A synopsis of Cohniella (Orchidaceae, Oncidiinae

A synopsis of Cohniella (Orchidaceae, Oncidiinae)

GERMAN CARNEVALI FERNÁNDEZ-CONCHA1, WILLIAM ROLANDO CETZAL IX2,

RICARDO BALAM NARVÁEZ1, AND GUSTAVO A. ROMERO-GONZÁLEZ

3

1Herbario CICY, Centro de Investigación Científica de Yucatán, Calle 43. No. 130. Col. Chuburnáde Hidalgo, 97200 Mérida, Yuc., México; e-mail: [email protected]; [email protected]

2El Colegio de la Frontera Sur, Unidad Chetumal, Av. del Centenario, km 5.5, Chetumal, QuintanaRoo, Mexico. C.P. 77000; email: [email protected]

3Orchid 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 updateswith a discusion of diagnostic features and biogeographical data. Thirteen species andone 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 amendedlectotype for Epidendrum cebolleta and several epitypifications and new synonyms.A key to the genera of the Trichocentrum complex is presented (Appendix) as well askeys 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, Cohniellapendula y Cohniella biorbicularis, se describen e ilustran. Se proponen las siguie-ntes combinaciones nuevas: Cohniella binotii, Cohniella brachyphylla, Cohniellacepula 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 ynuevos 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,Oncidiinae) is characterized by medium tolarge plants with inconspicuous to small pseu-dobulbs that bear a single, succulent, terete leaf.It includes 13 species that are known inhorticulture as the “rat-tail oncidiums”. Incommon with other groups within Oncidiinaethat bear yellow flowers that mimic malpighia-ceous blossoms, the species of Cohniella werefirst included within a broadly defined genusOncidium Sw. (Lindley, 1855; Reichenbach f.,

1863; Kränzlin, 1897; Garay & Stacy, 1974;Dunsterville & Garay, 1979; Foldats, 1970;Mora de Retana, 1999). Within Oncidium, rat-tails traditionally were referred to sectionCebolletae Lindl. (Lindley, 1842; Garay &Stacy, 1974) or to section Teretifolia (Lindley,1851, 1855; Reichenbach, 1863; Cogniaux,1906).Over time, it became evident that the rat-

tails were different from true Oncidium inseveral 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.

spicuous or small pseudobulbs bearing scar-ious sheaths and one terete leaf, their lowchromosome numbers (2n=26–36; Chase,1986; Chase & Palmer, 1989), and theirdistinctive leaf anatomy (Sandoval-Zapotitla& Terrazas, 2001). In contrast, species ofOncidium in a narrow sense are characterizedby conspicuous, laterally compressed pseu-dobulbs bearing one to several sheaths, ofwhich 1–3 may have foliar blades, and 1–3softly coriaceous, conduplicate leaves. These“typical oncidiums” also have higher chro-mosome numbers of 2n=56–60 (reviewed inTanaka & Kamemoto, 1984).Orchid hybridizers realized early on that it

was easier to obtain hybrids of the rat-tails withspecies of the genus Trichocentrum Poepp. &Endl. (Oncidiinae) and particular groups withinthe genus Oncidium than with the species oftypical oncidiums (Sanford, 1964). Thesegroups include the “mule ear oncidiums” (herereferred to the genus Lophiaris Raf.) and theOncidium splendidum A. Rich ex Duch. com-plex (for which the name Lophiarella Szlach.,Mytnik & Romowicz is available; Szlachetkoet al., 2006). Not coincidentally, the plants of allthe species that hybridize with the rat-tails havelow chromosome numbers and relatively smallpseudobulbs bearing scarious sheaths and asingle coriaceous or succulent leaf (hereaftercalled the “Trichocentrum syndrome”, TS),bespeaking of a relationship between themand the rat-tails.Recent phylogenetic analyses of nucleotide

sequences of both nuclear (Sosa et al., 2001;Williams et al., 2001a) and plastid (Chase &Palmer, 1989) DNA have shown that all thetaxa that exhibit TS form a highly supported,monophyletic entity. Some authors have pre-ferred to treat them as members of a broadlydefined Trichocentrum (e.g., Chase et al.,2005; Sandoval-Zapotitla & Terrazas, 2001;Sosa et al., 2001; Williams et al., 2001a, b;Chase et al., 2009) while others prefer torecognize several smaller, more narrowlydefined and easily diagnosable genera (e.g.,Braem, 1993; Pupulin, 1995; Königer &Pongratz, 1997, 1999; Jiménez-Machorro &Carnevali, 2001; Romero & Carnevali, 2000;Carnevali et al., 2001; Pupulin & Carnevali,2005). If recognized, the segregates of anarrowly defined Trichocentrum clade would

be the already mentioned Cohniella, Lophia-ris, Lophiarella, and Trichocentrum.The rationale for recognizing these segre-

gates follows several criteria outlined byBacklund and Bremer (1998) for the deriva-tion of classifications from phylogenies: (1) Tomerit recognition clades must be monophyleticand highly supported; (2) Nodes defininggenera preferably should include morpholog-ical synapomorphies that permit recognition ofmembers; and (3) The classification should beas consistent as possible (minimally disrup-tive) with previous classification systems.All these genera form highly supported,

monophyletic clades in phylogenetic analysesof both nuclear and plastid DNA (Sosa et al.,2001; Williams et al., 2001a, b; Cetzal et al.,in prep.), and of morphological and anatom-ical characters (Cetzal, 2007) . All of thesegroups are diagnosable using easily discern-able morphological characters (Cetzal, 2007;Cetzal et al., in preparation; see key to thegenera in the Appendix). There are alsomicromorphological and anatomical charac-ters that distinguish the segregates (Sandoval-Zapotitla & Terrazas, 2001; Cetzal et al., inprep.). The third criterion, that of consistencywith previous systems, is more problematic inthis case because an array of names andtreatments have been recently proposed inOncidiinae; some support a broadly definedTrichocentrum (e.g., Sandoval-Zapotitla &Terrazas, 2001; Sosa et al., 2001; Williamset al., 2001a, b; 2003; Chase et al., 2005;Chase et al., 2003; 2009) while others prefermore narrowly defined genera (e.g., Pupulin,1995; Romero & Carnevali, 2000; Carnevaliet al., 2001; Carnevali et al., in prep.;Jiménez-Machorro & Carnevali, 2001; Pupu-lin & Carnevali, 2005; Cetzal et al., 2008).Thus, names and classification systems forboth options are already available (i.e., practi-cally all the required combinations havealready been proposed) and which course ofaction is chosen must remain a matter of“practicality.”We have chosen to treat the major clades

within the Trichocentrum complex as distinctgenera primarily for two reasons, discussedbelow. In a subtribe such as Oncidiinae, wherethe generic circumscriptions are tenuous andconfusing at best, cryptic at worst (e.g., the

154 BRITTONIA [VOL 62

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generic limits within the clade that includesthe twig-epiphytes, see Chase, 1986), a nar-rowly defined Trichocentrum and Cohniella,Lophiaris, and Lophiarella are among the mostdistinctive species aggregates. Although allmembers of the clade share pseudobulbs bear-ing a single succulent leaf, all are easilyrecognizable using other characters. Trichocen-trum is unique due to its spurred flowers onshort, mostly successively-flowered racemes,and a consistently small vegetative habit. Theplants of all the other genera bear spurless,Oncidium-like flowers. Plants of Cohniella aremedium to large in size and bear terete leavesand erect to pendent inflorescences. Lophiarisis characterized by tiny pseudobubs bearingsucculent, relatively large leaves and arching totwinning inflorescences. Plants of Lophiarellahave relatively large pseudobulbs bearing thick,fleshy leaves and erect, stout inflorescencescovered with a thin coat of wax. We offer a keyto the four genera of the Trichocentrum com-plex that is entirely based on macroscopic,easily discernable characters (see Appendix).Furthermore, these four genera are horticultur-ally distinct (i.e., each has different culturalrequirements), a fact that orchid growers haverecognized (Sanders & Co., 1901). We predictthat, in this particular case, narrowly definedgenera will be better accepted by aficionados,the economically important horticultural trade,and in orchid conservation, rather than amorphologically and horticulturally heteroge-neous, broadly circumscribed Trichocentrum.The objective of this contribution is to provide

a synopsis of Cohniella, while clarifying manyof the problems associated with an orchid groupthat has a long history of cultivation andnumerous nomenclatural changes. Severalnames are typified, and a brief discussion isprovided for each species, stressing their distri-bution, variability, circumscription, and diagnos-tic characters. Furthermore, we describe andillustrate two new species and propose five newcombinations. Illustrations are also provided formost of the species treated in this synopsis.

Synopsis

Cohniella Pfitzer, Nat. Pflanzenfam. 2, 6: 194.1889. Cohnia Rchb. f., Bot. Zeit. 10: 928.1852, non Cohnia Kunth 1850 (Agavaceae).

Type: Cohnia quekettioides Rchb.f. [=Coh-niella ascendens (Lindl.) Christenson].

Oncidium sect. Cebolletae Lind., Bot. Reg.28: sub t .4. 1842. Type: Epidendrumcebolleta Jacq. [=Cohniella cebolleta(Jacq.) E. A. Christenson]

Oncidium sect. Teretifolia Lindl., Bot. Reg.32: sub t. 27. 1846. Type: Epidendrumcebolleta Jacq.

Oncidium sect, Teretoncidium Kuntze, Lex.Gen. Phan. 399. 1903. Type: Epidendrumcebolleta Jacq.

Stilifolium Königer & Pongratz, Arcula 7: 186.1997. Type: Epidendrum cebolleta Jacq.

Epiphytic or lithophytic herbs, caespitose;pseudobulbs small (relative to leaf length),suborbicular, bearing one leaf, clothed by threesmall, scarious sheaths; leaf terete, thicklyfleshy, often punctate or tinged with red orpurple; inflorescences lateral originating fromthe sides of the newest pseudobulbs, usuallypaniculate, rarely racemose, few to multi-flowered, with the flowers opening more orless simultaneously, peduncle clothed withsmall, remote, scarious bracts; flowers showy,resupinate, with the perianth segments subcor-iaceous, widely spreading, relatively longlasting (5–10 days); floral bracts inconspicu-ous; pedicellate ovary cylindrical; sepals sub-similar, clawed basally, the laterals somewhatoblique, usually green, chartreuse, or yellowwith red or maroon spots or blotches; petalssimilar to the sepals, the claw poorly devel-oped; labellum larger than the other perianthsegments, yellow (white in C. jonesiana) withred or maroon spots or blotches on or aroundthe callus area or the abaxial surface; con-spicuously 3-lobed, the central lobe usuallymuch larger than the laterals and joined to thedisk by a more or less well-developed claw,callus composed of a series of plates and/orteeth; column relatively short, subcylindrical,thicker at the base, stigmatic surface suborbic-ular, with well-developed stelidia at each side(almost absent in C. nuda), with a more or lesswell-developed tabula below the stigmaticsurface, rostellum transverse; anther terminal,operculate; pollinarium complex with 2 yellowobpyriform pollinia, an obovate or spathulatetegula, and a small, semi-hemispheric visci-dium. Fruit a capsule.

