Few orchid species have experienced such a
complicated taxonomic history as Ornithidium
pendulum (Poepp. & Endl.) Cogn. (Fig. 1). This
species has been described under six different
names from three (or four) countries. It has been
known by a misapplied name (Maxillaria
ramosa Ruiz & Pav.) for over 40 years, and it
has been confused with a hitherto undescribed
species from Venezuela and the Guianas.
McIllmurray and Oakeley (2004) unraveled
much of the confusion, but their paper has
remained relatively unknown among orchid tax-
onomists. The present paper aims to further clar-
ify the identity of O. pendulum.
FURTHER DISENTANGLING OFATAXONOMIC PUZZLE:
MAXILLARIA RAMOSA, ORNITHIDIUM PENDULUM,
ANDANEW SPECIES, O. ELIANAE (ORCHIDACEAE)
MARIO A. BLANCO,1,2 GERMÁN CARNEVALI,3 DIEGO BOGARÍN,2,4
AND RODRIGO B. SINGER5
Abstract.McIllmurray and Oakeley (2004) demonstrated that the nameMaxillaria ramosa has been misapplied
to Ornithidium pendulum since 1967, and possibly corresponds to M. cassapensis. We refer Ornithidium
ochraceum, O. loefgrenii, andMaxillaria spathulata to the synonymy of O. pendulum (in addition to the already
recognized synonyms O. dichotomum and Scaphyglottis tafallae), and designate a lectotype for O. dichotomum.
A new species from Venezuela and the Guianas (Ornithidium elianae), previously confused with O. pendulum, is
described. An updated description of O. pendulum is presented along with a review of its complicated taxonomic
history and the first record of this species for Costa Rica.
Resumen.McIllmurray y Oakeley (2004) demostraron que el nombre Maxillaria ramosa ha sido mal aplicado
a Ornithidium pendulum desde 1967, y posiblemente corresponde a M. cassapensis. Referimos los nombres
Ornithidium ochraceum, O. loefgrenii y Maxillaria spathulata a la sinonimia de O. pendulum (además de los
sinónimos ya reconocidos O. dichotomum y Scaphyglottis tafallae), y designamos un lectotipo para O. dichoto-
mum. Se describe una nueva especie de Venezuela y las Guyanas (O. elianae), la cual hasta ahora había sido con-
fundida conO. pendulum. Se presenta una descripción actualizada deO. pendulum, una revisión de su complicada
historia taxonómica, y el primer informe de esta especie para Costa Rica.
Keywords: Cymbidieae,Maxillaria, Maxillariinae, Ornithidium
Inês Cordeiro (SP), Phil Cribb and Clare Drinkell (K), Mar González Bausá (MA), Jonathan Gregson (BM), Jennifer
Gruhn (MO), Eric Hágsater and Miguel Angel Soto Arenas (AMO), Muriel Hecquet and Fred Stauffer (G), Wesley Higgins
(SEL), Henry Kesner and GustavoA. Romero-González (AMES, GH), Mayra Maldonado and Margaret Dix (UVAL), Eliana
Noguera (VEN), Gerardo Salazar (MEXU), Roberto Vásquez (Herbarium Vasquezianum, Bolivia), and Bruno Wallnöfer
(W) kindly provided images and assistance from their respective herbaria. We thank the curators and staff of BM, CAR, CR,
K, G, INB, M, MA, MO, MY, P, PAN, PORT, QCA, QCNE, SEL, SP, RENZ, USJ, W, WU, VEN, and Z, for assistance dur-
ing our visits. Robert Dressler (Jardín Botánico Lankester, Universidad de Costa Rica), Samantha Koehler (Escola Superior
de Agricultura Luiz de Queiroz, Universidade de São Paulo), and Norris Williams (FLAS) provided useful comments and
corrections on a preliminary version of this manuscript. Adam Karremans (Jardín Botánico Lankester) collected the first
record ofOrnithidium pendulum in Costa Rica, which precipitated this study. Funding was provided by a Furniss Foundation
graduate student fellowship from the American Orchid Society and a Kew Latin American Research Fellowship (Royal
Botanic Gardens, Kew) to M.A. Blanco for work in European herbaria; by CICY to G. Carnevali for the project “Orchidaceae
Neotropicales”; by the Darwin Initiative, UK, to D. Bogarín for the project “Conservation and Monitoring of Meso-American
Orchids” (Ref. 14001); by FAPESP to R. B. Singer for postdoctoral work (process No. 01/08958-1); and by the U.S. National
Science Foundation to Norris H. Williams and W. Mark Whitten (FLAS) for the project “Systematics of Maxillariinae
(Orchidaceae): Generic delimitation, pollinator rewards, and pollination” (grant no. DEB-0234064).
1Department of Botany, University of Florida, 220 Bartram Hall, Gainesville, Florida 32611-8526, U.S.A. Email:
mablanco@ufl.edu. Author for correspondence.
2Jardín Botánico Lankester, Universidad de Costa Rica, P. O. Box 1031-7050, Cartago, Costa Rica.
3Herbarium CICY, Centro de Investigación Científica de Yucatán, A. C., Calle 43, No. 130, Col. Chuburná de Hidalgo,
Mérida 97200, Yucatán, México.
4Centro de Investigación en Orquídeas de los Andes “Ángel Andreetta,” Universidad Alfredo Pérez Guerrero, Extensión
Gualaceo, Ecuador.
5Departamento de Botânica, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves
9500, Bloco IV Prédio 43432 Sala 207, Bairro Agronomia, CEP 91501-970, Porto Alegre, RS, Brazil.
Harvard Papers in Botany, Vol. 13, No. 1, 2008, pp. 137–154.
© President and Fellows of Harvard College, 2008.
Species ofOrnithidium Salisb. ex R. Br. have
until recently, been considered part of
Maxillaria Ruiz & Pav. by many authors (e.g.,
Foldats, 1970; Pabst and Dungs, 1977;
Dunsterville and Garay, 1979; Ortiz, 1988,
1995; Sprunger et al., 1996; Atwood, 1999,
2003a; Romero and Carnevali, 2000; Hamer,
2001; Christenson, 2002a, 2002b; Dodson,
2002; Carnevali and Ramírez-Morillo, 2003;
Govaerts et al., 2005). However, new phyloge-
netic analyses based on molecular data indicate
thatMaxillaria is grossly polyphyletic (Whitten
et al., 2007), and we have segregated and rein-
stalled several genera within subtribe
Maxillariinae, includingOrnithidium (Blanco et
al., 2007).
138 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
TAXONOMIC HISTORY OF ORNITHIDIUM PENDULUM AND ITS SYNONYMS
Table 1 provides a summary of the major his-
torical events in the taxonomy of O. pendulum,
including its multiple synonyms, misapplied
names, and species it has been confused with.
Details for each basionym are provided below.
Scaphyglottis pendula Poepp. & Endl.
This species was first described by Poeppig
and Endlicher in 1836, from a plant collected in
1830 in Peru by Poeppig himself. They pro-
posed the genus Scaphyglottis in the same pub-
lication (Poeppig and Endlicher, 1836), but most
species assigned to that genus by Poeppig and
Endlicher are currently placed in Fernandezia
Ruiz & Pav.6
Bentham (1881: 325) suggested that Scaphy-
glottis pendula should be placed inOrnithidium,
a genus created in 1813 by Robert Brown, and
typified by Epidendrum coccineum Jacq. In
1904, Cogniaux formally transferred Scaphy-
glottis pendula to Ornithidium. Both Bentham
and Cogniaux were correct: O. pendulum is
indeed closely related toO. coccineum (Whitten
et al., 2007). In 1945, Schweinfurth transferred
the species toMaxillaria, in line with the inclu-
sive circumscription of the latter genus preva-
lent at the time.
