A Synopsis of Cohniella

A synopsis of Cohniella (Orchidaceae, Oncidiinae)
GERMAN CARNEVALI FERNÁNDEZ-CONCHA1, WILLIAM ROLANDO CETZAL IX2,

RICARDO BALAM NARVÁEZ1, AND GUSTAVO A. ROMERO-GONZÁLEZ3
1
Herbario CICY, Centro de Investigación Científica de Yucatán, Calle 43. No. 130. Col. Chuburná
de Hidalgo, 97200 Mérida, Yuc., México; e-mail: [email protected]; [email protected]
2
El Colegio de la Frontera Sur, Unidad Chetumal, Av. del Centenario, km 5.5, Chetumal, Quintana
Roo, Mexico. C.P. 77000; email: [email protected]
3
Orchid Herbarium of Oakes Ames, Harvard University Herbaria, 22 Divinity Avenue, Cambridge,
Massachusetts 02138, USA; e-mail: [email protected]
Abstract. A synopsis of the genus Cohniella is presented with nomenclatural updates

with a discusion of diagnostic features and biogeographical data. Thirteen species and
one natural hybrid are recognized. Two new species, Cohniella pendula and Cohn-
iella biorbicularis, are described and illustrated. The following four new combina-
tions are proposed: Cohniella binotii, Cohniella brachyphylla, Cohniella cepula,
and Cohniella longifolia. Lectotypes are selected for Oncidium subulifolium, On-
cidium helicanthum, Oncidium humboldtii, Oncidium jonesianum var. phaean-
thum, Oncidium cepula, and Oncidium wittii. We also propose an amended
lectotype for Epidendrum cebolleta and several epitypifications and new synonyms.
A key to the genera of the Trichocentrum complex is presented (Appendix) as well as
keys to the species of Cohniella.
Key Words: Cohniella, Oncidiinae, Orchidaceae, new taxa, new combinations,
typifications.
Resumen. Se presenta una sinopsis del género Cohniella, con actualizaciones nom-
enclaturales y discusiones de caracteres diagnósticos y datos biogeográficos. Se ace-
ptan 13 especies y un híbrido natural en el género. Dos especies nuevas, Cohniella
pendula y Cohniella biorbicularis, se describen e ilustran. Se proponen las siguie-
ntes combinaciones nuevas: Cohniella binotii, Cohniella brachyphylla, Cohniella
cepula y Cohniella longifolia. Se seleccionan lectotipos para Oncidium subulifo-
lium, Oncidium helicanthum, Oncidium humboldtii, Oncidium jonesianum var.
phaeanthum, Oncidium cepula y Oncidium wittii. También proponemos una mo-
dificación a la lectotipificación de Epidendrum cebolleta y varias epitipificaciones y
nuevos sinónimos. Se presenta una clave para los géneros del complejo Trichocen-
trum en un apéndice, así como una clave para las especies de Cohniella.
The genus Cohniella Pfitzer (Orchidaceae, 1863; Kränzlin, 1897; Garay & Stacy, 1974;
Oncidiinae) is characterized by medium to Dunsterville & Garay, 1979; Foldats, 1970;
large plants with inconspicuous to small pseu- Mora de Retana, 1999). Within Oncidium, rat-
dobulbs that bear a single, succulent, terete leaf. tails traditionally were referred to section
It includes 13 species that are known in Cebolletae Lindl. (Lindley, 1842; Garay &
horticulture as the “rat-tail oncidiums”. In Stacy, 1974) or to section Teretifolia (Lindley,
common with other groups within Oncidiinae 1851, 1855; Reichenbach, 1863; Cogniaux,
that bear yellow flowers that mimic malpighia- 1906).
ceous blossoms, the species of Cohniella were Over time, it became evident that the rat-
first included within a broadly defined genus tails were different from true Oncidium in
Oncidium Sw. (Lindley, 1855; Reichenbach f., several important ways, such as their incon-
Brittonia, 62(2), 2010, pp. 153–177 ISSUED: 1 June 2010

© 2010, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
154 BRITTONIA [VOL 62
spicuous or small pseudobulbs bearing scar- be the already mentioned Cohniella, Lophia-
ious sheaths and one terete leaf, their low ris, Lophiarella, and Trichocentrum.
chromosome numbers (2n=26–36; Chase, The rationale for recognizing these segre-
1986; Chase & Palmer, 1989), and their gates follows several criteria outlined by
distinctive leaf anatomy (Sandoval-Zapotitla Backlund and Bremer (1998) for the deriva-
& Terrazas, 2001). In contrast, species of tion of classifications from phylogenies: (1) To
Oncidium in a narrow sense are characterized merit recognition clades must be monophyletic
by conspicuous, laterally compressed pseu- and highly supported; (2) Nodes defining
dobulbs bearing one to several sheaths, of genera preferably should include morpholog-
which 1–3 may have foliar blades, and 1–3 ical synapomorphies that permit recognition of
softly coriaceous, conduplicate leaves. These members; and (3) The classification should be
“typical oncidiums” also have higher chro- as consistent as possible (minimally disrup-
mosome numbers of 2n=56–60 (reviewed in tive) with previous classification systems.
Tanaka & Kamemoto, 1984). All these genera form highly supported,
Orchid hybridizers realized early on that it monophyletic clades in phylogenetic analyses
was easier to obtain hybrids of the rat-tails with of both nuclear and plastid DNA (Sosa et al.,
species of the genus Trichocentrum Poepp. & 2001; Williams et al., 2001a, b; Cetzal et al.,
Endl. (Oncidiinae) and particular groups within in prep.), and of morphological and anatom-
the genus Oncidium than with the species of ical characters (Cetzal, 2007) . All of these
typical oncidiums (Sanford, 1964). These groups are diagnosable using easily discern-
groups include the “mule ear oncidiums” (here able morphological characters (Cetzal, 2007;
referred to the genus Lophiaris Raf.) and the Cetzal et al., in preparation; see key to the
Oncidium splendidum A. Rich ex Duch. com- genera in the Appendix). There are also
plex (for which the name Lophiarella Szlach., micromorphological and anatomical charac-
Mytnik & Romowicz is available; Szlachetko ters that distinguish the segregates (Sandoval-
et al., 2006). Not coincidentally, the plants of all Zapotitla & Terrazas, 2001; Cetzal et al., in
the species that hybridize with the rat-tails have prep.). The third criterion, that of consistency
low chromosome numbers and relatively small with previous systems, is more problematic in
pseudobulbs bearing scarious sheaths and a this case because an array of names and
single coriaceous or succulent leaf (hereafter treatments have been recently proposed in
called the “Trichocentrum syndrome”, TS), Oncidiinae; some support a broadly defined
bespeaking of a relationship between them Trichocentrum (e.g., Sandoval-Zapotitla &
and the rat-tails. Terrazas, 2001; Sosa et al., 2001; Williams
Recent phylogenetic analyses of nucleotide et al., 2001a, b; 2003; Chase et al., 2005;
sequences of both nuclear (Sosa et al., 2001; Chase et al., 2003; 2009) while others prefer
Williams et al., 2001a) and plastid (Chase & more narrowly defined genera (e.g., Pupulin,
Palmer, 1989) DNA have shown that all the 1995; Romero & Carnevali, 2000; Carnevali
taxa that exhibit TS form a highly supported, et al., 2001; Carnevali et al., in prep.;
monophyletic entity. Some authors have pre- Jiménez-Machorro & Carnevali, 2001; Pupu-
ferred to treat them as members of a broadly lin & Carnevali, 2005; Cetzal et al., 2008).
defined Trichocentrum (e.g., Chase et al., Thus, names and classification systems for
2005; Sandoval-Zapotitla & Terrazas, 2001; both options are already available (i.e., practi-
Sosa et al., 2001; Williams et al., 2001a, b; cally all the required combinations have
Chase et al., 2009) while others prefer to already been proposed) and which course of
recognize several smaller, more narrowly action is chosen must remain a matter of
defined and easily diagnosable genera (e.g., “practicality.”
Braem, 1993; Pupulin, 1995; Königer & We have chosen to treat the major clades
Pongratz, 1997, 1999; Jiménez-Machorro & within the Trichocentrum complex as distinct
Carnevali, 2001; Romero & Carnevali, 2000; genera primarily for two reasons, discussed
Carnevali et al., 2001; Pupulin & Carnevali, below. In a subtribe such as Oncidiinae, where
2005). If recognized, the segregates of a the generic circumscriptions are tenuous and
narrowly defined Trichocentrum clade would confusing at best, cryptic at worst (e.g., the
2010] CARNEVALI ET AL.: COHNIELLA (ORCHIDACEAE) 155
generic limits within the clade that includes Type: Cohnia quekettioides Rchb.f. [=Coh-
the twig-epiphytes, see Chase, 1986), a nar- niella ascendens (Lindl.) Christenson].
rowly defined Trichocentrum and Cohniella, Oncidium sect. Cebolletae Lind., Bot. Reg.
Lophiaris, and Lophiarella are among the most 28: sub t .4. 1842. Type: Epidendrum
distinctive species aggregates. Although all cebolleta Jacq. [=Cohniella cebolleta
members of the clade share pseudobulbs bear- (Jacq.) E. A. Christenson]
ing a single succulent leaf, all are easily Oncidium sect. Teretifolia Lindl., Bot. Reg.
recognizable using other characters. Trichocen- 32: sub t. 27. 1846. Type: Epidendrum
trum is unique due to its spurred flowers on cebolleta Jacq.
short, mostly successively-flowered racemes, Oncidium sect, Teretoncidium Kuntze, Lex.
and a consistently small vegetative habit. The Gen. Phan. 399. 1903. Type: Epidendrum
plants of all the other genera bear spurless, cebolleta Jacq.
Oncidium-like flowers. Plants of Cohniella are Stilifolium Königer & Pongratz, Arcula 7: 186.
medium to large in size and bear terete leaves 1997. Type: Epidendrum cebolleta Jacq.
and erect to pendent inflorescences. Lophiaris
is characterized by tiny pseudobubs bearing Epiphytic or lithophytic herbs, caespitose;
succulent, relatively large leaves and arching to pseudobulbs small (relative to leaf length),
twinning inflorescences. Plants of Lophiarella suborbicular, bearing one leaf, clothed by three
have relatively large pseudobulbs bearing thick, small, scarious sheaths; leaf terete, thickly
fleshy leaves and erect, stout inflorescences fleshy, often punctate or tinged with red or
covered with a thin coat of wax. We offer a key purple; inflorescences lateral originating from
to the four genera of the Trichocentrum com- the sides of the newest pseudobulbs, usually
plex that is entirely based on macroscopic, paniculate, rarely racemose, few to multi-
easily discernable characters (see Appendix). flowered, with the flowers opening more or
Furthermore, these four genera are horticultur- less simultaneously, peduncle clothed with
ally distinct (i.e., each has different cultural small, remote, scarious bracts; flowers showy,
requirements), a fact that orchid growers have resupinate, with the perianth segments subcor-
recognized (Sanders & Co., 1901). We predict iaceous, widely spreading, relatively long
that, in this particular case, narrowly defined lasting (5–10 days); floral bracts inconspicu-
genera will be better accepted by aficionados, ous; pedicellate ovary cylindrical; sepals sub-
the economically important horticultural trade, similar, clawed basally, the laterals somewhat
and in orchid conservation, rather than a oblique, usually green, chartreuse, or yellow
morphologically and horticulturally heteroge- with red or maroon spots or blotches; petals
neous, broadly circumscribed Trichocentrum. similar to the sepals, the claw poorly devel-
The objective of this contribution is to provide oped; labellum larger than the other perianth
a synopsis of Cohniella, while clarifying many segments, yellow (white in C. jonesiana) with
of the problems associated with an orchid group red or maroon spots or blotches on or around
that has a long history of cultivation and the callus area or the abaxial surface; con-
numerous nomenclatural changes. Several spicuously 3-lobed, the central lobe usually
names are typified, and a brief discussion is much larger than the laterals and joined to the
provided for each species, stressing their distri- disk by a more or less well-developed claw,
bution, variability, circumscription, and diagnos- callus composed of a series of plates and/or
tic characters. Furthermore, we describe and teeth; column relatively short, subcylindrical,
illustrate two new species and propose five new thicker at the base, stigmatic surface suborbic-
combinations. Illustrations are also provided for ular, with well-developed stelidia at each side
most of the species treated in this synopsis. (almost absent in C. nuda), with a more or less
well-developed tabula below the stigmatic
surface, rostellum transverse; anther terminal,
Synopsis
operculate; pollinarium complex with 2 yellow
Cohniella Pfitzer, Nat. Pflanzenfam. 2, 6: 194. obpyriform pollinia, an obovate or spathulate
1889. Cohnia Rchb. f., Bot. Zeit. 10: 928. tegula, and a small, semi-hemispheric visci-
1852, non Cohnia Kunth 1850 (Agavaceae). dium. Fruit a capsule.
A Neotropical genus of 13 species, some vegetative maximum. In nature, any given