155CARNEVALI ET AL.: COHNIELLA (ORCHIDACEAE)2010]

A Neotropical genus of 13 species, some(e.g., Cohniella cebolleta and C. cepula) arerather broadly circumscribed in this synopsisand geographical variants may eventually berecognized as distinct taxa. The genus isdistributed widely from northern Mexico (C.brachyphylla reaching ca. 27° N in Sonora)into southern Brazil and northern Argentina(reaching ca. 26° S in Tucumán, Argentina),mostly in the lowlands. The species are mostcommon in seasonally dry forests. The genusis most diverse in Mexico where at least fivespecies occur, two or three of them endemic,and in Panama with four species.Cohniellawas based on Cohnia quekettiodes

Rchb.f. Reichenbach f. (1852) apparently mis-interpreted the relatively small size of the planthe described and the flower buds he examined(see Garay, 1963) and inferred a relationshipwith a totally unrelated clade of oncidioidorchids that also happens to bear terete leaves(he compared it to his genus Sigmatostalix andstated that “... Quekettia [Lindl.] was likely tobe closely related”: “Quekettia wird wohl auchsehr verwandt sein”.) The same author (1858)later provided a Latin description of his species(he had given only a brief description inGerman in the protologue) comparing it toOncidium ascendens Lindl. (“Planta habituOncidii ascendentis cui et vaginis amplisbrunneo fasciatis simile” [sic]) and an illustra-tion, which does appear to confirm that the typeplant bore only immature flowers. The use ofthe name Cohnia was, however, already occu-pied by a genus in the Agavaceae described byKunth in 1850. Pfitzer (1889) later proposed anew genus and a combination, Cohniellaquekettiodes (Rchb. f.) Pfitzer, which Garay(1973) referred to the synonymy of Oncidiumascendens Lindl.As treated here, most of the species of

Cohniella are rather variable entities. The natureof the variation is twofold, having both vegeta-tive and floral components. Most species reachflowering size at a fraction of their potential

vegetative maximum. In nature, any givenpopulation will include plants bearing flowersvarying in size from small to large, the latterwith leaves 2–3 times longer than those of thesmaller plants, thus rendering plant size unreli-able for species recognition. However, somespecies are, on average, smaller than others, andspecies in certain groups within the genus tendto be smaller than those in other groups (e.g., theplants of species in the Cohniella nuda complextend to be smaller than those of species in the C.cebolleta complex).There is also considerable variation in the

flowers of most species. InCohniella ascendens,the flowers vary to such an extent that it could beeasy to assume that one deals with more thanone species except that most of the variation isfound within a single population, even amongplants growing on the same phorophyte. Varia-tion involves in some measure flower size, butmostly resides in the outline of the labellumlobes, and the degree and depth of the colorpatterns. Despite all this variation, most speciesare easily diagnosed by the combinations offloral features that we discuss below.Inflorescence length, on the other hand,

though variable in absolute dimensionswithin species, is diagnostic for species orspecies groups and tends to remain within arange. For example, the inflorescence inspecies of the Cohniella nuda complex ofspecies is most commonly conspicuouslyshorter than the leaves while the opposite istrue for species related to C. cebolleta.The 13 species of Cohniella recognized in

this synopsis can be diagnosed with the use ofthe following set of keys. For ease ofidentification, we have chosen to offer threeseparate keys, which are geographically cir-cumscribed. Since Cohniella longifolia and itsvariation are not well-known, we have decidedto leave it out of the keys. For this key to workappropriately, users need good flowers andpreferably a portion of an inflorescence andthe subtending leaf.

Keys to species of Cohniella

1. Plants from Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subkey I1. Plants from Central America, South America, or the West Indies.

2. Plants from Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Subkey II2. 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 erectand partially enfolding the column; petals and sepals usually much longer than the lateral lobes of thelabellum; column wings linear, finger-like, much longer than wide; plants from tropical moist to tropicalsubdeciduous 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 thesame plane than the central lobe), flat and not enfolding the column; petals and sepals usually much shorter thanthe 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 andNayarit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 fromwestern 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 fromnorth of the Neovolcanic Transversal Axis in Sinaloa with outliers in Durango, Chihuahua, andSonora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brachyphylla

3. Leaves various, either rigidly erect to flexible, usually few (3–5) to a plant; inflorescences variousbut usually patent to somewhat pendent, more rarely erect, usually a panicle with 2 or morebranches in well-developed plants; plants from the Gulf coast of from the Pacific coast south of theTrans-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 orfrom 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 thedisk; 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 veryshallowly so and then the emargination making an obtuse angle; labellum usually bearingspots on most of the undersurface; leaves 6–10 mm wide, conspisuously wider basallybut abruptly tapering at base (forming a “neck” just above the pseudobulb) and moregradually tapering distally; plants from the Pacific states of mexico, south of theTehuantepec 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 thenthe 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” justabove 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) orlonger 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. ascendens4. Lateral lobes of the labellum flat, not enfolding the column; flowers resupinate or not

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. teres3. 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 keelflanked by longitudinal depressions; plants from Venezuela, Colombia (eastern side of theAndes), and Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nuda

5. Columnwings present, subterete; callus composed of a single, massive central teeth occupying thewhole 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 longor as long as wide; plants from western and central Venezuela and the eastern slopes of the Andes inColombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 Caucavalley and presumably from the Chocó region in Colombia . . . . . . . . . . . . . . . . . . . . . . . .C. teres

1. Inflorescences usually longer than the subtending leaves; flower diameter variable; column wings broaderthan 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 southof the Amazon river course, reaching SE Brazil and Paraguay and along the eastern side of theAndes in Peru and Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. cepula

4. Callus composed of 5 keels, provided with accessory cally at each side of the labellar isthmus; column ofequal width at base and apex: columnwings 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. stacyi5. 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 dentate6. 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, andneighbouring 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,Lindleyana 14(4): 177. 1999. Oncidiumascendens Lindl., Edwards’s Bot. Reg. 28:sub t. 4. 1842. Stilifolium ascendens (Lindl.)Königer & Pongratz, Arcula 7: 186. 1997.Trichocentrum ascendens (Lindl.) M. W.Chase & N. H. Williams, Lindleyana 16(2):137. 2001. Type: Guatemala. Without anyother locality, Apr 1841, K. T. Hartweg s.n.(holotype: K-Lindl.). (Fig. 1A–D)

Cohnia quekettioides Rchb. f., V. Schl. Bot.Zeitung (Berlin): 10: 928. 1852. Cohniellaquekettioides (Rchb. f.) Pfitzer, Nat. Pflan-zenfam 2(6): 194. 1889. Type: Guatemala.Chantalas, “Mons Espina”, 1841, E. R. vonFriedrichsthal 834 (holotype: presumably atW, not seen).

Oncidium subulifolium Schltr., Repert. Spec.Nov. Regni Veg. Beih. 10: 79. 1922.Oncidium bolivianense Oppenheim, Orchis10: 93. 1916. (non Oncidim bolivienseRolfe, 1907). Type: Bolivia. Río Itenez,O. N. Witt s.n. (holotype: B, destroyed;lectotype, here designated, Orchis 10, No.5, Tafel IV, Fig. 2. 1916).

Oncidium helicanthum Kraenzl., Pflanzenr.(Engler) 95: 281. 1922. Type: Colombia.Without any other locality or collector(holotype: B, destroyed; lectotype, heredesignated, Das Planzenreich (A. Angler)heft 80, 4, 50: 282, Fig. 24C, a–d. 1922).

Diagnostic features.—Cohniella ascendensis easily recognized by its relatively smalllateral lobes of the labellum that are held


FIG. 1. Flowers of Cohniella. A–D. Cohniella ascendens. A. Whole flowers, frontal view. B. Labellum, frontalview. 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. Columnwith 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 Carnevali7283, CICY; I–L from Carnevali 7311, CICY; M–P from Carnevali 7027, CICY).


erect and partially enclosing the column. Thecallus is relatively simple and consists of agreatly reduced proximal, transverse platewith or without small, lateral low teeth anda much larger distal, apically rounded and

laterally compressed tooth surrounded bylower, reduced lateral keels. The columnwings are narrow and point downward andinward, almost in a hook-like fashion. As inits hypothesized closest relatives, C. teres and

FIG. 2. Flowers of Cohniella. A–G. Cohniella biorbicularis. A.Whole flower, front view. B. Whole flower, front andback view.C. Labellum, front and back views.D. Sepals and petals, front and back views.E.Columnwith anther cap, frontand 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.).


C. nuda, C. ascendens is more of a shade-loving species and the leaves are relativelysoft and flexible. The inflorescences areusually shorter than the subtending leaves.Distribution.—Mexico southward to Pan-

ama; possibly northern Colombia and NWVenezuela. The Colombian and Venezuelanrecords of the species are somewhat doubtfuland better material is required to confirm them.Variation range.—This is a common spe-

cies with a great deal of floral variation,mostly in the shape and size of the laterallobes of the labellum. The color of the callusis also variable, ranging from almost con-colorous with the bright yellow labellum todeep purple or even guava-colored. Thepetals are usually oblong-spathulate, but insome clones they can be broadly obovate.Taxonomic commentary.—This is the type

of the genus Cohnia Rchb. f. (=Cohniella).The type locality is not mentioned in theprotologue but the author cited it a few yearslater (Reichenbach f., 1858). In the samepublication, he stated that he had described aunicate at the Naturhistorisches MuseumWien (“Das unicum im Herbar des WienerMuseums”), where the collections of E. R.von Friedrichstall are, but we have not beenable to examine it. However, Garay (1973)clarified the identity of the Cohniella queket-tioides as being identical to the earlierOncidium ascendens (Christenson, 1999).We have studied the original plate of Cohniaquekettioides and concluded that Garay wascorrect in its identity assessment. This platefeatures a terete leaved plant with the sche-matic of a flower which clearly resembles theentity we currently treat as Cohniella ascen-dens except for the reduced lateral lobes ofthe labellum, something resulting from thedissection of a flower bud.Oncidium subulifolium (O. bolivianense

Oppenheim 1922 non O. boliviense Rolfe1907) was described from a cultivated plantsupposedly collected in Bolivia, but nofurther material referable to this species (orto the complex of taxa related to C. ascen-dens) has ever been collected in Bolivia, andwe assume the type plant must have beencollected in another country. Known collec-tions of Cohniella ascendens reach theirsouthernmost limits in northern Colombia(possibly into NW Venezuela) and it is highly

unlikely that the species jumps the gap of theAndean ranges and the Amazon Basin to re-appear in Bolivia. The highly schematic typeillustration of Oncidium helicanthum is con-sistent with the overall morphology of C.ascendens and, in concordance with otherauthors (e,g., McLeish et al., 1995), it istentatively placed here.

