VOL. 8, No. 3 - Lankesteriana - Universidad de Costa Rica

ISSN 1409-3871 

VOL. 8, No. 3 DECEMBER 2008 

Vanilla planifolia, the first Mesoamerican orchid illustrated, 

and notes on the de la Cruz-Badiano Codex 

Luis D. Gómez P. 

A new Phragmipedium from Colombia 

WesLey e. HiGGins & PauLa ViVeros 

Listado de especies de la familia Orchidaceae 

para el estado Sucre, Venezuela 

CarLos LeoParDi & Luis J. Cumana 

Vegetative anatomy of Calypsoeae (Orchidaceae) 

WiLLiam Louis stern & BarBara s. CarLsWarD 

Book reviews 

Index of taxonomic novelties published in Lankesteriana, Vol. 1—8 

I N T E R N A T I O N A L J O U R N A L O N O R C H I D O L O G Y 

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89 

93 

105 

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InternatIonal Journal on orchIdology 

Copyright © 2008 Lankester Botanical Garden, University of Costa Rica 

Effective publication date: December 31, 2008 

Layout: Jardín Botánico Lankester. 

Cover: Flower of Phragmipedium manzurii W.E. Higgins & P. Viveros. Drawing by S. Dalström. 

Printer: Litografía Ediciones Sanabria S.A. 

Printed copies: 500 

Printed in Costa Rica / Impreso en Costa Rica 

R Lankesteriana / International Journal on Orchidology 

No. 1 (2001)-- . -- San José, Costa Rica: Editorial 

Universidad de Costa Rica, 2001-v. 

ISSN-1409-3871 

1. Botánica - Publicaciones periódicas, 2. Publicaciones 

periódicas costarricenses

LANKESTERIANA 8(1): 

81-88. 2008. 

Vanilla planifolia, the first MesoaMerican orchid 

illustrated, and notes on the de la cruz-Badiano codex 

Luis D. Gómez P. 

Academia Nacional de Ciencias, Costa Rica 

Apartado 676-2050, San Pedro, Costa Rica 

gomezp@ots.ac.cr 

AbstrAct. Abstract. The Codex Barberini lat. 241 or Codex Badianus is a manuscript by Martín de la Cruz, 

entitled Libellus Medicinalibus Indorum Herbis and the first New World herbal, written in 1552. It includes 

249 medicinal plants of which only 184 are illustrated. Among the plants depicted is tlilxochitl which is the 

Nahuatl name, even to this day, for Vanilla planifolia Andrews. Thus, the illustration of that plant in the 

Codex constitutes the first illustration of a Mesoamerican orchid done for European readers and antedates 

the publication of the genus and its species by two centuries. Some new historical aspects of the Codex are 

presented. 

Key worDs: Vanilla, tlilxochitl, Codex de la Cruz-Badiano 

The situation and conditions of the indigenous 

peoples in the early colonial times in the New World 

were of hopelessness and brutality. An extraordinary 

figure appears in Bartolomé de las Casas (1484 1566), 

considered by many as the father of anti-imperialism 

and anti-racism who, in 1520, presented a defense 

of the Indians to Emperor Charles V. In 1523 he 

commenced writing his Apologética historia de las 

Indias and Historia de las Indias (the latter not to be 

published until 1875!) and, finally, in 1542 he succeeds 

in obtaining from the Emperor the promulgation of the 

Nuevas Leyes prohibiting slavery, the beginning of the 

end for the infamous systems of “encomiendas”, and 

promotes the proper education of Native Americans. In 

1543 las Casas published his demolishing denunciation 

of the Spanish exploitation and barbaric cruelty, La 

Destrucción de los Indios. 

The sadness of those events have been treated by 

many authors. Of import for these notes Keen (1999), 

Pincherle (1952), Somolinos (1964), Viesca (1992). 

Whether with or without new laws, the Spanish settlers 

and the criollos born in the Americas, maintained a 

disdain for things indigenous, and it is sad to admit 

that those sentiments of inequality still prevail in many 

parts of Latin America. But de las Casas influenced 

many others in positions of power, both secular and 

ecclesiastical. 

It was in that atmosphere that under the auspices 

of the Viceroy of New Spain, Antonio de Mendoza 

and the Bishop of Mexico Juan de Zumárraga, both 

in favor of the protection of the Indians and scarcely 

thirty years after the fall of Tenochtitlan, that the 

Colegio de Santa Cruz de Tlatelolco, was founded in 

1536 by the Franciscan friars thus becoming the first 

institution of higher learning in the continent (Borgia, 

1944). However, Viceroy Mendoza was transferred 

to Perú and Bishop Zumárraga died in 1548. The 

Colegio taught Nahuatl, Latin, Greek as well as crafts 

such as illumination, bookbinding and other European 

arts to the descendants of the Aztec nobility and 

other principal people. Among its first teachers were 

some important figures in the recording of Mexican 

ethnography, anthropology and Nahuatl literature: 

Alonso de Molina, Andrés de Olmos, Bernardino de 

Sahagún. Even fray Juan de Torquemada, himself a 

fine historiographer, unfortunately later linked to the 

Inquisition, was a professor at Tlatelolco. As a student 

was one Juan Badiano and in some capacity a native 

healer, Martin de la Cruz, both from Xochimilco. 

Mason (2004) says both were convert priests while 

others supposed them both to be medics but we know 

nothing about these two individuals. 

As the opposition and ill-will of the peninsulares and 

criollos toward the Indians and the Colegio persisted,

82 

Antonio de Mendoza´s son, Francisco, requested that 

a compilation of local healing practices and herbs be 

produced for the King of Spain, by now Phillip II, 

son of Charles V, in another effort to demonstrate that 

the Indians were knowledgeable, capable of learning 

from Europeans and very worthy of support from the 

Viceroyalty and the protection of the King. Martin de 

la Cruz was commissioned to produce the volume and 

Juan Badiano was charged with its translation from 

Nahuatl into Latin. That is the origin of the Codex. 

The administration of the Colegio was given to the 

native Franciscans in 1554, financial support from 

government sources stopped in 1606 and by the middle 

of the 17 th century the Colegio de Tlatelolco ceased to 

exist. For more on its history Somolinos (1964, 1996). 

The original text in Nahuatl has disappeared but 

the project was under the tutelage of friar Jacobo 

de Grado, superior of the convent and the Colegio, 

who did oversee the production and translation of the 

manuscript for Francisco Mendoza who had it sent to 

Phillip or possibly took it himself to the court in Madrid 

(Goodman 1990, Martínez 1994) and the Codex must 

have been in the royal library at El Escorial at least 

until the 17 th century, and forgotten. That is until it was 

found in the possession of don Diego de Cortavila y 

Sanabria, Apothecary to the King and founder of a 

medicinal plant garden on the palatial premises, as well 

as developing one of his own. The Codex was later 

obtained by Cardinal Francesco Barberini who was 

sent as Nunzio to Spain by his uncle Maffeo Barberini 

who was pope Urban VIII. The Cardinal was an avid 

bibliophile with a penchant for Americana, specially 

early texts, and he incorporated the Codex into his 

library. 

There has been much doubt as to how the codex 

passed to Cortavila. Martínez Millán (1994) supposes 

that it was given by the Infanta Juana to the Convento 

de las Descalzas Reales which she had founded. Juana, 

youngest sister to Philip II, governed Spain in his 

absence and actually met with Francisco Mendoza to 

discuss, among other things the “capitulaciones” or 

agreements concerning for the commerce of medicinal 

plants between Spain and Nueva Espana. Cortavila 

was the pharmacist for Juana and all members of the 

royal house which also included Infanta Margarita de 

la Cruz, a niece of Juana and also a nun at said convent, 

LANKESTERIANA 

LANKESTERIANA 8(3), December 2008. © Universidad de Costa Rica, 2008. 

who might have presented her apothecary with the 

Libellus as an appropiate gift to a renowed herbalist. 

The frontispiece of the Codex bears a hand written 

inscription “ex libris didaçi Cortauila”, this book 

belongs to Diego Cortavila. 

One could easily suppose that Cortavila, very much 

in the favour of the King had access to the royal library 

and its massive and mostly uncatalogued treasures and 

decided to keep it for the better practice of his métier? 

In any case, Cortavila did not promote the Codex 

among his learned acquaintances and seems not to have 

been particularly interested in the Mexican manuscript. 

Otherwise, his close friend and collaborator Bartolomé 

de Cienfuegos, pharmacist, bibliophile and fine grower 

of herbs would have mentioned it in his careful notes 

(Arévalo 1935, Rey 2004). 

How it came to the hands of Barberini has been another 

mistery. His Eminence bought many things from many 

people being a patron of the Arts and Sciences. Did he 

also buy the Codex or pushed his ecclesiastical weight 

to obtain it? The recent publication of the personal 

secretary and factotum to the Cardinal, Cassiano dal 

Pozzo´s, detailed and meticulous notes of Barberini´s 

trip to Spain (Anselmi 2006) solves the problem. The 

entry for June 26,1626 records a visit of Barberini and 

his retinue to Cortavila´s botanical garden where the 

pharmacist presented His Eminence with “un libretto 

di Semplici diversi Indiani con le sue figure, e virtu 

appropiate alla maggior parte dell´l’ indispositioni di 

corpi humani”. Barberini simply walked away with a 

precious gift. What then? 

Francesco Barberini founded the Vatican Library 

in 1679, not long before his death, but he was also an 

illustrious member of the Accademia dei Lincei, so 

respected that at the demise of its patron founder Prince 

Federico Cesi, the Cardinal was suggested as his successor. 

(Gabrieli 1880).The Accademia, as most cognoscenti in 

Europe, was enthralled with the exotic new things from 

the New World (Guerrini 2008) and was contemplating 

the publication of a materia medica prepared by the 

Neapolitan physician Nardo Antonio Recchi, based on 

Francisco Hernández de Toledo’s Rerum medicarum 

Novae Hispaniae Thesaurus, the Tesoro Messicano. One 

would think that Barberini would have promoted the 

manuscript in his possession as an equal candidate but he 

did not, or there is no evidence that he tried.

Gómez — Vanilla planifolia and the La Cruz-Badiano Codex 83 

However, diligent dal Pozzo, also a member of the 

Lincei had a quasi-facsimilar copy done by the painter 

Vincenzo Leonardi of Grimignano in the winter of 

1626-27 (Mason 2004) and that copy became known 

to many people after it was used as a pictorial source of 

Mexican flora to accompany the illustration of Mexican 

fauna and a first version of Hernández’ Rerum, in 

Johan Faber´s Animalia Mexicana descriptionibus 

scholjistique exposita, thesauri rerum medicarum 

novae hispaniae, published in Rome in 1628 with 

the beneplacito of the Lincei. That is the copy that 

eventually Pozzo’s heirs sold to pope Clement XI 

who in turn sold it to his nephew Cardinal Alessandro 

Albani who in turn sold it to George III of England 

(Byland 2000) and it is now in the Royal Library at 

Windsor Palace. It is stamped with dal Pozzo´s Coat of 

Arms. Another quandary solved. 

It has been suggested that a second copy might have 

been done for Francesco de Stelluti, with Cesi, van 

Heeck and de Filiis one of the founders and Consigliere 

Maggiore of the Accademia, but if so it is lost. Why 

would Stelluti, the author of Melissographia, the first 

anatomical description of insects under Galileo’s 

microscope, wish for a copy? Because he was charged 

by Cesi with the edition of the Tesoro Messicano, on 

which he worked in close collaboration with dal Pozzo. 

In the original plan for this edition, Barberini was the 

dedicatee but when the Cardinal was banned from 

Rome in 1644 by pope Innocent X, Barberini’s name 

was deleted. The final edition by the Lincei appeared 

in 1651. 

Incidentally, the work of Hernández, may be 

considered as a report of the first scientific expedition 

to the New World in 1571 and was translated into 

Spanish by friar Francisco Ximénez and published in 

México in 1615. 

And what of the original Codex? It rested forgotten 

and ignored in the shelves of Barberini and later, when 

the entire Barberini collection, some 60000 plus items, 

was incorporated into the Vaticana by pope Leo XIII 

in 1902 (Major 1931), it rested under the signature 

Barberini lat. 241, forgotten and ignored a few more 

decades until a student of the Barberini entries, Charles 

Upson Clark, a historian of early American medicine 

at the Smithsonian Institution, discovered it and 

announced it to the world in 1929. According to Sharp 

(2005) Clark brought the manuscript to the attention 

of Dr. William Welch at Johns Hopkins University 

and Welch´s interest prompted the first facsimilar 

edition by Emmart. León Portilla (2002) writes in 

error that the discovery dates to 1925 and was almost 

simultaneous by Clark, Giuseppe Gabrieli an erudite 

Lincei y Lynd Thorndike, a historian of science. To 

León Portilla´s commentary I must add that Gabrieli 

did not discover the original Codex but the dal Pozzo 

copy at Windsor (Gabrieli 1929), Lynd Thorndike was 

in Rome trying to enumerate and describe the contents 

of the Barberini collection dealing with the history 

of science and obviously listed the Codex among 

the items (Thorndike 1929-1930). The true date of 

discovery remains as 1929. The original de la Cruz- 

Badiano manuscript was returned to Mexico by pope 

John Paul II in 1990 (Durazo et al. 1991). 

After its discovery by Clark there have been several 

facsimilar editions of the de la Cruz Badiano Codex as 

I the Libellus should be justly known: The W. Gates 

edition of 1939 as The de la Cruz-Badiano Aztec 

Herbal (Publications 22, 23 of the Maya Society, 

Baltimore) which does not reproduce the illustrations in 

color, the Emily Walcott Emmart edition of 1940, The 

Badianus manuscript (Codex Barberini, Latin 241). 

An Aztec herbal of 1552, published by Johns Hopkins, 

the edition by the Instituto Mexicano de Seguro 

Social in 1964, followed by the Fondo de Cultura 

Económica in Mexico in 1991 with a second printing 

in 1994, reprinted in 1996 which is the one I own, all 

as Libellus Medicinalibis Indorum Herbis. Francisco 

Guerra, a physician in Mexico who has written much 

about pre-Columbian medicine, published an edition 

without pictures in 1955, being the first translation into 

Spanish. In 2000 the publishing house of Dover has 

produced a new facsimile of Gates’ edition. For more 

on other editions and versions cf Somolinos (1996). 

The Codex has been described in the editions 

mentioned above and by Robertson (1959), Fernández 

(1964) and Stolz (1964). It was written in a format 

slightly smaller than quarto (15.2 x 20.6 cm) on 

European paper, as per the watermarks from the mill 

of Basili Accinelli of Genoa whose paper was rather 

frequent in the colonial New World, the folios sewn 

into eight quires or booklets, bound in red velvet with 

a six cords spine. The inks are ferric gall for the text 

LANKESTERIANA 8(3), December 2008. © Universidad de Costa Rica, 2008.