(e.g., Cohniella cebolleta and C. cepula) are population will include plants bearing flowers
rather broadly circumscribed in this synopsis varying in size from small to large, the latter
and geographical variants may eventually be with leaves 2–3 times longer than those of the
recognized as distinct taxa. The genus is smaller plants, thus rendering plant size unreli-
distributed widely from northern Mexico (C. able for species recognition. However, some
brachyphylla reaching ca. 27° N in Sonora) species are, on average, smaller than others, and
into southern Brazil and northern Argentina species in certain groups within the genus tend
(reaching ca. 26° S in Tucumán, Argentina), to be smaller than those in other groups (e.g., the
mostly in the lowlands. The species are most plants of species in the Cohniella nuda complex
common in seasonally dry forests. The genus tend to be smaller than those of species in the C.
is most diverse in Mexico where at least five cebolleta complex).
species occur, two or three of them endemic, There is also considerable variation in the
and in Panama with four species. flowers of most species. In Cohniella ascendens,
Cohniella was based on Cohnia quekettiodes the flowers vary to such an extent that it could be
Rchb.f. Reichenbach f. (1852) apparently mis- easy to assume that one deals with more than
interpreted the relatively small size of the plant one species except that most of the variation is
he described and the flower buds he examined found within a single population, even among
(see Garay, 1963) and inferred a relationship plants growing on the same phorophyte. Varia-
with a totally unrelated clade of oncidioid tion involves in some measure flower size, but
orchids that also happens to bear terete leaves mostly resides in the outline of the labellum
(he compared it to his genus Sigmatostalix and lobes, and the degree and depth of the color
stated that “... Quekettia [Lindl.] was likely to patterns. Despite all this variation, most species
be closely related”: “Quekettia wird wohl auch are easily diagnosed by the combinations of
sehr verwandt sein”.) The same author (1858) floral features that we discuss below.
later provided a Latin description of his species Inflorescence length, on the other hand,
(he had given only a brief description in though variable in absolute dimensions
German in the protologue) comparing it to within species, is diagnostic for species or
Oncidium ascendens Lindl. (“Planta habitu species groups and tends to remain within a
Oncidii ascendentis cui et vaginis amplis range. For example, the inflorescence in
brunneo fasciatis simile” [sic]) and an illustra- species of the Cohniella nuda complex of
tion, which does appear to confirm that the type species is most commonly conspicuously
plant bore only immature flowers. The use of shorter than the leaves while the opposite is
the name Cohnia was, however, already occu- true for species related to C. cebolleta.
pied by a genus in the Agavaceae described by The 13 species of Cohniella recognized in
Kunth in 1850. Pfitzer (1889) later proposed a this synopsis can be diagnosed with the use of
new genus and a combination, Cohniella the following set of keys. For ease of
quekettiodes (Rchb. f.) Pfitzer, which Garay identification, we have chosen to offer three
(1973) referred to the synonymy of Oncidium separate keys, which are geographically cir-
ascendens Lindl. cumscribed. Since Cohniella longifolia and its
As treated here, most of the species of variation are not well-known, we have decided
Cohniella are rather variable entities. The nature to leave it out of the keys. For this key to work
of the variation is twofold, having both vegeta- appropriately, users need good flowers and
tive and floral components. Most species reach preferably a portion of an inflorescence and
flowering size at a fraction of their potential the subtending leaf.
Keys to species of Cohniella

1. Plants from Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey I
1. Plants from Central America, South America, or the West Indies.
2. Plants from Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Subkey II
2. Plants from South America and the West Indies . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey III
Subkey I: Key to the species of Cohniella from Mexico

1. Inflorescences usually shorter than the subtending leaves, rarely as long; lateral lobes of the labellum erect
and partially enfolding the column; petals and sepals usually much longer than the lateral lobes of the
labellum; column wings linear, finger-like, much longer than wide; plants from tropical moist to tropical
subdeciduous forests on the Gulf Coast. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ascendens
1. Inflorescences usually longer to much longer than the subtending leaves; lateral lobes of the labellum flat (in the
same plane than the central lobe), flat and not enfolding the column; petals and sepals usually much shorter than
the lateral lobes of the labellum; column wings transversely reniform or elliptic, bilobed, broader than wide,
rarely slightly longer than wide; plants usually from tropical dry forests from all over Mexico.
2. Central lobe of the labellum similar in shape and size to the lateral lobes, 5.8–8 mm wide; leaves
pendent; plants from the western extreme of the Neovolcanic Transversal Axis in coastal Jalisco and
Nayarit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pendula
2. Central lobe of the labellum different in shape and size (usually larger) from the lateral lobes,
21 mm wide; leaves usually erect or patent, rarely pendent; plants from other areas, but not from
western extreme of the Neovolcanic Transversal Axis.
3. Leaves rigidly erect, 5.5–16 (−28) cm long, many on the plant simultaneously (5–15); inflorescences
rigidly erect, racemose, more rarely with a single lateral branch when well-developed; plants from
north of the Neovolcanic Transversal Axis in Sinaloa with outliers in Durango, Chihuahua, and
Sonora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brachyphylla
3. Leaves various, either rigidly erect to flexible, usually few (3–5) to a plant; inflorescences various
but usually patent to somewhat pendent, more rarely erect, usually a panicle with 2 or more
branches in well-developed plants; plants from the Gulf coast of from the Pacific coast south of the
Trans-MexicanVolcanic Belt.
4. Lateral lobes of the labellum as broad as long, almost as broad as the central lobe (length/
width=0.85/1–1/1); plants from the Gulf states W and north of the Tehuantepec isthmus
(Querétaro, San Luis Potosí, Tamaulipas, Veracruz) . . . . . . . . . . . . . . . . . . . . . . C. biorbicularis
4. Lateral lobes of the labellum always longer than wide; plants from the Yucatan peninsula or
from the Pacific Coast of Mexico.
5. Central lobe of the labellum deeply emarginate, the emargination making an acute angle;
labellum lacking spots on the undersurface, or if present only on the underside of the
disk; leaves 2.5–4 mm wide, of homogeneous width, not tapering basally and distally;
plants from the northern portion of the Yucatan Peninsula . . . . . . . . . . . . C. cebolleta
5. Central lobe of the labellum rounded to truncate, not emarginate at all or only very
shallowly so and then the emargination making an obtuse angle; labellum usually bearing
spots on most of the undersurface; leaves 6–10 mm wide, conspisuously wider basally
but abruptly tapering at base (forming a “neck” just above the pseudobulb) and more
gradually tapering distally; plants from the Pacific states of mexico, south of the
Tehuantepec isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brachyphylla
Subkey II: Key to the species of Cohniella from Central America
1. Inflorescences usually longer than the subtending leaves; column wings broader than their width, 2-lobed;
column tetragonal in cross-section.
2. Central lobe of the labellum deeply emarginate, the emargination making an acute angle; labellum
lacking spots on the undersurface, if present only on the underside of the disk; leaves 2.5–4 mm wide,
of homogeneous width, not tapering basally and distally . . . . . . . . . . . . . . . . . . . . . . .C. cebolleta
2. Central lobe of the labellum rounded to truncate, not emarginate at all or only very shallowly so and then
the emargination making an obtuse angle; labellum usually bearing spots on most of the undersurface;
leaves 6–10 mm wide, conspisuously wider basally but abruptly tapering at base (forming a “neck” just
above the pseudobulb) and more gradually tapering distally . . . . . . . . . . . . . . . . . . C. brachyphylla
1. Inflorescences usually shorter than the subtending leaves, rarely as long; column wings either absent (C. nuda) or
longer than their width, simple; column subterete in cross-section.
3. Labelar isthmus about as long as wide, at least half the width of the central lobe; callus complex,
composed of 5 teeth or keels.
4. Lateral lobes of the labellum erect and partially enfolding the column; flowers always resupinate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ascendens
4. Lateral lobes of the labellum flat, not enfolding the column; flowers resupinate or not
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. teres
3. Labelar isthmus much longer than wide, maximum 1/3 of the width of the central lobe; callus simple,
composed of 1–3 teeth or keels.
5. Column wings absent or almost; callus composed of a single, slightly raised central keel
flanked by longitudinal depressions; plants from Venezuela, Colombia (eastern side of the
Andes), and Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nuda
5. Column wings present, subterete; callus composed of a single, massive central teeth occupying the
whole width of the isthmus; plants from Costa Rica and Panama . . . . . . . . . . . . . . C. stipitata
Subkey III: Key to the species of Cohniella from South America and the West Indies
1. Inflorescences usually shorter than the subtending leaves, rarely as long; flower diameter not exceeding 3.5 cm;
column wings either absent (C. nuda) or longer than their width, simple; column subterete in cross-section.
2. Labelar isthmus much longer than wide, maximum 1/3 of the width of the central lobe; callus simple
composed of 1–3 teeth or keels; column wings almost absent, if marginally present broader than long
or as long as wide; plants from western and central Venezuela and the eastern slopes of the Andes in
Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nuda
2. Labelar isthmus about as long as wide, at least half the width of the central lobe; callus complex,
composed of 5 teeth or keels; column wings conspicuous, longer than wide; plants from the Cauca
valley and presumably from the Chocó region in Colombia . . . . . . . . . . . . . . . . . . . . . . . .C. teres
1. Inflorescences usually longer than the subtending leaves; flower diameter variable; column wings broader
than their width, 2-lobed; column tetragonal in cross-section.
3. Flower diameter not exceeding 3.5 cm; petals and sepals much shorter than the labellum; leaves erect or
pendent; plants from most of South America.
4. Callus composed of 3 keels, lacking accessory cally at each side of the ishtmus; column basally
inflated; column wings perpendicular to the proximal-distal axis of the column; plants from south
of the Amazon river course, reaching SE Brazil and Paraguay and along the eastern side of the
Andes in Peru and Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. cepula
4. Callus composed of 5 keels, provided with accessory cally at each side of the labellar isthmus; column of
equal width at base and apex: column wings on the same plane as the proximal-distal axis of the column;
plants from Venezuela, Colombia, the Guianas, and the West Indies. . . . . . . . . . . . . . . . . C. cebolleta
3. Flower diameter at least 3.5 cm; petals and sepals as long as the labellum; leaves almost always pendent;
plants from Bolivia, Paraguay, N. Argentina, and SE Brazil.
5. Labellum yellow with orange-brown spots or blotches; distal portions of the lateral lobes of the
labellum long fimbriate-ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stacyi
5. Labellum white or rarely with a few red spots along the proximal rim of the central lobe; distal
portions of the lobes of the labellum entire to dentate
6. Sepals and petals pale yellow-green or chartreuse colored with a few, widely separated red-
brown spots; leaves mostly erect; plants from Bolivia, Paraguay, Northern Argentina, and
neighbouring areas of Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. jonesiana
6. Sepals and petals pale yellow-green or chartreuse, almost entirely covered by confluent dark red-
brown blotches which render the perianth segments almost unicolor; leaves mostly pendent;
plants from Minas Gerais in SE Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. binotii
Cohniella ascendens (Lindl.) Christenson, Oncidium subulifolium Schltr., Repert. Spec.