Hoehne (1953: 338), not having seen the type
of Scaphyglottis pendula, suggested that this
species (asMaxillaria pendula) could be closely
related to Pseudomaxillaria chloroleuca (Barb.
Rodr.) Hoehne (a synonym of Maxillaria parv-
iflora (Poepp. & Endl.) Garay7). Brieger (1977),
who never saw the type of S. pendula either, was
probably misled by Hoehne’s opinion and
assigned Ornithidium anceps Rchb.f. (a syn-
onym of Camaridium anceps (Rchb.f.) M. A.
Blanco, a close relative of M. parviflora; see
Atwood, 1993, 1999) to the synonymy of S.
pendula, and transferred the latter to the genus
Pseudomaxillaria Hoehne (typified by P.
chloroleuca). The type of Scaphyglottis pendula
is very different from those of P. chloroleuca
and O. anceps; both Hoehne and Brieger relied
exclusively on the inadequate original descrip-
tion and drawing of S. pendula to reach their
conclusions. Brieger’s (1977) assertion that “no
other specimen of [Pseudomaxillaria pendula]
has been found in Peru during the last 150
years” is clearly based on his erroneous
synonymization.
The two known extant duplicates of the type
collection of Scaphyglottis pendula are in the
Naturhistorisches Museum in Vienna:W-Reich.-
Orch. No. 40118, which has only two leaves and
two drawings of the plant; andW-0007400, which
consists of a large specimen in good condition and
was not part of the Reichenbach f. herbarium
eventually bequeathed to W, but must have been
part of Poeppig’s personal herbarium. Despite an
exhaustive search, no duplicates were found in G,
which holds many Poeppig collections.
The name Camaridium pendulum Barb.
Rodr.8 belongs to a different species, also wide-
spread in SouthAmerica (illustrated in Hoehne,
1953; Sprunger et al., 1996). Surprisingly,
camaridium pendulum and Ornithidium pendu-
lum have never been confused despite being
6Scaphyglottis is a currently accepted genus in subtribe Laeliinae. Dressler (1960) designated Fernandezia graminifoliaRuiz
& Pav. as the generic type of Scaphyglottis to preserve its modern circumscription and to avoid the massive nomenclatural
changes that would have been required otherwise.
7Currently Camaridium micranthum M. A. Blanco. Blanco et al. (2007) had to propose a new name when transferring
Maxillaria parviflora to Camaridium, because the specific epithet was already occupied by Camaridium parviflorum Fawc.
(1910).
8Currently Ornithidium pendens (Pabst) Senghas. Pabst had to propose a new name when transferring Camaridium pen-
dulum toMaxillaria, because the specific epithet was already occupied byMaxillaria pendula (Poepp. & Endl.) C. Schweinf.
(1945). Senghas (1993) maintained the specific epithet “pendens” when transferring the name to Ornithidium because
“pendulum” was pre-occupied by Ornithidium pendulum (Poepp. & Endl.) Cogn. (1904). This species also belongs in the
Ornithidium clade (Whitten et al., 2007).
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 139
B
A
SI
O
N
Y
M
CO
LL
EC
TI
O
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CO
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TR
Y
ST
AT
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a
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iz
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vó
n
s.
n
.
Pe
ru
Pr
o
to
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gu
e
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u
iz
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v
ón
(17
98
)
(de
st
ro
ye
d?
K
?).
D
en
dr
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ra
m
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su
m
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rs
o
o
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(18
07
)
Sy
n
.
w
ith
O
rn
ith
id
iu
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G
ar
ay
(19
67
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Pa
in
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ál
v
ez
pe
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(m
is.
)
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o
ts
yn
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w
ith
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.
pe
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llm
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04
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sy
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w
ith
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ca
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sa
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ph
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la
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17
49
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36
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m
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la
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R
ei
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en
ba
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(18
54
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id
iu
m
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Co
gn
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(19
04
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hw
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Pr
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(18
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Ta
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(20
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(20
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ew
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(18
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a
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a
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lla
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a
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ar
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(19
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a)
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n
.
o
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Lö
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CG
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19
54
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ra
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T A
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1.
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as
sig
n
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to
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a
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e.
140 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
B
A
SI
O
N
Y
M
CO
LL
EC
TI
O
N
CO
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(19
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th
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la
M
.
A
.
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&
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.
,
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b
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.
TA
B
LE
1
CO
N
T.
closely related and sharing the same specific
epithet; therefore, Camaridium pendulum is not
included in Table 1.
Scaphyglottis tafallae Rchb.f.
Reichenbach f. published this name in 1849
based on a Peruvian collection made by Juan
Tafalla in 1797 (the earliest known collection of
Ornithidium pendulum). A few years later,
Reichenbach (1854) recognized Scaphyglottis
tafallae as conspecific with S. pendula, but still
transferred his species to Ornithidium and
treated S. pendula as a synonym. Schweinfurth
(1945), seemingly unaware of Reichenbach’s
synonymization, transferred both S. tafallae and
S. pendula toMaxillaria (apparently he did not
see their types either). In 1967, Garay put M.
tafallae and M. pendula back together, but this
time under the synonymy of M. ramosa (see
“Confusion with Maxillaria ramosa Ruiz &
Pav.” below).
The isotypes of Scaphyglottis tafallae at BM,
G, and MAhave the unpublished name “Orchys
ramosa” written on their labels (“Orchys” is a
misspelling of the genus Orchis L.). The type
collection was made by Juan Tafalla for Ruiz
and Pavón, but only Pavón’s name is written on
the labels (at BM, G, and W), and only Ruiz’s
name was mentioned in the protologue. The
date “Año 97” (year 1797) and the locality
“Chicoplaya” is written on the labels of the
specimens at G, MA, and W, and is mentioned
in the protologue. The isotype at G has an anno-
tation by Reichenbach f. as “Ornithidium ramo-
sum Rchb.f.,” likely based on “Orchys ramosa,”
but he never published that combination.
Reichenbach (1856) cited two duplicates of
Ornithidium tafallae, one in the Boissier herbar-
ium (G), and the other in Berlin; the latter was
undoubtedly destroyed during the allied bomb-
ings of 1943 (Ames, 1944). Reichenbach (1856)
misspelled the name as “O. Tabellae” (species
number 67) but made reference to his transfer in
Bonplandia 2: 18, and to the field data “Chico-
playa 1797.” According to Garay (1967: 260),
the specimen atW is made up of fragments (pos-
sibly taken by Reichenbach f.) from the speci-
men at G, but it is also possible that they were
taken from the now-destroyed specimen at B.
Mansfeld annotated the specimen at MA as
Ornithidium tafallae in 1934, and Carnevali and
Ramírez annotated it as the type of Maxillaria
ramosa in 1988; this second annotation is incor-
rect (derived from Garay’s misapplication of the
name; see “Confusion with Maxillaria ramosa
Ruiz & Pav.” below). In the same year, E.
Christenson also annotated the specimen at BM
as the type of M. ramosa.
Ornithidium ochraceum Rchb.f.
In 1887, Reichenbach f. described Ornithid-
ium ochraceum based on a plant from “New
Grenada” sent to him by Hermann Wendland,
then director of the Royal Gardens in Herren-
hausen (Hanover, Germany). Reichenbach even
compared his “new” species to O. tafallae, but
did not mention any differences. Garay trans-
ferred O. ochraceum to Maxillaria in 1968,
but did not recognize it as conspecific with
O. pendulum.
The type specimen of Ornithidium
ochraceum is depauperate; it consists of a hand-
ful of aggregate, leafless pseudobulbs with a
broken segment of rhizome at the base.