84 

and elsewhere and for the red rectangle created on 

each page a minium gall ink (Zetina et al 2008). The 

calligraphy is cancelleresca. The text is in a Latin of 

Pliny’s style, and although sufficient it nevertheless 

conveys the feeling that Badiano may not have been 

the most advantaged pupil of the Colegio. The folios 

are numbered in the upper right corner only of the front 

or recto of the page, the verso is not. The paintings 

are a fine example of syncretism of Renaissance style 

expressed through Aztec canons. Throughout several 

pages are blank. The text is organized in thirteen 

chapters, each dealing with medical conditions thought 

by de la Cruz to be somewhat related. 

The botanical aspects of the Codex have been 

very poorly treated by Gates (1939) and very well by 

Emmart (1940), Reko (1947), Miranda and Valdés 

(1964,1996). It is the last entry of Chapter 10, that has 

caught my attention (Fig.1). 

The verso of page 56 shows the paintings of 

four plants: mecaxochitl, tlilxochitl, hueynacaztli 

and copalxochitl. The depiction of the first two is 

somewhat confusing because the artist has chosen to 

show the plants intertwined. The second root system 

corresponds to tlilxochitl. Under the drawings is a title 

followed below by the materia medica for the plants 

illustrated, which read; 

Viatoris presidium. 

“Dessicati flores mecaxochitl, tlilxochitl, hueynacaztli, 

arborum copalxochitl et atoyaxocotl cortices, folia 

srboris axocotl, albus tus, ceratum xochiocotzotl, et 

yolloxochitl ita terantu ut puluerunt; puluerasta uero 

intra notissimi et fragantissimi floris huacalxochitl 

concauitatem ponito, quo huius floris odorem 

redolentissimum capiant, spirentque. Postremo 

laudatissimum florem yolloxochitl tamdem accipito 

quem decenter excauato, cuius factae concauitati 

salutiferum puuisculum condito, uasculam collo 

suspendito” 

Which I freely translate as: 

For the traveller a protection. 

“[The] dried flowers of mecaxochitl, tlilxochitl, 

hueynacaztli [and] the barks of the trees copalxochitl 

and atoyaxocotl, leaves of the tree axocotl, white 

LANKESTERIANA 


incense, [the] wax of xochiocotzotl, and yolloxochitl 

[all together] pound to a fine powder. Once powdered 

put into the cavity of the showy and very fragrant 

flowers of huacalxochitl, as in there the scents will 

intensify and [are] trapped the odours [of that flower]. 

All that done, one of the much-praised flowers of 

yolloxochitl is taken which has a concavity in its 

center, and in this hollow the salutiferous powder [is] 

put [and that] little basket is suspended from the neck 

[of the traveller]” 

It is, rather than a remedy for specific diseases of 

travellers, a charm to keep the traveller from potential 

harm and reminds one of the amulets, talismans and 

similar trinkets worn around the neck in the Middle 

Ages to avoid contagion. The word presidium, instead 

of the correct praesidium, is one of the many examples 

of Badiano´s faulty Latin. 

The second plant is illustrated without flowers but 

with its pods (vainillas, or small pods) and corresponds 

to Vanilla planifolia still called tlilxochitl, meaning 

“black flower” by today´s Nahuatl speakers and it is 

the only orchid mentioned in the Codex. 

Mecaxochitl or “plant of the strings” is an species 

of Piper as many of them are called “cordoncillos”; 

hueynacaztli or “large ears” was thought by Emmart 

and Reko to represent Cymbopogon penduliflorum 

(Dunal)Baill., a species with pendulous flowers 

on long pedicels, characters hard to be missed by 

keen native observers. I interpret it as Enterolobium 

cyclocarpon (Jacq.) Griseb., its fruits resembling 

big ears and, furthermore, the illustration suggests 

mimosoid flowers to me. This species is known as 

guanacaste in many parts of its range. Copalxochitl 

or “flower of copal” corresponds according to Reko 

to Cyrtocarpa procera Kunth could be any of the 

plants that yield aromatic resins or latices (Stross 

1997, Langenheim 2003); axocotl or “fruit of the 

water” Reko identifies this as Spondias purpurea 

L., and atoyaxocotl or fruit of the river could refer 

to Pachira aquatica Aubl., much used throughout 

its range as an aromatizer. Xochiocotzotl, qualified 

by ceratum, is the gummy resin or turpentine from 

Liquidambar styraciflua L. or even from Pinus 

spp. (Breedlove & Laughlin 1993). Yolloxochitl 

or “flower of the heart” is Talauma mexicana (DC) 

G.Don (53v); huacalxochitl or “flower in shape of a


FiGure 1. Page 56 verso. of Libellus Medicinalibus Indorum Herbis, the de la Cruz-Badiano Codex. The second plant (left 

to right) is tlilxochitl, Vanilla planifolia Andrews, the first depiction of a Mesoamerican orchid. 


86 

huacal” (or a vessel) is that of a species of Araceae, 

e.g. Philodendron with dentate or lobed leaves as 

it is illustrated elsewhere in the Codex. (18v). For 

the synonymy of Nahuatl words see Díaz (1976) and 

Garibay (1996). 

The genus Vanilla Plum. ex Mill. was described 

in 1754 (Gard. Dict. Abr. (ed.4), 3. 1754) based on 

Charles Plumier’s name of 1702. Francisco Hernández 

de Toledo, Royal Physician, records tlilxochitl in 

his material medica De rerum medicarum Novae 

Hispaniae Thesaurus, precisely the edition by the 

Accademia dei Lincei in Rome 1628 and gives it the 

name Araco aromatico with a description and use. 

Clusius in 1602 calls it Lobus largus aromaticus in 

LANKESTERIANA 

FiGure 2. Tlilxochitl, Vanilla planifolia, in Hernández´s Rerum Medicarum…Liber Secundun. Aromata Promit, Rome 

1651, p. 38. 


Rariorum plantarum historia. Andrews, who described 

V. planifolia (Bot. Repository, for new, and rare plants 

8: pl. 538. 1808) writes for the protologue and type: 

“the finest specimen in England and the only one that 

has blossomed, is in the choice collection of the Right 

Hon. Charles Greville, at Paddington, from which our 

drawing was taken. We are informed that it is native of 

the West Indies, and was introduced to this country by 

the Marquis of Blanford”. All indicates that the Codex 

contains the earliest iconography of this orchid genus 

and species. 

In Hernandez’ Rerum medicarum… Tesoro 

Messicani, Liber Secundum, Aromata Promit, on page 

38 of the 1651 final edition, tlilxochitl is very accurately


illustrated, perhaps the second oldest depiction of the 

genus (Fig. 2). 

AcKnowLeDGements. My thanks to Mahmood Sasa who 

presented me with the 1996 Mexican facsimilar edition of 

the Libellus. Rebeca Brenes for help in the editing of the 

final version of this paper. Neal Smith and Carlos Ossenbach 

made insightful suggestions to the earlier version of this 

paper as did two anonymous reviewers. 

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Congr. On NDA of Art, Jerusalem, Mayo 2008: 1-10.


89-92. 2008. 

A New PhragmiPedium (OrchidAceAe) frOm cOlOmbiA 

Wesley e. Higgins 1—3,5 & Paula ViVeros 4 

1 Center for Tropical Plant Research and Conservation, Marie Selby Botanical Gardens 

811 South Palm Avenue, Sarasota, FL 34236-7726 U.S.A. 

2 International Scientific Committee of Lankesteriana, Universidad de Costa Rica. 

3 Research Associate, Centro de Investigación en Orquídeas de los Andes “Ángel Andreetta” 

Universidad Alfredo Pérez Guerrero, Ecuador. 

4 School of Forest Resources and Conservation, University of Florida, Gainesville, FL 32653, U.S.A. 

5 Corresponding author: whiggins@selby.org 

abstract. A new species from Colombia in Phragmipedium section Micropetalum is described: Phragmipedium 

manzurii. 

resumen. Se describe una nueva especie de Phragmipedium sección Micropetalum para Colombia: 

Phragmipedium manzuri. 

Key Words: Orchidaceae, Cypripedioideae, Phragmipedieae, Phragmipediinae, Phragmipedium, Micropetalum, 

Colombia, new species, taxonomy 

introduction. In April 2008 David Manzur sent 

photographs of a Phragmipedium for identification 

to the Orchid Identification Center. The images 

appeared distinctive from Phragmipedium schlimii 

thus necessitating examination of the specimen by 

a taxonomist. Since Manzur was unable to send a 

specimen due to governmental restrictions Paula 

Viveros traveled to Colombia to examine the plant. 

Viveros examined eight specimens and confirmed 

that they represented a new species. 

David Manzur started collecting plants in 

Antioquia, Colombia, several years ago. At first he 

thought that they all corresponded to Phragmipedium 

schlimii, but he started noticing differences in floral 

structures and leaves that led him to suspect that 

they corresponded to different varieties or species. 

Consequently he started collecting phragmipediums 

from other regions in Colombia, growing them, 

and recording their variations. He keeps his living 

collections at his farm in Caldas, Colombia; 

the herbarium specimens are kept at Herbario 

Universidad de Caldas, FAUC. 

Taxonomic treatment 

Phragmipedium manzurii W.E. Higgins & P. Viveros, 

sp. nov. 

tyPe: Colombia. Santander: ex hort. D. A. Manzur. 

June 2008, D.A. Manzur 1501 (holotype: FAUC). 

Fig. 1—3. 

Species haec Phragmipedium fischeri Braem & 

Mohr et P. schlimii (Linden & Rchb.f.) Rolfe similis, 

sed staminodio circulari breve emarginato viridiflavo 

differt, sepalis et petalisque subviridis, ellipticis; petalis 

complanatis subviolaceis, et labello albido intus violaceo 

notato, ad apicem inflato incurvato porcato differt. 

Section Micropetalum (Hallier) Garay. 

Plant caespitose, herbaceous, terrestrial. Leaves 

6, basal distichous, blades linear, coriaceous, keeled 

beneath, acuminate, margin revolute, 29.5 to 36.6 cm 

long, 2.3 to 4 cm wide. Inflorescence erect, 12 to 27 cm 

tall. Ovary pubescent 4.8 to 5.8 cm long, subtended 

by an acute green floral bract. Flowers 3 successive, 

resupinate, 6 x 5.2 cm. Sepals yellow-green, elliptical,

90 

LANKESTERIANA 

Figure 1. Phragmipedium manzurii W.E.Higgins & P.Viveros. A. Flower, lateral view. b. Flower, frontal view. c. Dissected 

flower. d. Staminode, frontal and lateral view. Drawing by Stig Dalström, based on Manzur 1501 (FAUC). 


Higgins & ViVeros — A new Phragmipedium 91 

Figure 2. Phragmipedium manzurii W.E.Higgins & P.Viveros. A. Flower, frontal view. b. Flower, lateral view. c. Dissected 

pouch. d. Dissected flower. Photos by D. A. Manzur. 


92 

Figure 3. Phragmipedium manzurii W.E.Higgins & 

P.Viveros. A. Side view of ovary. b. Front view of 

staminode. Photos by D. A. Manzur. 

somewhat concave, pubescent on reverse; dorsal sepal 

elliptical, 2 cm long, 1 to 1.5 cm wide; synsepal 1.8 to 

2 cm long, 1.4 to 1.7 cm wide. Petals white, flushed 

with pale violet, elliptical, slightly recurved, hirsute 

pubescent, 2.2 to 2.8 cm long, 1.1 to 2 cm wide. Lip 

white flushed pale violet, yellow-green at base, with 

violet markings, saccate with the apex turned up in 

front and the margin infolded, opening marked with 

short violet ridges, interior marked with violet spots, 

central yellow/violet lines on interior, tomentose 

pubescent, 1.8 to 2.4 cm long, 1 to 1.7 cm wide. 

Staminode green-yellow, round, raised center, notched 

on bottom margin, tomentose pubescent, 0.6 to 0.8 cm 

long, 0.6 to 0.8 cm wide. 

etymology: Named for David Angel Manzur, a retired 

professor of the Facultad de Agronomía, Universidad 

de Caldas, Manizales, Colombia. Manzur has been 

doing field research, observing, collecting and 

growing orchids for 20 years, with special interest in 

LANKESTERIANA 


Figure 4. Department of Santander, Colombia. 

Miltoniopsis vexillaria varieties and Phragmipedium. 

He is a member of the Risaralda Orchid Society and 

has received two American Orchid Society awards: 

Bollea coelestis David Manzur La Aldea FCC/AOS 

and Miltonia David Manzur La Aldea AM/AOS. Prior 

to this description, one species had been described 

in his name: Chodrorhyncha manzurii P.Ortiz. 

Professor Manzur has authored one orchid publication: 

Miltoniopsis vexillaria distribution and variation in 

Colombia [Orchids 74(1): 26-31. 2005]. 

additional sPecimen seen: June 2008, D.A. Manzur 

1507 (FAUC). 

distribution. Reported from the Department of 

Santander, Colombia (Fig. 4). 

acKnoWledgments. The authors thank David Manzur and 

Leon Trujillo for bringing this species to the attention of the 

Orchid Identification Center and Phillip Cribb for revision 

of the Latin diagnosis.


93-103. 2008. 

Listado de especies de La famiLia orchidaceae 

para eL estado sucre, VenezueLa 

Carlos leopardi 1,2 & luis J. Cumana 1 

1 Herbario Isidro Ramón Bermúdez Romero, Departamento de Biología, Universidad de Oriente, 

Núcleo de Sucre, Cumaná, Sucre, 6101 Apdo. 245, Venezuela. 

2 Autor para correspondencia: leopardiverde@gmail.com 

resumen. La familia Orchidaceae es uno de los grupos de Magnoliophyta de mayor distribución a nivel 

mundial, ocupan una amplia gama de nichos, desde el medio terrestre hasta las copas de los árboles (epífitas). 