Lindleyana 14(4): 177. 1999. Oncidium Nov. Regni Veg. Beih. 10: 79. 1922.
ascendens Lindl., Edwards’s Bot. Reg. 28: Oncidium bolivianense Oppenheim, Orchis
sub t. 4. 1842. Stilifolium ascendens (Lindl.) 10: 93. 1916. (non Oncidim boliviense
Königer & Pongratz, Arcula 7: 186. 1997. Rolfe, 1907). Type: Bolivia. Río Itenez,
Trichocentrum ascendens (Lindl.) M. W. O. N. Witt s.n. (holotype: B, destroyed;
Chase & N. H. Williams, Lindleyana 16(2): lectotype, here designated, Orchis 10, No.
137. 2001. Type: Guatemala. Without any 5, Tafel IV, Fig. 2. 1916).
other locality, Apr 1841, K. T. Hartweg s.n. Oncidium helicanthum Kraenzl., Pflanzenr.
(holotype: K-Lindl.). (Fig. 1A–D) (Engler) 95: 281. 1922. Type: Colombia.
Without any other locality or collector
Cohnia quekettioides Rchb. f., V. Schl. Bot. (holotype: B, destroyed; lectotype, here
Zeitung (Berlin): 10: 928. 1852. Cohniella designated, Das Planzenreich (A. Angler)
quekettioides (Rchb. f.) Pfitzer, Nat. Pflan- heft 80, 4, 50: 282, Fig. 24C, a–d. 1922).
zenfam 2(6): 194. 1889. Type: Guatemala.
Chantalas, “Mons Espina”, 1841, E. R. von Diagnostic features.—Cohniella ascendens
Friedrichsthal 834 (holotype: presumably at is easily recognized by its relatively small
W, not seen). lateral lobes of the labellum that are held
FIG. 1. Flowers of Cohniella. A–D. Cohniella ascendens. A. Whole flowers, frontal view. B. Labellum, frontal
view. C. Labellum, dorsal view. D. Column with anther cap, frontal view. E–H. Cohniella nuda. E. Whole flowers,
frontal view. F. Labellum, frontal view. G. Labellum, dorsal view. H. Column with anther cap, frontal view. I–L.
Cohniella stipitata. I. Whole flowers, frontal view. J. Labellum, frontal view. K. Labellum, dorsal view. L. Column
with anther cap, frontal view. M–P. Cohniella teres. M. Whole flowers, frontal view. N. Labellum, front view. O.
Labellum, dorsal view. P. Column with anther cap, frontal view (A–D from Cetzal 6, CICY; E–H from Carnevali
7283, CICY; I–L from Carnevali 7311, CICY; M–P from Carnevali 7027, CICY).
FIG. 2. Flowers of Cohniella. A–G. Cohniella biorbicularis. A. Whole flower, front view. B. Whole flower, front and
back view. C. Labellum, front and back views. D. Sepals and petals, front and back views. E. Column with anther cap, front
and back views. F. Anther cap. G. Pollinia, front and back view. H–O. Cohniella pendula. H–J. Whole flower, front view.
K. Habit with inflorescence. L. Labellum, front and back views. M. Column with anther cap, front and back views.
N. Anther cap. O. Sepals and petals, front and back views. (A from Cetzal 5, CICY; B–G from Carnevali & Ramírez 7308,
AMES, AMO, CICY; H, L–O Carnevali & Ramírez 6897, CICY; I–K Carnevali 7302, CICY.).
erect and partially enclosing the column. The laterally compressed tooth surrounded by
callus is relatively simple and consists of a lower, reduced lateral keels. The column
greatly reduced proximal, transverse plate wings are narrow and point downward and
with or without small, lateral low teeth and inward, almost in a hook-like fashion. As in
a much larger distal, apically rounded and its hypothesized closest relatives, C. teres and
C. nuda, C. ascendens is more of a shade- unlikely that the species jumps the gap of the
loving species and the leaves are relatively Andean ranges and the Amazon Basin to re-
soft and flexible. The inflorescences are appear in Bolivia. The highly schematic type
usually shorter than the subtending leaves. illustration of Oncidium helicanthum is con-
Distribution.—Mexico southward to Pan- sistent with the overall morphology of C.
ama; possibly northern Colombia and NW ascendens and, in concordance with other
Venezuela. The Colombian and Venezuelan authors (e,g., McLeish et al., 1995), it is
records of the species are somewhat doubtful tentatively placed here.
and better material is required to confirm them.
Variation range.—This is a common spe- Additional specimens examined. MEXICO.
cies with a great deal of floral variation, Campeche: a 65 km al S de Conhuas en el centro
mostly in the shape and size of the lateral ceremonial de Calakmul, Límite N del Petén Guatemalteco,
lobes of the labellum. The color of the callus 16 Mar 1983, Cabrera 4423 (MEXU). Chiapas: Mpio.
Catazajá, orilla oeste de laguna de Catazajá, 20 Abr 1999,
is also variable, ranging from almost con- Gutiérrez & Balam 6437 (CICY). Oaxaca: Juchitán, 9 km
colorous with the bright yellow labellum to al NE de Lázaro Cárdenas camino a Sta. María Chimalapa,
deep purple or even guava-colored. The 250 m, 22 Feb 1982, Cedillo & Torres 1114 (MEXU).
petals are usually oblong-spathulate, but in Quintana Roo: Othón P. Blanco, Ejido Graciano Sánchez,
some clones they can be broadly obovate. área forestal ca. 6 km al W de Valle Hermoso, 5 Mar 1995,
Trejo & Olmsted 311 (CICY). YUCATÁN: Mpio. de
Taxonomic commentary.—This is the type Valladolid, rehollada cerca de Valladolid, 27 Mar 1997,
of the genus Cohnia Rchb. f. (=Cohniella). Carnevali & Ramírez 4386 (CICY).
The type locality is not mentioned in the BELIZE. Corozal: Lower slopes of Richardson
protologue but the author cited it a few years Peak, Maya Mountains, directly N of the Junction of
later (Reichenbach f., 1858). In the same Richardson Creek and Bladen Branch, 88º46′30″ N,
16º33′35″ W, 300–620 m, 4, 6, 8 Mar 1987, Davidse
publication, he stated that he had described a & Brant 32522 (MEXU, MO).
unicate at the Naturhistorisches Museum GUATEMALA. Petén: Ruinenstadt Tikal (ca. 60 km
Wien (“Das unicum im Herbar des Wiener NE Flores/Santa Elena), ca. 3 km W von Tempel IV,
Museums”), where the collections of E. R. 17º13.387′ N, 89º37.683′ W, 440 m, Förther et al. 9986 a
von Friedrichstall are, but we have not been (M).
able to examine it. However, Garay (1973) EL SALVADOR. Ahuachapán: Plantas florecidas en
cultivo en la ciudad de Santa Ana, traídas de la orilla del
clarified the identity of the Cohniella queket-
río Nejapa, Cantón La Pandeadura, 8 km al Sur de la
tioides as being identical to the earlier ciudad de Ahuachapán, 800 m, Linares 1175 (MEXU).
Oncidium ascendens (Christenson, 1999). HONDURAS. El Paraíso: Dpto. El Paraíso. Mpio.
We have studied the original plate of Cohnia de Morocelli, Loc. Quebrada El Carrizal cerca El Plan,
quekettioides and concluded that Garay was ±2 km al E de El Plan, 17 Feb 2002, Linares 5692
correct in its identity assessment. This plate (MEXU).
features a terete leaved plant with the sche- NICARAGUA. Zelaya: Colonia Kururia, 14º41′ N,
84º04′ W, 50 m, 3 Mar 1975, Pipoly 3895 (MEXU, MO).
matic of a flower which clearly resembles the
COSTA RICA. Alajuela, Los Chiles, Refugio
entity we currently treat as Cohniella ascen- Nacional de Vida Silvestre Caño Negro, Llanura de
dens except for the reduced lateral lobes of Guatuso, Playuelas, 40 m, 3 Feb 1993, K. Martinez,
the labellum, something resulting from the Zamora, Chavarría & Flores 50 (UCR, MO).
dissection of a flower bud.
Oncidium subulifolium (O. bolivianense Cohniella binotii (Pabst) G. A. Romero &
Oppenheim 1922 non O. boliviense Rolfe Carnevali, stat. nov. Oncidium jonesianum
1907) was described from a cultivated plant var. binotii Pabst, Bradea 2: 170. 1977.
supposedly collected in Bolivia, but no Type: Brazil. Minas Gerais: regione Montes
further material referable to this species (or Claros, 24 Feb 1972, Casa Binot s.n.
to the complex of taxa related to C. ascen- (holotype: HB).
dens) has ever been collected in Bolivia, and
we assume the type plant must have been Diagnostic features and variation range.—
collected in another country. Known collec- Cohniella binotii is closely related to C.
tions of Cohniella ascendens reach their jonesiana. It differs, however, in the larger
southernmost limits in northern Colombia flowers (sepals and petals to 2.5 cm long and
(possibly into NW Venezuela) and it is highly labellum to 2.8 cm wide) and in the large,
irregular blotches in the petals and sepals that half, the claw 1.22–3.17 ×0.76–0.95 mm;
are almost confluent and cover most of the lateral sepals partially fused at the very base,
surface of the perianth segments. It also ovate-lanceolate, 6.34–8.44 × 2.07–4.33 mm;
occupies a disjunct range in SE Brazil. petals 6.70–8.13×3–4 mm, oblanceolate, the
Distribution.—Endemic to Brazil, known apex rounded, somewhat repand and reflexed
only in the area of Montes Claros in Minas in natural position; labellum 3-lobed, 10.82–
Gerais. 15.36 mm long from the base to the apex of
the central lobe, 11.81–18.91 mm wide across
Additional specimen examined. BRAZIL. Distrito the apices of the lateral lobes, the lateral lobes
Federal: Brasilia, Schumburg 19/705 (AMES). in the same plane as the central lobe and
+/−perpendicular to it; central lobe 10–11 ×
Cohniella biorbicularis Balam & Cetzal, sp.
17–18 mm, transverssely obdeltoid to sub-
nov. Type: Mexico. Querétaro: Municipio
quadrate in outline, apically rounded to
Landa de Matamoros, Camino de Matzacintla
subquadrate, basally produced into a short
al Río Moctezuma, 21°20′04″ N, 99°20′04″
isthmus, 2 × 2–3 mm; lateral lobes 13–19×5–
W, 1100 m, cañada orientada SE con vegeta- 6 mm, suborbicular to very broadly obovate,
ción de bosque tropical caducifolio sobre
apically rounded; disc large, ca. 3.5×5.5 mm,
laderas de roca caliza; colectada original-
bearing a well-developed callus, ca. 5×3 mm,
mente en el año 2006 por I. M. Ramírez consisting of a large, elevated, +/−flat, oblon-
(#1432); floreciendo en cultivo el 10 Marzo
goid, apically with two teeth, the central one
2008, G. Carnevali & I. M. Ramírez 7308
laterally compressed, the laterals very larger,
(holotype: CICY; isotypes: AMES, AMO,
rhombic, the basal portion of the callus with
MEXU, QMEX, SEL, US). (Fig. 2A–G)
sharp upper margins; column 3–3.5 × 1 mm,
tabula infrastigmatica obdeltoid to subqua-
Species haec Cohniellae brachyphyllae similis sed
floribus majoribus, lobis labelli lateralibus suborbicularibus drate, stigmatic surface, sub-rounded; column
vel latissime obovatis proportione majoris recedit. wings short, ca. 1 × 1 mm, asymmetrically
bilobed; anther 3 × 2 mm, apical, operculate,
Epiphytic herbs, caespitose; rhizome short, ellipsoid; pollinarium typical for the genus.
thin, brittle; roots 1–1.5 mm thick, white Distribution and ecology.—Endemic to
when old; pseudobulbs 5.6–9.9×4.8–6.3 mm, Mexico. Cohniella biorbicularis is restrict-
subspherical to broadly ovoid, apically 1- ed to the Gulf Coast states, west and north
leaved, green, totally enclosed by 3 imbricate of the Tehuantepec Ithsmus, namely Tam-
sheaths, eventually deciduous; leaves terete, aulipas, San Luis Potosí, Querétaro, and
thickly fleshy-coriaceous 10–27 cm long, 2– Veracruz. It grows at elevations of 0–
5 mm thick, dark green; inflorescences 1400 m, usually in tropical dry forests or
solitary from the base of the pseudobulbs, in the ecotones between tropical dry forests
13.50–112.20 cm long, a 27–53 flowered and pine-oak forests. The plants always
raceme or panicle with 1–3 short branches, grow on trees.
the branches 3–6 flowered; peduncle and Diagnostic features.—Cohniella biorbicu-
rachis green-purple; peduncle more or less laris is easily recognized by its relatively
erect, 3–4 mm thick, terete, with 6–12 shortly large, suborbicular to very broadly obovate
bracted internodes, the basal-most longest, lateral labellum lobes. These are almost as
oblanceolate, acuminate, tubular; floral bracts wide as the midlobe in some specimens
3.31–5.10 mm long, narrowly elliptic, acu- (Nagel 6734, MEXU; Carnevali & Ramírez
minate; flowers with perianth segments 6372, CICY), but more commonly they are
widely opening and the petals and sepals somewhat narrower but always at least 75%
somewhat reflexed; ovary with pedicel 15.5– as wide as the width of the midlobe. The
22 mm long, of which ca. 3.71–7.29 mm plants look very similar to those of the
correspond to the ovary, this 0.60–1.33 mm western Cohniella brachyphylla, but can be
thick; sepals basally clawed, flat or somewhat distinctly larger.
reflexed; dorsal sepal 6.27–7.50 × 2.52– Variation range.—Plants from Veracruz
4.51 mm, obovate-lanceolate, apically obtuse have the broadest lateral lobes while those
and minutely apiculate, concave in the upper from Querétaro and San Luis Potosi have the
narrowest. Aside from this, Cohniella bio- spread lateral lobes than across the apical