Additional specimens examined. MEXICO.Campeche: a 65 km al S de Conhuas en el centroceremonial de Calakmul, Límite N del Petén Guatemalteco,16 Mar 1983, Cabrera 4423 (MEXU). Chiapas: Mpio.Catazajá, orilla oeste de laguna de Catazajá, 20 Abr 1999,Gutiérrez & Balam 6437 (CICY). Oaxaca: Juchitán, 9 kmal NE de Lázaro Cárdenas camino a Sta. María Chimalapa,250 m, 22 Feb 1982, Cedillo & Torres 1114 (MEXU).Quintana Roo: Othón P. Blanco, Ejido Graciano Sánchez,área forestal ca. 6 km al W de Valle Hermoso, 5 Mar 1995,Trejo & Olmsted 311 (CICY). YUCATÁN: Mpio. deValladolid, rehollada cerca de Valladolid, 27 Mar 1997,Carnevali & Ramírez 4386 (CICY).

BELIZE. Corozal: Lower slopes of RichardsonPeak, Maya Mountains, directly N of the Junction ofRichardson Creek and Bladen Branch, 88º46′30″ N,16º33′35″ W, 300–620 m, 4, 6, 8 Mar 1987, Davidse& Brant 32522 (MEXU, MO).

GUATEMALA. Petén: Ruinenstadt Tikal (ca. 60 kmNE Flores/Santa Elena), ca. 3 km W von Tempel IV,17º13.387′ N, 89º37.683′ W, 440 m, Förther et al. 9986 a(M).

EL SALVADOR. Ahuachapán: Plantas florecidas encultivo en la ciudad de Santa Ana, traídas de la orilla delrío Nejapa, Cantón La Pandeadura, 8 km al Sur de laciudad de Ahuachapán, 800 m, Linares 1175 (MEXU).

HONDURAS. El Paraíso: Dpto. El Paraíso. Mpio.de Morocelli, Loc. Quebrada El Carrizal cerca El Plan,±2 km al E de El Plan, 17 Feb 2002, Linares 5692(MEXU).

NICARAGUA. Zelaya: Colonia Kururia, 14º41′ N,84º04′ W, 50 m, 3 Mar 1975, Pipoly 3895 (MEXU, MO).

COSTA RICA. Alajuela, Los Chiles, RefugioNacional de Vida Silvestre Caño Negro, Llanura deGuatuso, Playuelas, 40 m, 3 Feb 1993, K. Martinez,Zamora, Chavarría & Flores 50 (UCR, MO).

Cohniella binotii (Pabst) G. A. Romero &Carnevali, stat. nov. Oncidium jonesianumvar. binotii Pabst, Bradea 2: 170. 1977.Type: Brazil. Minas Gerais: regione MontesClaros, 24 Feb 1972, Casa Binot s.n.(holotype: HB).

Diagnostic features and variation range.—Cohniella binotii is closely related to C.jonesiana. It differs, however, in the largerflowers (sepals and petals to 2.5 cm long andlabellum to 2.8 cm wide) and in the large,


irregular blotches in the petals and sepals thatare almost confluent and cover most of thesurface of the perianth segments. It alsooccupies a disjunct range in SE Brazil.Distribution.—Endemic to Brazil, known

only in the area of Montes Claros in MinasGerais.

Additional specimen examined. BRAZIL. DistritoFederal: Brasilia, Schumburg 19/705 (AMES).

Cohniella biorbicularis Balam & Cetzal, sp.nov. Type: Mexico. Querétaro: MunicipioLanda deMatamoros, Camino deMatzacintlaal Río Moctezuma, 21°20′04″ N, 99°20′04″W, 1100 m, cañada orientada SE con vegeta-ción de bosque tropical caducifolio sobreladeras de roca caliza; colectada original-mente en el año 2006 por I. M. Ramírez(#1432); floreciendo en cultivo el 10 Marzo2008, G. Carnevali & I. M. Ramírez 7308(holotype: CICY; isotypes: AMES, AMO,MEXU, QMEX, SEL, US). (Fig. 2A–G)

Species haec Cohniellae brachyphyllae similis sedfloribus majoribus, lobis labelli lateralibus suborbicularibusvel latissime obovatis proportione majoris recedit.

Epiphytic herbs, caespitose; rhizome short,thin, brittle; roots 1–1.5 mm thick, whitewhen old; pseudobulbs 5.6–9.9×4.8–6.3 mm,subspherical to broadly ovoid, apically 1-leaved, green, totally enclosed by 3 imbricatesheaths, eventually deciduous; leaves terete,thickly fleshy-coriaceous 10–27 cm long, 2–5 mm thick, dark green; inflorescencessolitary from the base of the pseudobulbs,13.50–112.20 cm long, a 27–53 floweredraceme or panicle with 1–3 short branches,the branches 3–6 flowered; peduncle andrachis green-purple; peduncle more or lesserect, 3–4 mm thick, terete, with 6–12 shortlybracted internodes, the basal-most longest,oblanceolate, acuminate, tubular; floral bracts3.31–5.10 mm long, narrowly elliptic, acu-minate; flowers with perianth segmentswidely opening and the petals and sepalssomewhat reflexed; ovary with pedicel 15.5–22 mm long, of which ca. 3.71–7.29 mmcorrespond to the ovary, this 0.60–1.33 mmthick; sepals basally clawed, flat or somewhatreflexed; dorsal sepal 6.27–7.50 × 2.52–4.51 mm, obovate-lanceolate, apically obtuseand minutely apiculate, concave in the upper

half, the claw 1.22–3.17×0.76–0.95 mm;lateral sepals partially fused at the very base,ovate-lanceolate, 6.34–8.44 × 2.07–4.33 mm;petals 6.70–8.13×3–4 mm, oblanceolate, theapex rounded, somewhat repand and reflexedin natural position; labellum 3-lobed, 10.82–15.36 mm long from the base to the apex ofthe central lobe, 11.81–18.91 mm wide acrossthe apices of the lateral lobes, the lateral lobesin the same plane as the central lobe and+/−perpendicular to it; central lobe 10–11 ×17–18 mm, transverssely obdeltoid to sub-quadrate in outline, apically rounded tosubquadrate, basally produced into a shortisthmus, 2 × 2–3 mm; lateral lobes 13–19×5–6 mm, suborbicular to very broadly obovate,apically rounded; disc large, ca. 3.5×5.5 mm,bearing a well-developed callus, ca. 5×3 mm,consisting of a large, elevated, +/−flat, oblon-goid, apically with two teeth, the central onelaterally compressed, the laterals very larger,rhombic, the basal portion of the callus withsharp upper margins; column 3–3.5 × 1 mm,tabula infrastigmatica obdeltoid to subqua-drate, stigmatic surface, sub-rounded; columnwings short, ca. 1 × 1 mm, asymmetricallybilobed; anther 3 × 2 mm, apical, operculate,ellipsoid; pollinarium typical for the genus.Distribution and ecology.—Endemic to

Mexico. Cohniella biorbicularis is restrict-ed to the Gulf Coast states, west and northof the Tehuantepec Ithsmus, namely Tam-aulipas, San Luis Potosí, Querétaro, andVeracruz. It grows at elevations of 0–1400 m, usually in tropical dry forests orin the ecotones between tropical dry forestsand pine-oak forests. The plants alwaysgrow on trees.Diagnostic features.—Cohniella biorbicu-

laris is easily recognized by its relativelylarge, suborbicular to very broadly obovatelateral labellum lobes. These are almost aswide as the midlobe in some specimens(Nagel 6734, MEXU; Carnevali & Ramírez6372, CICY), but more commonly they aresomewhat narrower but always at least 75%as wide as the width of the midlobe. Theplants look very similar to those of thewestern Cohniella brachyphylla, but can bedistinctly larger.Variation range.—Plants from Veracruz

have the broadest lateral lobes while thosefrom Querétaro and San Luis Potosi have the


narrowest. Aside from this, Cohniella bio-rbicularis is a species with relatively littlevariation in floral characters.Taxonomic commentary.—As with most

taxa in the Cohniella cebolleta complex, C.biorbicularis has been confused with thatspecies. However, the broad lateral lobes ofthe labellum easily distinguish this speciesfrom any other Cohniella.

Additional specimens examined. MEXICO. Querétaro:Mpio. Arroyo Seco, 6 km al N de Concá, 600 m, 18 Mar1985, Fernández N. 2778 (US); río Santa María, 1.5 km alNE del puente de Concá, 600 m, 6 Mar 1988, Herrera 61(XAL); Mpio. Jalpan, km 194 carretera hacia Landa deMatamoros, 1490m, 16 Apr 1984,Aguirre 41–638 (AMO);oriente de Tanchanaquito, 300 m, 8 Apr 1992, López 986(XAL); Mpio. Landa de Matamoros, S de El Capulín,Puerto Blanco, camino al Río Moctezuma, 920 m, 13 Mar1988, Herrera 103 (AMO). San Luis Potosí: Mpio. ElNaranjo, 5 Apr 1960, Dressler 2606 (US). Tamaulipas:Mpio. Antiguo Morelos, about 7 km N of Antiguo Morelos(km 540), 28 Mar 1961, Dressler 2632 (US); Mpio.Aldama, pueblo El Plomo, Sierra de Tamaulipas, 550 m,19 Jan 1991, Jiménez, O. Rocha & C. Rocha 1090 (AMO).Veracruz: Mpio. Atoyac, 1.5 km al N de Potrero Nuevo-Miraflores, 18º53′58″ N, 96º49′21″W, 425 m, 7 Mar 1986,Acevedo & Acosta 880 (XAL); Mpio. Jilotepec, compradaen las calles de Xalapa, 26 Apr 2001, Carnevali & Ramírez6372 (CICY, 3 sheets);Mpio.Martínez de la Torre, 20º03′N,97º03′W, 300 m, 30 Mar 1935, Nagel 4656 (US); Mpio.Naolinco, ca. SanAntonio Paso del Toro, 19º35′N, 96º49′W,600 m, 23 Mar 1975, Ortega 745 (XAL); Mpio. Actopan,Trapiche 400 m, 15 Feb 1971, Ventura A. 3114 (US).