Históricamente esta familia ha recibido mucha atención dada la belleza de las flores de algunos de sus miembros, 

lo que ha llevado con el tiempo a buscar el conocimiento, entre otras cosas, sobre su distribución; por lo que 

han aparecido listas de especies para varios lugares de América y el mundo. En Venezuela, exceptuando los 

tratamientos que a nivel general se ha dado a la familia, son escasos los estudios a nivel local y en particular 

en la cordillera de la costa; razón por la que en este trabajo se presenta una lista actualizada de las especies de 

Orchidaceae conocidas hasta el presente en la flora del estado Sucre. Para ello se revisaron entre los años 2006- 

2007 los herbarios VEN e IRBR, las bases de datos en línea de los herbarios K, MO, AMES, NY y F; aunado 

a ello, se realizaron exploraciones en 14 localidades del estado Sucre, distribuidas entre el extremo occidental 

de la Península de Araya hasta el extremo oriental de la Península de Paria, los alrededores de Cumaná, el 

Parque Nacional Mochima y parte del Macizo del Turimiquire. De lo anterior deriva una lista de 121 especies 

distribuidas en 63 géneros, los más importantes son: Epidendrum L. (16 spp.), Habenaria Willd. (8 spp.), 

Pleurothallis R. Br. s. l. (7 spp.), Maxillaria Ruiz & Pav. s. str. (4 spp.) y Scaphyglottis Poepp. & Endl. (4 spp.). 

Adicional a éstas, existen 12 especies, señaladas en la literatura, de las que seis cuentan con respaldo de colector 

y número de colección. Del total de las especies referidas para el estado Sucre, 15 son mencionadas en el Libro 

Rojo de la Flora Venezolana; de éstas, dos están en peligro crítico (Cattleya gaskelliana Rchb. f. y Psychopsis 

papilio (Lindl.) H.G. Jones), una en peligro, dos vulnerables y las restantes en la categoría menor riesgo. 

abstraCt. The family Orchidaceae is one of the most widespread groups of Magnoliophyta, they occur in 

a broad range of habitats, from the terrestrial habit to the treetops (epiphytes). Historically, this family had 

received a great deal of attention because of the beauty of the flowers of some species, which has lead to study 

its distribution; as a consequence, lists of species have been published for various places of America and the 

world. In Venezuela, except for treatments, of the family on a general level, there are few studies of local areas, 

and very little on the coastal cordillera; for that reason, we offer an updated list of the species of Orchidaceae 

so far know in the flora of Sucre state. To achieve this, the herbaria VEN and IRBR were studied during 2006- 

2007, and the on-line database of herbaria K, MO, AMES, NY and F were consulted. Field exploration was 

undertaken in 14 localities in Sucre state, divided between the Peninsulas of Araya and Paria, around of Cumaná 

and the Macizo del Turimiquire. As results, we present a list of 121 species of 63 genera. The most important 

are: Epidendrum L. (16 spp.), Habenaria Willd. (8 spp.), Pleurothallis R. Br. s. l. (7 spp.), Maxillaria Ruiz 

& Pav. s. str. (4 spp.) y Scaphyglottis Poepp. & Endl. (4 spp.). Additionally, 12 species were mentioned in the 

literature, of which six have data (collector and collection number). Of the total of the species know from Sucre 

state, 15 was mentioned in the Red Book of the Venenezuelan flora, two of these are in critic danger (Cattleya 

gaskelliana Rchb. f. y Psychopsis papilio (Lindl.) H.G. Jones), one in danger, two vulnerable, the rest are in the 

category minor risk. 

palabras Clave / Key words: Orchidaceae, lista de especies, species checklist, Sucre, Venezuela

94 

La familia Orchidaceae es el grupo con la mayor 

diversidad entre las Magnoliophyta, con más de 19,000 

especies (Judd et al. 1999). Esta familia se distribuye 

por todo el mundo, alcanzando su máxima diversidad 

en las regiones tropicales, especialmente en los andes 

americanos (Dressler 1981). Esta amplia distribución y 

diversidad se asocian a una alta plasticidad adaptativa 

que le ha permitido colonizar una variedad de nichos, 

desde el medio terrestre, incluyendo formas litófitas y 

rupícolas hasta especies de vida aérea (epífitas) en las 

copas de los árboles (Chase et al. 2003). 

Históricamente, por la belleza de las flores de 

algunos de sus miembros, esta familia ha recibido gran 

atención; como muestra la publicación de monografías 

sobre el grupo durante los siglos XVIII--XXI (v. gr. 

Lindey 1830, Bateman 1874, Dressler 1981; Prigdeon 

et al. 2005); así, como la aparición de sociedades 

hortícolas que pagaban y pagan importantes sumas 

por estas plantas, lo que ha permitido financiar las 

expediciones de los llamados “cazadores de orquídeas”, 

como Schomburgk, Linden, entre otros, que visitaban 

las tierras americanas en busca de estas plantas (Texera 

1991, Romero 1999). 

Este interés histórico por las orquídeas, ha tenido gran 

influencia en la elaboración de listados de orquídeas u 

orquideofloras para Brasil, Trinidad y Tobago, Chile, 

Guyana, Surinam, Guyana Francesa, Colombia, Cuba, 

Panamá, Costa Rica y Venezuela, entre otros (Cogniaux 

1965, Schultes 1967, Lehnebach 2003, Funk et al. 

2007, Huber et al. 1998); sin embargo, estos trabajos 

en su mayoría son de carácter general, siendo escasos 

los estudios para áreas específicas, como el elaborado 

en el Parque Nacional Manuel Antonio en Costa Rica 

(Populin 1998). 

Venezuela no escapa a esta realidad, a nivel general 

los trabajos de mayor envergadura que se han realizado 

son los de Dunsterville y Garay (1959, 1961, 1965, 

1966, 1972, 1976), Foldats (1969, 1970a,b,c,d) y 

Romero y Carnevali (2000); a nivel local, la zona 

que ha recibido la mayor atención ha sido Guayana, 

donde se han realizado una serie de estudios menores 

como el el reporte elaborado por Steyermark de las 

exploraciones para el Ptari-Tepuy, Roraima cerro Duida 

(Schweinfurth 1957), el informe sobre las orquídeas de 

Cerro Autana (Dunsterville 1975), la aclaratoria sobre 

el complejo de especies de Epidendrum nocturnum 

de la Guayana (Carnevali & Romero 1996) y otros 

LANKESTERIANA 


estudios mayores que llevaron a la publicación de la 

orquideoflora de la zona (Carnevali et al. 2003). 

En el nororiente de Venezuela, la familia Orchidaceae 

ha recibido poca atención, a pesar de la importancia de 

ésta en los resultados de las exploraciones botánicas 

como las realizadas por Steyermark al cerro Turimiquire 

y la región oriental adyacente (Schweinfurth 1957, 

Steyermark 1966) y a la Península de Paria (Steyermark 

& Agostini 1967), los resultados de Lárez (2003) para 

el Parque Nacional el Guácharo, entre otros. 

En el estado Sucre, el interés sobre esta familia se 

ha incrementado en los últimos años, publicándose una 

lista preliminar de especies basada sólo en información 

bibliográfica (Leopardi & Véliz, 2006) y algunas listas 

especificas locales, como las presentadas para el Parque 

Nacional Mochima (Leopardi et al. 2007, Cumana, en 

prensa). El objetivo de este trabajo es mostrar la lista 

actualizada de especies de la familia Orchidaceae para 

el estado Sucre, basada en la revisión de herbarios y en 

trabajo de campo. 

metodología 

El listado presentado es el producto de la revisión, 

entre los años 2006 y 2007, de los Herbarios Isidro 

Ramón Bermúdez Romero (IRBR) y del Herbario 

Nacional de Venezuela (VEN); así, como de las 

bases de datos disponibles en línea de los herbarios 

Royal Botanical Garden Herbarium (K) (K) (http:// 

www.kew.org/herbcat); Missouri Botanical Garden 

Herbarium (MO) (http://www.mobot.org/mobot/ 

research/herbarium.shtml); Oakes Ames Orchid 

Herbarium (AMES) (http://asaweb.huh.harvard. 

edu:8080/databases/specimen_index.html); New 

York Botanical Garden Herbarium (NY) (http:// 

sciweb.nybg.org/science2/hcol/vasc/index.asp); Field 

Museum of Natural History (F) (http://emuweb. 

fieldmuseum.org/botany/detailed.php); demás, de una 

serie de exploraciones realizadas en 14 localidades 

del estado Sucre, distribuidas de la siguiente forma: 

Península de Araya y adyacencias (Campoma, Cariaco, 

Guacarapo, Guayacán), Península de Paria (Río El 

Pilar, Yaguaraparo), alrededores de Cumaná (Parque 

Nacional Mochima, El Tacal, San Juan de Macarapana, 

Río Brito, Humo Negro, Cerro Imposible), Macizo del 

Turimiquire (Catuaro, Las Pideras de Cocollar). En el 

apéndice 1 se muestra una selección de los ejemplares 

revisados y/o colectados.

leopardi & Cumana — Listado de Orchidaceae para Sucre, Venezuela 95 

La colección de las muestras siguió el protocolo 

tradicional para el procesamiento de material de 

herbario (Lindorf et al. 1999), tomando de uno a tres 

individuos (dependiendo de la abundancia); así mismo, 

en caso de que el material estuviese en fenofase 

reproductiva, porciones de las inflorescencias fueron 

conservadas en una mezcla de formol, ácido acético 

y etanol (FAA) en proporción 1:1:3; en caso de que 

el material estuviese en fenofase vegetativa se tomó 

una parte y se mantuvo en condiciones de vivero hasta 

alcanzar la floración, colectándose luego material para 

herborizar y conservándose algunas flores en FAA. El 

material colectado está depositado en IRBR. 

El listado de especies presentado se ajusta en forma 

general al sistema propuesto por Chase et al. (2003) 

y sigue, en forma amplia, la sinonimia propuesta en 

tropicos.org (http://www.tropicos.org) y por Govaerts 

(2005). Algunos casos especiales sensu stricto son: 

Cohniella Pfitzer y Lophiaris Raf. Los Maxillarinae 

siguen el tratamiento sugerido por Blanco et al. (2007). 

resultados y discusión 

La familia Orchidaceae, en el estado Sucre, está 

representada por 121 especies, de 63 géneros, los más 

numerosos son: Epidendrum (16 spp.), Habenaria (8 

spp.), Pleurothallis s. l. (7 spp.), Maxillaria s. str. (4 

spp.) y Scaphyglottis (4 spp.) (tabla 1). Al considerar 

la riqueza por zona, se encuentra que la más pobre es 

la Península de Araya y áreas adyacentes (11 spp.); 

mientras que, la zona más rica es la Península de Paria 

(83 spp.), seguida del macizo del Turimiquire (36 spp.) 

y los Alrededores de Cumaná (33 spp.). 

Estos resultados se relacionan con lo que ha 

sido estimado para el país, siendo los géneros más 

numerosos a nivel nacional Epidendrum (162 spp.), 

Pleurothallis s.l. (125 spp.) y Maxillaria s. l. (100 

spp.) (Huber et al. 1998, Carnevali et al. 2007). 

La coincidencia en la dominancia genérica puede 

asociarse a la diversidad de ambientes presentes en el 

estado Sucre: arbustales xerófilos, sabanas, manglares, 

bosques tropófilos, bosques húmedos, entre otros. 

También es notable que estos géneros alcancen su 

máxima diversidad en el estado en las vertientes de la 

Península de Paria y del Macizo del Turimiquire (tabla 

1), que son las zonas más frescas y húmedas, lo que 

coincide con el hecho de que esta familia tiende a ser 

más diversa en zonas con características similares a la 

andina, que es donde alcanza su máxima diversidad 

(Dressler 1981, Carnevali et al. 2007). 

La pobreza de especies en la Península de Araya se 

debe, básicamente, a las escasas precipitaciones (son 

inferiores a 260 mm anuales) y a la acción del viento, 

que en las zonas de menores elevaciones, hacia la 

región occidental de la península, alcanza velocidad 

suficiente como para impedir el establecimiento no 

sólo de los representantes de esta familia, sino de casi 

todos los grupos vegetales (Cumana 1999) 

Tres especies son endémicas para el estado 

Sucre (Acianthera pariaënsis (Carnevali & G. A. 

Romero) Carnevali & G. A. Romero, Epidendrum 

dunstervilleorum Foldats y Masdevallia irapana H. 

R. Sweet). Por otro lado, en el Libro Rojo de la Flora 

Venezolana (Llamozas et al. 2003), se mencionan 

15 de las especies reportadas para el estado Sucre, 

distribuidas de la siguiente forma: Cattleya gaskelliana 

Rchb. f. y Psychopsis papilio (Lindl.) H. G. Jones 

son señaladas como especies en peligro crítico de 

extinción, Chysis aurea Lindl. como especie en peligro 

y Encyclia cordigera (Kunth) Dressler y Huntleya 

lucida (Rolfe) Rolfe como especies vulnerables, las 

siete especies restantes aparecen bajo la categoría de 

menor riesgo (tabla 1). 

Adicionalmente, en los trabajos de Steyermark 

(1966), Steyermark y Agostini (1967) y Foldats (1969, 

1970b,c,d), son referidas seis especies con colector 

y número de colección: Cycnoches lodigessii Lindl 

(Moritz, 611); Dichaea pendula (Aubl.) Cogn. (J. 

Steyermark, 91932), Epidendrum cardioglossum 

Rchb. f. (J. Steyermark y cols. 62596), Gomphichis 

costaricensis (Schltr.) Ames (J. Steyermark y cols, 

62517), Liparis neuroglossa Rchb. f. (J. Steyermark, 

62645), Oncidium bicolor Lindl. (Dunsterville 378); 

sin embargo, estas exsiccata no fueron encontradas en 

los herbarios revisados. 