However, there is a drawing of the plant
attached to the sheet that clearly shows the char-
acteristic leaf shape of O. pendulum, with one
flower emerging from within the bracts at the
base of the pseudobulb. It also shows the lip
with a subtriangular, reflexed epichile with dark
warts (mauve-colored spots, according to the
protologue). No duplicates of the type collection
have been located in GOET (J. Heinrichs, pers.
comm.), where Wendland’s main collection
resides.
Both Schlechter (1920: 274) and Garay
(1968: 235) assumed that the type of
Ornithidium ochraceum was collected in mod-
ern-day Colombia. However, Panama was also
part of the Republic of Nueva Granada at the
time of publication of the protologue, and it is
also possible that the type came from there. The
specimen must have been prepared by
Wendland from a plant cultivated at
Herrenhausen, but not collected by him in the
field. It is known that Wendland collected in
Belize, Guatemala, Honduras, El Salvador, and
Costa Rica (Wittmack, 1903; Stafleu and
Cowan, 1988; Vegter, 1988), but there is no indi-
cation that he ever collected in Panama or
Colombia.
Ornithidium loefgrenii Cogn.
Cogniaux described Ornithidium loefgrenii
in 1904, in the same publication where he trans-
ferred Scaphyglottis pendula to Ornithidium,
and thus failed to recognize both as conspecific
(Cogniaux used the spelling “löfgrenii,” which
was corrected to “loefgrenii”; article 60.6 in
McNeill et al., 2006). In 1942, Hoehne trans-
ferred O. loefgrenii to Camaridium, failing to
recognize it as conspecific withMaxillaria pen-
dula, even though he transcribed the original
description of the latter species in the same pub-
lication. He repeated the same information in
Flora Brasílica (Hoehne, 1953). Pabst trans-
ferred O. loefgrenii to Maxillaria in 1972. The
nameMaxillaria loefgrenii (Cogn.) Pabst there-
fore became widely used in Brazil, as this com-
bination was used in the influentialOrchidaceae
Brasilienses (Pabst and Dungs, 1977).
The type specimen ofOrnithidium loefgrenii
was collected by Löfgren in São Paulo, Brazil.
Hoehne (1953: t. 90) published a drawing of the
type in which the shape of the labellum is almost
identical to that illustrated by Lehmann for O.
dichotomum Schltr. (Fig. 2). In an extensive
molecular phylogeny of subtribe Maxillariinae
(Whitten et al., 2007), O. pendulum (from
Ecuador) is strongly supported as sister to O.
loefgrenii (as Maxillaria pendula and M. loef-
grenii, respectively), which provides support for
their merging.
Ornithidium dichotomum Schltr.
Schlechter (1920) described Ornithidium
dichotomum based on a plant collected by
Friederich C. Lehmann in Popayán, Colombia,
and compared it to O. tafallae. Schweinfurth
(1945) referred Schlechter’s species to the syn-
onymy of Maxillaria tafallae (Rchb.f.) C.
Schweinf.
There is a watercolor in Kew (F. C. Lehmann
Icones No. 1005) based on Lehmann s.n. (B.T.
230, K), also from Popayán, that portrays the
plant in life (Fig. 2).
The name Camaridium dichotomum Schltr.
(synonym:Maxillaria dichotoma (Schltr.) L. O.
Williams) belongs to a different species in the
Camaridium clade, which occurs from Costa
Rica to Peru (Atwood, 1999; Whitten et al.,
2007).
Maxillaria spathulata C. Schweinf.
Schweinfurth described Maxillaria spathu-
lata in 1952 from a Peruvian collection by Julio
C. Vargas-Calderón. Schweinfurth even
acknowledged a close relationship withM. tafal-
lae, but distinguishedM. spathulata by its larger
flowers and differently shaped lip. We view
these differences as part of the natural variation
of O. pendulum, and possibly as artifacts from
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 141
pressing, likely to happen in a flower bearing
such a rigidly recurved labellum, which is almost
impossible to flatten without any distortion.
Pabst referred Maxillaria spathulata to the
synonymy of M. loefgrenii in 1972. Here, we
place these two names in the synonymy of
O. pendulum for the first time.
Unpublished names
In the 1960’s, A. H. Heller collected plants
ofOrnithidium pendulum in Nicaragua. Initially
unaware of their identity, he intended to describe
them as “Ornithidium nicaraguensis” and pre-
pared an illustration and description (now in the
archives at SEL). The prospective holotype
(Heller 8403 at F, but not the duplicate at SEL),
was annotated as “Ornithidium tafallae var.
nicaraguensis Heller.” Fortunately, these two
names remained unpublished (the combination
O. nicaraguense (Hamer & Garay) M. A.
Blanco & Ojeda was coined for the species
known until recently asMaxillaria nicaraguen-
sis (Hamer & Garay) J. T.Atwood; Blanco et al.,
2007). A few years after Heller’s death, Hamer
(1983, 1990) annotated the specimens and pub-
lished the illustration as Maxillaria ramosa.
142 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
CONFUSION WITH MAXILLARIA RAMOSA RUIZ & PAV.
The genus Maxillaria was established in
1794 by Ruiz and Pavón, and four years later
they formally described 16 species in this genus,
including M. ramosa. As in Scaphyglottis, the
original characterization of the genus was vague
and species descriptions were very generalized
and therefore easily applicable to many other
species currently known. Most of those original
species of Maxillaria were eventually trans-
ferred to other genera. Only M. prolifera, M.
platypetala, andM. ramosa remained within the
genus until recently. For several years there was
a heated debate on which of these should be
regarded as the type ofMaxillaria (Brieger and
Hunt, 1969; Garay and Sweet, 1972; Ortiz,
1988; Senghas, 1993). After painstaking analy-
ses, Maxillaria platypetala was finally chosen
as the generic type (Garay, 1997; McIllmurray
and Oakeley, 2001), a decision that has been
widely accepted.
The type ofMaxillaria ramosawas collected
in the vicinity of Chinchao in Peru (Department
Huánuco), and has the number 16 assigned to it.
In the Delessert herbarium in Geneva there is an
isotype of Scaphyglottis tafallae (=Ornithidium
pendulum) with the unpublished name “Orchys
ramosa,” the number 16 and the name “Pavón”
written on the label. Garay (1967) examined this
specimen and assumed that it was the type of
Maxillaria ramosa—an apparently logical, but
erroneous conclusion. The name Maxillaria
ramosa was widely misapplied to Ornithidium
pendulum from then on (and soon afterward to
O. elianae Carnevali & M. A. Blanco as well;
see under “Maxillaria ramosa auct., non Ruiz &
Pavón,” at the end of the synonymy of O. pen-
dulum, and in the usage synonymy of O.
elianae, below). At one point, Garay and Sweet
(1972) even designated M. ramosa as the
generic type of Maxillaria, based on the con-
fused type of Scaphyglottis tafallae.
Garay’s confusion was unearthed by
McIllmurray and Oakeley (2001) when they
found a painting prepared by Isidro Gálvez (one
of the illustrators in the Ruiz and Pavón expedi-
tion to Peru) in the archives of the Real Jardín
Botánico in Madrid (a photo of this painting was
published by McIllmurray and Oakeley, 2001,
2004). The painting has the name Maxillaria
ramosa on it, but depicts a plant clearly differ-
ent from Scaphyglottis tafallae. McIllmurray
and Oakeley (2001) wrongfully stated that the
painting and the herbarium specimen corre-
sponded to each other. This error was quickly
pointed out by Atwood (2001) who, in a flawed
attempt to stabilize the nomenclature, desig-
nated the herbarium specimen at Madrid as a
lectotype ofM. ramosa (following Garay’s mis-
application of the name).