Cohniella brachyphylla (Lindl.) Cetzal &Carnevali, comb. nov. Oncidium brachy-phyllum Lindl., Edwards’s Bot. Reg. 28:sub t. 4. 1842. Type: Mexico. Withoutprecise locality,K. T. Hartweg s.n. (holotype:K-Lindl.). (Fig. 3A–D, H)

Diagnostic features.—Cohniella brachy-phylla is distinguished by its relatively short,thick leaves on top of relatively conspicuouspseudobulbs. The leaves are typically rigidlyerect and thickened on the lower half; theyare also almost always somewhat to stronglyfalcate. The inflorescences are either erect orhorizontal, usually not as long as they are inother species of the C. cebolleta complex.The isthmus of the labellum is short andbroad, wider than its length or about as longas wide as opposed to the longer isthsmusfound in populations here referred to C.cebolleta. Along with C. pendula and C.biorbicularis, the labellum is wider across the

spread lateral lobes than across the apicallobe, as opposed to the labellum in C.cebolleta, which is wider across the spreadapical lobe of the labellum. The lateral lobesof the labellum are obliquely oblong toobliquely obovate and are always more orless retrorse. The midlobe of the labellum isproportionally smaller than in the speciesrelated to C. cebolleta. The disk of thelabellum is suborbicular to obliquely pentag-onal, proportionally large, and about as longas the width of the midlobe. At each side ofthe isthmus, anterior margins are providedwith one tooth on each one.Distribution and ecology.—Mexico, possi-

bly into El Salvador, Guatemala, Honduras,and Nicaragua. In Mexico, it is widelydistributed along the Pacific coast of thecountry and into the intermountain valleys.It has been collected as far north as ca.Álamos, in Sonora (ca. 27°N) and thensouthwards to Oaxaca, along the westernslopes of the Sierra Madre Occidental. Onthe eastern side of the Tehuantepec Isthmus,the species or a similar entity reappears inChiapas and then ranges southwards into atleast Costa Rica at 0–800 (−1500) m; theseplants are somewhat different (see below).Cohniella brachyphylla is usually found inseveral kinds of dry forests, even into thornscrub, and is the Cohniella from the driestenvironments. It is almost always an epiphyteon thick branches, very rarely on rocks. It isoften found fully exposed to the sun.Variation range.—Cohniella brachyphylla

is a common and widespread species, and it isconsequently variable. The fact that thespecies occurs as a conglomerate of popula-tions isolated in intermountain valleys hasapparently created barriers to gene flow,favoring the establishment of local forms orraces. Since these populations are also linkedby neighboring, often intermediate popula-tions occurring along the coastal lowlands oftropical western Mexico, there has not beenextensive speciation associated with the pop-ulations currently referred to this species.Exceptions are the populations from coastalJalisco here treated as C. pendula which arediscrete in the variation of several features(see discussion under C. pendula).The great amount of variation found within

Cohniella brachyphylla occurs both between


and within populations. The shape of theseveral sections of the labellum, as well as itsgeneral outline are variable. The lateral lobes ofthe labellum, which are always somewhatreflexed, range from obovate-suborbicular tosubquadrate to obliquely elliptical. The mid-lobe ranges from almost suborbicular to trans-versely subquadrate. In some populations (e.g.,

some in the state of Morelos), the central lobeof the labellum is shortly but distinctly apicu-late. The calli and disk are always somewhattinged in various hues of red, but the extent andpatterns of these colorations are variable. Theadaxial side of the labellum is almost alwaysprovided with dull red-rose spots that arevariable in size, distribution, and density.

FIG. 3. Flowers of Cohniella. A–D, H. Cohniella brachyphylla. A–D. Whole flower, front views. H. Column withanther cap, front view. E–G. Cohniella cebolleta. E. Whole flower, front view. F. Labellum, front views G. Columnwith anther cap, front view. I–M. Cohniella cepula. I. Column with anther cap, front view. J, L. Whole flower, frontviews. 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.).


Populations here referred to Cohniellabrachyphylla from the southernmost extremeof the distribution from Chiapas southwardinto Guatemala are atypical in having broadlyobovate lateral lobes but otherwise conformto the characters of the species. On thenorthwestern extreme of the range ofthe species, the plants are smaller and theinflorescences tend to be racemose while theflowers are somewhat similar to those seen inthe Chiapas plants.

Selected specimens examined.MEXICO. Chihuahua:Mpio. Batopilas, arroyo Guimivo between Río Batopilasand Guimivo, 762–915 m, 24 Mar 1979, Bye et al. 9235(NY, MEXU). Durango: Glacala, 23 Feb 1899, Goldman327 (US). Estado de México: Temascaltepec, Guayabal, 9Feb 1933, Hinton 3362 (AMES). Guerrero: Near SanNicolas del Oro, 1200 m, 13 Jan 1938, Mexia 9106(AMES). Jalisco: Mpio. Autlán, Puerto los Mazos, a10 km antes de Autlán a 7 km antes de la desviación aGuadalajara, 1000 m, 4 Oct 1987, R. Jiménez et. al 756(AMO). Michoacán: Coalcoman, 4 Apr 1939, Hinton13647 (US, AMES, NY). Morelos: Tlaquiltenango 2 kmal N de la desviación a San José Pala, sobre el camino aHuatla, 7 Nov 1996, A. Espejo et al. (AMES). Oaxaca:Microonda, San Cristóbal, Carr. Oaxaca-Tehuantepec, km144, desviación hacia la derecha aprox. 1 km, 1100 m, R.Jiménez et al. 3169 (AMO). Sinaloa: Mpio. de Culiacán, amás o menos 46 km al N de Culiacán, camino deBadiraguato a la Pitayita, 25º02′00″ N, 107º23′00″ W,100 m, Vega, Gutierrez G. & Hernández 8104 (MEXU).Sonora: Álamo Gordo, Carnevali & Gómez-Juárez 6803(CICY).

GUATEMALA. Huehuetenango: Mpio. Nentón, km10 a Gracias a Dios, 15º53′22.8′ N, 91º44′0.4″ W,1099 m, 24 Mar 2007, Velásquez et al. 184 (BIGU);km 11 a Gracias a Dios, 15º53′2.7″ N, 91º44′16.4″ W,1074 m, 21 Sep 2006, Véliz et al. 17461 (BIGU); km 11hacia Gracias a Dios, 15º53′0.63″ N, 91º44'21.0″ W,1061 m, 21 Sep 2006, Véliz et al. 18454 (BIGU). Jalapa:El Rancho, 28 Dec 1907, Kellerman 7002 (NY).Progreso: San Agustín AC Transecto El Rancho Norte,14º53′28″ N, 90º01′17″ W, 200–300 m, 20 Sep 2003,Cobar et al. (BIGU).

EL SALVADOR. Santa Ana: Mpio. Santa Ana,salida a Metapan, ca. 8 km al N del centro de la ciudadde Santa Ana, 10 Jan 2000, Linares s.n. (MEXU); Mpio.Metapán, ca. 6 km al NO de Metapán, por el camino aldespoblado, 12 Jan 2000, Linares et al.4755 (MEXU).

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,15 Jan 1925, Alfaro s.n. (AMES, US). Guanacaste: LaCruz, 15 Jan 1930, Jiménez 7909 (AMES).

Cohniella cebolleta (Jacq.) Christenson,Lindleyana 14: 177. 1999. Epidendrumcebolleta Jacq., Enum. Syst. Pl. 30. 1760.Oncidium cebolleta (Jacq.) Sw., Kongl.

Vetensk. Acad. Nya Handl. 21: 240.1800. Stilifolium cebolleta (Jacq.) Königer& Pongratz, Arcula 7: 186, 187. 1997.Trichocentrum cebolleta (Jacq.) M. W.Chase & N. H. Williams, Lindleyana 16:137. 2001. Type: Colombia. Cartagena[Carthagenae]. 1758, N. J. Jacquin s.n.(holotype: not located and most likely lost;lectotype: Select. Stirp. Amer. Hist., ed. 2,t. 217 [text on page 111], 1781, designated,albeit incorrectly, by Garay & Sweet,1974: 205; amended lectotype: Select.Stirp. Amer. Hist. [text on pages 230–231] t. 131, Fig. 2, 1763). (Fig. 3E–G)

Epidendrum juncifolium L., Sp. Pl. (ed. 2)1351. 1763. Cymbidium juncifolium (L.)Willd., Sp. Pl. 4: 102. 1805. Oncidiumjuncifolium (L.) Lindl., Coll. Bot. 27.1821. Type: presumably collected in Haiti[“habitat in America”], Plant. Amer. fasc. 8:t. 184, f. 2. 1759; epitype, here designated:Martinica, P. Dusse 2078 (NY).

Oncidium humboldtii Schltr., Repert. Spec.Nov. Regni Veg. 23: 65.1926. Oncidiumottonis Rchb.f. ex Kraenzl., Pflanzenr.(Engler) IV, Fam. 50: 92, Fig. 10. 1922.(non Schltr., 1914). Type: Venezuela.Without locality, 1840, K. F. E. Otto 997(syntype: B, destroyed; isosyntype: W-Reichenbach 27863); Guárico: Orituco,K. F. E. Otto 541 (syntype: B, destroyed;isosyntypes: W-Reichenbach 27869 &27872); Carabobo: Puerto Cabello, A. vonHumboldt & A. Bonpland s.n. (syntype:B-W); lectotype, here designated: A. vonHumboldt & A. Bonpland s.n. (B–W).

Oncidium ultrajectinum Pulle, Rec. Trav. Bot.Neerl. 4: 121. 1907. Type: Surinam.“Cultivated at the Utrech Botanical Gardenfrom a plant coming from Surinam as a giftfrom v. Asch v. Wijck, the Governor of thecolony, to our University” (holotype: pre-sumably at U, not seen).

Diagnostic features.—Cohniella cebolletais characterized by the small, relativelyinconspicuous pseudobulbs that bear a rela-tively narrow but fairly rigid leaf. The flowershave a complicated callus with a relativelylong basal oblong plate and three apical teethor keels, the central one being the largest.There are one or two additional teeth on each


side of this basal plate. From its Mexicanrelatives, C. cebolleta can be distinguished bythe general outline of the labellum as indi-cated in the discussion under C. brachy-phylla. The column of Cohniella cebolleta isnarrower than in C. cepula and the columnwings are pure yellow (very rarely with a hintof reddish on the margins) while in C. cepulathese are always maculated with large redspots.Distribution and ecology.—As understood

here, Cohniella cebolleta is a species wide-spread in the Caribbean, ranging from theYucatan Peninsula in southeastern Mexico,throughout Central America and into northernColombia and Venezuela. It also occurs inJamaica and the Lesser Antilles. Cohniellacebolleta grows in the driest environmentsoccupied by any Cohniella (except somepopulations of C. brachyphylla), typicallytropical dry forests, or even coastal dunes orthorn forests. In Venezuela and Colombia it isnot an uncommon sight on large branches ofold saman [Albizia saman (Jacq.) F. Muell.],guatacaro [Bourreria cumanensis (Loefl.) O.E.Schulz] or caro or guanacaste [Enterolobiumcyclocarpum (Jacq.) Griseb.] trees. On aregional scale, C. cebolleta is often allopatricor parapatric with other species of the genusthat usually grow in more humid environments.In all of Central America, where C. ascendensandC. cebolleta are parapatric or sympatric, thesecond grows only on the driest vegetationstypes, whereas the latter occurs in more humidecosystems. Cohniella cebolleta reaches itssouthernmost distribution at the mid course ofthe Orinoco River in Venezuela. Then, there is agap in the distribution of the genus Cohniella,which reappears at the southern margin of theAmazonas River, where it is represented byC. cepula.Variation range.—As here circumscribed

and even after excluding several taxa heretreated as distinct species, Cohniella cebolletaremains a very variable entity, mainly inflower size and labellum shape. The labellumis usually widest across the apical (central)lobe but in some populations it can be asbroad or even slightly broader across thespread lateral lobes. The labellum isthmus isalmost always conspicuous (about as long asthe disk), but in some individuals it isshorter. Flower size is also variable. Plants

from central and coastal Venezuela as well asthose from eastern and coastal Colombia aretypical of the species (type from ca. Cartagena,Department of Bolívar) and feature relativelylarge flowers with a long isthmus. Theseforms are well depicted in the iconographyof this paper and by illustrations by Dunster-ville (e.g., featured in Romero & Carnevali,2000).In populations of the northern rim of the

