MBML). Amazonas: Igarapé do Tarumã, near
Manaus, alt. 200 m. collected by M. Madison, lowered in
cultivation 19 March 1979, C. Luer 4038 (SEL).
LiTerATure ciTed
BArros, F. 2004. Taxonomic and nomenclatural notes on Brazilian
Orchidaceae. Pages 7–22 in s. mAniLAL And c. sATish KumAr,
eds. Orchid Memories: A Tribute to Gunnar Seidenfaden.
Mentor Books and Indian Association for Angiosperm
Taxonomy (IAAT), Calcutta.
–––––– And F. F. V. A. BArBArenA. 2010. Notas nomenclaturais e
novas combinações em Anathallis e Specklinia (Orchidaceae).
Rodriguésia 61(1): 127–131.
–––––– And L. r. s. guimArães. 2010. new combinations and a
new name in Brazilian Orchidaceae. Neodiversity 5: 26–33.
––––––, F. Vinhos, V. T. rodrigues, F. F. V. A. BArBerenA, c. n.
FrAgA, e. m. pessoA, w. ForsTer, L. menini neTo, s. g. FurTAdo,
c. nArdy, c. o. AzeVedo, And L. r. s. guimArães. 2015.
Orchidaceae in Lista de Espécies da Flora do Brasil. Jardim
Botânico do Rio de Janeiro. http://relora.jbrj.gov.br/jabot/
loradobrasil/FB10986 (accessed April 25, 2015, 16:07 GMT).
BuzATTo, c. r., r. B. singer, And g. A. romero-gonzáLez. 2011.
Typiications and new synonymies in Capanemia (Orchidaceae,
Oncidiinae). Novon 21: 28–33.
chiron, g. r. And V. p. cAsTro neTo. 2008. Contribution à la
connaissance des orchidées du Brésil. XIV : trois nouvelles
espèces d’Orchidaceae de l’état d’Espírito Santo (Brésil).
Richardiana 9(1): 3–20
–––––– And n. sAnson. 2010. Deux nouvelles espèces de
Pleurothallidinae (Orchidaceae) d’Espírito Santo (Brésil).
Richardiana 10(3): 152–60.
–––––– And A. peTini-BeneLLi. 2012. Une nouvelle espèce
d’Acianthera (Orchidaceae, Pleurothallidinae) du Mato Grosso
(Brésil). Richardiana 13: 66–70.
cogniAux, A. 1896. Orchidaceae. Fl. Bras. (Mart.) 3(4): 320–651.
FerreirA, A. w. c., d. h. BApTisTA, And e. r. pAnsArin. 2013.
Anathallis marginata (Orchidaceae, Pleurothallidinae): A new
record from the state of São Paulo, Brazil. Richardiana
13: 156–163.
goVAerTs, r., p. BerneT, K. KrATochViL, g. gerLAch, g. cArr,
p. ALrich, A. m. pridgeon, J. pFAhL, m. A. cAmpAcci, d. h.
BApTisTA, h.Tigges, J. shAw, p. criBB, A. george, K. Kreuz,
And J. wood. 2015. World Checklist of Selected Plant Families.
Orchidaceae. The Board of Trustees of the Royal Botanic
Gardens, Kew; http://apps.kew.org/wcsp/qsearch.do (accessed
April 25, 2015, 16:21 GMT).
hoehne, F. c. 1929. Contribuições para o conhecimento da lora
Orchidologica brasilica I. Arch. Inst. Biol. (São Paulo) 2: 1–52.
––––––. 1936. Orchidaceae dos herbarios de Alexandre Curt Brade
e do Museu Nacional. Bol. Mus. Nac. Rio de Janeiro, Bot. 12(2):
1–37.
KArremAns, A. p. 2015. Visualizing Pleurothallids: Lankesteriana.
Orchids 84(5): 305–312.
Luer, c. A. 2004. Icones Pleurothallidinarum XXVI. Systematics
of Pleurothallis subgenus Acianthera (Orchidaceae). Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 1–114.
mAnsFeLd, r. 1930. Blütenanalysen neuer Orchideen von R.
Schltechter. I. Südamerikanische Orchideen. Repert. Spec. Nov.
Regni Veg. 58, tab. 35, ig. 138.
spAnnAgeL, c. 1940. Uma nova orchidea do genero Pleurothallis.
An. Prim. Reun. Sul-Amer. Bot. 3: 171, ig. 1.
sprunger, s., p. J. criBB, And A. L. V. ToscAno de BriTo, eds. 1996.
João Barbosa Rodrigues–Iconographie des orchidées du Brésil,
Vol. 1: The illustrations. Friedrich Reinhardt Verlag, Basle.
ToscAno de BriTo, A. L. V. And c. A. Luer. 2015. New species and
nomenclatural notes in Acianthera from Brazil. Lankesteriana
15(1): 77–92.
Vásquez ch., r. And c. h. dodson. 1982. Icones Plantarum
Tropicarum Fasc. 6. Orchids of Bolivia, plate 575. The Marie
Selby Botanical Gardens, U.S.A.
–––––– And p. L. iBisch. 2000. Orchids of Bolivia. Diversity and
Conservation Status. Vol. 1. Editorial F.A.N, Santa Cruz de la
Sierra.