Finalmente, otras seis especies son mencionadas, 

principalmente por Foldats (1970a, b, c, d), pero 

sin ningún soporte localizable, a saber: Aspasia 

variegata Lindl., Brassia caudata Lindl., Epidendrum 

purpurascens Focke, Lockhartia acuta (Lindl.) Rchb. 

f.), L. oblongicallosa Carnevali & G.A. Romero y 

Pleurothallis testaefolia (Sw.) Lindl. Por otro lado, 

existe una serie de especies que han sido encontradas 

en la zona limítrofe con el estado Monagas (del lado 

de Monagas) que, potencialmente, también podrían 


96 

LANKESTERIANA 

tabla 1. Lista de especies de la familia Orchidaceae presentes en el estado Sucre, Venezuela. 

especie a a p c t dr b L c 

Acianthera lanceana (Lodd. ex Lindl.) Pridgeon & M. W. Chase - + - + · MR/ca 

Acianthera pariaënsis (Carnevali & G. A. Romero) Carnevali & G. A. Romero - + - - ES · 

Beloglottis costaricensis (Rchb. f.) Schltr. - + - - · · 

Brassavola cucullata (L.) R. Br. + + - - · · 

Campylocentrum micranthum (Lindl.) Rolfe - + + + · · 

Campylocentrum schneeanum Foldats - + - - EC · 

Catasetum macrocarpum Rich. ex Kunth - + + - · · 

Catasetum planiceps Lindl. - + + - · · 

Cattleya gaskelliana Rchb. f. - - - + EO CR 

Caularthron bicornutum (Hook.) Raf. + + - - · · 

Caularthron bilamellatum (Rchb. f.) R. E. Schult. + + + - · · 

Chysis aurea Lindl. - + - - · EN 

Cohniella cebolleta (Jacq.) Christenson + + + - · MR/ca 

Cyclopogon elatus (Sw.) Schltr. - - - + · · 

Cyrtopodium willmorei Knowles & Westc. + - + + · · 

Dichaea hystricina Rchb. f. - - - - · · 

Dichaea muricata (Sw.) Lindl. - - - - · · 

Dimerandra emarginata (G. Mey.) Hoehne + + + - · · 

Elleanthus arpophyllostachys Rchb. f. - + - - EC · 

Elleanthus furfuraceus (Lindl.) Rchb.f. - - - + · · 

Encyclia cordigera (Kunth) Dressler + - + + · VU 

Encyclia isochila (Rchb. f.) Dodson - + - - · · 

Epidendrum anceps Jacq. - + - + · · 

Epidendrum cernuum Kunth - - - + · · 

Epidendrum ciliare var. squamatum Schnee + - + - RV · 

Epidendrum coronatum Ruiz & Pav. - - - + · MR/ca 

Epidendrum dunstervilleorum Foldats - + - - ES · 

Epidendrum ferrugineum Ruiz & Pav. - + - + · · 

Epidendrum miserrimum Rchb. f. - + - - · · 

Epidendrum pseudoramosum Schltr. - + - - · · 

Epidendrum purum Lindl. - + - - · · 

Epidendrum ramosum Jacq. - + - - · · 

Epidendrum rigidum Jacq. - + + + · · 

Epidendrum rostratum Garay & Dunst. - + - - · · 

Epidendrum secundum Jacq. - + - + · · 

Epidendrum strobiliferum Rchb. f. - + - - · · 

Epidendrum carpophorum Barb. Rodr. - + - + · · 

Epidendrum vincentinum Lindl. - + - - · MR/ca 

Eulophia alta (L.) Fawc. & Rendle - + - - · · 

Gongora quinquenervis Rchb. f. - - + + · · 

Govenia utriculata (Sw.) Lindl. - - + + · · 

Habenaria alata Hook. - - + - · MR/ca 




Habenaria bractescens Lindl. - - + - · · 

Habenaria entomantha (Lex.) Lindl. · · · · · · 

Habenaria heptadactyla Rchb. f. - - + - · · 

Habenaria monorrhiza (Sw.) Rchb. f. - + + - · MR/ca 

Habenaria obtusa Lindl. - - + - · · 

Habenaria petalodes Lindl. - + - - · · 

Habenaria trifida Kunth - - + - · · 

Heterotaxis discolor (G. Lodd.) Ojeda & Carnevali - + - - · · 

Houlletia odoratissima Linden ex Lindl. - + - - · · 

Huntleya lucida (Rolfe) Rolfe - + - - · VU 

Isochilus linearis (Jacq.) R. Br. - + - + · · 

Jacquiniella globosa (Jacq.) Schltr. - + - + · · 

Jacquiniella teretifolia (Sw.) Britton & Wilson - - - + · · 

Laelia undulata (Lindl.) L. O. Williams + - + - · · 

Leochilus labiatus (Sw.) Kuntze - - - + · · 

Lepanthes pariaënsis Foldats - + - - RV · 

Lepanthes wageneri Rchb. f. - + - - · · 

Lepanthopsis astrophora (Rchb. f.) Garay - + - - · · 

Liparis nervosa subsp. nervosa Thunb. - - + - · · 

Lophiaris lanceana (Lindl.) Braem + + + - · MR/ca 

Lophiaris lurida (Lindl.) Braem + + + + · · 

Macroclinium wullschlaegelianum (H. Focke) Dodson - + - - · · 

Masdevallia irapana H. R. Sweet - + - - ES · 

Masdevallia kyponantha H. R. Sweet - + - - · · 

Masdevallia lansbergii Rchb. f. - + - - · · 

Maxillaria brachybulbon Schltr. - + - - · · 

Maxillaria melina Lindl. - + - - · · 

Maxillaria nasuta Rchb. f. - + - - · · 

Maxillaria porrecta Lindl. - + - + · · 

Microchilus paleaceus (Schltr.) Omerod. - + - + · · 

Microchilus venezuelanus (Garay & Dunst.) Ormerod. - + - - RV · 

Myoxanthus exasperatus (Lindl.) Luer - - - + · · 

Notylia rhombilabia C. Schweinf. · · · · · · 

Octomeria apiculata (Lindl.) Kuntze - + - - · · 

Oeceoclades maculata (Lindl.) Lindl. - + + + · · 

Oncidium klotzschianum Rchb. f. - + - + · · 

Ornithocephalus bonplandii Rchb. f. - - + - · · 

Ornithidium patulum (C. Schweinf.) M. A. Blanco & Ojeda - - - + · · 

Otoglossum globuliferum (Kunth) L. O.Williams & M. W. Chase - + - - · · 

Otoglossum scansor (Rchb. f.) Carnevali & I. Ramírez - + - - · · 

Pachyphyllum pastii Rchb. f. - - - + · · 

Platystele oxyglossa (Schltr.) Garay - + - - · · 

Platystele stenostachya (Rchb. f.) Garay - + - - · · 

Pleurothallis discoidea Lindl. - + - - · · 


98 

tabla 1. (continúa) 

LANKESTERIANA 


Pleurothallis macrophylla Kunth - + - - · · 

Pleurothallis pruinosa Lindl. - - + - · · 

Pleurothallis revoluta (Ruíz & Pav.) Garay - + - - · · 

Pleurothallis ruscifolia (Jacq.) R. Br. - + - - · · 

Pleurothallis aff. secunda Poepp. & Endl. - - + - · · 

Polystachya foliosa (Hook.) Rchb. f. - + - + · · 

Prosthechea cochleata (L.) W. E. Higgins - - - + · MR/ca 

Prosthechea aemula (Lindl.) W. E. Higgins - + - - · · 

Prosthechea livida (Lindl.) W. E. Higgins - - + + · · 

Psilochilus modestus Barb. Rodr. - + - - · · 

Psychopsis papilio (Lindl.) H. G. Jones - - + - · CR 

Psygmorchis glossomystax (Rchb. f.) Dodson & Dressler - + - - · · 

Psygmorchis pusilla (L.) Dodson & Dressler - - + - · · 

Rhetinantha notylioglossa (Rchb. f.) M. A. Blanco - + - - · · 

Rodriguezia lanceolata Ruiz & Pav. - + - + · MR/ca 

Sauvetrea aff. alpestris (Lindl.) Szlach. - - - + · · 

Scaphosepalum breve (Rchb.f.) Rolfe - + - - · · 

Scaphyglottis grandiflora Ames & C. Schweinf. - + - - · · 

Scaphyglottis modesta (Rchb. f.) Schltr. - + - - · · 

Scaphyglottis prolifera Cogn. - - - + · · 

Scaphyglottis reflexa Lindl. - + - - · · 

Sobralia cf. ciliata (Presl) C. Schweinf. ex Foldats - + - + · · 

Sobralia aff. violacea Linden ex Lindl. - - + - · · 

Stelis argentata Lindl. - - + - · · 

Stelis papaquerensis Rchb. f. - + - - · · 

Stelis tristyla Lindl. - + - - · · 

Stenia pallida Lindl. - + - - · · 

Trichosalpinx ciliaris (Lindl.) Luer - + - - · · 

Trichosalpinx intricata (Lindl.) Luer - + - - · · 

Triphora gentianoides (Sw.) Nutt. ex Ames & Schltr. - + - - · · 

Trizeuxis falcata Lindl. - + + + · · 

Vanilla pompona Schiede - + + - · MR/pm 

Wullschlaegelia calcarata Benth. - + - - · · 

Xylobium pallidiflorum (Hook.) G. Nicholson - + - - · · 

total de especies: 11 83 33 36 

aA: Península de Araya y adyacencias; P: Península de Paria; C: alrededores de Cumaná; T: Macizo Montañoso del 

Turimiquire (sólo la sección que está en el estado Sucre). 

bDR: Especies de distribución restringida; ES: endémica estado Sucre; EC: endémica Cordillera de la Costa; EO: endémica 

Cordillera de la Costa, región oriental; RV: restringida a Venezuela. 

cL: estado de conservación según Llamozas et al. (2003); MR/ca: menor riesgo, casi amenazada; CR: peligro crítico; EN: 

en peligro; VU: vulnerable; MR/pm: menor riesgo, preocupación menor; ·: dato no disponible. 



estar en el estado Sucre, entre las que cabe destacar: 

Acianthera rubroviridis (Lindl.) Pridgeon & M.W. 

Chase, Epidendrum subpurum Rchb. f., Ionopsis 

utricularioides (Sw.) Lindl., Mormodes buccinator 

Lindl., entre otras. 

agradeCimientos. Los autores desean expresar su 

agradecimiento al Consejo de Investigación de la 

Universidad de Oriente (CI-2-010101-1224/05 y CI-2-0101- 

01-1374/07) y a PROVITA (2007-08) por el financiamiento 

otorgado. A la Dra Leyda Rodríguez (curadora de VEN) por 

el permitir la revisión de la colección de Orchidaceae allí 

depositada y por el préstamo de algunos de los ejemplares. 

A Carmen Malavé, Freddy Subero y Christian Marchán 

por toda la colaboración prestada durante el trabajo de 

campo. A José Imery y a los dos revisores anónimos por sus 

sugerencias y críticas al manuscrito. Al personal del IRBR 

por su colaboración durante la ejecución de este trabajo. 

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apéndiCe 1. seleCCión del material examinado 

VENEZUELA. sucre. Acianthera, A. lanceana: 

municipio Montes, Las Piedras de Cocollar, 31-03-2007, C. 

Leopardi et al. 103 (IRBR). A. pariaënsis: municipio 

Valdez, Península de Paria, Cerro Patao, 19-07-1962, J. 

Steyermark y G. Agostini 91087 (VEN). Beloglottis, B. 

costaricensis: municipio Cajigal, Yaguaraparo, 18-02-2007, 

C. Leopardi et al. (IRBR). Brassavola, B. cucullata: 

municipio Cruz Salmerón Acosta, Guayacán, Península de 

Araya, 28-01-2006, C. Leopardi y J. Véliz 01 (IRBR). 

Campylocentrum, C. micranthum: municipio Cajigal, 

Yaguaraparo, 19-02-2007- C. Leopardi y F. Subero 99 

(IRBR). C. schneeanum: municipio Mariño, Península de 

Paria, este de Cerro Humo, 24-02-1980, J. Steyermark y R. 

Liesner 121705 (VEN). Catasetum, C. macrocarpum: 

municipio Cajigal, Yaguaraparo, 18-02-2007, C. Leopardi 

et al. 76 (IRBR). C. planiceps: municipio Montes, Humo 

Negro, 12-10-2006, C. Leopardi y J. Véliz 49 (IRBR). 

Cattleya, C. gaskelliana: Entre Cerro El Diablo y La 

Trinidad, 1945, J. Steyermark 62765 (VEN). Caularthron, 

C. bicornutum: municipio Valdez, Península de Paria, 

noreste de Puerto Hierro, 05-09-1984, Milliken et al. 132 

(VEN). C. bilamellatum: municipio Cajigal, Yaguaraparo, 

17-02-2007, C. Leopardi et al. 68 (IRBR). Chysis, C. aurea: 

municipio Mariño, Península de Paria, Río Grande, 02-01- 

1981, J. Steyermark S/N (VEN). Cohniella, C. cebolleta: 

municipio Sucre, Vega grande, Parque Nacional Mochima, 

22-01-1994, N. Ortega y N. Medina 70 (IRBR). Cyclopogon, 

C. elatus: municipio Montes, Las Piedras de Cocollar, 02- 

04-2007, C. Leopardi et al. 129 (IRBR). Cyrtopodium: C. 

willmorei: Sabanas entre Sucre y Monagas, 28-04-2007, C. 

Leopardi y J. Véliz 140 (IRBR). Dichaea, D. hystricina: 

Sucre sin localidad exacta, S/F, Sin colector S/N. (IRBR). 

D. latifolia var longa: Sucre sin localidad exacta, S/F, Sin 

colector S/N. (IRBR). Dimerandra, D. emarginata: 

municipio Benítez, Río El Pilar, 06-05-2007, C. Leopardi y 

C. Malavé 141 (IRBR). Elleanthus, E. arpophyllostachys: 

municipio Mariño, Península de Paria, entre Los Pocitos y 

La Roma, 11-02-1966, J. Steyermark y M. Rabe 96345 

(VEN). E. furfuraceus: municipio Montes, Las Piedras de 

Cocollar, 02-04-2007, C. Leopardi et al. 135 (IRBR). 

Encyclia, E. cordigera: municipio Cruz Salmerón Acosta, 

Península de Araya, Guarapo-Oturo, 15-12-1984, M. y W. 

Lampe 28 (IRBR). E. isochila: municipio Mariño, Península 

de Paria, Las Melenas, 01-12-1979, J. Steyermark y R. 

Liesner 120828 (VEN). Epidendrum, E. anceps: municipio 

Montes, Las Piedras de Cocollar, 01-04-2007, C. Leopardi 

et al. 121 (IRBR). E. carpophorum: municipio Montes, 

Macizo del Turimiquire, (25-31)-03-1995, L. Cumana et al. 

6202 (IRBR). E. cernuum: municipio Montes, Cara norte 

del Cerro Turimiquire, cerca de la cumbre, (06-08)-05- 

1945, J. Steyermark 62596 (VEN). E. ciliare var. 

Squamatum: municipio Cruz Salmerón Acosta, Laguna de 

Chacopata, , 18-12-2006, C. Leopardi y J. Véliz 57 (IRBR). 

E. coronatum: municipio Benítez, Río El Pilar, Hacienda El 

Otro Lado, 06-05-2007, C. Leopardi y C. Malavé 151 

(IRBR). E. dunstervilleorum: municipio Mariño, Península 

de Paria, Cerro Humo, 03-03-1966, J. Steyermark 94972 

(VEN). E. ferrugineum: municipio Mariño, Península de 

Paria, cerca de Río Seco, 11-08-1967, V. Carreño S/N 

(VEN). E. miserrimum: municipio Mariño, Península de 

Paria, Cerro Humo, entre Los Pocitos y la Roma, 11-08- 

1966, J. Steyermark y M. Rabe 96347 (VEN). E. 

pseudoramosum: municipio Mariño, Cerro Humo , S/F, J. 