Guayana Highlands, the flowers are smallwith central lobes of the labellum about10 mm wide, while populations along theAndean foothills in Colombia and Venezuelaoften have large labella with midlobes 15–20 mm wide. In eastern Venezuela, thepopulations feature the largest flowers of thiscomplex (up to 35 mm diam.) and mayrepresent a different taxon. In the Guianas,the flowers are small (18–20 mm diameter)and have a distinctive callus. Should theseGuianan populations need to be treated as adifferent species, the name Oncidium ultra-jectinum is available. The Lesser Antillespopulations of the complex (including thetype of Cymbidium juncifolium) have smallflowers (18–20 mm) with narrow lateral lobesto the labellum associated with relativelylarge petals and sepals heavily spotted.Taxonomic commentary.—Christenson

(1999) cited the wrong basionym (“Dendrobiumcebolleta”, supposedly published in 1760,although the genus Dendrobium was not pub-lished by O. Swartz until 1799). This wrongcitation is considered a “correctable error” sincethe author cited the right publication, page, andyear (K. Gandhi, pers. comm.). The combinationDendrobium cebolleta, as far as we have beenable to ascertain, has never been formallypublished. Nonetheless, it does appear in numer-ous publications and at least one major databaseon the Internet (i.e., Kew’s World Checklist ofMonocotyledons at http://apps.kew.org).Garay and Sweet (1974: 205), proposed

plate 217 of the second edition of Jacquin’sSelectarum Stirpium Americanarum Historia...(1781) as the lectotype of Epidendrum cebol-leta (de facto, originally as “holotype”). Thisplate shows a plant with a paniculate inflor-escence, which is much longer than the leaves,bearing multiple fruits. However, plate 131,Fig. 1 of the first edition of this work,published in 1763, unambiguously shows the


http://apps.kew.org

vegetative parts and a separate, cut-out inflor-escence (clearly showing a peduncle longerthan the leaves) of Epidendrum cebolleta, andit clearly agrees with the 1760 protologue andhas priority over the plate in the secondedition. Stafleu and Cowan (1979: 408) statedthat “The 1763 publication (i.e., SelectarumStirpium Americanarum Historia) is an impor-tant complement to the 1760 Enumeratio (i.e.,the source of the protologue) and shouldalways be consulted with it”.Furthermore, both the 1763 and 1781

editions of this work clearly state that thetype was collected in Cartagena, Colombia(Carthagenae in sylvis praesertim maritimis),and not in Martinique, as stated in Garay andSweet (1974: 205) and later in Nir (2000:263), nor in Brazil, as stated in Foldats (1970:298). We have exhaustively searched for thetype material but have not been able to locate it.It could be argued that Jacquin might have

collected either Cohniella cebolleta or C.nuda (Bateman ex Lindl.) Christenson inCartagena, since Jacquin’s original collectionand description did not include floral charac-ters that distinguish them, and these twospecies are vegetatively undistinguishableand possibly could occur in habitats whereJacquin collected. Cohniella cebolleta, how-ever, always bears inflorescences much lon-ger than the leaves, versus inflorescencesabout as long or, more commonly, shorterthan the leaves in C. nuda.With the exception of the large-flowered

species of South America (Cohniella jonesi-ana and C. stacyi) and the species related toC. ascendens, almost all other taxa of thegenus have at one time or the other beenconsidered synonyms of Cohniella cebolleta.As a result, published distributions for Coh-niella cebolleta report, albeit incorrectly, aspecies ranging from northern Mexico intoSE Brazil and Argentina. Oncidium juncifo-lium was based on a plant without a preciselocality in America, but being a speciesnamed by Jacquin is most likely of a WestIndian origin. The holotype is a line drawingby Plumier, published simultaneously withthe original description, featuring an idealisedCohniella with non-resupinate flowers. Thisillustration may represent almost any memberof the genus, but the West Indian originindicates C. cebolleta. Oncidium humboldtii

is a typical Cohniella cebolleta, representa-tive of the forms found in coastal Venezuela.Oncidium ultrajectinum represents a small-flowered population growing along the north-ern rim of the Guianas.

Additional specimens examined. MEXICO. Cam-peche: Municipio Calkiní, El Remate, unos 8.5 km aloeste de Tankuché, 20º32′30″ N, 90º19′20″ W, 31 Jan2000, Carnevali et al. 6016 (CICY). Quintana Roo:Mpio. Benito Juárez, 3 km al sur del aeropuerto deCancún, 16 Feb 1999, Carnevali, Gómez & Piven 5333(CICY). Yucatán: Mpio Mérida, Dzitya, 13 Mar 2009,Carnevali 7424 (CICY).

GUATEMALA. Jalapa: El Rancho, 28 Dec 1907,Kellerman 7002, (NY).

NICARAGUA. Acoyapa: 2 km south of Acoyapa, 4Jan 1969, Atwood 1656 (NY).

COSTA RICA. San José: Santa María Dota, 5 Mar1924, Alfaro 36617 (AMES).

DOMINIQUE. Gran Savannah, Lloyd 826 (NY).

SANTA LUCIA. Middle W slope of Gros Piton,Proctor 18062 (AMES).

COLOMBIA. Magdalena: Santa Marta, Near Borda,12 Jan 1898, Smith 2353 (NY).

VENEZUELA. Anzoátegui: Woods along Río Leónby Quebrada Danta, tributary to Río Neverí, northeast ofBergantin, 500 m, 20 Feb 1945, Steyermark 61022(AMES). Bolivar: Ciudad Bolivar and vicinity, on theOrinoco, 8º10′ N, 63°31′17″W, 200 m, 27 Feb 1921, L. H.Bailey & E. Z. Bailey 1350 (NY). Delta Amacuro: Dpto.Antonio Díaz, Isla Curiapo, entre caños Obaruvaca yNaguabanoco, 8º31′ N, 61º04′ W, 0 m, Fernández 3820(NY). Río Maniamo, Vuelta Triste, 20 Feb 1911, Bond,Gillin & Brown 164 (AMES). Zulia: Kunana, 30 Feb2008, 10°03′3″N, 72°41′12″W, 300–400 m, Carnevali7222 (AMES, CICY, VEN).

GUYANA. Rupununi: Ca. 5 miles from Karasabi (4–5 hours walk) along Yurora river; 4º00′ N, 59º21′ W,300 m, 3 Jan 1982, Knapp & Mallet 2811 (NY).

Cohniella cepula (Hoffmanns.) Carnevali &G. Romero, comb. nov. Oncidium cepulaHoffmanns., Verz. Orchid. Ed. 2, 56. 1843.Type: Brazil. Río de Janeiro: ex icon. [“N.Icon. ined”] (holotype: B, destroyed; lec-totype, here designated, tracing in Herba-rium Reichenbach Nr. 15230, upper leftcorner, W). (Fig. 3I–M)

Oncidium sprucei Lindl., Fol. Orchid. 56.1855. Cohniella sprucei (Lindl.) Königer& Pongratz, Arcula 10: 280. 2000. Tricho-centrum sprucei (Lindl.) M.W. Chase & N.H. Williams, Lindleyana 16: 218. 2001.Type: Brazil. R[io] Negro and Solimões:R. Spruce 1526 (holotype: K-Lindl., iso-types: AMES, NY, photograph).


Oncidium glaziovii Cogn., Fl. Bras. 3, 6: 440.1906. Type. Brazil. Goias: A. F. M. Glaziou22179 (holotype: BR, isotype: G; photo:AMES, NY).

Oncidium ostenianum Schltr., Repert. Spec.Nov. Regni Veg. 21: 341. 1925. Cohniellaosteniana (Schltr.) Christenson, Lindleyana14: 177. 1999. Stilifolium ostenianum(Schltr.) Königer & Pongratz, Arcula 7:189. 1997. Trichocentrum ostenianum(Schltr.) M. W. Chase & N. H. Williams,Lindleyana 16: 138. 2001. Type: Paraguay.Epiphyt auf Bäumen am Río Salado, beiSan Bernardino, Rojas, Marz 1916 (holo-type: Osten 8557, herbarium Ostenianum;isotype: AMES).

Oncidium cebolleta var. purum L. C.Menezes, Schlechteriana 2: 132. 1991.Type: Brazil. Minas Gerais: Arinos,700 m, March 1991, C. García s.n.(holotype: UB-14).

Oncidium wittii Oppenheim, Orchis 10: 93.1916. Lophiaris wittii (Oppenheim) Braem,Schlechteriana 4: 21. 1993. Stilifolium wittii(Oppenheim)Königer & Pongratz, Arcula 7:190. 1997. Cohniella wittii (Oppenheim)Senghas, Orchideen (Schlechter) 173–181.2001. Trichocentrum wittii (Oppenheim) M.W. Chase & N. H. Williams, Lindleyana16: 138. 2001. Type: Bolivia. Río Iténez,Ost s.n. holotype B, destroyed; lectotype,here designated, plate 4, Orchis 10, No. 5,Tafel IV, Fig. 1. 1916; epitype, heredesignated, Bolivia. Santa Cruz: ProvinciaAndrés Ibañez, 12 km de Santa Cruz, 11Aug 1987, M. Nee 35623 (NY).

Stilifolium pongratzianum Königer, Arcula 9:261. 1999. Cohniella pongratziana(Königer) Königer, Arcula 10: 280. 2000.Trichocentrum pongratzianum (Königer) M.W. Chase & N. H. Williams, Lindleyana 16:218. 2001. Type: Peru. Departamento SanMartin: Juanjui, ca. 300 m, from M. Arias,Lima, cultived at Rosenheim, W. KönigerWK-99 (holotype: M; isotypes: USM,UNALM, Herb. Königer [not seen]).