rbicularis is a species with relatively little lobe, as opposed to the labellum in C.
variation in floral characters. cebolleta, which is wider across the spread
Taxonomic commentary.—As with most apical lobe of the labellum. The lateral lobes
taxa in the Cohniella cebolleta complex, C. of the labellum are obliquely oblong to
biorbicularis has been confused with that obliquely obovate and are always more or
species. However, the broad lateral lobes of less retrorse. The midlobe of the labellum is
the labellum easily distinguish this species proportionally smaller than in the species
from any other Cohniella. related to C. cebolleta. The disk of the
labellum is suborbicular to obliquely pentag-
Additional specimens examined. MEXICO. Querétaro: onal, proportionally large, and about as long
Mpio. Arroyo Seco, 6 km al N de Concá, 600 m, 18 Mar as the width of the midlobe. At each side of
1985, Fernández N. 2778 (US); río Santa María, 1.5 km al
NE del puente de Concá, 600 m, 6 Mar 1988, Herrera 61
the isthmus, anterior margins are provided
(XAL); Mpio. Jalpan, km 194 carretera hacia Landa de with one tooth on each one.
Matamoros, 1490 m, 16 Apr 1984, Aguirre 41–638 (AMO); Distribution and ecology.—Mexico, possi-
oriente de Tanchanaquito, 300 m, 8 Apr 1992, López 986 bly into El Salvador, Guatemala, Honduras,
(XAL); Mpio. Landa de Matamoros, S de El Capulín, and Nicaragua. In Mexico, it is widely
Puerto Blanco, camino al Río Moctezuma, 920 m, 13 Mar
1988, Herrera 103 (AMO). San Luis Potosí: Mpio. El distributed along the Pacific coast of the
Naranjo, 5 Apr 1960, Dressler 2606 (US). Tamaulipas: country and into the intermountain valleys.
Mpio. Antiguo Morelos, about 7 km N of Antiguo Morelos It has been collected as far north as ca.
(km 540), 28 Mar 1961, Dressler 2632 (US); Mpio. Álamos, in Sonora (ca. 27°N) and then
Aldama, pueblo El Plomo, Sierra de Tamaulipas, 550 m, southwards to Oaxaca, along the western
19 Jan 1991, Jiménez, O. Rocha & C. Rocha 1090 (AMO).
Veracruz: Mpio. Atoyac, 1.5 km al N de Potrero Nuevo- slopes of the Sierra Madre Occidental. On
Miraflores, 18º53′58″ N, 96º49′21″ W, 425 m, 7 Mar 1986, the eastern side of the Tehuantepec Isthmus,
Acevedo & Acosta 880 (XAL); Mpio. Jilotepec, comprada the species or a similar entity reappears in
en las calles de Xalapa, 26 Apr 2001, Carnevali & Ramírez Chiapas and then ranges southwards into at
6372 (CICY, 3 sheets); Mpio. Martínez de la Torre, 20º03′N,
97º03′W, 300 m, 30 Mar 1935, Nagel 4656 (US); Mpio. least Costa Rica at 0–800 (−1500) m; these
Naolinco, ca. San Antonio Paso del Toro, 19º35′ N, 96º49′ W, plants are somewhat different (see below).
600 m, 23 Mar 1975, Ortega 745 (XAL); Mpio. Actopan, Cohniella brachyphylla is usually found in
Trapiche 400 m, 15 Feb 1971, Ventura A. 3114 (US). several kinds of dry forests, even into thorn
scrub, and is the Cohniella from the driest
Cohniella brachyphylla (Lindl.) Cetzal & environments. It is almost always an epiphyte
Carnevali, comb. nov. Oncidium brachy- on thick branches, very rarely on rocks. It is
phyllum Lindl., Edwards’s Bot. Reg. 28: often found fully exposed to the sun.
sub t. 4. 1842. Type: Mexico. Without Variation range.—Cohniella brachyphylla
precise locality, K. T. Hartweg s.n. (holotype: is a common and widespread species, and it is
K-Lindl.). (Fig. 3A–D, H) consequently variable. The fact that the
species occurs as a conglomerate of popula-
Diagnostic features.—Cohniella brachy- tions isolated in intermountain valleys has
phylla is distinguished by its relatively short, apparently created barriers to gene flow,
thick leaves on top of relatively conspicuous favoring the establishment of local forms or
pseudobulbs. The leaves are typically rigidly races. Since these populations are also linked
erect and thickened on the lower half; they by neighboring, often intermediate popula-
are also almost always somewhat to strongly tions occurring along the coastal lowlands of
falcate. The inflorescences are either erect or tropical western Mexico, there has not been
horizontal, usually not as long as they are in extensive speciation associated with the pop-
other species of the C. cebolleta complex. ulations currently referred to this species.
The isthmus of the labellum is short and Exceptions are the populations from coastal
broad, wider than its length or about as long Jalisco here treated as C. pendula which are
as wide as opposed to the longer isthsmus discrete in the variation of several features
found in populations here referred to C. (see discussion under C. pendula).
cebolleta. Along with C. pendula and C. The great amount of variation found within
biorbicularis, the labellum is wider across the Cohniella brachyphylla occurs both between
FIG. 3. Flowers of Cohniella. A–D, H. Cohniella brachyphylla. A–D. Whole flower, front views. H. Column with
anther cap, front view. E–G. Cohniella cebolleta. E. Whole flower, front view. F. Labellum, front views G. Column
with anther cap, front view. I–M. Cohniella cepula. I. Column with anther cap, front view. J, L. Whole flower, front
views. K, M. Labellum, front views. (A, H from Carnevali & Gómez Juárez 6803, CICY; B from Carnevali
&Ramírez 6552, CICY; C from Carnevali 7297, CICY; D from Carnevali 7310, CICY; E–G from Carnevali 7222,
CICY; J–K from Paiva s.n (photo), CICY; I, L–M from Carnevali 6382, CICY.).
and within populations. The shape of the some in the state of Morelos), the central lobe
several sections of the labellum, as well as its of the labellum is shortly but distinctly apicu-
general outline are variable. The lateral lobes of late. The calli and disk are always somewhat
the labellum, which are always somewhat tinged in various hues of red, but the extent and
reflexed, range from obovate-suborbicular to patterns of these colorations are variable. The
subquadrate to obliquely elliptical. The mid- adaxial side of the labellum is almost always
lobe ranges from almost suborbicular to trans- provided with dull red-rose spots that are
versely subquadrate. In some populations (e.g., variable in size, distribution, and density.
Populations here referred to Cohniella Vetensk. Acad. Nya Handl. 21: 240.
brachyphylla from the southernmost extreme 1800. Stilifolium cebolleta (Jacq.) Königer
of the distribution from Chiapas southward & Pongratz, Arcula 7: 186, 187. 1997.
into Guatemala are atypical in having broadly Trichocentrum cebolleta (Jacq.) M. W.
obovate lateral lobes but otherwise conform Chase & N. H. Williams, Lindleyana 16:
to the characters of the species. On the 137. 2001. Type: Colombia. Cartagena
northwestern extreme of the range of [Carthagenae]. 1758, N. J. Jacquin s.n.
the species, the plants are smaller and the (holotype: not located and most likely lost;
inflorescences tend to be racemose while the lectotype: Select. Stirp. Amer. Hist., ed. 2,
flowers are somewhat similar to those seen in t. 217 [text on page 111], 1781, designated,
the Chiapas plants. albeit incorrectly, by Garay & Sweet,
1974: 205; amended lectotype: Select.
Selected specimens examined. MEXICO. Chihuahua: Stirp. Amer. Hist. [text on pages 230–
Mpio. Batopilas, arroyo Guimivo between Río Batopilas 231] t. 131, Fig. 2, 1763). (Fig. 3E–G)
and Guimivo, 762–915 m, 24 Mar 1979, Bye et al. 9235
(NY, MEXU). Durango: Glacala, 23 Feb 1899, Goldman
327 (US). Estado de México: Temascaltepec, Guayabal, 9 Epidendrum juncifolium L., Sp. Pl. (ed. 2)
Feb 1933, Hinton 3362 (AMES). Guerrero: Near San 1351. 1763. Cymbidium juncifolium (L.)
Nicolas del Oro, 1200 m, 13 Jan 1938, Mexia 9106 Willd., Sp. Pl. 4: 102. 1805. Oncidium
(AMES). Jalisco: Mpio. Autlán, Puerto los Mazos, a juncifolium (L.) Lindl., Coll. Bot. 27.
10 km antes de Autlán a 7 km antes de la desviación a
Guadalajara, 1000 m, 4 Oct 1987, R. Jiménez et. al 756 1821. Type: presumably collected in Haiti
(AMO). Michoacán: Coalcoman, 4 Apr 1939, Hinton [“habitat in America”], Plant. Amer. fasc. 8:
13647 (US, AMES, NY). Morelos: Tlaquiltenango 2 km t. 184, f. 2. 1759; epitype, here designated:
al N de la desviación a San José Pala, sobre el camino a Martinica, P. Dusse 2078 (NY).
Huatla, 7 Nov 1996, A. Espejo et al. (AMES). Oaxaca: Oncidium humboldtii Schltr., Repert. Spec.
Microonda, San Cristóbal, Carr. Oaxaca-Tehuantepec, km
144, desviación hacia la derecha aprox. 1 km, 1100 m, R. Nov. Regni Veg. 23: 65.1926. Oncidium
Jiménez et al. 3169 (AMO). Sinaloa: Mpio. de Culiacán, a ottonis Rchb.f. ex Kraenzl., Pflanzenr.
más o menos 46 km al N de Culiacán, camino de (Engler) IV, Fam. 50: 92, Fig. 10. 1922.
Badiraguato a la Pitayita, 25º02′00″ N, 107º23′00″ W, (non Schltr., 1914). Type: Venezuela.
100 m, Vega, Gutierrez G. & Hernández 8104 (MEXU).
Sonora: Álamo Gordo, Carnevali & Gómez-Juárez 6803
Without locality, 1840, K. F. E. Otto 997
(CICY). (syntype: B, destroyed; isosyntype: W-
GUATEMALA. Huehuetenango: Mpio. Nentón, km Reichenbach 27863); Guárico: Orituco,
10 a Gracias a Dios, 15º53′22.8′ N, 91º44′0.4″ W, K. F. E. Otto 541 (syntype: B, destroyed;
1099 m, 24 Mar 2007, Velásquez et al. 184 (BIGU); isosyntypes: W-Reichenbach 27869 &
km 11 a Gracias a Dios, 15º53′2.7″ N, 91º44′16.4″ W, 27872); Carabobo: Puerto Cabello, A. von
1074 m, 21 Sep 2006, Véliz et al. 17461 (BIGU); km 11
hacia Gracias a Dios, 15º53′0.63″ N, 91º44'21.0″ W, Humboldt & A. Bonpland s.n. (syntype:
1061 m, 21 Sep 2006, Véliz et al. 18454 (BIGU). Jalapa: B-W); lectotype, here designated: A. von
El Rancho, 28 Dec 1907, Kellerman 7002 (NY). Humboldt & A. Bonpland s.n. (B–W).
Progreso: San Agustín AC Transecto El Rancho Norte, Oncidium ultrajectinum Pulle, Rec. Trav. Bot.
14º53′28″ N, 90º01′17″ W, 200–300 m, 20 Sep 2003,
Cobar et al. (BIGU).
Neerl. 4: 121. 1907. Type: Surinam.
EL SALVADOR. Santa Ana: Mpio. Santa Ana, “Cultivated at the Utrech Botanical Garden
salida a Metapan, ca. 8 km al N del centro de la ciudad from a plant coming from Surinam as a gift
de Santa Ana, 10 Jan 2000, Linares s.n. (MEXU); Mpio. from v. Asch v. Wijck, the Governor of the
Metapán, ca. 6 km al NO de Metapán, por el camino al colony, to our University” (holotype: pre-
despoblado, 12 Jan 2000, Linares et al.4755 (MEXU). sumably at U, not seen).
COSTA RICA. San José: Monte Redondo, 1400 m,
1 May 1926, Alfaro 270 (AMES); Santa María de Dota,
5 Mar 1924, Alfaro 36617 (AMES); Turrúcares, 650 m, Diagnostic features.—Cohniella cebolleta
15 Jan 1925, Alfaro s.n. (AMES, US). Guanacaste: La is characterized by the small, relatively
Cruz, 15 Jan 1930, Jiménez 7909 (AMES). inconspicuous pseudobulbs that bear a rela-
tively narrow but fairly rigid leaf. The flowers
Cohniella cebolleta (Jacq.) Christenson, have a complicated callus with a relatively
Lindleyana 14: 177. 1999. Epidendrum long basal oblong plate and three apical teeth
cebolleta Jacq., Enum. Syst. Pl. 30. 1760. or keels, the central one being the largest.
Oncidium cebolleta (Jacq.) Sw., Kongl. There are one or two additional teeth on each
side of this basal plate. From its Mexican from central and coastal Venezuela as well as
relatives, C. cebolleta can be distinguished by those from eastern and coastal Colombia are
the general outline of the labellum as indi- typical of the species (type from ca. Cartagena,
cated in the discussion under C. brachy- Department of Bolívar) and feature relatively
phylla. The column of Cohniella cebolleta is large flowers with a long isthmus. These
narrower than in C. cepula and the column forms are well depicted in the iconography
wings are pure yellow (very rarely with a hint of this paper and by illustrations by Dunster-
of reddish on the margins) while in C. cepula ville (e.g., featured in Romero & Carnevali,
these are always maculated with large red 2000).
spots. In populations of the northern rim of the
Distribution and ecology.—As understood Guayana Highlands, the flowers are small
here, Cohniella cebolleta is a species wide- with central lobes of the labellum about
spread in the Caribbean, ranging from the 10 mm wide, while populations along the
Yucatan Peninsula in southeastern Mexico, Andean foothills in Colombia and Venezuela
throughout Central America and into northern often have large labella with midlobes 15–
Colombia and Venezuela. It also occurs in 20 mm wide. In eastern Venezuela, the
Jamaica and the Lesser Antilles. Cohniella populations feature the largest flowers of this