Soon afterwards, McIllmurray and Oakeley
(2004) demonstrated that the original description
ofMaxillaria ramosa and Gálvez’s painting cor-
responded to each other, but not to the herbarium
specimen inMadrid designated as a lectotype by
Atwood (2001). Ruiz, Pavón, and Gálvez
returned to Spain in 1788; some of their paid col-
lectors, including Juan Tafalla, stayed in Peru
and continued to send plants to Spain for Ruiz
and Pavón’s Flora Peruviana et Chilensis, and
for Tafalla’s own Flora Huayaquilensis (Estrella,
1991). It was Tafalla who collected the speci-
mens labeled as “Orchys ramosa,” which
Reichenbach f. later used to describe
Scaphyglottis tafallae in his honor. That Tafalla
made this collection (and not Ruiz or Pavón) is
evident by the year written down on the speci-
men labels at G, MA, and W (1797, nine years
after the return of Ruiz, Pavón, and Gálvez to
Spain), and by the annotation “F. P.” (present in
the duplicates at MA and G) that Tafalla made
on the labels of the plants collected for Flora
Peruviana from 1793 onward (Estrella, 1991).
Furthermore, Chicoplaya (the locality written on
the label) was never visited by Ruiz and Pavón
(Ruiz, 1940, 1998; McIllmurray and Oakeley,
2004), but it was visited by Tafalla between 1797
and 1798 (Estrella, 1995: 54). It seems that both
Ruiz and Tafalla, by an unfortunate coincidence,
used the same specific epithet (“ramosa”) and
the same number (16) for their collections of
related but different species, both of which
became types. Furthermore, Tafalla did not write
his name on the labels but Pavón did, probably
because it was collected for him.All these factors
contributed to the identity confusion.Additional
evidence for this argument is presented by
McIllmurray and Oakeley (2004).
McIllmurray and Oakeley (2004) correctly
concluded that Tafalla’s “Orchys ramosa” is con-
specific with Scaphyglottis pendula and S. tafal-
lae. They also suggested that the name
Maxillaria ramosa corresponds toM. cassapen-
sis Rchb.f. (now Maxillariella cassapensis
(Rchb.f.) M. A. Blanco & Carnevali, a member
of the Maxillaria graminifolia (Kunth) Rchb.f.
suballiance sensuAtwood, 2003b), a conclusion
we agree with. Therefore, Maxillaria ramosa is
likely an older synonym of Maxillariella cass-
apensis (but see below). A fruiting specimen of
M. cassapensis from the Ruiz and Pavón collec-
tions (annotated “Orchys, Ex Herb. de R & P,
Lima”) and later incorporated in Hooker’s
herbarium (now in the general collection at K),
might represent type material of M. ramosa.
Thus, Atwood’s (2001) lectotypification of M.
ramosa must be rescinded. McIllmurray and
Oakeley (2004: 35) claimed that they designated
the painting of Gálvez as the lectotype of M.
ramosa in their previous (2001) publication, but
they did not comply with article 7.11 of the Code
(to include the phrase “designated here” or an
equivalent, a requirement since 1 January 2001)
and thus their lectotypification is invalid.
In any case, McIllmurray and Oakeley (2004)
conclusively demonstrated that Maxillaria
ramosa and Ornithidium pendulum are het-
erotypic names that correspond to separate
species. However, their paper has been over-
looked by some taxonomists of neotropical
Orchidaceae. Only Christenson (2002b; but not
in the original English version, 2002a) used the
name M. ramosa in its correct, clarified new
sense, when he assigned it toMaxillaria section
Ebulbes Pfitz. (in theM. graminifolia suballiance
sensuAtwood, 2003b). Given the long history of
misapplication of the nameM. ramosa, however,
a case can be made for its rejection (see articles
56 and 57 in McNeill et al., 2006; M.A. Blanco,
in prep.), and we opted for not transferring the
name toMaxillariella (Blanco et al., 2007).
Whitten et al. (2007) used the name
Maxillaria pendula, and Blanco et al. (2007)
used the name Ornithidium pendulum in the
sense used here. Sitko et al. (2006) used the
misapplied name Maxillaria ramosa for
Ornithidium pendulum.
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 143
TAXONOMIC TREATMENT OF ORNITHIDIUM PENDULUM AND O. ELIANAE
Ornithidium pendulum (Poepp. & Endl.)
Cogn., Fl. Bras. (Martius) 3(6): 92. 1904.
Fig. 1–2.
Basionym: Scaphyglottis pendula Poepp. &
Endl., Nov. Gen. Sp. Pl. (Poeppig and
Endlicher) 1: 58, t. 98. 1836; Maxillaria
pendula (Poepp. & Endl.) C. Schweinf.,
Bot. Mus. Leafl. 11: 285. 1945;
Pseudomaxillaria pendula (Poepp. &
Endl.) Brieger, Bot. Jahrb. Syst. 97: 556.
1977; not Camaridium pendulum Barb.
Rodr. TYPE: PERU. [Huánuco:] Cuchero,
February 1830, E. F. Poeppig 1749
(Holotype: W-0007400; Isotype: W-
Reich.-Orch. 40118).
Synonyms:Scaphyglottis tafallaeRchb.f., Linnaea
22: 855. 1849; Ornithidium tafallae
(Rchb.f.) Rchb.f., Bonplandia 2: 18. 1854.
Maxillaria tafallae (Rchb.f.) C. Schweinf.,
Bot. Mus. Leafl. 11: 288. 1945. TYPE:
PERU. [Huánuco:] Chicoplaya, 1797, J.
Tafalla sub Pavón s.n. (Holotype: W-
Reich.-Orch. [40121]; Isotypes: B
[destroyed], BM, MA, SEL [photo, not
seen] , US [photo, not seen], G; Drawings:
W-Reich.-Orch., AMES 38598).
Ornithidium ochraceum Rchb.f., Gard.
Chron. 1: 209. 1887;Maxillaria ochracea
(Rchb.f.) Garay, Caldasia 10: 235. 1968.
Syn. nov. TYPE: NEW GRENADA
144 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
FIGURE 1.Ornithidium pendulum (Poepp. & Endl.) Cogn.A, plant habit; B, flower, side view;C, dissected peri-
anth;D, labellum and column attached to ovary, sepals and petals removed, side view; E, column, side (left) and
ventral (right) views. Drawn by D. Bogarín from Karremans 448 (CR).
(COLOMBIA or PANAMA). Ex Hort.
Royal Gardens in Herrenhausen,
Germany,H. Wendland s.n. (Holotype:W-
Reich.-Orch. 40122).
Ornithidium loefgrenii Cogn., Fl. Bras.
(Martius) 3 (6): 92. 1904, ‘löfgrenii’;
Camaridium loefgrenii (Cogn.) Hoehne,
Arq. Bot. Estado São Paulo n.s., formato
maior 2(4): 72. 1947; Maxillaria loef-
grenii (Cogn.) Pabst, Bradea 1(19): 175.
1972. Syn. nov. TYPE: BRAZIL. [São
Paulo, Campo Grande; fide protologue],
cult. na Capital [São Paulo city, capital of
São Paulo state; fide Hoehne, 1953], 20
January 1894, A. Löfgren CGG 1954
(Holotype: SP).
Ornithidium dichotomum Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 7: 178. 1920;
illustration in Repert. Spec. Nov. Regni
Veg. Beih. 57: t. 63, nr. 245. 1929; not
Camaridium dichotomum Schltr. TYPE:
COLOMBIA. Cauca: An Bäumen auf
dem Hochland von Popayan, 1400–1800
m, [1884–1900], F. C. Lehmann 8114
(Holotype: B [destroyed]; Lectotype, des-
ignated here: K-79237; Isolectotype: K-
79236).
Maxillaria spathulata C. Schweinf., Bot.
Mus. Leafl. 15: 164, t. 54. 1952. Syn. nov.