Diagnostic features.—Cohniella cepula isextremely variable but can always be recog-nized by its relatively short and broad columnand simple, three-toothed callus. The columnfeatures a massive tabula infrastigmatica thatis deeply bilobed and cleft and the proximal

portion of these lobes is appressed to the basalplate of the callus. Since the column of thisspecies is short and bent backwards (and thusappears stalkless), the attachment point of thepetals spreads all across the column height,while in C. cebolleta and relatives the stalk ofthe column is below the base of the attachmentpoint of the petals. The callus consists of ahorizontal platform ending in three teeth, ofwhich the central is much larger. Anotherdistinctive feature of this species is the columnwings that always have red or red-purple spots(except in rare albino forms) as opposed to theentirely yellow column wings of the membersof the C. cebolleta complex north of theAmazon Basin. In C. cepula the column wingsare also almost perpendicular to or at least at avery broad angle to the main axis of thecolumn, while in C. cebolleta and relatives,the wings are essentially parallel to the mainaxis of the column.Distribution.—Argentina, Bolivia, Brazil,

Paraguay, and Peru. This species rangeswidely south of the Amazon basin.Variation range.—As with many other

Cohniella species, C. cepula is extremelyvariable and at this time how this variationcorrelates with the geographical distributionof the species is not well understood. Thus,we refer all the specimens from of the AmazonBasin and southwards to a broadly circum-scribed and variable C. cepula. The variationof this species involves basically the shape andwidth of the apical lobe of the labellum. Thisvariation follows the pattern found in most ofthe species of the genus, where the narrowestmidlobes correlate with the proportionallylongest isthmuses and narrower lateral lobes(morph A, Fig. 3 J) while the broadest label-lum midlobes correlate with a proportionallyshorter isthmus and broader lateral lobes(morph B, Fig. 3 L). Both Oncidium cepulaand O. ostenianum are referable to morph A,while Oncidium wittii and Stilifolium pongrat-zianum to morph B. The latter two speciesmight eventually deserve recognition at thespecies level, in which case the name Cohniellawittii would be available. Plants from thenorthern Amazon Basin (the type of Oncidiumsprucei among them) are also referable tomorph B and have the largest flowers of thecomplex. This complex of forms obviouslyneeds a more detailed examination.


Taxonomic commentary.—All citations andrecords of Cohniella (Oncidium) cebolletafrom south of the Amazon River belong tothis species.

Additional specimens examined. BRAZIL. Amazonas:Río Solimões, near 20 km abovemouth of Río Negro, Paranádo Xiboreninha, 15 Jun 1992, Mori & Gracie 22412 (NY).Rondônia: Mpio. de Cacoal, BR 364, Rodovia Cuiabá-PortoVelho, Km 234, ao Norte de cidade, morro da Torre daEmbratel, 11º12′ S, 61º 62′W, Cid, Lima, Guedes & Coelho4749 (NY). Ceará: Near Umari, 15 m, 7 Jul 1945, Cutler8376 (AMES). Maranhão: between Viana & Banderante,back road from Viana to Pínheiro, 3º0′ S, 45º10′ W, 17 Oct1980, Daly, Campbell, Silva, Bahia, & dos SantosD653 (NY). Mato Grosso: 12º49′ S, 51º46′ W, 3 Aug1968, Richards 6586 (AMES, NY). Minas Gerais: Mpio.Porteirinha, near city of Porteirinha, 25 May 1945, Williams&Assis 7052 (AMES).Pará: Tucuruí, Breu Branco, igapó àsmargens do rio Tocantins. 14 Aug 1983, Revilla, Miranda,Lima & Silva 8691 (NY).

PERU. Lima: Surquillo, unknown procedence, 22 Jul1997,Fernández s.n. (NY). SanMartin: nearMoyobambo,approx. 6°02′ 60″S, 76°58′00″W, 700–800 m, 25 Feb 2005,Carnevali & Ramírez 7367 (CICY); Juanjui, cultived atRosenheim, 300 m, Feb 2000, Königer WK99 (M).

BOLIVIA. Santa Cruz: Ichilo, near 30 km SE ofBuena Vista along Río Surutú, 17º36′ S, 63º36′ W, 400 m,1 Sep 1985, Solomon 14194 (NY).

ARGENTINA. Tucumán: Formosa, Jan 1918, Löse-man 2072 (AMES).

Cohniella jonesiana (Rchb. f.) Christenson,Lindleyana 14: 177. 1999. Oncidium jone-sianum Rchb.f., Gard. Chron. N.S. 20: 781.1883. Stilifolium jonesianum (Rchb. f.)Königer & Pongratz, Arcula 7: 189. 1997.Trichocentrum jonesianum (Rchb.f.) M. W.Chase & N. H. Williams, Lindleyana 16:137. 2001. Type: Paraguay. Without anyother locality or collector, ex Hort., FredHorsman & Company s.n. (holotype: W-Reichenbach 27551). (Fig. 4A–E)

Oncidium jonesianum var. flavens Reichb. f.,Gard. Chron. S. 3., 4: 234, V. 237, 1888.Type: Without locality, but presumably fromParaguay, ex Hort. B. S. Williams (holotype:W-Reichenbach 27550 [upper left figure]).

Oncidium jonesianum var. phaeanthum Sander,Reichenbachia S. I.,1: 47, t. 21, Fig. 2,1886–1891. Type: Paraguay. Without pre-cise locality, L. Saint-Leger ex Hort. SirTrevor Lawrence (holotype: presumably atW-Reichenbach, not seen).

Diagnostic features.—Cohniella jonesianais easily distinguished from all other members

of the genus by the combination of a largeflower (petals to 2 cm long, labellum to 2.2 cmwide) and a white central labellum lobe withyellow, finely fimbriate, reduced lateral lobes.Distribution.—Argentina, Bolivia, Brazil,

and Paraguay.Variation range.—Cohniella jonesiana

varies in the color of the sepals and petalsthat range from very pale green to creamcolored. They are always spotted with rela-tively conspicuous brown-red spots orblotches that are, however, never confluent.As in other variable species, pale or albino aswell as dark-colored forms are known (var.flavens and var. phaeanthum, respectively).Taxonomic commentary.—This species is

so distinct that it has never been mistakenwith any other member of the genus.

Additional specimens examined. BOLIVIA. SantaCruz: Andrés Ibáñez, between Pedro Lorenzo and Peji,Feb 1982, Vásquez 659 (Herbarium Vazquezianun).

PARAGUAY. Paraguarí: Cerro de Acahay, Mar1919, Rojas 3503 (AMES). Asunción: Cordillera deAltos, Apr 1940, Rojas 8822 (AMES).

ARGENTINA. Corrientes: Ituzaingo, Estancia SantaRita, 27º3′ S, 56º4′ W, 16 Feb 1991, Tressens, Ferrucci &Radovancich 3957 (AMES).

Cohniella longifolia (Lindley) Cetzal &Carnevali, comb. nov. Oncidium longifo-lium Lindl., Edwards’s Bot. Reg. 27: 22.1841. Type: Mexico. Without preciselocality, collected by K. T. Hartweg, exHort. Royal Horticultural Society andLoddiges (holotype: K-Lindl.).

Diagnostic features.—According to theprotologue, Cohniella longifolia is easilydistinguished from other Mexican Cohniellasby its long (“ ... often three feet long ...”),pendent leaves of apparently soft texture (“ ...long, whiplike ...”) and by its elongateisthmus subtending a relatively small centrallobe, this feature easily observable in the typematerial. The inflorescences were describedas “ ... forming dense panicles three feet longof very large and showy yellow and brownflowers.” Later, Lindley (1842), in the text inLatin that accompanied a color plate of thisspecies, stated that the leaves could be 3–4feet long.Distribution.—Known only from the type

collection.


Variation range.—Known only from the type.Taxonomic commentary.—According to the

protologue, Lindley was aware of livingplants of C. longifolia cultivated both by theRoyal Horticultural Society and the Loddigesestablishment, but a watercolor of a singleflower and his type at Kew (a panicle withfive branches) do not indicate provenance. NoMexican Cohniella resembling the type ofOncidium longifolium has ever been col-lected. The plants and flowers of the typematerial are suspiciously similar to those ofseveral populations of Cohniella cebolletafrom Perijá, Venezuela (e.g., Carnevali 7222,AMES, CICY, VEN) with which it shares thelong, often pendent leaves, the large panicu-late inflorescences and the diagnostic featuresof the flowers (the long isthmus and “ ... bird

like ...” lateral lobes of the labellum).Although it is not impossible that Mexicanplants resembling the type of Oncidiumlongifolium may be found in the future, wesuspect this may be yet another orchid speciesreported from the wrong locality.

Cohniella×marvraganii (Lückel) Christenson,Lindleyana 14: 177. 1999. Stilifolium marv-raganii Lückel, Orchidee (Hamburg) 49: 90.1998. (as “marvreganii”)—Trichocentrummarvraganii (Lückel) M. W. Chase &N. H. Williams, Lindleyana 16: 137. 2001.Type: Bolivia. Santa Cruz: M. E. Ragan s.n.(holotype: USF).

Diagnostic features.—Cohniella×marvra-ganii is intermediate between the putative

FIG. 4. Flowers of Cohniella. A–E. Cohniella jonesiana. A. Labellum, front view.. B. lateral view of the callusand 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–Efrom Paiva s.n. (photo) CICY; F–H from Kennedy s.n. (photo) AMES, CICY).


parents (C. jonesiana and C. stacyi) in severalrespects. The general outline of the labellumis similar to that of C. stacyi but it is coloredwhite or very pale yellow as in C. jonesiana;the basal lobes of the labellum are mostsimilar to those of C. jonesiana, the calli aresimilar to those of C. stacyi. Column wingsare similar to those of C. stacyi as well as theshape and disposition of the petals.Distribution.—This nothospecies is known

from NW of Santa Cruz, Bolivia.Variation range.—Only known from the

type material.Taxonomic commentary.—This nothospe-

cies was originally described as OncidiumJason Fuchs by Ragan & Sauleda (1982). Itwas formally proposed as a nothospecies byLückel (as Stilifolium “marvreganii”). Ragan& Sauleda (1982) commented “The ranges ofOncidium stacyi and O. jonesianum overlapjust north of Santa Cruz, precisely in the areawhere this natural hybrid was discovered.Vegetatively the hybrid is intermediatebetween the parents. The flowers are slightlysmaller than Oncidium stacyi with a lighteryellow labellum”.

Cohniella nuda (Bateman ex Lindl.) Chris-tenson, Lindleyana 14: 177. 1999.Oncidiumnudum Bateman ex Lindl., Edwards’s Bot.Reg. 23: t. 1994. 1837. Stilifolium nudum(Bateman ex Lindl.) Königer & Pongratz,Arcula 7: 189. 1997. Trichocentrum nudum(Bateman ex Lindl.) M. W. Chase & N. H.Williams, Lindleyana 16: 138. 2001. Type:Venezuela. Distrito Capital: Caracas, 1837,ex Hort. Bateman (holotype: K-Lindl.).(Fig. 1E–H)

Oncidium ebrachiatum Ames & C. Schweinf.Sched. Orch. 2: 75. 1923. Type: Panama.Cana and vicinity, 4 Apr 1908, R. S.Williams 975 (holotype: AMES).