Steyermark y M. Rabe 96325 (VEN). E. purum: municipio 

Mariño, Península de Paria, Manacal, 20-02-1967, V. 

Carreño S/N (VEN). E. ramosum, Sucre, S/F, J. Steyermark 

et al. 121724 (MO, no visto). E. rigidum: municipio Cajigal, 

Río Grande, Yaguaraparo, 19-02-2007, C. Leopardi y F. 

Subero 92 (IRBR). E. rostratum: municipio Cajigal, 

Península de Paria, 21-02-1980, J. Steyermark et al. 121468 

(VEN). E. secundum: municipio Montes, Río Totucual, El 

Guamal, macizo montañoso del Turimiquire, (03-06)-08- 

1993, L. Cumana et al. 5011(IRBR). E. strobiliferum: 

municipio Benítez, Serranía La Paloma, 19-02-1980, J. 

Steyermark et al. 121379 (VEN). E. vincentinum: municipio 

Mariño, Península de Paria, Cerro de Humo, 01-03-1966, J. 

Steyermark 94828 (VEN). Eulophia, E. alta: municipio 

Arismendi, Península de Paria, entre Río Grande y 

Mejillones, 02-08-1961, L. Aristeguieta y G. Agostini 4792 

(VEN). Gongora, G. quinquenervis: municipio Sucre, 

Alrededores de Los Altos de Santa Fe, feb-1994, D. 

Hamenstein 1112/92 (VEN). Govenia, G. utriculata: 

municipio Montes, Las Piedras de Cocollar, 03-04-2007, C. 

Leopardi et al. 136 (IRBR). Habenaria, H. alata: municipio 

Sucre, San Juan de Macarapana, 06-09-2006, C. Leopardi y 

J. Véliz 46 (IRBR). H. bractescens: municipio Montes, 

Cerro Grande, La Fragua, 28-09-1974, W. Lampe 18 

(IRBR). H. entomantha: Sucre, sin localidad más exacta, 

S/C S/N (IRBR). H. heptadactyla: municipio Sucre, Bella 

Vista, Parque Nacional Mochima, 24-08-1990, L. Cumana 

y J. Véliz 4418 (IRBR). H. monorrhiza: municipio Sucre, 

La Guayuta (Altos de Santa Fe), 27-12-1990, L. Cumana y 

P. Cabeza 4581 (IRBR). H. obtusa: municipio Sucre, Bella 

Vista, Parque Nacional Mochima, 24-08-1990, L. Cumana 

y J. Véliz 4472 (IRBR). H. petalodes: municipio Mariño, 

Península de Paria, cumbre Las Estrellas, 30-11-1979, J. 

Steyermark y R. Liesner 120715 (VEN). H. trifida: 

municipio Sucre, Cerro imposible, 07-08-2000, L. Cumana 

et al. 6697 (IRBR). Heterotaxis, H. discolor: municipio 

Mariño, Mundo Nuevo, oeste de Cerro Humo, 07-08-1966, 


102 

J. Steyermark y M. Rabe 96146 (VEN). Houlletia, H. 

odoratissima: municipio Mariño, Península de Paria, Cerro 

Humo, 04-03-1966, J. Steyermark 95020 (VEN). Huntleya, 

H. lucida: municipio Valdez, Cerro Patao, norte de Puerto 

Hierro, S/F, J. Steyermark y G. Agostini 91387 (VEN). 

Isochilus, I. linearis: municipio Montes, Las Piedras de 

Cocollar, 02-04-2007, C. Leopardi et al. 134 (IRBR). 

Jacquiniella, J. globosa: municipio Montes, Cocollar, 04- 

02-2007, L. Cumana, et al. 6978 (IRBR). J. teretifolia: 

municipio Montes, Cerro Turimiquire, 1945, J. Steyermark 

62663 (VEN). Laelia, L. undulata: municipio Montes, 

Adyacencias de Cumanacoa, feb-2007, R. Coronado S/N 

(IRBR). Leochilus, L. labiatus: municipio Montes, Río 

Arriba, Las Piedras de Cocollar, 01-04-2007, C. Leopardi et 

al. 108 (IRBR). Lepanthes, L. pariaënsis: municipio 

Mariño, Península de Paria, cerca de La Tacarigua, 23-02- 

1979, J. Steyermark et al. 121640 (VEN). Lepanthes, L. 

pectinata: municipio Mariño, Península de Paria: Cerro de 

Humo, 24-02-1980, J. Steyermark y M. Rabe 96349 (VEN). 

L. wageneri: municipio Mariño, Península de Paria, Cerro 

Humo, mar-1966, J. Steyermark 94819 (VEN). 

Lepanthopsis, L. astrophora: municipio Mariño, Península 

de Paria, Las Melenas, 01-12-1979, J. Steyermark y R. 

Liesner 120926 (VEN). Liparis, L. nervosa subsp. nervosa: 

municipio Sucre, Altos de Santa Fe, 23-07-1990, L. Cumana 

4133 (IRBR). Lophiaris, L. lanceana: municipio Benítez, 

Entre Guaraúnos y Los Ajíes, J. Steyermark et al. 121294 

(VEN). L. lurida: municipio Mariño, La Montana, cerca 

Irapa, 23-05-1968, Carreño S/N (VEN). Macroclinium, M. 

wullschlaegelianum: municipio Mariño, Península de Paria, 

cercanías de Manacal, S/F, J. Steyermark y R. Liesner 

120628 (VEN). Masdevallia, M. irapana: Sucre, Venezuela, 

05-03-1966. J. Steyermark 95079 (AMES, no visto). M. 

kyponantha: municipio Mariño, Península de Paria, Cerro 

de Humo, 05-03-1966, J. Steyermark 95079 (VEN). M. 

lansbergii: entre municipios Mariño y Arismendi, Península 

de Paria, descendiendo a Las Melenas, feb-1980, J. 

Steyermark et al. 121757 (VEN). Maxillaria, M. 

brachybulbon: municipio Mariño, Península de Paria, 

Cerro de Río Arriba, 09-08-1966, J. Steyermark y M. Rabe 

96206 (VEN). M. melina: municipio Cajigal, Península de 

Paria, 21-02-1980, J. Steyermark et al. 121475 (VEN). M. 

nasuta: municipio Valdez, Montaña de Falcón (camino a 

Los Chorros), Cristobal Colón, S/F, F. Fernández 307 

(VEN). M. porrecta: municipio Mariño, Península de Paria, 

Cerro Humo, 04-04-1966, J. Steyermark 95029-A (VEN). 

Microchilus, M. paleaceus: municipio Montes, Las Piedras 

de Cocollar, 02-04-2007, C. Leopardi et al. 133 (IRBR). M. 

venezuelanus: municipio Mariño, Península de Paria, 

carretera a El Paujil, 20-02-1980, J. Steyermark et al. 

342151-VEN. (VEN). Myoxanthus, M. exasperatus: 

municipio Montes, Turimiquire, (25-31)-03-1995, L. 

LANKESTERIANA 


Cumana et al. 6151 (IRBR). Notylia, N. rhombilabia: 

Sucre, abr-1977, Carreño S/N (VEN). Octomeria, O. 

apiculata: municipio Valdez, Península de Paria, Cerro 

Patao, 19-07-1962, J. Steyermark y G. Agostini 91143 

(VEN). Oeceoclades, O. maculata: municipio Benítez, Río 

El Pilar, Hacienda El Otro Lado, 06-05-2007, C. Leopardi y 

C. Malavé 152 (IRBR). Oncidium, O. klotzschianum: 

municipio Cajigal, Península de Paria, 21-02-1980, J. 

Steyermark et al. 121474 (VEN). Ornithocephalus, O. 

bonplandii: municipio Sucre, Altos de Santa Fé, 03-04- 

2007, Leopardi 63 (IRBR). Ornitidium, O. patulum: Entre 

la parte superior de Río Colorado y Cerro Turimiquire, S/F, 

J. Steyermark 62650 (VEN). Otoglossum, O. globuliferum: 

municipio Mariño, camino a Los Pocitos de Santa Isabel, 

Cerro Humo, 12-07-1972, G. Morillo 2591(VEN). 

Otoglossum, O. scansor: municipios Mariño y Arismendi, 

Península de Paria, 22-02-1980, J. Steyermark et al. 121542 

(VEN). Pachyphyllum, P. pastii: municipio Montes, Cerro 

Turimiquire, 06-05-1945, J. Steyermark 62568 (VEN). 

Platystele, P. oxyglossa: municipio Mariño, Península de 

Paria, Cerro Humo, 23-02-1980, J. Steyermark et al. 

121692 (VEN). Platystele stenostachya: municipio Mariño, 

Península de Paria, Cerro de Humo, oeste de Santa Isabel, 

mar-1966, J. Steyermark 94831-A (VEN). Pleurothallis, P. 

discoidea: municipio Mariño, Península de Paria, oeste de 

Cerro Humo, 07-08-1966, J. Steyermark y M. Rabe 96419 

(VEN). P. macrophylla: municipio Mariño, Los Pocitos de 

Santa Isabel, Cerro de Humo, 12-07-1972, Morillo 2610 

(VEN). P. pruinosa: municipio Sucre, El Guayabito, (20- 

22)-11-1981, G. Davidse y A. González 19161 (VEN). P. 

revoluta: municipio Mariño, Península de Paria, cumbre de 

Las Estrellas, oeste de Manacal, 30-11-1979, J. Steyermark 

y R. Liesner 120815 (VEN). P. ruscifolia: municipio 

Mariño, Cerro de Humo, entre Los Pocitos y La Roma, 11- 

08-1966, J. Steyermark y M. Rabe 96330 (VEN). P. aff. 

secunda: municipio Montes, Macizo montañoso del 

Turimiquire, fila La Pesjua, (16-20)-03-1994, L. Cumana et 

al. 5860 (IRBR). P. velaticaulis, municipio Montes, Las 

Piedras de Cocollar, macizo montañoso del Turimiquire, 

23-12-2007, C. Leopardi 156 (IRBR). Polystachya, P. 

foliosa: municipio Cajigal, Yaguaraparo, 19-02-2007, C. 

Leopardi y F. Subero 97 (IRBR). Prostechea, P. cochleata: 

municipio Montes, Río Totucual – El Guamal, macizo 

Turimiquire, (03-06)-08-1993, L. Cumana et al. 5576 

(IRBR). P. aemula: municipio Benítez, Río El Pilar, 06-05- 

2007, C. Leopardi y C. Malavé 145 (IRBR). P. livida: 

municipio Sucre, Altos de Santa Fe, 21-05-2006, C. 

Leopardi S/N (IRBR). Psilochilus, P. modestus: municipio 

Mariño, Península de Paria, entre Manacal y La Cuchilla, 

13-08-1966, J. Steyermark y M. Rabe 96415 (VEN). 

Psychopsis, P. papilio, municipio Sucre, Altos de Santa Fe, 

22-01-2007, C. Leopardi 64. (IRBR). Psygmorchis, P.


glossomystax: municipio Sucre, Altos de Santa Fe, 22-01- 

2007, C. Leopardi 61 (IRBR). P. pusilla: municipio Bolívar, 

Marigüitar, Río Frío, 20-02-1989, L. Cumana y P. Cabeza 

3560 (IRBR). Rhetinantha, R. notylioglossa: municipio 

Valdez, Península de Paria, Cerro Patao, norte de Puerto 

Hierro, 19-07-1972, J. Steyermark y G. Agostini 91144 

(VEN). Rodriguezia, R. lanceolata: municipio Benítez, Río 

El Pilar, Hacienda El Otro Lado, 06-05-2007, C. Leopardi y 

C. Malavé 148 (IRBR). Sauvetrea, S. aff. alpestris: 

municipio Montes, Macizo montañoso del Turimiquire, 

(25-31)-03-1995, L. Cumana et al. 6180 (IRBR). 

Scaphosepalum, S. breve: municipio Cajigal, Península de 

Paria, 21-02-1980, J. Steyermark et al. 121477 (VEN). 

Scaphyglottis, S. grandiflora: municipio Mariño, Península 

de Paria, Cerro Humo, entre El Paujil y El Brasil, 11-08- 

1966, J. Steyermark y M. Rabe 96351 (VEN). S. modesta: 

municipio Valdez, Península de Paria, Cerro Patao, (25- 

26)-07-1962, J. Steyermark y G. Agostini 91393 (VEN). S. 

prolifera: municipio Montes, Palomar, Periquitos- 

Turimiquire, 10-11-1996, L. Cumana 6328 (IRBR). S. 

reflexa, municipio Arismendi, Península de Paria, entre La 

Tacarigua y las cabeceras del Río Tacarigua, 21-02-1980, J. 

Steyermark et al. 121465 (VEN). Sobralia, S. cf. ciliata: 

municipio Montes, El Guamal, fila La Pesjua, Macizo 

Montañoso del Turimiquire, (23-31)-03-1995, L. Cumana 

et al. 6208 (IRBR). S. aff. violacea: municipio Sucre, Altos 

de Santa Fe, 22-01-2007, Leopardi 66 (IRBR). Stelis, S. 

argentata: municipio Sucre, El Guayabito, (20-22)-11- 

1981, G. Davidse y A. González 19142 (VEN). S. 

papaquerensis: municipio Mariño, Península de Paria, 

camino de Los Pocitos de Santa Isabel a La Roma, 12-07- 

1972, Dumont et al. VE-7642 (VEN). S. tristyla: municipio 

Mariño, Península de Paria, Las Melenas, 01-12-1979, J. 

Steyermark y R. Liesner 120890 (VEN). Stenia, S. pallida: 

municipio Mariño, Península de Paria, Cerro de Humo, 11- 

08-1966, J. Steyermark y M. Rabe 96326-A (VEN). 

Trichosalpinx, T. ciliaris: municipio Cajigal, Península de 

Paria, entre El Paujil y El Brasil, 21-02-1980, J. Steyermark 

et al. 121507 (VEN). T. intricata: municipios Mariño y 

Arismendi, Península de Paria, Las Melenas, 24-02-1980, 

J. Steyermark et al. 121757 (VEN). Triphora, T. 

gentianoides: municipio Valdez, Península de Paria, Cerro 

Patao, 20-07-1962, J. Steyermark y G. Agostini 67582 

(VEN). Trizeuxis, T. falcata: municipio Cajigal, 

Yaguaraparo, 19-02-2007, C. Leopardi y F. Subero 82 

(IRBR). Vanilla, V. pompona: municipio Sucre, Sabilar, 

nov-1984, Cumana 2596 (IRBR). Wullschlaegelia, W. 

calcarata: municipio Valdez, Península de Paria, Cerro 

Patao, (25-26)-07-1962, J Seyermark y G. Agostini 91334 

(VEN). Xylobium, X. pallidiflorum: municipio Benítez, 

Serranía de La Paloma, 19-02-1980, J. Steyermark et al. 