TYPE: PERU. Cuzco: Prov. Paucartambo,
between Santa Isabel andAsunción, 1800
m, 4 January 1946, J. C. Vargas-Calderón
5532 (Holotype: AMES; Isotype: CUZ
[not seen]).
“Maxillaria ramosa” auct. non Ruiz &
Pav.: Garay in Bot. Mus. Leafl. 21: 259.
1967; Schweinfurth in Fieldiana, Bot. 33:
64, 65. 1970; Garay and Sweet in J.
ArnoldArbor. 53: 524. 1972; Dodson and
Gentry in Selbyana 4: 170. 1978; Dodson
and Dodson in Icon. Pl. Trop. 2: 161.
1980; Hamer in Ic. Pl. Trop. 9: 865. 1983;
Siegerist in Selbyana 7: 298. 1984; Ortiz
in Orquideología 17: 237. 1988; Hamer in
Selbyana 11 (Suppl.): 486. 1990; Brako
and Zarucchi in Monogr. Syst. Bot.
Missouri Bot. Gard. 45: 820. 1993;
Senghas in Orchideen (Schlechter), ed. 3,
1B: 1771. 1993; Ortiz in Orquídeas de
Colombia, ed. 2: 284. 1995; Jørgensen
and León-Yánez in Cat. Vasc. Pl.
Ecuador: 706. 1999; Dix and Dix in
Monogr. Syst. Bot. Missouri Bot. Gard.
78: 33. 2000; Atwood in Orch. Rev. 109:
316. 2001; Hamer in Monogr. Syst. Bot.
Missouri Bot. Gard. 85: 1756–1757.
2001; Dodson in Native Ecuadorian
Orchids 3: 562. 2002; Dodson in Native
Ecuadorian Orchids 5: 1134. 2004;
Ossenbach et al. in Orquídeas del Istmo
Centroamericano: 96, 214. 2007.
“Maxillaria repens” auct. non L. O.
Williams: Dix and Dix in Monogr. Syst.
Bot. Missouri Bot. Gard. 78: 33. 2000;
Govaerts et al. in World Checklist of
Orchidaceae, 2005 (in both as synonym of
M. ramosa).
“Ornithidium tafallae var. nicaraguensis”
Heller, in sched. (A. H. Heller 8403, F).
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 145
FIGURE 2. Ornithidium pendulum (Poepp. & Endl.)
Cogn. Photo of painting No. 1005 by F. C. Lehmann,
based on Lehmann s.n. (B.T. 230, K), from the type
locality of O. dichotomum Schltr. Reproduced with
the kind permission of the Trustees, Royal Botanic
Gardens, Kew.
Epiphytic or lithophytic herbs, to 2 m long,
most commonly to 70 cm long or less; plants
pendent or scandent, with stems branching at the
bases of pseudobulbs. Roots cylindrical, 1 mm
in diameter. Stems sympodial, always termi-
nated by a pseudobulb. Rhizome to 3–4 mm
diameter, first covered with thin, scarious, acute,
green sheaths, eventually brownish or gray with
age; branches divaricate, usually 2, produced
from the axils of consecutive non-foliar bracts
immediately behind the pseudobulb; the seg-
ments of rhizome between pseudobulbs made
up of few, elongate internodes, the pseudobulbs
5–20 cm apart on the rhizome, occasionally
groups of 2–3 pseudobulbs growing close
together. Pseudobulbs 2.5 cm long, 1.5–5.0 cm
wide, 0.7–1.5 cm thick, brownish to gray-green
at the base, grading to silvery green at apex;
plump and smooth when young, slightly wrin-
kled when old, apically 1-leaved, ellipsoid,
ovoid to (rarely) suborbicular, basally clothed
by several imbricate sheaths of which the (1–
)2(–4) innermost bear foliar blades, these even-
tually caducous. Leaves and blades of sheaths
bright green, subcoriaceous, smooth, fleshy,
conduplicate, articulate, twisted 90 degrees at
base so that all the leaves face to the same side;
elliptic to oblong or linear-elliptic, the margins
often suffused with purple and slightly revolute,
the apex acute to shortly acuminate, oblique
because one of the halves is conspicuously
shorter than the other, keeled abaxially, 8–20 cm
long, 2.4–5.0 cm wide, the innermost sheath
blade larger than the apical leaf and the outer-
most smaller, the sheaths 2–5 cm long.
Inflorescences numerous, 1-flowered, with thin
peduncles, borne singly or several from the axils
of the sheaths enveloping the pseudobulbs and
from the bracts at any point along the rhizome,
several flowers open simultaneously along any
given rhizome segment; peduncle 1–2 cm long,
cylindrical, basally with 1–3 thin, soft, brown
bracts; ovary with pedicel 6–8 mm long; floral
bracts tubular, acuminate. Flowers small and
inconspicuous, to 1 cm long, usually resupinate,
sepals and petals greenish-white, often suffused
with pink, and sometimes deep gray-brown; lip
white to ochre-yellow, frequently with purple
spots, more rarely with pink tinges, the column
greenish. Sepals 5.5–10.0 mm long, 1.7 mm
wide, rectangular, oblong, acute, concave
basally, flat to convex distally, the lateral sepals
slightly oblique, somewhat longer and wider
than dorsal sepal, basally produced into a small,
obtuse mentum. Petals 5–9 mm long, 1.0–1.5
mm wide, linear-elliptic, oblong to oblong-
oblanceolate to narrowly obovate oblanceolate,
acute. Labellum 6 mm long, 3 mm wide, rec-
tangular and narrow at the base, 3-lobed, the lat-
eral lobes 4.0–6.0 mm long, 0.5–8.0 mm wide,
rectangular, straight and parallel to the column
in natural position, partially surrounding the col-
umn; the central (apical) lobe sharply to
obscurely 4-lobulate, 3–2 mm long, 3.6–8.0 mm
wide, the margin finely denticulate, apically
emarginate, thickened and often verrucose in the
central part, strongly reflexed in natural posi-
tion; the callus to 4 mm long, rectangular and
shorter than the lateral lobes, consisting of a
transverse plate at the union of the lateral lobes
with the central lobe. Column subcylindric, ven-
trally gibbous, basally produced into a short
foot, to 4.7 mm long; anther cap cucullate, pale-
brown; pollinia 4, in two unequal pairs attached
to a short ligulate stipe and a tiny semilunar vis-
cidium. Fruit an ovate, pendent, dehiscent cap-
sule, 9–12 mm long, 7–8 mm wide, valves
separating apically upon maturity.
Habitat and ecology: plants grow as hanging
epiphytes or lithophytes and can become large,
pendent mats. The species occurs at 400–1800
m in montane and lower montane, wet and cloud
forests. The small green to yellowish flowers
suggest that pollinators are flies or small bees,
but no floral visitors have ever been documented.
Phenology: flowering occurs sporadically
throughout the year.
Conservation status: a widespread species
and common along the eastern tropical Andes.
It is rare in other parts of its distribution, but this
rarity appears to be natural, not anthropogenic.
This species is not threatened.
Illustrations: detailed analytical drawings
have been published under the namesMaxillaria
spathulata (Schweinfurth, 1945), Camaridium
loefgrenii (Hoehne, 1953),Maxillaria ochracea
(Dunsterville and Garay, 1976; Bennett and
Christenson, 1993; Romero and Carnevali,
2000) and Maxillaria ramosa (Dodson and
Gentry, 1978; Dodson and Dodson, 1980;
Hamer, 1983, 1990; Senghas, 1993; Dodson,
2002). Some of these (e.g., Bennett and
Christenson, 1993) depict the plant as erect,
although it is normally pendent.
146 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
Photographs of Ornithidium pendulum
appear in Senghas (1993, asM. ramosa), Miller
and Warren (1994, 1996, as M. loefgrenii), and
Fernández (2004, as M. ochracea).