cebolleta grows in the driest environments complex (up to 35 mm diam.) and may
occupied by any Cohniella (except some represent a different taxon. In the Guianas,
populations of C. brachyphylla), typically the flowers are small (18–20 mm diameter)
tropical dry forests, or even coastal dunes or and have a distinctive callus. Should these
thorn forests. In Venezuela and Colombia it is Guianan populations need to be treated as a
not an uncommon sight on large branches of different species, the name Oncidium ultra-
old saman [Albizia saman (Jacq.) F. Muell.], jectinum is available. The Lesser Antilles
guatacaro [Bourreria cumanensis (Loefl.) O.E. populations of the complex (including the
Schulz] or caro or guanacaste [Enterolobium type of Cymbidium juncifolium) have small
cyclocarpum (Jacq.) Griseb.] trees. On a flowers (18–20 mm) with narrow lateral lobes
regional scale, C. cebolleta is often allopatric to the labellum associated with relatively
or parapatric with other species of the genus large petals and sepals heavily spotted.
that usually grow in more humid environments. Taxonomic commentary.—Christenson
In all of Central America, where C. ascendens (1999) cited the wrong basionym (“Dendrobium
and C. cebolleta are parapatric or sympatric, the cebolleta”, supposedly published in 1760,
second grows only on the driest vegetations although the genus Dendrobium was not pub-
types, whereas the latter occurs in more humid lished by O. Swartz until 1799). This wrong
ecosystems. Cohniella cebolleta reaches its citation is considered a “correctable error” since
southernmost distribution at the mid course of the author cited the right publication, page, and
the Orinoco River in Venezuela. Then, there is a year (K. Gandhi, pers. comm.). The combination
gap in the distribution of the genus Cohniella, Dendrobium cebolleta, as far as we have been
which reappears at the southern margin of the able to ascertain, has never been formally
Amazonas River, where it is represented by published. Nonetheless, it does appear in numer-
C. cepula. ous publications and at least one major database
Variation range.—As here circumscribed on the Internet (i.e., Kew’s World Checklist of
and even after excluding several taxa here Monocotyledons at http://apps.kew.org).
treated as distinct species, Cohniella cebolleta Garay and Sweet (1974: 205), proposed
remains a very variable entity, mainly in plate 217 of the second edition of Jacquin’s
flower size and labellum shape. The labellum Selectarum Stirpium Americanarum Historia...
is usually widest across the apical (central) (1781) as the lectotype of Epidendrum cebol-
lobe but in some populations it can be as leta (de facto, originally as “holotype”). This
broad or even slightly broader across the plate shows a plant with a paniculate inflor-
spread lateral lobes. The labellum isthmus is escence, which is much longer than the leaves,
almost always conspicuous (about as long as bearing multiple fruits. However, plate 131,
the disk), but in some individuals it is Fig. 1 of the first edition of this work,
shorter. Flower size is also variable. Plants published in 1763, unambiguously shows the
vegetative parts and a separate, cut-out inflor- is a typical Cohniella cebolleta, representa-
escence (clearly showing a peduncle longer tive of the forms found in coastal Venezuela.
than the leaves) of Epidendrum cebolleta, and Oncidium ultrajectinum represents a small-
it clearly agrees with the 1760 protologue and flowered population growing along the north-
has priority over the plate in the second ern rim of the Guianas.
edition. Stafleu and Cowan (1979: 408) stated
that “The 1763 publication (i.e., Selectarum Additional specimens examined. MEXICO. Cam-
peche: Municipio Calkiní, El Remate, unos 8.5 km al
Stirpium Americanarum Historia) is an impor- oeste de Tankuché, 20º32′30″ N, 90º19′20″ W, 31 Jan
tant complement to the 1760 Enumeratio (i.e., 2000, Carnevali et al. 6016 (CICY). Quintana Roo:
the source of the protologue) and should Mpio. Benito Juárez, 3 km al sur del aeropuerto de
always be consulted with it”. Cancún, 16 Feb 1999, Carnevali, Gómez & Piven 5333
Furthermore, both the 1763 and 1781 (CICY). Yucatán: Mpio Mérida, Dzitya, 13 Mar 2009,
Carnevali 7424 (CICY).
editions of this work clearly state that the GUATEMALA. Jalapa: El Rancho, 28 Dec 1907,
type was collected in Cartagena, Colombia Kellerman 7002, (NY).
(Carthagenae in sylvis praesertim maritimis), NICARAGUA. Acoyapa: 2 km south of Acoyapa, 4
and not in Martinique, as stated in Garay and Jan 1969, Atwood 1656 (NY).
Sweet (1974: 205) and later in Nir (2000: COSTA RICA. San José: Santa María Dota, 5 Mar
263), nor in Brazil, as stated in Foldats (1970: 1924, Alfaro 36617 (AMES).
298). We have exhaustively searched for the DOMINIQUE. Gran Savannah, Lloyd 826 (NY).
type material but have not been able to locate it. SANTA LUCIA. Middle W slope of Gros Piton,
It could be argued that Jacquin might have Proctor 18062 (AMES).
collected either Cohniella cebolleta or C. COLOMBIA. Magdalena: Santa Marta, Near Borda,
12 Jan 1898, Smith 2353 (NY).
nuda (Bateman ex Lindl.) Christenson in
VENEZUELA. Anzoátegui: Woods along Río León
Cartagena, since Jacquin’s original collection by Quebrada Danta, tributary to Río Neverí, northeast of
and description did not include floral charac- Bergantin, 500 m, 20 Feb 1945, Steyermark 61022
ters that distinguish them, and these two (AMES). Bolivar: Ciudad Bolivar and vicinity, on the
species are vegetatively undistinguishable Orinoco, 8º10′ N, 63°31′17″W, 200 m, 27 Feb 1921, L. H.
and possibly could occur in habitats where Bailey & E. Z. Bailey 1350 (NY). Delta Amacuro: Dpto.
Antonio Díaz, Isla Curiapo, entre caños Obaruvaca y
Jacquin collected. Cohniella cebolleta, how- Naguabanoco, 8º31′ N, 61º04′ W, 0 m, Fernández 3820
ever, always bears inflorescences much lon- (NY). Río Maniamo, Vuelta Triste, 20 Feb 1911, Bond,
ger than the leaves, versus inflorescences Gillin & Brown 164 (AMES). Zulia: Kunana, 30 Feb
about as long or, more commonly, shorter 2008, 10°03′3″N, 72°41′12″W, 300–400 m, Carnevali
7222 (AMES, CICY, VEN).
than the leaves in C. nuda.
GUYANA. Rupununi: Ca. 5 miles from Karasabi (4–
With the exception of the large-flowered 5 hours walk) along Yurora river; 4º00′ N, 59º21′ W,
species of South America (Cohniella jonesi- 300 m, 3 Jan 1982, Knapp & Mallet 2811 (NY).
ana and C. stacyi) and the species related to
C. ascendens, almost all other taxa of the Cohniella cepula (Hoffmanns.) Carnevali &
genus have at one time or the other been G. Romero, comb. nov. Oncidium cepula
considered synonyms of Cohniella cebolleta. Hoffmanns., Verz. Orchid. Ed. 2, 56. 1843.
As a result, published distributions for Coh- Type: Brazil. Río de Janeiro: ex icon. [“N.
niella cebolleta report, albeit incorrectly, a Icon. ined”] (holotype: B, destroyed; lec-
species ranging from northern Mexico into totype, here designated, tracing in Herba-
SE Brazil and Argentina. Oncidium juncifo- rium Reichenbach Nr. 15230, upper left
lium was based on a plant without a precise corner, W). (Fig. 3I–M)
locality in America, but being a species
named by Jacquin is most likely of a West Oncidium sprucei Lindl., Fol. Orchid. 56.
Indian origin. The holotype is a line drawing 1855. Cohniella sprucei (Lindl.) Königer
by Plumier, published simultaneously with & Pongratz, Arcula 10: 280. 2000. Tricho-
the original description, featuring an idealised centrum sprucei (Lindl.) M.W. Chase & N.
Cohniella with non-resupinate flowers. This H. Williams, Lindleyana 16: 218. 2001.
illustration may represent almost any member Type: Brazil. R[io] Negro and Solimões:
of the genus, but the West Indian origin R. Spruce 1526 (holotype: K-Lindl., iso-
indicates C. cebolleta. Oncidium humboldtii types: AMES, NY, photograph).
Oncidium glaziovii Cogn., Fl. Bras. 3, 6: 440. portion of these lobes is appressed to the basal
1906. Type. Brazil. Goias: A. F. M. Glaziou plate of the callus. Since the column of this
22179 (holotype: BR, isotype: G; photo: species is short and bent backwards (and thus
AMES, NY). appears stalkless), the attachment point of the
Oncidium ostenianum Schltr., Repert. Spec. petals spreads all across the column height,
Nov. Regni Veg. 21: 341. 1925. Cohniella while in C. cebolleta and relatives the stalk of
osteniana (Schltr.) Christenson, Lindleyana the column is below the base of the attachment
14: 177. 1999. Stilifolium ostenianum point of the petals. The callus consists of a
(Schltr.) Königer & Pongratz, Arcula 7: horizontal platform ending in three teeth, of
189. 1997. Trichocentrum ostenianum which the central is much larger. Another
(Schltr.) M. W. Chase & N. H. Williams, distinctive feature of this species is the column
Lindleyana 16: 138. 2001. Type: Paraguay. wings that always have red or red-purple spots
Epiphyt auf Bäumen am Río Salado, bei (except in rare albino forms) as opposed to the
San Bernardino, Rojas, Marz 1916 (holo- entirely yellow column wings of the members
type: Osten 8557, herbarium Ostenianum; of the C. cebolleta complex north of the
isotype: AMES). Amazon Basin. In C. cepula the column wings
Oncidium cebolleta var. purum L. C. are also almost perpendicular to or at least at a
Menezes, Schlechteriana 2: 132. 1991. very broad angle to the main axis of the
Type: Brazil. Minas Gerais: Arinos, column, while in C. cebolleta and relatives,
700 m, March 1991, C. García s.n. the wings are essentially parallel to the main
(holotype: UB-14). axis of the column.
Oncidium wittii Oppenheim, Orchis 10: 93. Distribution.—Argentina, Bolivia, Brazil,
1916. Lophiaris wittii (Oppenheim) Braem, Paraguay, and Peru. This species ranges
Schlechteriana 4: 21. 1993. Stilifolium wittii widely south of the Amazon basin.
(Oppenheim)Königer & Pongratz, Arcula 7: Variation range.—As with many other
190. 1997. Cohniella wittii (Oppenheim) Cohniella species, C. cepula is extremely
Senghas, Orchideen (Schlechter) 173–181. variable and at this time how this variation
2001. Trichocentrum wittii (Oppenheim) M. correlates with the geographical distribution
W. Chase & N. H. Williams, Lindleyana of the species is not well understood. Thus,
16: 138. 2001. Type: Bolivia. Río Iténez, we refer all the specimens from of the Amazon
Ost s.n. holotype B, destroyed; lectotype, Basin and southwards to a broadly circum-
here designated, plate 4, Orchis 10, No. 5, scribed and variable C. cepula. The variation
Tafel IV, Fig. 1. 1916; epitype, here of this species involves basically the shape and
designated, Bolivia. Santa Cruz: Provincia width of the apical lobe of the labellum. This
Andrés Ibañez, 12 km de Santa Cruz, 11 variation follows the pattern found in most of
Aug 1987, M. Nee 35623 (NY). the species of the genus, where the narrowest
Stilifolium pongratzianum Königer, Arcula 9: midlobes correlate with the proportionally
261. 1999. Cohniella pongratziana longest isthmuses and narrower lateral lobes
(Königer) Königer, Arcula 10: 280. 2000. (morph A, Fig. 3 J) while the broadest label-
Trichocentrum pongratzianum (Königer) M. lum midlobes correlate with a proportionally
W. Chase & N. H. Williams, Lindleyana 16: shorter isthmus and broader lateral lobes
218. 2001. Type: Peru. Departamento San (morph B, Fig. 3 L). Both Oncidium cepula
Martin: Juanjui, ca. 300 m, from M. Arias, and O. ostenianum are referable to morph A,
Lima, cultived at Rosenheim, W. Königer while Oncidium wittii and Stilifolium pongrat-
WK-99 (holotype: M; isotypes: USM, zianum to morph B. The latter two species
UNALM, Herb. Königer [not seen]). might eventually deserve recognition at the
species level, in which case the name Cohniella
Diagnostic features.—Cohniella cepula is wittii would be available. Plants from the
extremely variable but can always be recog- northern Amazon Basin (the type of Oncidium
nized by its relatively short and broad column sprucei among them) are also referable to
and simple, three-toothed callus. The column morph B and have the largest flowers of the
features a massive tabula infrastigmatica that complex. This complex of forms obviously
is deeply bilobed and cleft and the proximal needs a more detailed examination.
Taxonomic commentary.—All citations and of the genus by the combination of a large