Diagnostic features.—Cohniella nuda ischaracterized by its minute column wings,often almost absent. Furthermore, the mor-phology of the small callus is unmistakable. Itconsists of three low ridges flanking twodepressions, which look almost wet. The twoexternal ridges are close to, and parallel to themargins of the labellar disk. The labellum ischaracterized by its long, narrow isthmus andthe relatively small lateral lobes, which are

retrorse upon flattening. The leaves of thisspecies are usually rigidly pendent. Theinflorescences are always shorter than theleaves and very dense with flowers crowdedas compared to other Cohniella species to theextent that the flowers partially overlap.Distribution.—Panama, Colombia, and

Venezuela.Variation range.—Despite its relatively

restricted distribution, Cohniella nuda isextremely variable in the size and morphol-ogy of the flowers. Flowers range from smallwith the labellum ca. 7 mm long to largerwith the labellum ca. 13 cm long. The apicallobe of the labellum varies from transverselyelliptic to transversely oblong to rhomboid; itsapical emargination ranges from inconspicuousthrough deep and with overlapping lobes. Theplants start flowering at a very early age; somespecimens flower with leaves less than 8 cmlong while fully mature plants feature thick,usually pendent leaves of up to 50 cm long.Taxonomic commentary.—Despite its vari-

ability, Cohniella nuda is relatively easy todistinguish from related species and itssynonymy is limited. However, it seems tograde into the Panamanian taxon Cohniellastipitata near Panama City and at the PearlArchipielago (Dressler & Williams, 2003).The type of Oncidium ebrachiatum is identi-cal to populations of C. nuda from westernVenezuela and is here treated as a synonym ofthat species.

Additional specimens examined. COLOMBIA.Atlántico: Puerto Colombia, 10–60 m, 10 Apr 1906,Maxon 3843 (NY). Córdoba: Morrocoquiel, on RioSinu, 11 Mar 1918, Pennell 4693 (NY).

PANAMA. Chiriqui: halfway between Progreso andPuerto Armuelles, 16 Feb 1973, Croat 21878 (MEXU).Panamá: Torti, área cercana al Darien, 50 m, Abr 2002,Carnevali 7283 (CICY).

VENEZUELA. Miranda: Guarenas, 1846, Funck &.Schlim 481 (K-Lindl.); along Quebrada Chaguarama, 6 kmSE of Cúpira, 10º8′ N, 65º41′ W, 50–150 m, 5 Mar 1980,Liesner & González 9217 (NY). Guinand Estate(Cárdenas), Siquire Valley, 500 m, 19–24Mar 1913, Pittier5980 (NY). ZULIA: orillas del río Kunana, unos 4.5 Km. alW de La Sierra y unos 15 Km. al W de Machiques, 300–400 m., 5 Feb 2008, Carnevali 7283 (CICY).

Cohniella pendula Carnevali & Cetzal, sp.nov. Type: Mexico. Jalisco: Municipio LaHuerta, Loma Alta, 40 km. de La Huertahacia Barra de Navidad, aprox. 19°22′0″


N, 104°41′59″ W , aprox. 350–450 m,collected by G. Carnevali and G. Salazar, 3Nov 1997, flowered in cultivation 10 Mar2004, G. Carnevali & I. Ramírez 6897(holotype: CICY; isotypes: AMES, AMO,MO, NY). (Fig. 2H–O)

Species haec Cohniellae brachyphyllae (Lindley)Cetzal & Carnevali similis sed foliis pendentibus, lobocentrali minore, quam lobulis lateralibus subaequalis aliscolumnae conspicue minoribus recedit.

Epiphytic pendent herbs, sun-loving to semi-umbrophyllous, shortly creeping to caespitose;rhizome short, thin, brittle; roots 1–2 mm thick,white; pseudobulbs 12–14 mm long, 6–8 mmthick, subspherical to broadly ovoid, apically 1-leaved, red-purple tinged, totally enclosed by 3imbricate sheaths, 2.5–6.4 × 12–20 mm uponspreading, eventually deciduous; leaves terete,thickly fleshy-coriaceous, 15–36 cm long, 6–13 mm thick, dark green, usually purplespotted, when fresh abruptly constricted prox-imally, broadest at its lower 1/5, graduallyattenuated distally into a sharp apex, oftensomewhat falciform; inflorescences solitaryfrom the base of the pseudobulbs, 35–45 cmlong, a (8–)15–30-flowered raceme or paniclewith 1 or 2 short branches 4.5–5 cm long, thebranches 5–9-flowered; peduncle and rachisdark green, variably purple tinged to totallypurple; peduncle first pendent, then arching tohorizontal, 3–4 mm thick, terete, with 7–10internodes, peduncle bracts 14–21 × 3–4 mm,the basal most longest, oblanceolate, acumi-nate, tubular; bracts subtending the lateralbranches 5–7 × 2.5 mm, elliptic, acuminate;floral bracts 2–4 mm long, narrowly elliptic,acuminate; flowers resupinate, small ormediumsized for the genus, with perianth segmentswidely opening and the petals and sepalssomewhat reflexed; ovary with pedicel 12–14 mm long, of which ca. 5 mm correspond tothe ovary, this 2 mm thick; sepals basallyclawed for about 1/3 of total length of the sepal,flat or somewhat reflexed, dorsal sepal 5.5–6.5 × 1.8–2.2 mm, in general outline oblanceo-late, apically obtuse and minutely apiculate,concave in the upper half, the claw 8–11 mmwide; lateral sepals partially fused at the verybase, then free, similar the dorsal, 7.5–8.5x 2.8–3 mm; petals 9.5–10.5 × 2.3–2.6 mm, oblan-ceolate, somewhat oblique, the apex rounded to

subtruncate, somewhat reflexed in naturalposition; labellum deeply 3-lobed, 8–12 mmlong from the base to the apex of the centrallobe, 11–16 mm wide across the apices of thelateral lobes, the lateral lobes in the same planeas the central lobe and±perpendicular to it;central lobe 8–11 mmx 4–7 mm, spathulate-oblongoid to transversely elliptic or subquadratein outline, apically rounded to subquadrate,basally produced into a short isthmus, 2 ×1 mm; lateral lobes 4.2–7×4–5 mm, oblong tobroadly obovate, apically truncate-rounded tosharply obliquely truncate, the upper margin ofthe lateral lobes flat to rounded, the lowermargin straight; disc relatively large, ca. 4×5 mm, bearing a well-developed callus, ca. 2 ×1 mm, consisting of a large, elevated, ± flat,oblongoid platform, apically with three teeth,the central one laterally compressed, the lateralssmaller, divergent, somewhat pointing upward,conical, the basal portion of the callus withsharp upper margins; column 4 × 2 mm, theventral face in the same plane as the labellumlobes, oblongoid, tabula infrastigmatica subqua-drate, stigmatic surface, sub-rounded, ca. 1.7 ×1.7 mm; column wings relatively small, ca.0.5 × 1 mm, reniform; anther 1.3 × 1 mm,apical, operculate, ellipsoid; pollinarium typicalfor the genus, ca. 1.5 mm long, tegula spathu-late, 0.8×ca. 0.4 mm at the subtruncate apex;viscidium disc-like, small, pollinia 0.7–1 mmlong, yellow.

Additional specimens examined. MEXICO.Jalisco: Mpio. Cabo Corrientes, Las Juntas de Tuito,aprox. 450 m, 16 Mar 1984, Salazar & Soto 580 (AMO);Mpio. Chiquilistan, 1300 m, Feb 1973, Rosillo deVelasco 131 (AMO); Municipio Cocula, 1700 m, Mar1973, Rosillo de Velasco 140 (AMO), 141 (AMO);Mpio. Tecalitlán, 1500 m, Mar 1981, Rosillo de Velascos.n. (AMO). Nayarit: Mpio. de Ruíz, km 56.3 delcamino de la carretera México 15 (Tepic-Mazatán) aJesús María, 6.3 Km adelante del poblado de El Naranjo,22°01′ N, 104°50′ W, 320 m, 25 Jul 1998, Soto 86888(AMO); Mpio. de San Blas, Barranca N.W. east of Tepic-Navarrete, 21°14′ N, 104°32′ W, 1350 m, 28 Aug 1948,Dressler 350 (US); Mpio. Santa María del Oro, VolcanoCeboruco near Tequepexpan, 4 May 1936, Nagel & JuanS. 5115 (US).

Cohniella pendula is closely related to C.brachyphylla with which it shares the rela-tively conspicuous pseudobulbs. It also fea-tures a rigid, recurved leaf that is distinctlyswollen in the lowermost ¼ of its length andconstricted just above the junction with the


pseudobulb, and then gradually attenuatedtoward the apex. The apex of the leaf isrigidly pungent. However, the most strikingvegetative difference is the pendent habit.The leaves are rigid and, even when the plantis cultivated upright, new leaves will growpendent. The inflorescences point down atfirst, then become horizontal or, more rarely,pendent. The inflorescence itself is relativelyfew-flowered and lax when compared toplants of C. brachyphylla of similar size andvigour. The flowers are also distinctive due totheir small size. Furthermore, the central lobeof the labellum is subquadrate to transverselyelliptic, about as long as wide or not morethan 1.5 times wider that long, as opposed todistinctly transverse (at least twice as wide aslong) as in C. brachyphylla and otherCohniella taxa. These proportions make thecentral lobe of the labellum in C. pendula aswide as the labellum disc, while the apicallobes are as broad as the expanded apices ofthe lateral lobes in C. brachyphylla and mostother Cohniella species. Another distinctivefeature of Cohniella pendula is the lowermargin of the disc that is cartilaginose-callosedue to lateral extensions of the callus.Although this same condition is also presentin C. brachyphylla, it is much more pro-nounced in this new entity. Furthermore, thecolumn wings are conspicuously smaller inthis new species, being 2–3 times smallerthan they are in C. brachyphylla.Cohniella pendula seems to be restricted to

sea-facing slopes along the Pacific coast onthe Sierra Madre Occidental where it growsin “selva baja caducifolia” or “selva medianasubcaducifolia”. It occurs with epiphytes suchas Myrmecophila galeottiana (Reichb. f.)Rolfe, Encyclia adenocarpon (La Llave &Lex.) Schltr., and Tillandsia caput-medusaeE. Morren. At the type locality, it grew in alarge colony in the lower branches of a tree,in an exposed position. The plants wereheavily tinged with purple.The flowers of Cohniella pendula are

amongst the smallest in the genus and arecertainly the smallest in the C. cebolletacomplex. The sepals and petals are of a pale,dull creamish green with dark red-purplespots, while the labellum is yellow with somepale, dull orange-red on the callus and alongthe front margin of the disk and the bases of

lateral lobes. The base of the column is dullorange-red, the tabula infrastigmatica isbright yellow, the stigmatic surface is paleyellow, and the anther cap is dark red-purple.