Distribution: one of the most widely dis-
tributed species in the genus (as circumscribed
byWhitten et al., 2007, and Blanco et al., 2007).
In South America it is recorded from western
Venezuela, Colombia, Ecuador, Peru, and south-
eastern Brazil; it is locally common in parts of
Rio de Janeiro state in Brazil (Miller and
Warren, 1994, 1996; Miller et al., 2007; as
Maxillaria loefgrenii) and along the eastern
Andes from Colombia to northern Peru, but it is
rare and patchily distributed elsewhere. It
appears to be absent in the Guiana Shield and
most of the Amazonian lowlands.
In Venezuela, Ornithidium pendulum is only
known in the Andean (western) states of Lara,
Mérida, Táchira, Trujillo, and Zulia where it is
local and rare. Brazilian populations (hitherto
classified asMaxillaria loefgrenii) appear to be
disjunct from those in the Andes, and are con-
fined to the Atlantic coastal mountain range in
the states of Espírito Santo, Rio de Janeiro, and
São Paulo (possibly also in southern Bahia
state), where they are also patchily distributed.
Hamer (1983) mentions this species (as
Maxillaria ramosa) as being present in Bolivia,
but he did not cite any specimens; his record is
probably based on the closely related
Ornithidium sillarense (Dodson &Vásquez) M.
A. Blanco & Ojeda (see below).
We also present the first record of
Ornithidium pendulum for Costa Rica (see spec-
imens examined). In Central America, this
species had been previously recorded (as M.
ramosa) for Nicaragua (Hamer, 1983, 1990,
2001) and Guatemala (Dix and Dix, 2000).
Additional specimens examined: BRAZIL.
Espírito Santo: Santa Teresa, morro da estação
repetidora de TV, 14 November 1985, Boone
880 (MO). COLOMBIA. [Cauca:] Popayán,
Lehmann s.n. (B.T. 230, K). Region of Popayán,
flowered in cultivation, SEL 72-1200, 9 May
1979, Kennedy s.n. (SEL [26960]). COSTA
RICA. Cartago: Turrialba, Centro Agronómico
Tropical de Investigación y Enseñanza (CATIE),
600 m, flowered in cultivation at Lankester
Botanical Garden, 31 October 2004, Karremans
448 (CR). ECUADOR. El Oro: 10 km W of
Piñas along the new road from Piñas to
Machala, 900 m, 19 July 1979, Dodson et al.
8439 (MO, SEL). Esmeraldas: km 16
Esmeraldas to Santo Domingo, 300 m, 13
September 1980, Dodson et al. 10436 (SEL).
Quninde. Bilsa Biological Station, Montañas de
Mache, 35 kmW of Quinindé, 5 kmW of Santa
Isabel, 00˚21'N, 79˚44'W, 400–600 m, 5 May
1995, Clark and Watt 742 (MO, QCNE, US).
Los Rios: Rio Palenque Science Center, km 56
Quevedo-Santo Domingo, 220 m, 20 September
1973, Dodson and Tan 5398 (RPSC, SEL);
flowered in cultivation, 9 May 1979, Dodson et
al. 9275 (SEL). Morona-Santiago: above Sucua,
800 m, 23 April 1982, Dalström 217 (SEL).
Road from El Pangui to Chiguinda, km 18,
03˚19'29"S, 78˚38'59"W, 1200 m, 3 October
2003, Whitten et al. 2513 (FLAS, QCA).
Cultivated at Ecuagenera greenhouses in El
Pangui, origin uncertain, 3 October 2003,
Whitten et al. 2487 (FLAS, QCA). Napo:
Archidona, Reserva de Biósfera Sumaco,
00˚40'03"S, 77˚35'40"W, 18 February 2003,
Farfán 431 (MO, QCNE); Cordillera de
Galeras, 00˚49'50"S, 77˚33'28"W, 1180 m, 4
March 2003, Farfán 460 (MO). La Cruz,
Arajuno, Puerto Misahualli, Rio Napo, 500 m,
July-September 1984, Suarez and Lindberg de
Suarez 44 (RPSC). Pastaza: Km 7 Puyo to Mera
on road Baños to Puyo, 1000 m, 17 June 1989,
Dodson and Niell 17413 (QCNE, RPSC).
Pichincha: Cooperativa Santa Marta #2, km 3W
of bypass around Santo Domingo, 530 m, 22
July 1979, Dodson et al. 8514 (SEL).
Tungurahua: Topo, at junction of Rio Topo and
Rio Pastaza, 1300 m, 10 December 1986,
Dodson and Hagsater 16741 (RPSC). Zamora-
Chichinpe: Rio Bombuscara, 2 km E of Zamora
city, 900 m, 18 May 1967, Sparre 16389 (MO).
Road Los Encuentros to Rio Machinaza, NW
portion of Cordillera del Condor, 1450–1650 m,
19 May 1988, Hirtz 3792 (QCNE, RPSC).
Zamora–Cenepa, Rio Zamora, 1100 m, 26 July
1960, Dodson 161 (SEL). Zumbi, N border of
Rio Zamora, 900 m, 18 May 1967, Sparre
16467 (MO). GUATEMALA. Alta Verapaz:
Cobán, Rio Sanchichaj, February 1990, Dix et
al. 6986 (UVAL [flowers in liquid]).
NICARAGUA. [Rivas: Ometepe Island, Volcán
Maderas, 4000 ft (fide Heller’s notes at SEL)],
Heller 8403 (F, SEL). PERU. Amazonas: Rio
Cenepa, creek flowing into Nahim, which flows
into the Huampami, trail E of Huampami, 1 day
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 147
walk to Shaim, 2000 ft, 27 November 1972,
Berlin 402 (MO, SEL). Huánuco: Huánuco,
Tingo María, Rio Huallaga, 19 July 1940,
Asplund 12373 (AMES). Leoncio Prado, Distr.
Damaso Beraun, Las Palmas, 880 m, UTM 18L
0394213-894790, Trujillo 316 (HAO, URP).
Junín: Rio Pinedo, N of La Merced, 700-900 m,
30 May 1929, Killip and Smith 23650 (AMES).
San Ramón, 900–1300 m, 9 June 1929, Killip
and Smith 24762 (AMES). Pasco: Oxapampa,
headwaters of Rio Tunqui, trail to Chuchurras-
Palcazu, 10˚14'S, 75˚28'W, 1850 m, 2 January
1984, Foster et al. 7783 (MO). Between La
Merced and Oxapampa, 1400 m, 30 January
1979, Luer 3809 (SEL). San Martín: Lamas,
distr. Alonso de Alvarado, San Juan de
Pacaizapa, km 72 carretera Tarapoto-
Moyobamba, 1000–1050 m, 8 June 1977,
Schunke 9661 (MO). Zepelacio, near
Moyobamba, 1200–1600 m, January 1934, Klug
3544 (MO); 1100 m, June 1934, Klug 3694
(AMES). VENEZUELA. Lara: 1350 m, 18
October 1952, Renz 7839 (RENZ). Mérida:
1600 m, 26 March 1949, Renz 5119 (RENZ).
Miranda: Quinta Colibría, 1400 m, cultivated in
orchidarium of Mr. and Mrs. G. C. K.
Dunsterville [plant originally from Zulia, Sierra
de Perijá], 13 October 1963, Steyermark and
Dunsterville 56254 (VEN). Táchira: 12 March
1951, Renz 6667 (RENZ). Zulia: Sierra de
Perijá, entre Pishicaco y la frontera con
Colombia hacia Socorpa, 1500 m, floreciendo
en Caracas, December 1974, Dunsterville s.n.
(VEN).
Ornithidium pendulum is vegetatively indis-
tinguishable from the BolivianO. sillarense, and
both species are obviously very closely related.