121345 (VEN). 


LANKESTERIANA

LANKESTERIANA 8(1): 105-112. 2008. 

VegetatiVe anatomy of Calypsoeae (orChidaCeae) 

William louis stern 1 & BarBara s. CarlsWard 2,3 

1Department of Biological Sciences, Biscayne Bay Campus, MSB 357, Florida International University, 

North Miami, Florida 33181, USA 

2Department of Biological Sciences, Eastern Illinois University, Charleston, Illinois 61920-3099, USA 

3 Corresponding author: bscarlsward@eiu.edu 

aBstraCt. Calypsoeae represent a small tribe of anatomically little-known orchids with a wide distribution in 

the Western Hemisphere. Leaves are present in all genera, except Corallorhiza and Wullschlaegelia both of 

which are subterranean taxa. Stomata are abaxial (ad- and abaxial in Aplectrum) and tetracytic (anomocytic 

in Calypso). Fiber bundles are absent in leaves of all taxa examined except Govenia tingens. Stegmata 

are present in leaves of only Cremastra and Govenia. Roots are velamentous, except in filiform roots of 

Wullschlaegelia. Vegetative anatomy supports a relationship between Wullschlaegelia and Corallorhiza but 

does not support the grouping of winter-leaved Aplectrum and Tipularia nor proposed groupings of genera 

based on pollinarium features. 

additional KeyWords: Leaf, stem, root, fiber bundle, velamen, stegmata 

introduction 

Calypsoeae (Camus) Dressler [Corallorhizinae 

in Dressler (1981) except Calypso] is a small tribe 

consisting of approximately 12 genera and 62 species 

(Freudenstein 2005). Plants are terrestrial, mostly 

cormous and/or rhizomatous, leafless and rootless 

in some genera, and usually mycorrhizal. Several of 

these genera are monotypic or oligotypic, Govenia 

being the richest with about 30 species. Plants are 

distributed widely from Europe, northern Asia, and 

North America to tropical Central America and the 

Caribbean, Brazil, and Argentina. Calypsoeae are 

absent from Africa, Australia, and islands of the East 

Indies and Pacific Ocean. Yoania australis, a New 

Zealand endemic, was renamed Danhatchia australis 

by Garay and Christenson, and placed in Cranichideae 

(Freudenstein 2005). 

Little is known of the anatomy of the groups and 

except for Corallorhiza (Carlsward & Stern 2008), 

Wullschlaegelia (Stern 1999), and to a certain extent 

Yoania (Campbell 1970), only brief mention is made 

of anatomy for Aplectrum by Holm (1904), Solereder 

& Meyer (1930), Porembski & Barthlott (1988); for 

Calypso by Holm (1904), Møller & Rasmussen (1984), 

Porembski & Barthlott (1988); for Corallorhiza by 

Johow (1885), Solereder & Meyer (1930), Møller 

& Rasmussen (1984); Govenia by Pridgeon, Stern 

& Benzing (1983), Porembski & Barthlott (1988); 

for Oreorchis by Porembski & Barthlott (1988); for 

Tipularia by Holm (1904), Solereder & Meyer (1930), 

Porembski & Barthlott (1988); and for Wullschlaegelia 

by Johow (1885), Solereder & Meyer (1930). 

The rhizome of Yoania australis is covered 

with scale leaves, but the plant lacks expanded, 

chlorophyllous leaves (Campbell 1970) as do 

Corallorhiza and Wullschlaegelia. The rhizome 

has short conical projections bearing long hairs, 

resembling Corallorhiza and Cremastra. There is 

a parenchymatous cortex bounded internally by an 

endodermis with Casparian strips. Vascular tissue 

consists of two to six collateral bundles embedded in 

parenchyma. Among these genera, the only anatomical 

features of any substantive value are the foliar stegmata 

in Cremastra and Govenia reported in this study and 

the spiranthosomes in cortical parenchyma cells from 

fusiform roots of Wullschlaegelia, noted by Stern 

(1999). Other characters occur widely in Orchidaceae. 

Two groups of Calypsoeae have been identified 

based upon the origin of the stalk supporting the 

pollinia and molecular data (Freudenstein 2005).

106 

Pollinia in Aplectrum, Cremastra, Corallorhiza, 

Govenia, and Oreorchis are attached to a stalk (stipe) 

derived from the apex of the rostellum, i.e., a hamulus. 

Pollinia in Calypso, Yoania, and possibly Changnienia, 

are tegular, i.e., the stalk is derived from tissues of the 

anther bed on the column. There is no stalk supporting 

the pollinia of Wullschlaegelia. Along with Tipularia, 

the last three genera also have spurred lips. Winter 

leaves appear in Aplectrum and Tipularia, genera 

that occur in deciduous woodlands, and depend upon 

sunlight reaching leaves during winter. This feature 

may also be indicative of a close relationship between 

these two genera (Freudenstein 2005). 

material and methods 

We had available for study representatives of 

seven of the 12 genera included in Calypsoeae by 

Freudenstein (2005). Corallorhiza and Wullschlaegelia 

are treated in two other publications (Stern 1999, 

Carlsward & Stern 2008). Binomials and authorities, 

abbreviated according to Brummitt and Powell 

(1992), representing these genera appear in Table 1, 

along with organs available for study. Methods and 

descriptive conventions have appeared in recently 

published papers (e.g., Stern & Carlsward 2006), and 

we have followed similar procedures here. “Periclinal” 

and “anticlinal” have been used as shorthand for 

“periclinally orientated” and “anticlinally orientated.” 

Plant parts were preserved in FAA (70% ethanolglacial 

acetic acid-commercial formalin, 9.0:0.5:0.5) 

and stored in 70% ethanol. Transverse and longitudinal 

LANKESTERIANA 

taBle 1. Species of Calypsoeae studied. L, leaf; S, stem; R, root; Rh, rhizome, C, corm. 


sections of leaves and transverse sections of stems and 

roots were cut unembedded as thinly as possible with 

a Reichert sliding microtome, stained in Heidenhain’s 

iron-alum hematoxylin, and counter-stained with 

safranin. Leaf scrapings followed Cutler’s method 

(Cutler 1978) and were stained with safranin. Sections 

and scrapings were mounted on glass slides with 

Canada balsam. Observations were made using a 

Nikon Optiphot microscope, and photographs were 

taken with a PixeraPro 150es digital camera attached 

to a Zeiss Axioskop 40 microscope. Measurements 

of the lengths and widths of ten guard-cell pairs were 

made, and these are given in Table 2. 

anatomy results 

leaf — Surface: HAIRS absent in Aplectrum, 

Cremastra, and Govenia tingens. In Calypso two-celled 

thin-walled hairs ad- and abaxial, the base embedded 

among cells in a nest of small epidermal cells; adaxial: 

basal cell clear, apical cell bulbous, darkly staining 

(Fig. 1A); abaxial: basal cell much shorter than blunttipped 

darkly staining, elongated apical cell (Fig. 1B). 

Hairs ad- and abaxial, two-celled in Govenia superba, 

thin-walled arising from a cluster of small epidermal 

cells; apical cell blunt-tipped. Tipularia has twocelled 

(?) hairs on both leaf surfaces arising from a 

group of small epidermal cells. Hair bases embedded 

among epidermal cells. STOMATA abaxial, except 

ad- and abaxial in Aplectrum. Occasionally a few 

stomata may appear adaxially on leaves in taxa that 

normally bear abaxial stomata. Stomatal apparatus is 

taxon Collector/Voucher parts available 

Aplectrum hyemale (Muhl. ex Willd.) Nutt. B. Carlsward s.n./SEL L, S, R, Rh, C 

Calypso bulbosa (L.) Oakes R. Halse 7141/OSC L, S, R, C 

Corallorhiza maculata (Raf.) Raf. K. Chambers 5597/OSC Rh 

C. odontorhiza (Willd.) Nutt. M. W. Morris & R. Carter 4098/FLAS Rh 

C. wisteriana Conrad M. W. & M. S. Morris 4102/FLAS Rh 

Cremastra appendiculata (D. Don) Makino R. Determann s.n./SEL L, S, R, Rh, C 

Govenia superba (La Llave & Lex.) Lindl. Hort./SEL L, S, R, C 

G. tingens Poepp. & Endl. W. M. Whitten 91284/FLAS L, S, R, C 

Tipularia discolor (Pursh) Nutt. W. L. Stern s.n./FLAS L, S, R, C 

Wullschlaegelia aphylla (Sw.) Rchb.f. R. L. Dressler 4940/FLAS S, R 

W. calcarata Benth. R. L. Dressler 4646/FLAS S, R 

W. calcarata Benth. M. G. Born 41/U S, R

taBle 2. Stomatal dimensions (μm) 

stern & CarlsWard – Vegetative anatomy of Calypsoeae 107 

Average Range 

taxon length Width length Width 

Aplectrum hyemale ad. 45 40 42-47 35-45 

Aplectrum hyemale ab. 46 39 31-52 32-42 

Calypso bulbosa 50 45 42-55 40-50 

Cremastra appendiculata 42 35 37-47 27-37 

Govenia superba 59 48 50-80 42-50 

Govenia tingens 62 52 57-67 47-62 

Tipularia discolor 58 45 50-62 37-50 

basically tetracytic in Aplectrum, Cremastra, Govenia, 

and Tipularia with a few anomocytic configurations. 

Lateral cells of the stomatal apparatus in G. tingens 

often elongated serpent-like to intruded between 

adjoining epidermal cells (Fig. 1C); in Tipularia lateral 

cells and sometimes apical cells protrude among other 

epidermal cells. In Calypso, stomatal apparatuses 

entirely anomocytic (Fig. 1D). Guard cells typically 

reniform and stomata parallel the long axis of the 

guard-cell pair. Average stomatal lengths range from 

figure 1. Leaf scrapings. Scale bars = 50 mm. A. Calypso bulbosa. Adaxial: hairs two-celled, basal cell clear, shorter 

than dark-staining, bulbous apical cell. B. Calypso bulbosa. Abaxial: hairs two-celled, basal cell clear, shorter than 

dark-staining elongated apical cell. C. Govenia tingens. Lateral cells of tetracytic stomatal apparatuses intrude among 

adjacent epidermal cells. D. Calypso bulbosa. Anomocytic stomatal apparatuses. 


108 

figure 2. T-S leaf. Scale bars = 100 mm. A. Tipularia 

discolor with chlorophyllous, upright hypodermal cells 

and spongy mesophyll as in a eudicotyledonous leaf. 

B. Aplectrum hyemale. T-S leaf showing pronounced 

abaxial mid-vein sclerenchyma, xylem sclerenchyma 

clusters resulting in adaxial surface ribbing, and crystal 

idioblasts. 

42 μm in Cremastra to 62 μm in Govenia tingens and 

widths from 39 μm in Aplectrum to 52 μm in Govenia 

tingens. Epidermal cells polygonal on both surfaces; 

abaxial cells in Calypso may be elongated; walls 

straight-sided or curvilinear. 

Section: CUTICLE smooth, somewhat granulose in 

Calypso; 2.5 μm to less than 2.5 μm thick. Epidermal 

cells mostly isodiametric to periclinal in Govenia and 

to a certain extent in Tipularia. STOMATA superficial; 

substomatal chambers large in Tipularia, Govenia, 

and Calypso; moderate in Aplectrum and Cremastra. 

Stomatal ledges usually poorly defined in section; outer 

ledges apparent in Aplectrum, Calypso, Cremastra, 

Govenia, and Tipularia but most pronounced in 

Aplectrum. Inner ledges obscure but apiculate in G. 

tingens and Tipularia. FIBER BUNDLES absent 

throughout, except in G. tingens. HYPODERMIS 

uniseriate adaxially, cells globose or inflated, sparsely 

provided with chloroplasts in Aplectrum; biseriate 

adaxially in Tipularia, outer layer of more or less inflated 

upright cells (Fig. 2A), inner layer cells isodiametric; 

LANKESTERIANA 


cells of both layers rich in chloroplasts. Hypodermis 

absent in Calypso, Cremastra, and Govenia. 

MESOPHYLL homogeneous, 4-7 cells wide, cells thinwalled, 

mostly oval and circular with small triangular 

and polyhedral intercellular spaces; in Tipularia, cells 

and associated intercellular spaces are organized as in 

a eudicotyledon (Fig. 2A). Raphide-bearing idioblasts 

circular in TS (Fig. 2B), saccate, blunt-ended in LS. 

VASCULAR BUNDLES collateral in a single series. 

In larger vascular bundles of Aplectrum, Cremastra, 

and Govenia both xylem and phloem subtended by 

patches of thin-walled sclerenchyma. On the xylem 

side in Aplectrum and Govenia, these produce bulges 

(TS) resulting in ridges on the adaxial leaf surface 

(Fig. 2B) but not on the phloem side. Midvein in 

Aplectrum, Cremastra, and Govenia is subtended 

opposite phloem by a massive cluster of sclerenchyma 

cells creating a pronounced keel (Fig. 2B). Vascular 

bundles in Calypso not associated with sclerenchyma; 

in Tipularia sclerenchyma is associated only with the 

xylem. STEGMATA absent from Aplectrum, Calypso, 

and Tipularia but present in Cremastra and Govenia. 

Conical, rough-surfaced silica bodies in stegmata 

occur along sclerenchyma opposite xylem and phloem 

in Cremastra, along phloem sclerenchyma only in 

Govenia, and associated with fiber bundles in G. 

tingens. Bundle sheath cells circular, thin-walled, and 

chloroplast-bearing in all taxa; chloroplasts do not 

occur in some bundle sheath cells of Calypso. 

stem — Subterranean storage, perennating, and 

connective organs (rhizomes) several in Calypsoeae, 

as noted by Freudenstein (2005), and represented in 

our research materials. These organs correspond to 

cauline structure in that vascular bundles, containing 

adnate strands of xylem and phloem, are associated in 

a usually parenchymatous matrix. They are arranged 

in different configurations in contrast with root 

morphology where conductive tissues are typically 

concentrated in a central vascular cylinder. There 

are a rhizome and corm in Aplectrum and Cremastra 

and a corm in Calypso, Govenia, and Tipularia. The 

organizational pattern of each organ is described 

separately for each taxon. 