Cohniella stacyi (Garay) Christenson,Lindleyana 14: 177. 1999. Oncidium stacyiGaray, Bot. Mus. Leafl. 23: 301. 1973.Trichocentrum stacyi (Garay) M. W. Chase&N. H.Williams, Lindleyana 16: 138. 2001.Type: Bolivia. Naranjillos, road to Cocha-bamba, 11 km south west from Santa Cruz, J.Stacy s.n. (holotype: AMES). (Fig. 4F–H)

Diagnostic features.—Cohniella stacyi iseasy to distinguish among other Cohniellaspecies due to its large flowers (> 50 mmdiam.). The petals and sepals are dark yellow-greenish heavily covered with irregular darkred-brown blotches. The labellum is brightyellow with scattered orange-brown blotches,denser toward the margins; there is always abroad orange-brown band across the isthmus.The callus is white, heavily orange-brownspotted. The morphology of the labellum of thisspecies is most distinctive due to its reducedlateral lobes which are laciniated at their distalmargin; the central lobe has fimbriate or coarselyerose-dentate margins. Its closest relative seemsto be C. jonesiana (a relationship stronglysupported by the phylogenetic analyses of Sosaet al., 2001; Williams et al. 2001b, Cetzal,2007), also from southern South America, withwhich it shares the large flowers with smalllateral lobes.Cohniella stacyi has larger flowersof different colors, and a much longer (abouttwice as long) isthmus to the central lobe.Distribution.—Cohniella stacyi is only

known from northern Bolivia.Variation range.—Cohniella stacyi is

known from only a few wild collections.However, the species is currently common inthe orchid trade since it has been reproducedmassively from seed. The progeny of thispropagation program are morphologicallyuniform but variable in color, mainly in thedepth and density of the red-brown blotching.Taxonomic commentary.—Due to its rela-

tively recent description that was widelypublicized (e.g., Fuchs, 1975; Garay, 1976)and its horticultural appeal, Cohniella stacyiis well known and has not been subject tonomenclatural wrangling.


Cohniella stipitata (Lindl.) Christenson,Lindleyana 14: 177. 1999. Oncidium stip-itatum Lindl., Bot. Voy. Sulphur 172.1843. Stilifolium stipitatum (Lindl.)Königer & Pongratz, Arcula 7: 189.1997. Trichocentrum stipitatum (Batemanex Lindl.) M. W. Chase & N. H. Williams,Lindleyana 16: 138. 2001. Trichocentrumnudum (Bateman ex Lindl.) M. W.Chase &N. H.Williams subsp. stipitatum (Lindl.)Dressler & N. H.Williams, Selbyana 24:45. 2003 [21 Oct 2003]. Type: Panama.Without any other locality, G. W. Barclay958 (holotype: BM). (Fig. 1I–L)

Oncidium lacerum Lindl., Bot. Reg. 30. Misc.38. 1844. Type: Panama. Without preciselocality, ex Hort. Loddigges (holotype: K-Lindl.).

Oncidium stipitatum Lindl. var. platyonyxRchb. f., Gard. Chron. N.S. 9: 788. 1878.Type: Panama. Without precise locality, exHort. W. Bull (holotype: W-Reichenbach27554, flowers in envelope in the upper left).

Diagnostic features.—Related to Cohniellanuda but it is easily distinguished by theproportionally longer and narrower isthmusof the labellum and the truncate, rounded tosubcordate base of the central lobe of thelabellum. The column wings in Cohniellastipitata are small but conspicuous, triangu-lar, narrowly triangular to subquadrate. In C.nuda, the column wings are almost absent,reduced to two narrow flaps of columnartissue at each side of the stigmatic surface(thus the name Oncidium ebrachiatum). Fur-thermore, the callus is strikingly differentbetween the two subspecies. In C. nuda, thecallus is low and composed as describedabove under that subspecies. In C. stipitatathe callus is composed of a single hemi-spherical callus that is somewhat laterallycompressed.Distribution.—Chiefly known from Pan-

ama where it is found in the Canal Zone andin the Provinces of Panama and Colón. Itoccurs only in tropical evergreen and tropicalsubdeciduous forests at elevations from sealevel up to 300 m. It has also been reportedfrom Costa Rica (Dressler, 1993; Mora deRetana, 1999) but the it is now apparent thatthe plant that served as the voucher for theinclusion of Oncidium nudum in the flora of

Costa Rica (USJ) was imported from Panama,according to the grower who cultivated thisspecimen (F. Pupulin, pers. comm.)Variation range.—Cohniella stipitata is a

fairly homogeneus taxon; most variationseem to be restricted to the shape of thecentral lobe of the labellum, whose basevaries from truncate to subcordate. Also, themargins of the labellum are somewhat varia-ble as to their degree of fimbriation.Taxonomic commentary.—Oncidium stipi-

tatum var. platyonix was based on a specimenwith somewhat dentate margins of the isth-mus, at each side of the callus. This feature issomewhat variable but is is present in allspecimens that we examined. Oncidium lac-erum was based upon a specimen with adeeply lacerated, crisp central lobe of thelabellum but it is otherwise identical totypical forms of the species.

Additional specimens examined. PANAMA. With-out any other locality, cultivated in the New YorkBotanical Garden Conservatory, 2 Feb 1917, Nash44955 (NY). Colón: Panama, Chilibre, 35 m , 15 Feb2005, Carnevali 7311 (CICY). Canal Area: ChagresRiver, about 3 miles above Gamboa Bridge along riverbank, 50–100 m, 09º08′32″ N, 79º40′49″ W, 07 Feb1973, Kennedy, von Chong & Steiner 2299 (MO).

Cohniella teres (Ames & C. Schweinf.)Christenson, Lindleyana 14: 177. 1999.Oncidium teres Ames & C. Schweinf.,Sched. Orch. 8: 78. 1925. Stilifolium teres(Ames & C. Schweinf.) Königer & Pongratz,Arcula 7: 190. 1997. Trichocentrum teres(Ames & C. Schweinf.) M. W. Chase & N.H. Williams, Lindleyana 16: 138. 2001.Type. Panama. Veraguas: San Francisco,1000 feet [350 m], C. W. Powell 383(holotype: AMES). (Fig. 1M–P)

Cohniella aguirrei (Königer) Königer, Arcula10: 280. 2000. Stilifolium aguirrei Königer,Arcula 9: 259. 1999. Trichocentrum aguirrei(Königer) M. W. Chase & N. H. Williams,Lindleyana 16: 218. 2001. Type: Colombia.Huila: Departamento Girardot, 1200 m, W.Köninger 95 (holotype: M; isotypes, JAUM ,K, Herb. Köninger [isotypes not seen]).

Diagnostic features.—Cohniella teres isvery similar to C. ascendens and additionalcollections from Panama and northernColombia are needed to understand the nature


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of their relationship. Cohniella teres hasshorter, narrower lateral lobes to the labellumthat are not erect nor projected forward asthey are in C. ascendens. Furthermore, theflowers of C. teres are non–resupinate, asopposed to the resupinate flowers featured byC. ascendens. The distal portion of thecolumn in C. teres is bent forward, thusplacing the stigmatic surface almost perpen-dicular to the anther, which is erect in C.ascendens. Some of the specimens of C. teresdisplay a lax covering of small papillaeclothing the lateral lobes and the disk of thelabellum.Distribution.—Nicaragua, Panama, and

Colombia. It probably also occurs in CostaRica.Variation range.—For a species known

from so few specimens, Cohniella teres dis-plays a surprising amount of variation. Everyspecimen studied by us is different from theothers. Flower size ranges from 10 mm in thetype of Cohniella teres to 18 mm in that of C.aguirrei (a cultivated plant). The type of thespecies features linear-oblong, acute toshortly acuminate lateral lobes (3–4 mmlong), while other specimens (e.g., G. Carne-vali 7027, CICY) have very small (ca. 2 mmlong) linear lobes. Some specimens, includ-ing the type, have subquadrate, sessile petalsbut the type of C. aguirrei displays narrower(albeit still sessile) petals. Although some-what different from the type material (nar-rower petals, a callus that is somewhatdifferent in its form) we have chosen to treatC. aguirrei as a synonym of C. teres becauseit shares the diagnostic features of C. teres.Many additional collections of the C. ascen-dens/teres/aguirrei-complex are required tobetter understand whether more than onetaxon is required to explain the variation inthe complex. We especially need collectionsbridging the geographical gaps presentlyfound in the distribution of the C. teres.Cohniella teres is known to us from a singlecollection from Nicaragua (featured in IconesPlantarum Tropicarum I: 1063), a series oflive plants from around Soná, on the Pacificcoast of Panama, and a single collection fromwestern Colombia, the type of C. aguirrei.The plant illustrated in Mora de Retana &Atwood (1999: 98, plate 35 b) is not thisspecies but Cohniella stipitata.

Additional specimens examined. NICARAGUA.Jinotega: Rápido Samaska, Río Bocay, ca. 100 m, Stevens16450 (MO).

PANAMA. Veraguas: Soná, 25 m, 18 Feb 2005,Carnevali 7027 (CICY).

Acknowledgments

The authors are indebted to all of theherbaria that loaned us material and/or kindlyassisted us while we visited their institutions:AMO, BIGU, M, MEXU, MO, MY, QMEX,US, NY, XAL, VEN. Norris H. Williams(FLAS), Franco Pupulin (JBL) reviewed thefirst submitted draft of the manuscript; theircomments highly improved the quality of thiscontribution. Lisa Campbell (NY), DonnaTremonte (HUH), Rodrigo Duno, Ivón M.Ramírez, José Luis Tapia (CICY), ElianaNoguera Savelli (CICY), Lizandro PerazaFlores (CICY), and Carlos Leopardi Verde(CICY) provided comments and suggestionson earlier drafts of this manuscript. GunterGerlach (M) helped us with texts in German.Victorino Paiva Castro contributed images,data and discussion of the Brazilian cohniellas.Gaspar and Katia Silvera provided material ofCohniella nuda, C. stipitata and C. teres.Óscar Moreno (CICY) provided material ofC. biorbicularis. Silvia Hernández-Aguilar andLilia Can-Itzá (CICY) helped with handlingand databasing of loans. CONACyT partiallyfunded this project via grant 49980-Q “Filo-genia molecular y morfológica, revisión siste-mática y una exploración de cuatro regionesno-codificantes del genoma del cloroplastopara estudios filogeográficos en el complejoTrichocentrum (Orchidaceae: Cymbidieae:Oncidiinae)”, awarded to the senior author.The generous support of the Orchid Society ofArizona and the Massachusetts Orchid Society(to GAR-G) is here acknowledged.

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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 andconspicuous.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 . . . . Trichocentrum2. 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 lesssimultaneous, 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 rachisnon-glaucous; plants usually epiphytic, rarely lithophytic . . . . . . . . . . . . . . . . . . . . . . . . . Lophiaris


A synopsis of Cohniella (Orchidaceae, Oncidiinae

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