The flowers are very similar, but the labellum of
O. sillarense is distinct enough to warrant spe-
cific recognition: the midlobe is almost twice as
long as the rest of the labellum (vs. subequal or
smaller than the rest of the labellum in O. pen-
dulum), is not reflexed, and has a different
shape. We have not seen intermediate forms in
terms of labellum structure. As far as we know,
Ornithidium pendulum has not been collected in
Bolivia. According to the protologue, the holo-
type of O. sillarense was deposited in MO.
However, it is presently housed in Herbarium
Vasquezianum in Bolivia (R. Vásquez, pers.
comm., 2006). A previously unreported isotype
of O. sillarense was recently found in SEL.
Both Dix and Dix (2000) and Govaerts et al.
(2005) treatedMaxillaria repens L. O.Williams
(now Ornithidium repens (L. O. Williams)
M.A. Blanco & Ojeda) as a synonym ofO. pen-
dulum (as M. ramosa). However, O. repens is
a different species endemic to Panama, easily
distinguished from O. pendulum by its
more robust, ascending rhizomes devoid of
pseudobulbs.
Herbarium specimens of Ornithidium pen-
dulum have been commonly annotated as
Maxillaria loefgrenii (those from Brazil), M.
ochracea, or M. ramosa.
Ornithidium elianae Carnevali & M. A.
Blanco, sp. nov. TYPE: VENEZUELA. Estado
Carabobo: Municipio Autónomo Mora, cuenca
hidrográfica del rio Morón, parte alta, bosque
nublado, 10˚17–28'N, 68˚10–16'W, 700–1100
m, 13–15April 1991 (flowers),Wilmer Díaz 110
(Holotype: VEN; Isotypes: MO, PORT). Fig. 3.
Usage synonyms: Maxillaria taphallae (sic.)
auct. non Rchb.f.: Dunsterville and Garay
in Venez. Orch. Ill. 1: 242–243. 1959;
Dunsterville and Dunsterville in Amer.
Orchid Soc. Bull. 36: 794. 1967; 38: 496.
1969.
Maxillaria ramosa auct. non Ruiz &
Pavón: Foldats in Fl. Venez. 15(4): 516.
1970; Dunsterville and Garay in Venez.
Orch. Ill. 6: 37. 1976; Steyermark and
Huber in Fl. Avila: 680, 697. 1978;
Dunsterville and Garay in Orchids
Venezuela: 545. 1979; Cremers and Hoff
in Invent. Taxon. Pl. Guyane Franc. II
Orchidac.: 54. 1992; Boggan et al. in
Checkl. Pl. Guianas, ed. 2: 158. 1997;
Romero and Carnevali in Orchids
Venezuela, ed. 2: 581. 2000; Clarke et al.
in Sida, Bot. Misc. 21: 50. 2001;
Carnevali and Ramírez in Fl. Venez.
Guayana 7: 442. 2003; Chiron and
Bellone in Orch. Guyane Franc.: 264.
2005; Funk et al. in Contr. U.S. Natl.
Herb. 55: 127. 2007.
Species haec Ornithidio pendulo (Poepp. &
Endl.) Cogn. similis, foliis angustioribus, labelli
lobulo apicale oblongo concavo (vs. ovato vel
ovato oblongo convexo recurvo) abhorret.
Epiphytic or rarely lithophytic herbs, to 1.5
m long, most commonly to 50 cm long or less;
plants first suberect or sprawling to creeping,
148 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 149
FIGURE 3. Ornithidium elianae Carnevali & M.A. Blanco. A, plant habit; B, flower, front view; C, flower, side
view; D, dissected perianth; E, labellum and column attached to ovary, sepals and petals removed, side view.
Drawn by G. C. K. Dunsterville from Dunsterville 204 (voucher not found), from Guatopo (Estado Miranda,
Venezuela). This illustration was published asMaxillaria “taphallae” Rchb.f. (sic.) in Dunsterville and Garay
(1959), and asM. ramosa Ruiz & Pav. in Dunsterville and Garay (1976: 37; 1979) and Romero and Carnevali
(2000).
eventually arching to pendent, usually growing
on thick branches or tree boughs in cloud
forests. Roots cylindrical, 1 mm in diameter.
Stems sympodial, always terminated by a
pseudobulb. Rhizome to 4 mm diameter, first
covered with thin, scarious, eventually evanes-
cent sheaths, becoming naked and brownish;
branches divaricate, usually two, produced from
the axils of consecutive non-foliar bracts imme-
diately behind pseudobulb; the segments of rhi-
zome between pseudobulbs made up of few,
elongated internodes, the pseudobulbs 5–20 cm
apart on the rhizome. Pseudobulbs 1.5–4.2 cm
long, 1.4–2.5 cm wide, silvery gray-green, grad-
ing to silvery-brown, smooth and slightly wrin-
kled, apically 1-leaved, ellipsoid, ovoid to
(rarely) suborbicular, slightly laterally com-
pressed, clothed by several imbricate sheaths of
which the 1(–2) innermost bear foliar blades;
leaves and sheath blades early caducous (mature
pseudobulbs usually devoid of them). Leaves
and the blades of the sheaths bright green, sub-
coriaceous, conduplicate, articulate, linear-ellip-
tic, narrowly elliptic to narrowly oblong-elliptic,
the margins slightly revolute, the apex acute to
shortly acuminate, oblique due to the fact that
one of the halves is conspicuously shorter than
the other, keeled abaxially, 3.5–17.0 cm long,
0.5–1.6(–2.3) cm wide, the sheaths 2–3 cm long.
Inflorescences 1-flowered, borne singly or sev-
eral from the axils of the sheaths enveloping the
developing or youngest pseudobulbs, up to 20
flowers produced successively per shoot over a
long period, only 1–3(–4) flowers are open on
any given shoot simultaneously; peduncle 10–
15 mm long, cylindrical, basally with 1–2 thin,
soft, brown bracts; ovary with pedicel 5.5–6.5
mm long, floral bracts ca. 3 mm long, tubular,
acuminate. Flowers small and inconspicuous,
resupinate, subcampanulate, perianth segments
white or greenish-white, more rarely with pink
tinges, the column greenish-yellow; the petals
and sepals thin-membranous, almost translu-
cent, with a heavily thickened midnerve dorsally
which is slightly sulcate on the inner face.
Sepals 4.5–6.0 mm long, 1.7–2.2 mm wide,
oblong elliptic to lanceolate, obtuse to acute,
basally concave, flat to convex distally, the lat-
eral sepals slightly oblique, somewhat longer
and wider than dorsal sepal, basally produced
into a small, obtuse mentum. Petals 3.5–5.0 mm
long, 1.2–1.7 mm wide, oblong to oblong-
oblanceolate to narrowly obovate-oblanceolate,
acute to obtuse. Labellum 4–7 mm long, 2.5–4.0
mm wide upon flattening, in general outline
elliptic to obovate-elliptic from a subcuneate
base; rigidly attached to the column foot, 3-
lobed below middle, middle lobe 2.5–3.1 mm
long, 2.2–2.9 mm wide, oblong subquadrate to
almost suborbicular, smooth, apically emar-
ginate to bilobed, the margins erect (thus the
lobe deeply concave), the margin finely dentate
to irregularly crenate; lateral lobes 1.0–1.5 mm
long, ca. 0.7 mm wide, erect, porrect in natural
position and enfolding the column, the free por-
tions suborbicular to elliptic; the disk provided
with a callus consisting of a transverse plate
between the bases of the lateral lobes. Column
subcylindric, relatively short and thick, basally
produced into a short foot, 2.8–4.1 mm long;
pollinia not seen. Fruit an ovate, pendent, dehis-
cent capsule, 9 mm long, 7 mm wide, valves
separating apically upon maturity.