records of Cohniella (Oncidium) cebolleta flower (petals to 2 cm long, labellum to 2.2 cm
from south of the Amazon River belong to wide) and a white central labellum lobe with
this species. yellow, finely fimbriate, reduced lateral lobes.
Distribution.—Argentina, Bolivia, Brazil,
Additional specimens examined. BRAZIL. Amazonas: and Paraguay.
Río Solimões, near 20 km above mouth of Río Negro, Paraná Variation range.—Cohniella jonesiana
do Xiboreninha, 15 Jun 1992, Mori & Gracie 22412 (NY). varies in the color of the sepals and petals
Rondônia: Mpio. de Cacoal, BR 364, Rodovia Cuiabá-Porto
Velho, Km 234, ao Norte de cidade, morro da Torre da
that range from very pale green to cream
Embratel, 11º12′ S, 61º 62′ W, Cid, Lima, Guedes & Coelho colored. They are always spotted with rela-
4749 (NY). Ceará: Near Umari, 15 m, 7 Jul 1945, Cutler tively conspicuous brown-red spots or
8376 (AMES). Maranhão: between Viana & Banderante, blotches that are, however, never confluent.
back road from Viana to Pínheiro, 3º0′ S, 45º10′ W, 17 Oct
1980, Daly, Campbell, Silva, Bahia, & dos Santos
As in other variable species, pale or albino as
D653 (NY). Mato Grosso: 12º49′ S, 51º46′ W, 3 Aug well as dark-colored forms are known (var.
1968, Richards 6586 (AMES, NY). Minas Gerais: Mpio. flavens and var. phaeanthum, respectively).
Porteirinha, near city of Porteirinha, 25 May 1945, Williams Taxonomic commentary.—This species is
& Assis 7052 (AMES). Pará: Tucuruí, Breu Branco, igapó às so distinct that it has never been mistaken
margens do rio Tocantins. 14 Aug 1983, Revilla, Miranda,
Lima & Silva 8691 (NY). with any other member of the genus.
PERU. Lima: Surquillo, unknown procedence, 22 Jul
1997, Fernández s.n. (NY). San Martin: near Moyobambo, Additional specimens examined. BOLIVIA. Santa
approx. 6°02′ 60″S, 76°58′00″W, 700–800 m, 25 Feb 2005, Cruz: Andrés Ibáñez, between Pedro Lorenzo and Peji,
Carnevali & Ramírez 7367 (CICY); Juanjui, cultived at Feb 1982, Vásquez 659 (Herbarium Vazquezianun).
Rosenheim, 300 m, Feb 2000, Königer WK99 (M). PARAGUAY. Paraguarí: Cerro de Acahay, Mar
BOLIVIA. Santa Cruz: Ichilo, near 30 km SE of 1919, Rojas 3503 (AMES). Asunción: Cordillera de
Buena Vista along Río Surutú, 17º36′ S, 63º36′ W, 400 m, Altos, Apr 1940, Rojas 8822 (AMES).
1 Sep 1985, Solomon 14194 (NY). ARGENTINA. Corrientes: Ituzaingo, Estancia Santa
ARGENTINA. Tucumán: Formosa, Jan 1918, Löse- Rita, 27º3′ S, 56º4′ W, 16 Feb 1991, Tressens, Ferrucci &
man 2072 (AMES). Radovancich 3957 (AMES).
Cohniella jonesiana (Rchb. f.) Christenson, Cohniella longifolia (Lindley) Cetzal &
Lindleyana 14: 177. 1999. Oncidium jone- Carnevali, comb. nov. Oncidium longifo-
sianum Rchb.f., Gard. Chron. N.S. 20: 781. lium Lindl., Edwards’s Bot. Reg. 27: 22.
1883. Stilifolium jonesianum (Rchb. f.) 1841. Type: Mexico. Without precise
Königer & Pongratz, Arcula 7: 189. 1997. locality, collected by K. T. Hartweg, ex
Trichocentrum jonesianum (Rchb.f.) M. W. Hort. Royal Horticultural Society and
Chase & N. H. Williams, Lindleyana 16: Loddiges (holotype: K-Lindl.).
137. 2001. Type: Paraguay. Without any
other locality or collector, ex Hort., Fred Diagnostic features.—According to the
Horsman & Company s.n. (holotype: W- protologue, Cohniella longifolia is easily
Reichenbach 27551). (Fig. 4A–E) distinguished from other Mexican Cohniellas
Oncidium jonesianum var. flavens Reichb. f., by its long (“ ... often three feet long ...”),
Gard. Chron. S. 3., 4: 234, V. 237, 1888. pendent leaves of apparently soft texture (“ ...
Type: Without locality, but presumably from long, whiplike ...”) and by its elongate
Paraguay, ex Hort. B. S. Williams (holotype: isthmus subtending a relatively small central
W-Reichenbach 27550 [upper left figure]). lobe, this feature easily observable in the type
Oncidium jonesianum var. phaeanthum Sander, material. The inflorescences were described
Reichenbachia S. I.,1: 47, t. 21, Fig. 2, as “ ... forming dense panicles three feet long
1886–1891. Type: Paraguay. Without pre- of very large and showy yellow and brown
cise locality, L. Saint-Leger ex Hort. Sir flowers.” Later, Lindley (1842), in the text in
Trevor Lawrence (holotype: presumably at Latin that accompanied a color plate of this
W-Reichenbach, not seen). species, stated that the leaves could be 3–4
feet long.
Diagnostic features.—Cohniella jonesiana Distribution.—Known only from the type
is easily distinguished from all other members collection.
FIG. 4. Flowers of Cohniella. A–E. Cohniella jonesiana. A. Labellum, front view.. B. lateral view of the callus
and the column. C. Column with anther cap, front view. D. Anther cap. E. Sepals and petals, front view. F–H.
Cohniella stacyi. F. Whole flower, front view. G. Labellum, front views H. Column with anther cap, front view. (A–E
from Paiva s.n. (photo) CICY; F–H from Kennedy s.n. (photo) AMES, CICY).
Variation range.—Known only from the type. like ...” lateral lobes of the labellum).
Taxonomic commentary.—According to the Although it is not impossible that Mexican
protologue, Lindley was aware of living plants resembling the type of Oncidium
plants of C. longifolia cultivated both by the longifolium may be found in the future, we
Royal Horticultural Society and the Loddiges suspect this may be yet another orchid species
establishment, but a watercolor of a single reported from the wrong locality.
flower and his type at Kew (a panicle with
five branches) do not indicate provenance. No Cohniella×marvraganii (Lückel) Christenson,
Mexican Cohniella resembling the type of Lindleyana 14: 177. 1999. Stilifolium marv-
Oncidium longifolium has ever been col- raganii Lückel, Orchidee (Hamburg) 49: 90.
lected. The plants and flowers of the type 1998. (as “marvreganii”)—Trichocentrum
material are suspiciously similar to those of marvraganii (Lückel) M. W. Chase &
several populations of Cohniella cebolleta N. H. Williams, Lindleyana 16: 137. 2001.
from Perijá, Venezuela (e.g., Carnevali 7222, Type: Bolivia. Santa Cruz: M. E. Ragan s.n.
AMES, CICY, VEN) with which it shares the (holotype: USF).
long, often pendent leaves, the large panicu-
late inflorescences and the diagnostic features Diagnostic features.—Cohniella×marvra-
of the flowers (the long isthmus and “ ... bird ganii is intermediate between the putative
parents (C. jonesiana and C. stacyi) in several retrorse upon flattening. The leaves of this
respects. The general outline of the labellum species are usually rigidly pendent. The
is similar to that of C. stacyi but it is colored inflorescences are always shorter than the
white or very pale yellow as in C. jonesiana; leaves and very dense with flowers crowded
the basal lobes of the labellum are most as compared to other Cohniella species to the
similar to those of C. jonesiana, the calli are extent that the flowers partially overlap.
similar to those of C. stacyi. Column wings Distribution.—Panama, Colombia, and
are similar to those of C. stacyi as well as the Venezuela.
shape and disposition of the petals. Variation range.—Despite its relatively
Distribution.—This nothospecies is known restricted distribution, Cohniella nuda is
from NW of Santa Cruz, Bolivia. extremely variable in the size and morphol-
Variation range.—Only known from the ogy of the flowers. Flowers range from small
type material. with the labellum ca. 7 mm long to larger
Taxonomic commentary.—This nothospe- with the labellum ca. 13 cm long. The apical
cies was originally described as Oncidium lobe of the labellum varies from transversely
Jason Fuchs by Ragan & Sauleda (1982). It elliptic to transversely oblong to rhomboid; its
was formally proposed as a nothospecies by apical emargination ranges from inconspicuous
Lückel (as Stilifolium “marvreganii”). Ragan through deep and with overlapping lobes. The
& Sauleda (1982) commented “The ranges of plants start flowering at a very early age; some
Oncidium stacyi and O. jonesianum overlap specimens flower with leaves less than 8 cm
just north of Santa Cruz, precisely in the area long while fully mature plants feature thick,
where this natural hybrid was discovered. usually pendent leaves of up to 50 cm long.
Vegetatively the hybrid is intermediate Taxonomic commentary.—Despite its vari-
between the parents. The flowers are slightly ability, Cohniella nuda is relatively easy to
smaller than Oncidium stacyi with a lighter distinguish from related species and its
yellow labellum”. synonymy is limited. However, it seems to
grade into the Panamanian taxon Cohniella
Cohniella nuda (Bateman ex Lindl.) Chris- stipitata near Panama City and at the Pearl
tenson, Lindleyana 14: 177. 1999. Oncidium Archipielago (Dressler & Williams, 2003).
nudum Bateman ex Lindl., Edwards’s Bot. The type of Oncidium ebrachiatum is identi-
Reg. 23: t. 1994. 1837. Stilifolium nudum cal to populations of C. nuda from western
(Bateman ex Lindl.) Königer & Pongratz, Venezuela and is here treated as a synonym of
Arcula 7: 189. 1997. Trichocentrum nudum that species.
(Bateman ex Lindl.) M. W. Chase & N. H.
Williams, Lindleyana 16: 138. 2001. Type: Additional specimens examined. COLOMBIA.
Venezuela. Distrito Capital: Caracas, 1837, Atlántico: Puerto Colombia, 10–60 m, 10 Apr 1906,
ex Hort. Bateman (holotype: K-Lindl.). Maxon 3843 (NY). Córdoba: Morrocoquiel, on Rio
(Fig. 1E–H) Sinu, 11 Mar 1918, Pennell 4693 (NY).
Oncidium ebrachiatum Ames & C. Schweinf. PANAMA. Chiriqui: halfway between Progreso and
Puerto Armuelles, 16 Feb 1973, Croat 21878 (MEXU).
Sched. Orch. 2: 75. 1923. Type: Panama. Panamá: Torti, área cercana al Darien, 50 m, Abr 2002,
Cana and vicinity, 4 Apr 1908, R. S. Carnevali 7283 (CICY).
Williams 975 (holotype: AMES). VENEZUELA. Miranda: Guarenas, 1846, Funck &.
Schlim 481 (K-Lindl.); along Quebrada Chaguarama, 6 km
Diagnostic features.—Cohniella nuda is SE of Cúpira, 10º8′ N, 65º41′ W, 50–150 m, 5 Mar 1980,
characterized by its minute column wings, Liesner & González 9217 (NY). Guinand Estate
(Cárdenas), Siquire Valley, 500 m, 19–24 Mar 1913, Pittier
often almost absent. Furthermore, the mor- 5980 (NY). ZULIA: orillas del río Kunana, unos 4.5 Km. al
phology of the small callus is unmistakable. It W de La Sierra y unos 15 Km. al W de Machiques, 300–
consists of three low ridges flanking two 400 m., 5 Feb 2008, Carnevali 7283 (CICY).
depressions, which look almost wet. The two
external ridges are close to, and parallel to the Cohniella pendula Carnevali & Cetzal, sp.
margins of the labellar disk. The labellum is nov. Type: Mexico. Jalisco: Municipio La
characterized by its long, narrow isthmus and Huerta, Loma Alta, 40 km. de La Huerta
the relatively small lateral lobes, which are hacia Barra de Navidad, aprox. 19°22′0″
N, 104°41′59″ W , aprox. 350–450 m, subtruncate, somewhat reflexed in natural