Aplectrum rhizome — HAIRS none. STOMATA 

present, superficial, substomatal chamber large.


figure 3. T-S underground stem. A. Aplectrum hyemale corm with starch grains in smaller, nucleated, assimilatory cells and 

larger enucleate, water-storage cells under polarized light. Scale bar = 100 mm. B. Calypso bulbosa corm bearing twocelled 

hairs. Scale bar = 50 mm. C. Cremastra appendiculata rhizome with excrescences bearing tufts of unicellular 

hairs. Scale bar = 100 mm. D. Cremastra appendiculata corm, vascular bundles surrounded by an endodermis. Scale 

bar = 100 mm. 

CUTICLE smooth to rugose, 2.5 μm thick. 

EPIDERMAL CELLS mostly isodiametric, subtended 

by a uniseriate HYPODERMIS in some areas; cells 

larger than epidermal cells but smaller than ground 

tissue cells. CORTEX none. ENDODERMIS 

discontinuous around the vascular cylinder; 

cells isodiametric, entirely thin-walled, lacking 

intercellular spaces; there are possibly Casparian 

strips. Presence of a PERICYCLE is questionable. 

GROUND TISSUE cells thin-walled, circular, 

oval, and variably shaped; cruciate starch grains in 

assimilatory cells; thin-walled, circular; enucleate 

water-storage cells scattered about, intercellular 

spaces triangular (Fig. 3A). Twenty-five to 30 widely 

separated VASCULAR BUNDLES distributed within 

the endodermis surrounded by a parenchymatous 

matrix of thin-walled, nucleated, small angular cells 

lacking intercellular spaces. SCLERENCHYMA and 

STEGMATA absent. 

Aplectrum corm — HAIRS and STOMATA none. 

CUTICLE absent. EPIDERMAL CELLS isodiametric. 

CORTEX two or three cells wide; cells variably shaped. 

ENDODERMIS and PERICYCLE absent. GROUND 

TISSUE with numerous, circular, and elliptical, large 

enucleate water-storage cells surrounded by much 

smaller, variably shaped, numerous assimilatory 

cells with cruciate starch grains. Intercellular spaces 

triangular. VASCULAR BUNDLES many, collateral, 

widely scattered; xylem unitary, binary, trinary in each 

bundle. SCLERENCHYMA and STEGMATA none. 

Calypso corm — HAIRS two-celled, thickwalled, 

apical cell clavate, darkly-staining (Fig. 

3B). STOMATA absent. CUTICLE smooth, 5.0 μm 

thick. EPIDERMAL CELLS squarish, isodiametric. 

HYPODERMIS uniseriate, cells tending toward 

periclinal. CORTEX, ENDODERMIS, PERICYCLE 

absent. GROUND TISSUE cells with thick and thin 

walls, oval, angular and circular; larger, enucleate 

water-storage cells and smaller, nucleated assimilatory 

cells lacking starch grains; intercellular spaces few. 

VASCULAR BUNDLES many, collateral, widely 

scattered. Xylem arcuate in some bundles, phloem 

centered in the arms of the arc. SCLERENCHYMA 

and STEGMATA none. 

Cremastra rhizome — HAIRS emerge as tufts of 

unicellular strands from raised, truncate, pyramidal, 

multicellular cushions along the periphery of the stem 

(Fig. 3C). STOMATA and CUTICLE indistinguishable. 

EPIDERMAL CELLS squarish, isodiametric. 


110 

CORTEX many cells wide; cells crowded, various, 

angular, oval, circular, irregular; walls thickish; 

water-storage cells circular, empty. Cruciate starch 

grains in assimilatory cells. ENDODERMAL CELLS 

surrounding vascular bundles, entirely thin-walled, 

rectangular, square, some roundish, isodiametric, 

with Casparian strips (Fig. 3D). PERICYCLE 

discontinuous, cells like endodermal cells without 

Casparian strips. GROUND TISSUE of oval and 

angular thin- and thick-walled parenchyma cells. 

VASCULAR TISSUE organized as an irregular series 

of discontinuous arcs in which collateral vascular 

bundles are interspersed. SCLERENCHYMA and 


Cremastra corm — HAIRS and STOMATA absent. 

CUTICLE none. EPIDERMAL CELLS periclinal. 

ENDODERMAL CELLS angular, thin-walled, 

surrounding each vascular bundle. GROUND TISSUE 

with larger, almost circular, empty, water-storage cells 

and smaller, thin-walled, oval and variously shaped, 

assimilatory cells containing cruciate starch grains; 

intercellular spaces tiny, triangular, and various. 

VASCULAR BUNDLES many, collateral, scattered. 

SCLERENCHYMA and STEGMATA absent. 

Govenia superba corm — HAIRS and CUTICLE 

absent. STOMATA present in G. tingens, subtended 

by small substomatal chambers. EPIDERMAL 

CELLS isodiametric. CORTEX, ENDODERMIS, 

PERICYCLE none. GROUND TISSUE: Large 

numbers of wide, circular, thin-walled water-storage 

cells surrounded by much smaller, thin-walled, oval, 

rectangular, square, and variously shaped assimilatory 

cells lacking starch grains. VASCULAR BUNDLES 

collateral, many, scattered. SCLERENCHYMA and 


Tipularia corm — HAIRS, STOMATA, and 

CUTICLE absent. EPIDERMAL CELLS isodiametric. 

CORTEX, ENDODERMIS, and PERICYCLE none. 

GROUND TISSUE of larger, circular, thick-walled 

water-storage cells and much smaller, irregularly 

shaped assimilatory cells with cruciate starch grains. 

Intercellular spaces absent. VASCULAR BUNDLES 

collateral, many, scattered. SCLERENCHYMA and 


LANKESTERIANA 


root — VELAMEN one cell layer wide in Aplectrum 

(Fig. 4A) and Govenia, one or two cell layers wide 

in Cremastra, two cell layers wide in Calypso (Fig. 

4B), four cell layers wide in Tipularia (Fig. 4C). 

Cells thin-walled without secondary thickenings, 

isodiametric in Aplectrum, Govenia, Cremastra, and 

Tipularia; epivelamen cells periclinal, endovelamen 

cells isodiametric in Calypso. Unicellular hairs present 

in all taxa. TILOSOMES absent. EXODERMAL 

CELLS square and polygonal to anticlinal, thin-walled 

throughout; passage cells intermittent. CORTEX up to 

10 cells wide in Aplectrum, 8-10 cells wide in Calypso, 

7 cells wide in Cremastra, 7-9 cells wide in Govenia, 

and 7 or 8 cells wide in Tipularia; cells thin-walled. 

Hyphae, dead cell masses, or clots, and pelotons occur 

in cortices of all taxa (Fig. 4D) except Cremastra 

and Govenia tingens. ENDODERMAL CELLS 

isodiametric, rectangular in Cremastra, periclinal in 

Govenia tingens, entirely thin-walled, Casparian strips 

in Aplectrum and Tipularia. PERICYCLE CELLS thinwalled 

throughout, mostly isodiametric, smaller than 

endodermal cells. VASCULAR CYLINDER 9-arch 

in Aplectrum, 3-arch in Calypso (Fig. 4D), 5-arch in 

Cremastra, 6-arch in Govenia, 4-arch in Tipularia. 

Xylem in short rays; cells clustered in Govenia; several 

intramedullary xylem clusters in Aplectrum; xylem 

alternates with groups of phloem cells. Vascular tissue 

embedded in thin-walled sclerenchyma or thick-walled 

parenchyma. PITH parenchymatous, sometimes with 

thick walls in Aplectrum; cells polygonal, intercellular 

spaces absent. 

discussion 

Aplectrum stands alone among Calypsoeae, owing 

to its combination of amphistomatal leaves, lack 

of foliar hairs, and absence of stegmata. Calypso, 

too, is unique with its exclusively anomocytic 

stomatal apparatuses, absence of vascular bundle 

sclerenchyma and lack of stegmata. Corallorhiza and 

Wullschlaegelia are characterized by their leaflessness 

and subterranean habit. Govenia tingens lacks foliar 

hairs and is the only taxon studied with foliar fiber 

bundles lined with stegmata. Wullschlaegelia has roots 

of two distinct kinds, fusiform and filiform. The latter 

lack a velamen. Cremastra lacks hairs on leaves but


figure 4. T-S root. A. Aplectrum hyemale with one-cell wide velamen. B. Calypso bulbosa with two-cell-wide velamen. 

C. Tipularia discolor with four-cell-wide velamen. Scale bars = 50 mm. D. Calypso bulbosa . T-S root, cortical cells 

with hyphal pelotons and dead hyphal masses. Triarch vascular cylinder. Scale bar = 100 mm. 

has tufts of unicellular hairs that arise from cushions 

along the rhizome. 

Cremastra, Govenia, and Tipularia have abaxial 

tetracytic stomata; Calypso has mostly abaxial, and a 

few adaxial anomocytic stomatal apparatuses. Foliar 

hairs are present on both surfaces in Calypso, Govenia 

superba, and Tipularia. A foliar hypodermis occurs in 

Aplectrum and Tipularia. Sclerenchyma appears along 

both xylem and phloem sides of vascular bundles in 

Aplectrum, Cremastra, and Govenia, but only on the 

xylem side in Tipularia. Govenia and Tipularia have 

the largest stomatal dimensions; measurements for the 

other taxa are substantially smaller (Table 2). 

Freudenstein (2005) suggested that the appearance 

of winter leaves in Aplectrum and Tipularia may be 

evidence of a close relationship between these two 

genera. Indeed, among the taxa studied, only these 

two are represented by a foliar hypodermis consisting 

of enlarged, thin-walled, globose or expanded 

cells with chloroplasts. Evert (2006) observed that 

cells of the inner layer (s) of a multiple epidermis 

(i.e., a hypodermis) commonly contain few or no 

chloroplasts. Indeed, the cells of hypodermises 

sometimes appear to lack contents entirely, especially 

when they serve as water-storage cells. In the 

cases of winter leaves of Aplectrum and Tipularia, 

however, the hypodermal cells are chlorophyllous, 

especially in Tipularia. The hypotheses that establish 

two groups of genera in Calypsoeae, based upon 

origins of the pollen stipes and presence of spurred 

lips, cannot be substantiated by vegetative anatomy. 

Dressler’s Wullschlaegelieae (1980) and Lindley’s 

Corallorhizidae (1853) are groupings more congruent 

with our findings. 


112 

aCKnoWledgements. The authors thank Kenton L. 

Chambers, Oregon State University, for the superb specimen 

of Calypso bulbosa that he provided for our research. 

For various facilities and use of research equipment, we 

acknowledge with appreciation George Bowes and the 

Department of Botany, University of Florida. David Lee, 

formerly of Florida International University, provided 

laboratory space for the early phases of this research. The 

Kampong of the National Tropical Botanical Garden in 

Coconut Grove, Florida, provided laboratory room. Finally, 

we’d like to thank Alec Pridgeon, an anonymous reviewer, 

and J. Richard Abbott for their help with the manuscript. 

literature Cited 

Brummitt, R. K. & C. E. Powell. 1992. Authors of plant 

names. Royal Botanic Gardens, Kew. 

Campbell, E. O. 1970. The fungal association of Yoania 

australis. Trans. Roy. Soc. New Zealand 12: 5-12. 

Carlsward, B. S. & W. L. Stern. 2008. Corallorhiza, a 

rootless, leafless terrestrial. Orchid Rev. 116: 334-339. 

Cutler, D. F. 1978. Applied plant anatomy. Longman Group 

Ltd., London. 

Dressler, R. L. 1980. Orchídeas huérfanas I. Wullschlaegelia 

una nueva tribu Wullschlaegelieae. Orquídea (Méx.) 7: 

277-282. 

Dressler, R. L. 1981. The orchids: natural history and 

classification. Harvard University Press, Cambridge, 

Massachusetts. 

Evert, R. F. 2006. Esau’s plant anatomy. John Wiley & 

Sons, Hoboken, New Jersey. 

LANKESTERIANA 


Freudenstein, J. V. 2005. Tribe Calypsoeae. In: A. M. 

Pridgeon, P. J. Cribb, M. W. Chase, & F. Rasmussen [eds.], 

Genera Orchidacearum. Volume 4. Epidendroideae 

(Part one). Oxford University Press, Oxford. 

Holm, T. 1904. The root structure of North American 

Orchideae. Amer. J. Sc., series 4 18: 197-212. 

Johow, F. 1885. Die chlorophyllfreien Humusbewohner 

West-Indians, biologisch-morphologisch dargestellt. 

Jahrb. Wissensch. Bot. 16: 415-449. 

Lindley, J. 1853. The vegetable kingdom. Bradbury & 

Evans, London. 

Møller, J. D. & H. Rasmussen. 1984. Stegmata in 

Orchidales: character state distribution and polarity. 

Bot. J. Linn. Soc. 89: 53-76. 

Porembski, S. & W. Barthlott. 1988. Velamen radicum 

micromorphology and classification of Orchidaceae. 

Nordic J. Bot. 8: 117-137. 

Pridgeon, A. M., W. L. Stern & D. H. Benzing. 1983. 

Tilosomes in roots of Orchidaceae: morphology and 

systematic occurrence. Amer. J. Bot. 70: 1365-1377. 

Solereder, H. & F. J. Meyer. 1930. Systematische Anatomie 

der Monokotyledonen. VI. Scitamineae-Microspermae. 

Verlag von Gebrüder Bornträger, Berlin. 

Stern, W. L. 1999. Comparative vegetative anatomy 

of two saprophytic orchids from tropical America: 

Wullschlaegelia and Uleiorchis. Lindleyana 14: 136- 

146. 

Stern, W. L. & B. S. Carlsward. 2006. Comparative 

vegetative anatomy and systematics of Oncidiinae 

(Maxillareae, Orchidaceae). Bot. J. Linn. Soc. 152: 91- 

107.

LANKESTERIANA 8(3): 113. 2008. 

BOOK REVIEW 

Alrich, P. & W.E. Higgins. 2008. The Marie Selby Botanical Garden Illustrated Dictionary 

of Orchid Genera. Comstock Publishing Association. 

As I am listed as one of the editors of this dictionary, 

I should, perhaps, be disqualified as a reviewer. In my 

defense, I reviewed an early version of the manuscript, 

and it has clearly grown and evolved a great deal since 

I last checked over the list of names. 

The authors estimate that there are about 850 

accepted generic names treated here, but synonyms, 

orthographic variants, and names published before 

1753 bring the Dictionary to more than three times 

that number. The casual reader might prefer a shorter 

treatment of the 850 or so valid and accepted names. 