Habitat and ecology: locally common in
many places of the Venezuelan Coastal Range
at 600–1600 m, frequently in cloud forests. The
plants grow as creeping epiphytes first, but
eventually become huge, heavy mats and their
long stems become arching and pendent. Plants
are often found fallen on the forest floor after
storms or severe rainfall, but are incapable of
surviving in the deep shade of the cloud forest
understory and eventually die (G. Carnevali,
pers. obs.).
Phenology: data from herbarium specimens
and from records published by Dunsterville and
Dunsterville (1967; asMaxillaria ramosa) indi-
cate that flowering occurs sporadically through-
out the year.
Conservation status: common only in the
coastal cordillera of northern Venezuela, but it
is protected in several national parks in that area
(Canaima, El Avila, Guatopo, Macarao, San
Esteban, and Yurubí; G. Carnevali, pers. obs.).
This species is not threatened.
Illustrations: first illustrated by Dunsterville
and Garay (1959) as Maxillaria “taphallae”
(sic.); this illustration is reproduced here (Fig.
3). After Garay’s (1967) unfortunate confusion,
Dunsterville and Garay (1976: 37) changed the
name to Maxillaria ramosa, and illustrated
O. pendulum under its synonym Maxillaria
ochracea. These confused determinations
were perpetuated in Orchids of Venezuela: An
150 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
Illustrated Field Guide (Dunsterville and Garay,
1979) and its recent revised edition (Romero
and Carnevali, 2000). Foldats (1970) also
cited and illustrated O. elianae as Maxillaria
ramosa.
Eponymy: named after Eliana Noguera,
curator of Orchidaceae in the Venezuelan
National Herbarium (VEN) who kindly pro-
vided us with material, data, and images of this
novelty, O. pendulum, and other related species
from Venezuela.
Distribution: known from northern
Venezuela (where it is relatively common and
widespread) and the Guiana Shield (in southern
Venezuela, Guyana, Surinam, and French
Guiana, where it is local and rare). In Venezuela,
it occurs in the Sierra de San Luis, the Coastal
Range (including Caracas; Dunsterville and
Dunsterville, 1969; as Maxillaria ramosa), and
from two collections in the Venezuelan Andes
(where both O. elianae and O. pendulum are
rare but potentially sympatric). There is an
unconfirmed report of this species from Cerro
Auyantepui in the Venezuelan Guayana
(Foldats, 1970; Carnevali and Ramírez, 2003;
Funk et al., 2007). It is also known from a sin-
gle collection in western Guyana, and is
reported from French Guiana and Surinam
(Cremers and Hoff, 1992; Boggan et al., 1997;
Chiron and Bellone, 2005; Funk et al., 2007; all
as M. ramosa, vouchers not seen by us); these
reports are from areas adjacent to the known dis-
tribution ofO. elianae and most likely represent
this species instead of O. pendulum, which has
not been collected in the Guiana Shield.
Ortiz’s illustration of “Maxillaria ramosa”
(Ortiz, 1988, 1995) is a crude tracing of
Dunsterville’s line drawing of O. elianae (Fig.
3). As far as we know, however,O. elianae does
not occur in Colombia.
Additional specimens examined:GUYANA.
Cuyami-Mazaruni, Paruima, 9 km W, Ararata
scrub area, 05˚49' N, 61˚08'W, 800 m, 3 July
1997 (fruit), Clarke et al. 5252 (US).
VENEZUELA.Aragua: Dpto. Girardot, Rancho
Grande bei Maracay, Parque National
Regenwald, 1400 m, 22 May 1963 (flowers),
Renz 10203 (RENZ). Carabobo: cabeceras del
Rio San Gián, arriba de La Toma, al sur de
Borburata, 800 m, 30 March 1966 (flowers),
Steyermark and Steyermark 95631 (VEN).
Distrito Federal: Cerro Naiguatá, arriba del
pueblo de Naiguatá, Lomas de Las Delicias,
entre Quebrada de Basenilla y Quebrada
Guayoyo, 9–12 km suroeste de Hacienda
Cocuizal. 1500–1635 m, 15–19 November 1963
(sterile), Steyermark 92010 (AMES, ECON,
VEN). Falcón: Sierra de San Luis, entre La
Chapa y Uria, 1400 m, 19 July 1967 (sterile),
Steyermark 99199 (AMES). Mérida: Pico
Espejo, 18 January 1964 (flowers), Ehiendorfer
s.n. (WU). Bei Rodeo Grande, collected by
Ehiendorfer, cultivated in the Botanisches
Institut der Universität Wien, 26 January 1971
(flowers), Vöth s.n. (WU). Trujillo: Quebrada
Palmichero, between Escuque andMt. Carmelo,
1400 m, flowered in cultivation in Trujillo, 15
April 1949 (flowers), Renz 5378 (RENZ).
Yaracuy: Distrito San Felipe, cabeceras del rio
Taria, 12 km al norte de Salom, El Amparo,
10˚15'N, 68˚29'W, 1050 m, 7 December 1980
(sterile), Steyermark and Carreño Espinoza
123788 (SEL).
Ornithidium elianae is very similar to O.
pendulum and O. sillarense but is distinguished
by its more elongate pseudobulbs, and its thin-
ner, narrower leaves and sheath blades, 0.5–
1.6(–2.3) cm wide (vs. 2.2–4.5 cm wide) that are
shed upon maturation of the pseudobulb (vs.
thicker, relatively wider, and persistent in O.
pendulum and O. sillarense). The flowers of O.
elianae are produced a few at a time per growth
(vs.O. pendulum andO. sillarense, that can pro-
duce several to many flowers simultaneously),
and each flower has a straight, concave, and
smooth labellum midlobe (vs. reflexed, convex,
and verrucose in O. pendulum).
Ornithidium elianae could potentially be
confused also withO. histrionicum Rchb.f. (syn-
onyms: Maxillaria histrionica (Rchb.f.) L. O.
Williams and M. aristeguietae Foldats), which
also has long rhizome segments separating the
narrowly ovate pseudobulbs, and small, green-
ish flowers (Dunsterville and Garay, 1976,
1979; Romero and Carnevali, 2000). However,
the leaves ofO. histrionicum are generally much
shorter, narrower, and more rigid, the flowers
are slightly larger and fleshier, and the labellum
midlobe is markedly convex and has a promi-
nent, subapical mucron abaxially (vs. concave
and emarginate in O. elianae). Furthermore, in
O. histrionicum pseudobulbs tend to be pro-
duced only at or near the base of the plant, and
they become increasingly smaller and absent
2008 BLANCO ETAL., A TAXONOMIC PUZZLE (ORCHIDACEAE) 151
toward the distal part of the long branches.
In contrast, O. elianae pseudobulbs of approxi-
mately equal size are always produced at the end
of each sympodium. Both species occur sym-
patrically in some areas.
Ornithidium lasallei (Foldats) M. A. Blanco
& Ojeda is very similar to O. histrionicum and
thus also similar to O. elianae. However, the
flowers are larger (sepals 20–27 mm long), and
the more membranous labellum has proportion-
ally less well-developed lateral lobes and
a much longer central lobe than those of O.
pendulum and O. elianae. Ornithidium lasallei
appears to be restricted to the eastern part of the
Guayana area in Venezuela and western Guyana
at elevations of 700–1500 m (Carnevali and
Ramírez, 2003), where it is probably sympatric
with O. elianae, which is rare in this area.
Ornithidium lasallei has thicker rhizomes than
O. elianae, which also tend to creep over the
phorophyte’s bark for most of its length and
only eventually become pendulous, as opposed
to O. pendulum and O. elianae, whose long rhi-
zomes become pendent early in the development
of the plant.
Herbarium specimens ofOrnithidium elianae
have commonly been misidentified as
Maxillaria tafallae or M. ramosa.
152 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
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