collected by G. Carnevali and G. Salazar, 3 position; labellum deeply 3-lobed, 8–12 mm
Nov 1997, flowered in cultivation 10 Mar long from the base to the apex of the central
2004, G. Carnevali & I. Ramírez 6897 lobe, 11–16 mm wide across the apices of the
(holotype: CICY; isotypes: AMES, AMO, lateral lobes, the lateral lobes in the same plane
MO, NY). (Fig. 2H–O) as the central lobe and±perpendicular to it;
central lobe 8–11 mmx 4–7 mm, spathulate-
Species haec Cohniellae brachyphyllae (Lindley) oblongoid to transversely elliptic or subquadrate
Cetzal & Carnevali similis sed foliis pendentibus, lobo in outline, apically rounded to subquadrate,
centrali minore, quam lobulis lateralibus subaequalis alis basally produced into a short isthmus, 2 ×
columnae conspicue minoribus recedit. 1 mm; lateral lobes 4.2–7×4–5 mm, oblong to
broadly obovate, apically truncate-rounded to
Epiphytic pendent herbs, sun-loving to semi- sharply obliquely truncate, the upper margin of
umbrophyllous, shortly creeping to caespitose; the lateral lobes flat to rounded, the lower
rhizome short, thin, brittle; roots 1–2 mm thick, margin straight; disc relatively large, ca. 4×
white; pseudobulbs 12–14 mm long, 6–8 mm 5 mm, bearing a well-developed callus, ca. 2 ×
thick, subspherical to broadly ovoid, apically 1- 1 mm, consisting of a large, elevated, ± flat,
leaved, red-purple tinged, totally enclosed by 3 oblongoid platform, apically with three teeth,
imbricate sheaths, 2.5–6.4 × 12–20 mm upon the central one laterally compressed, the laterals
spreading, eventually deciduous; leaves terete, smaller, divergent, somewhat pointing upward,
thickly fleshy-coriaceous, 15–36 cm long, 6– conical, the basal portion of the callus with
13 mm thick, dark green, usually purple sharp upper margins; column 4 × 2 mm, the
spotted, when fresh abruptly constricted prox- ventral face in the same plane as the labellum
imally, broadest at its lower 1/5, gradually lobes, oblongoid, tabula infrastigmatica subqua-
attenuated distally into a sharp apex, often drate, stigmatic surface, sub-rounded, ca. 1.7 ×
somewhat falciform; inflorescences solitary 1.7 mm; column wings relatively small, ca.
from the base of the pseudobulbs, 35–45 cm 0.5 × 1 mm, reniform; anther 1.3 × 1 mm,
long, a (8–)15–30-flowered raceme or panicle apical, operculate, ellipsoid; pollinarium typical
with 1 or 2 short branches 4.5–5 cm long, the for the genus, ca. 1.5 mm long, tegula spathu-
branches 5–9-flowered; peduncle and rachis late, 0.8×ca. 0.4 mm at the subtruncate apex;
dark green, variably purple tinged to totally viscidium disc-like, small, pollinia 0.7–1 mm
purple; peduncle first pendent, then arching to long, yellow.
horizontal, 3–4 mm thick, terete, with 7–10
internodes, peduncle bracts 14–21 × 3–4 mm, Additional specimens examined. MEXICO.
the basal most longest, oblanceolate, acumi- Jalisco: Mpio. Cabo Corrientes, Las Juntas de Tuito,
aprox. 450 m, 16 Mar 1984, Salazar & Soto 580 (AMO);
nate, tubular; bracts subtending the lateral Mpio. Chiquilistan, 1300 m, Feb 1973, Rosillo de
branches 5–7 × 2.5 mm, elliptic, acuminate; Velasco 131 (AMO); Municipio Cocula, 1700 m, Mar
floral bracts 2–4 mm long, narrowly elliptic, 1973, Rosillo de Velasco 140 (AMO), 141 (AMO);
acuminate; flowers resupinate, small or medium Mpio. Tecalitlán, 1500 m, Mar 1981, Rosillo de Velasco
s.n. (AMO). Nayarit: Mpio. de Ruíz, km 56.3 del
sized for the genus, with perianth segments
camino de la carretera México 15 (Tepic-Mazatán) a
widely opening and the petals and sepals Jesús María, 6.3 Km adelante del poblado de El Naranjo,
somewhat reflexed; ovary with pedicel 12– 22°01′ N, 104°50′ W, 320 m, 25 Jul 1998, Soto 86888
14 mm long, of which ca. 5 mm correspond to (AMO); Mpio. de San Blas, Barranca N.W. east of Tepic-
the ovary, this 2 mm thick; sepals basally Navarrete, 21°14′ N, 104°32′ W, 1350 m, 28 Aug 1948,
Dressler 350 (US); Mpio. Santa María del Oro, Volcano
clawed for about 1/3 of total length of the sepal, Ceboruco near Tequepexpan, 4 May 1936, Nagel & Juan
flat or somewhat reflexed, dorsal sepal 5.5– S. 5115 (US).
6.5 × 1.8–2.2 mm, in general outline oblanceo-
late, apically obtuse and minutely apiculate, Cohniella pendula is closely related to C.
concave in the upper half, the claw 8–11 mm brachyphylla with which it shares the rela-
wide; lateral sepals partially fused at the very tively conspicuous pseudobulbs. It also fea-
base, then free, similar the dorsal, 7.5–8.5x 2.8– tures a rigid, recurved leaf that is distinctly
3 mm; petals 9.5–10.5 × 2.3–2.6 mm, oblan- swollen in the lowermost ¼ of its length and
ceolate, somewhat oblique, the apex rounded to constricted just above the junction with the
pseudobulb, and then gradually attenuated lateral lobes. The base of the column is dull
toward the apex. The apex of the leaf is orange-red, the tabula infrastigmatica is
rigidly pungent. However, the most striking bright yellow, the stigmatic surface is pale
vegetative difference is the pendent habit. yellow, and the anther cap is dark red-purple.
The leaves are rigid and, even when the plant
is cultivated upright, new leaves will grow Cohniella stacyi (Garay) Christenson,
pendent. The inflorescences point down at Lindleyana 14: 177. 1999. Oncidium stacyi
first, then become horizontal or, more rarely, Garay, Bot. Mus. Leafl. 23: 301. 1973.
pendent. The inflorescence itself is relatively Trichocentrum stacyi (Garay) M. W. Chase
few-flowered and lax when compared to & N. H. Williams, Lindleyana 16: 138. 2001.
plants of C. brachyphylla of similar size and Type: Bolivia. Naranjillos, road to Cocha-
vigour. The flowers are also distinctive due to bamba, 11 km south west from Santa Cruz, J.
their small size. Furthermore, the central lobe Stacy s.n. (holotype: AMES). (Fig. 4F–H)
of the labellum is subquadrate to transversely
elliptic, about as long as wide or not more Diagnostic features.—Cohniella stacyi is
than 1.5 times wider that long, as opposed to easy to distinguish among other Cohniella
distinctly transverse (at least twice as wide as species due to its large flowers (> 50 mm
long) as in C. brachyphylla and other diam.). The petals and sepals are dark yellow-
Cohniella taxa. These proportions make the greenish heavily covered with irregular dark
central lobe of the labellum in C. pendula as red-brown blotches. The labellum is bright
wide as the labellum disc, while the apical yellow with scattered orange-brown blotches,
lobes are as broad as the expanded apices of denser toward the margins; there is always a
the lateral lobes in C. brachyphylla and most broad orange-brown band across the isthmus.
other Cohniella species. Another distinctive The callus is white, heavily orange-brown
feature of Cohniella pendula is the lower spotted. The morphology of the labellum of this
margin of the disc that is cartilaginose-callose species is most distinctive due to its reduced
due to lateral extensions of the callus. lateral lobes which are laciniated at their distal
Although this same condition is also present margin; the central lobe has fimbriate or coarsely
in C. brachyphylla, it is much more pro- erose-dentate margins. Its closest relative seems
nounced in this new entity. Furthermore, the to be C. jonesiana (a relationship strongly
column wings are conspicuously smaller in supported by the phylogenetic analyses of Sosa
this new species, being 2–3 times smaller et al., 2001; Williams et al. 2001b, Cetzal,
than they are in C. brachyphylla. 2007), also from southern South America, with
Cohniella pendula seems to be restricted to which it shares the large flowers with small
sea-facing slopes along the Pacific coast on lateral lobes. Cohniella stacyi has larger flowers
the Sierra Madre Occidental where it grows of different colors, and a much longer (about
in “selva baja caducifolia” or “selva mediana twice as long) isthmus to the central lobe.
subcaducifolia”. It occurs with epiphytes such Distribution.—Cohniella stacyi is only
as Myrmecophila galeottiana (Reichb. f.) known from northern Bolivia.
Rolfe, Encyclia adenocarpon (La Llave & Variation range.—Cohniella stacyi is
Lex.) Schltr., and Tillandsia caput-medusae known from only a few wild collections.
E. Morren. At the type locality, it grew in a However, the species is currently common in
large colony in the lower branches of a tree, the orchid trade since it has been reproduced
in an exposed position. The plants were massively from seed. The progeny of this
heavily tinged with purple. propagation program are morphologically
The flowers of Cohniella pendula are uniform but variable in color, mainly in the
amongst the smallest in the genus and are depth and density of the red-brown blotching.
certainly the smallest in the C. cebolleta Taxonomic commentary.—Due to its rela-
complex. The sepals and petals are of a pale, tively recent description that was widely
dull creamish green with dark red-purple publicized (e.g., Fuchs, 1975; Garay, 1976)
spots, while the labellum is yellow with some and its horticultural appeal, Cohniella stacyi
pale, dull orange-red on the callus and along is well known and has not been subject to
the front margin of the disk and the bases of nomenclatural wrangling.
Cohniella stipitata (Lindl.) Christenson, Costa Rica (USJ) was imported from Panama,
Lindleyana 14: 177. 1999. Oncidium stip- according to the grower who cultivated this
itatum Lindl., Bot. Voy. Sulphur 172. specimen (F. Pupulin, pers. comm.)
1843. Stilifolium stipitatum (Lindl.) Variation range.—Cohniella stipitata is a
Königer & Pongratz, Arcula 7: 189. fairly homogeneus taxon; most variation
1997. Trichocentrum stipitatum (Bateman seem to be restricted to the shape of the
ex Lindl.) M. W. Chase & N. H. Williams, central lobe of the labellum, whose base
Lindleyana 16: 138. 2001. Trichocentrum varies from truncate to subcordate. Also, the
nudum (Bateman ex Lindl.) M. W.Chase & margins of the labellum are somewhat varia-
N. H.Williams subsp. stipitatum (Lindl.) ble as to their degree of fimbriation.
Dressler & N. H.Williams, Selbyana 24: Taxonomic commentary.—Oncidium stipi-
45. 2003 [21 Oct 2003]. Type: Panama. tatum var. platyonix was based on a specimen
Without any other locality, G. W. Barclay with somewhat dentate margins of the isth-
958 (holotype: BM). (Fig. 1I–L) mus, at each side of the callus. This feature is
Oncidium lacerum Lindl., Bot. Reg. 30. Misc. somewhat variable but is is present in all
38. 1844. Type: Panama. Without precise specimens that we examined. Oncidium lac-
locality, ex Hort. Loddigges (holotype: K- erum was based upon a specimen with a
Lindl.). deeply lacerated, crisp central lobe of the
Oncidium stipitatum Lindl. var. platyonyx labellum but it is otherwise identical to
Rchb. f., Gard. Chron. N.S. 9: 788. 1878. typical forms of the species.
Type: Panama. Without precise locality, ex
Hort. W. Bull (holotype: W-Reichenbach Additional specimens examined. PANAMA. With-
27554, flowers in envelope in the upper left). out any other locality, cultivated in the New York
Botanical Garden Conservatory, 2 Feb 1917, Nash
Diagnostic features.—Related to Cohniella 44955 (NY). Colón: Panama, Chilibre, 35 m , 15 Feb
2005, Carnevali 7311 (CICY). Canal Area: Chagres
nuda but it is easily distinguished by the River, about 3 miles above Gamboa Bridge along river
proportionally longer and narrower isthmus bank, 50–100 m, 09º08′32″ N, 79º40′49″ W, 07 Feb
of the labellum and the truncate, rounded to 1973, Kennedy, von Chong & Steiner 2299 (MO).
subcordate base of the central lobe of the
labellum. The column wings in Cohniella Cohniella teres (Ames & C. Schweinf.)
stipitata are small but conspicuous, triangu- Christenson, Lindleyana 14: 177. 1999.
lar, narrowly triangular to subquadrate. In C. Oncidium teres Ames & C. Schweinf.,
nuda, the column wings are almost absent, Sched. Orch. 8: 78. 1925. Stilifolium teres
reduced to two narrow flaps of columnar (Ames & C. Schweinf.) Königer & Pongratz,
tissue at each side of the stigmatic surface Arcula 7: 190. 1997. Trichocentrum teres
(thus the name Oncidium ebrachiatum). Fur- (Ames & C. Schweinf.) M. W. Chase & N.
thermore, the callus is strikingly different H. Williams, Lindleyana 16: 138. 2001.
between the two subspecies. In C. nuda, the Type. Panama. Veraguas: San Francisco,
callus is low and composed as described 1000 feet [350 m], C. W. Powell 383
above under that subspecies. In C. stipitata (holotype: AMES). (Fig. 1M–P)
the callus is composed of a single hemi- Cohniella aguirrei (Königer) Königer, Arcula
spherical callus that is somewhat laterally 10: 280. 2000. Stilifolium aguirrei Königer,
compressed. Arcula 9: 259. 1999. Trichocentrum aguirrei
Distribution.—Chiefly known from Pan- (Königer) M. W. Chase & N. H. Williams,
ama where it is found in the Canal Zone and Lindleyana 16: 218. 2001. Type: Colombia.
in the Provinces of Panama and Colón. It Huila: Departamento Girardot, 1200 m, W.
occurs only in tropical evergreen and tropical Köninger 95 (holotype: M; isotypes, JAUM ,
subdeciduous forests at elevations from sea K, Herb. Köninger [isotypes not seen]).
level up to 300 m. It has also been reported
from Costa Rica (Dressler, 1993; Mora de Diagnostic features.—Cohniella teres is
Retana, 1999) but the it is now apparent that very similar to C. ascendens and additional
the plant that served as the voucher for the collections from Panama and northern
inclusion of Oncidium nudum in the flora of Colombia are needed to understand the nature
of their relationship. Cohniella teres has Additional specimens examined. NICARAGUA.

shorter, narrower lateral lobes to the labellum Jinotega: Rápido Samaska, Río Bocay, ca. 100 m, Stevens
16450 (MO).
that are not erect nor projected forward as
they are in C. ascendens. Furthermore, the PANAMA. Veraguas: Soná, 25 m, 18 Feb 2005,
Carnevali 7027 (CICY).
flowers of C. teres are non–resupinate, as
opposed to the resupinate flowers featured by
C. ascendens. The distal portion of the
column in C. teres is bent forward, thus Acknowledgments
placing the stigmatic surface almost perpen- The authors are indebted to all of the
dicular to the anther, which is erect in C. herbaria that loaned us material and/or kindly
ascendens. Some of the specimens of C. teres assisted us while we visited their institutions:
display a lax covering of small papillae AMO, BIGU, M, MEXU, MO, MY, QMEX,
clothing the lateral lobes and the disk of the US, NY, XAL, VEN. Norris H. Williams
labellum. (FLAS), Franco Pupulin (JBL) reviewed the
Distribution.—Nicaragua, Panama, and first submitted draft of the manuscript; their
Colombia. It probably also occurs in Costa comments highly improved the quality of this
Rica. contribution. Lisa Campbell (NY), Donna
Variation range.—For a species known Tremonte (HUH), Rodrigo Duno, Ivón M.
from so few specimens, Cohniella teres dis- Ramírez, José Luis Tapia (CICY), Eliana
plays a surprising amount of variation. Every Noguera Savelli (CICY), Lizandro Peraza
specimen studied by us is different from the Flores (CICY), and Carlos Leopardi Verde
others. Flower size ranges from 10 mm in the (CICY) provided comments and suggestions
type of Cohniella teres to 18 mm in that of C. on earlier drafts of this manuscript. Gunter
aguirrei (a cultivated plant). The type of the Gerlach (M) helped us with texts in German.
species features linear-oblong, acute to Victorino Paiva Castro contributed images,
shortly acuminate lateral lobes (3–4 mm data and discussion of the Brazilian cohniellas.
long), while other specimens (e.g., G. Carne- Gaspar and Katia Silvera provided material of
vali 7027, CICY) have very small (ca. 2 mm Cohniella nuda, C. stipitata and C. teres.
long) linear lobes. Some specimens, includ- Óscar Moreno (CICY) provided material of
ing the type, have subquadrate, sessile petals C. biorbicularis. Silvia Hernández-Aguilar and
but the type of C. aguirrei displays narrower Lilia Can-Itzá (CICY) helped with handling
(albeit still sessile) petals. Although some- and databasing of loans. CONACyT partially
what different from the type material (nar- funded this project via grant 49980-Q “Filo-
rower petals, a callus that is somewhat genia molecular y morfológica, revisión siste-
different in its form) we have chosen to treat mática y una exploración de cuatro regiones
C. aguirrei as a synonym of C. teres because no-codificantes del genoma del cloroplasto
it shares the diagnostic features of C. teres. para estudios filogeográficos en el complejo
Many additional collections of the C. ascen- Trichocentrum (Orchidaceae: Cymbidieae:
dens/teres/aguirrei-complex are required to Oncidiinae)”, awarded to the senior author.
better understand whether more than one The generous support of the Orchid Society of
taxon is required to explain the variation in Arizona and the Massachusetts Orchid Society
the complex. We especially need collections (to GAR-G) is here acknowledged.
bridging the geographical gaps presently
found in the distribution of the C. teres.
Cohniella teres is known to us from a single Literature Cited
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Appendix
Key to the genera of the Trichocentrum clade
1. Leaves terete, fleshy coriaceous; pseudobulbs relatively small and inconspicuous . . . . . . . . . . . . . . . . . Cohniella
1. Leaves conduplicate, either rigidly fleshy or coriaceous; pseudobulbs small, inconspicuous, to relatively large and
conspicuous.
2. Plants small (leaves rarely exceeding 10 cm long); inflorescences shorter than the subtending leaves, mature
plants bearing few [1–3(–5)], succesive flowers; labellum basally produced into a spur . . . . Trichocentrum
2. Plants usually larger (leaves usually exceeding 12 cm long; however, they may be smaller in Lophiaris
pumila and relatives but flowers lacking a spur); inflorescences usually longer than subtending leaves
(shorter in Lophiaris pumila and relatives), mature plants bearing many [(5−)10–50(−150)], more or less
simultaneous, rarely succesive (e.g., Lophiaris lindenii) flowers; labellum lacking a spur.
3. Leaves rigidly and thickly fleshy coriaceous; pseudobulbs large and conspicuous, at least 2 cm long, but
up to 4 cm long; inflorescences stiffly erect, peduncle and rachis glaucous (with a thin film of wax),
covered with a thin film of wax; plants usually lithophytic . . . . . . . . . . . . . . . . . . . . . . . Lophiarella
3. Leaves coriaceous or fleshy coriaceous, rarely rigid; pseudobulbs small, rarely exceeding 1.5 cm long;
inflorescences more commonly ascendent or arching to nutant, never stiffly erect; peduncle and rachis
non-glaucous; plants usually epiphytic, rarely lithophytic . . . . . . . . . . . . . . . . . . . . . . . . . Lophiaris

A Synopsis of Cohniella

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A synopsis of Cohniella (Orchidaceae, Oncidiinae)

GERMAN CARNEVALI FERNÁNDEZ-CONCHA1, WILLIAM ROLANDO CETZAL IX2,

Abstract. A synopsis of the genus Cohniella is presented with nomenclatural updates

Brittonia, 62(2), 2010, pp. 153–177 ISSUED: 1 June 2010

A Neotropical genus of 13 species, some vegetative maximum. In nature, any given

Keys to species of Cohniella

Subkey I: Key to the species of Cohniella from Mexico

Subkey II: Key to the species of Cohniella from Central America

Cohniella ascendens (Lindl.) Christenson, Oncidium subulifolium Schltr., Repert. Spec.

Taxonomic commentary.—All citations and of the genus by the combination of a large

N, 104°41′59″ W , aprox. 350–450 m, subtruncate, somewhat reﬂexed in natural

of their relationship. Cohniella teres has Additional specimens examined. NICARAGUA.

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