Some names published before 1753 and never used 

again are of merely historical interest (if that). However, 

our understanding of orchid classification is constantly 

changing with the growth of molecular systematics 

(analysis of DNA) and continuing study using various 

other techniques. No one can say which of the validly 

published names we now consider synonyms may 

eventually prove to represent distinct groups in need 

of their own names. By including every generic orchid 

name or variant ever published (as nearly as is humanly 

possible), the authors have produced a book that will 

continue to be useful for many years. 


To aid the reader, the validly published names 

that represent currently accepted genera (from Aa to 

Zygostates) are dark green, while paler green is used 

for validly published names that are not currently in 

use, and dark purple represents “superfluous” names, 

orthographic variants are shown in lilac, names 

published before 1753 are printed in brown, and the 

few names that (may) represent fossil orchid genera 

are shown in blue. Further the currently accepted or 

validly published names are each illustrated by a 

colored image of a flower. 

The Dictionary includes an informative profile of 

the family Orchidaceae by D. H. Benzing, a list of 

selected references and a list of orchid taxonomists, 

from Ackerman to Zollinger, a very useful list of 

book/periodical abbreviations, a brief summary of 

nomenclatural rules, and a glossary. 

The Dictionary is attractive and informative, and 

should continue to be very useful for many years to 

come. 

Robert L. Dressler 

Lankester Botanical Garden

LANKESTERIANA

LANKESTERIANA 8(3): 115-126. 2008. 

SeptobaSidiaceae 

INDEX OF NEW TAXA AND COMBINATIONS 

PUBLISHED IN LANKESTERIANA, VOL. 1—8 

BASIDIOMyCOTA 

Septobasidium alni torrend var. brasiliense couch, var. nov. 4(1): 77. 2004. 

Septobasidium alni torrend var. squamosum couch, var. nov. 4(1): 77. 2004. 

Septobasidium alveomarginatum couch, sp. nov. 4(1): 77. 2004. 

Septobasidium apiculatum couch, sp. nov. 4(1): 77. 2004. 

Septobasidium boedijnii couch, sp. nov. 4(1): 78. 2004. 

Septobasidium boedijnii couch, sp. nov. 4(1): 78. 2004. 

Septobasidium burtii Lloyd var. acerinum couch, var. nov. 4(1): 78. 2004. 

Septobasidium burtii Lloyd var. acerinum couch, var. nov. 4(1): 78. 2004. 

Septobasidium carestianum bres. var. natalense couch, var. nov. 4(1): 78. 2004. 

Septobasidium carestianum bres. var. natalense couch, var. nov. 4(1): 78. 2004. 

Septobasidium cervicolor couch, sp. nov. 4(1): 78. 2004. 

Septobasidium cervicolor couch, sp. nov. 4(1): 78. 2004. 

Septobasidium cokeri couch, sp. nov. 4(1): 80. 2004. 

Septobasidium conidiophorum couch, sp. nov. 4(1): 80. 2004. 

Septobasidium cremeum couch, sp. nov. 4(1): 80. 2004. 

Septobasidium crustaceum couch, sp. nov. 4(1): 80. 2004. 

Septobasidium cupressi couch, sp. nov. 4(1): 81. 2004. 

Septobasidium elatostemae couch, sp. nov. 4(1): 81. 2004. 

Septobasidium ficicolum pat. ex couch, sp. nov. 4(1): 81. 2004. 

Septobasidium filiforme couch, sp. nov. 4(1): 81. 2004. 

Septobasidium formosense couch, sp. nov. 4(1): 82. 2004. 

Septobasidium fragile couch, sp. nov. 4(1): 82. 2004. 

Septobasidium fusco-cinereum bresadola ex couch, sp. nov. 4(1): 82. 2004. 

Septobasidium fuscum couch, sp. nov. 4(1): 82. 2004. 

Septobasidium grandispinosum couch, sp. nov. 4(1): 82. 2004. 

Septobasidium grandisporum couch, sp. nov. 4(1): 83. 2004. 

Septobasidium griseopurpureum couch, sp. nov. 4(1): 83. 2004. 

Septobasidium griseum couch, sp. nov. 4(1): 83. 2004. 

Septobasidium hakgalanum couch & petch, sp. nov. 4(1): 83. 2004. 

Septobasidium hesleri couch, sp. nov. 4(1): 84. 2004. 

Septobasidium heveae couch, sp. nov. 4(1): 84. 2004. 

Septobasidium indigophorum couch, sp. nov. 4(1): 84. 2004. 

Septobasidium irregulare couch, sp. nov. 4(1): 84. 2004. 

Septobasidium lacunosum couch, sp. nov. 4(1): 84. 2004. 

Septobasidium lagerheimii couch, sp. nov. 4(1): 84. 2004. 

Septobasidium lepidosaphis couch, sp. nov. 4(1): 84. 2004. 

Septobasidium leprosum couch, sp. nov. 4(1): 84. 2004.

116 

LANKESTERIANA 

Septobasidium lilacinoalbum couch, sp. nov. 4(1): 86. 2004. 

Septobasidium linderi couch, sp. nov. 4(1): 86. 2004. 

Septobasidium macadamiae couch, sp. nov. 4(1): 86. 2004. 

Septobasidium mariani (bres. ex Sacc.) bres. var. japonicum couch, var. nov. 4(1): 86. 2004. 

Septobasidium molle couch, sp. nov. 4(1): 87. 2004. 

Septobasidium muelleri couch, sp. nov. 4(1): 87. 2004. 

Septobasidium myrsinae couch, sp. nov. 4(1): 87. 2004. 

Septobasidium natalense couch, sp. nov. 4(1): 87. 2004. 

Septobasidium pachydermum couch, sp. nov. 4(1): 88. 2004. 

Septobasidium pallidum couch, sp. nov. sp. nov. 4(1): 88. 2004. 

Septobasidium peckii couch, sp. nov. 4(1): 88. 2004. 

Septobasidium perforatum couch, sp. nov. 4(1): 88. 2004. 

Septobasidium petchii couch, sp. nov. 4(1): 89. 2004. 

Septobasidium philippinense couch, sp. nov. 4(1): 89. 2004. 

Septobasidium piperis p. Henn. ex couch, sp. nov. 4(1): 89. 2004. 

Septobasidium prunophilum couch, sp. nov. 4(1): 89. 2004. 

Septobasidium punctatum couch, sp. nov. 4(1): 90. 2004. 

Septobasidium reikingii pat. ex couch, sp. nov. 4(1): 90. 2004. 

Septobasidium rickii couch, sp. nov. 4(1): 90. 2004. 

Septobasidium rimulosum petch & couch, sp. nov. 4(1): 90. 2004. 

Septobasidium rugulosum couch, sp. nov. 4(1): 90. 2004. 

Septobasidium sabalis couch, sp. nov. 4(1): 90. 2004. 

Septobasidium sabalis-minoris couch, sp. nov. 4(1): 90. 2004. 

Septobasidium scabiosum couch & petch, sp. nov. 4(1): 90. 2004. 

Septobasidium schizostachyi couch, sp. nov. 4(1): 91. 2004 

Septobasidium sclerotioides couch, sp. nov. 4(1): 92. 2004. 

Septobasidium separans couch, sp. nov. 4(1): 92. 2004. 

Septobasidium simmondsii couch, sp. nov. 4(1): 92. 2004. 

Septobasidium sinense couch, sp. nov. 4(1): 92. 2004. 

Septobasidium sinuosum couch, sp. nov. 4(1): 93. 2004. 

Septobasidium stevensonii couch, sp. nov. 4(1): 93. 2004. 

Septobasidium stratosum couch, sp. nov. 4(1): 93. 2004. 

Septobasidium subcarbonaceum (berk. & br.) couch, comb. nov. 4(1): 93. 2004. 

Septobasidium sydowii couch, sp. nov. 4(1): 94. 2004. 

Septobasidium taxodii couch, sp. nov. 4(1): 94. 2004. 

Septobasidium tenue couch, sp. nov. 4(1): 94. 2004. 

Septobasidium tomentosum couch, sp. nov. 4(1): 94. 2004. 

Septobasidium ugandae couch, sp. nov. 4(1): 95. 2004. 

Septobasidium verrucosum couch, sp. nov. 4(1): 95. 2004. 

Septobasidium westonii couch, sp. nov. 4(1): 95. 2004. 

LyCOPHyTA 

Lycopodiaceae 

Huperzia oellgaardii a. Rojas, sp. nov. 5(2): 110. 2005. 


LecHnaceae 

index of taxa and combinations 117 

PTERyDOPHyTA 

Blechnum fuscosquamosum a. Rojas, sp. nov. 5(1): 49. 2005. 

cyatHeaceae 

Cnemidaria chiricana (Maxon) R.M. tryon var. contigua (Underw. ex Maxon) a. Rojas, var. nov. 5(3): 191. 2005. 

Cyathea povedae a. Rojas, sp. nov. 5(3): 192. 2005. 

Cyathea x smithiana a. Rojas. nothosp. nov. 5(3): 195. 2005. 

dRyopteRidaceae 

Polybotrya aureisquama a. Rojas, sp. nov. 7(3): 558. 2007. 

Polybotrya insularis a. Rojas, sp. nov. 7(3): 560. 2007. 

Tectaria dressleri a. Rojas, sp. nov. 6(1): 15. 2006. 

Tectaria x chaconiana a. Rojas, nothosp. nov. 4(2): 149. 2004. 

GRaMMitidaceae 

Enterosora bishopii a. Rojas, sp. nov. 6(1): 9. 2006. 

Enterosora enterosoroides (H. christ) a. Rojas, comb. nov. 6(1): 11. 2006. 

Lellingeria brenesii a. Rojas, sp. nov. 7(3): 553. 2007. 

Lellingeria pinnata a. Rojas, sp. nov. 6(3): 95. 2006. 

Terpsichore glandulifera a. Rojas, sp. nov. 6(3): 96. 2006. 

HyMenopHyLLaceae 

Hymenophyllum talamancanum a. Rojas, sp. nov. 4(2): 143. 2004. 

LoMaRiopSidaceae 

Elaphoglossum lenticulatum a. Rojas, sp. nov. 5(3): 185. 2005. 

Lycopodiaceae 

Huperzia oellgaardii a. Rojas, sp. nov. 5(2): 110. 2005. 

poLypodiaceae 

Campyloneurum gracile a. Rojas, sp. nov. 5(1): 41. 2005. 

ScHizaeaceae 

Anetium citrifolium (L.) Splitg. var. pendulum (Leprieur in Fée) L.d. Gómez, var. nov. 6(1): 6. 2006. 

acantHaceae 

SPERMATOPHyTA 

Justicia chaconii Gómez-Laur, sp. nov. 6(3): 155. 2006. 

apocynaceae 

Allotoonia woodsoniana (Monac.) J.F. Morales & J.K. Williams, comb. nov. 5(2): 119. 2005. 

aSteRaceae 

Neomirandea pendulissima al. Rodr, sp. nov. 5(3): 207. 2005. 

caneLLaceae 

Pleodendron costaricense n. zamora, Hammel & R. aguilar, sp. nov. 5(3): 211. 2005. 


118 

dicHapetaLaceae 

LANKESTERIANA 

Dichapetalum inopinatum al. Rodr. & Kriebel, sp. nov. 5(2): 127. 2005. 

Dichapetalum reliquum Kriebel & al. Rodr, sp. nov. 5(2): 135. 2005. 

Fabaceae-caeSaLpiniaceae 

Swartzia maquenqueana n. zamora & d. Solano, sp. nov. 6(3): 133. 2006. 

GeSneRiaceae 

Drymonia glandulosa Kriebel, sp. nov. 5(1): 81. 2005. 

Drymonia tomentulifera Kriebel, sp. nov. 6(2): 44. 2006. 

LaMiaceae 

Ruyschia moralesii Hammel, sp. nov. 6(2): 75. 2006. 

LaURaceae 

Licaria leonis Gómez-Laur. & estrada, sp. nov. 3: 5. 2002. 

LentibULaRiaceae 

Utricularia uxoris Gómez-Laur., sp. nov. 5(2): 137. 2005. 

MaRcGRaViaceae 

Marcgravia glandulosomarginata Hammel, sp. nov. 6(2): 73. 2006. 

Schwartzia tarrazuensis Hammel, sp. nov. 6(2): 76. 2006. 

MeLiaceae 

Guarea adenophylla al Rodr., sp. nov. 6(3): 102. 2006. 

Guarea aguilarii al. Rodr., sp. nov. 6(3): 103. 2006. 

Guarea ciliata al. Rodr., sp. nov. 6(3): 105. 2006. 

Guarea constricta al. Rodr., sp. nov. 6(3): 106. 2006. 

Guarea corticosa al. Rodr., sp. nov. 6(3): 107. 2006. 

Guarea inesiana al Rodr., sp. nov. 6(3): 109. 2006. 

Guarea macrocalyx al. Rodr., sp. nov. 6(3): 110. 2006. 

Guarea montana al. Rodr., sp. nov. 6(3): 111. 2006. 

Guarea pilosa al. Rodr., sp. nov. 6(3): 113. 2006. 

Guarea tafae-malekui al. Rodr., sp. nov. 6(3): 115. 2006. 

MoRaceae 

Ficus lasiosyce J. a. González & poveda, sp. nov. 8: 13. 2003. 

MyRtaceae 

Eugenia earthiana p.e. Sánchez, sp. nov. 4(3): 179. 2004. 

Plinia cuspidata Gómez-Laur. & Valverde, sp. nov. 3: 11. 2002. 

oRcHidaceae 

Acianthera aberrans pupulin & bogarín, comb. nov. 8(2): 53. 2008. 

Aetheorhyncha dressler, gen. nov. 5(2): 94. 2005. 

Aetheorhyncha andreettae (Jenny) dressler, comb. nov. 5(2): 95. 2005. 

Benzingia caudata (ackerman) dressler, comb. nov. 5(2): 93. 2005. 



Benzingia cornuta (Garay) dressler, comb. nov. 5(2): 93. 2005. 

Benzingia estradae (dodson) dodson, comb. nov. 5(2): 93. 2005. 

Benzingia hajekii (d.e. benn. & christenson) dressler, comb. nov. 5(2): 93. 2005. 

Benzingia jarae (d.e.benn. & christenson) dressler, comb. nov. 5(2): 93. 2005. 

Benzingia palorae (dodson & Hirtz) dressler, comb. nov. 5(2): 93. 2005. 

Benzingia reichenbachiana (Schltr.) dressler, comb. nov. 5(2): 93. 2005. 

Camaridium alfaroi (ames & c. Schweinf.) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium allenii (L. o. Williams) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium amabile (J. t. atwood) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium ampliflorum (c. Schweinf.) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium anceps (Rchb.f.) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium atratum (Lex.) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium aurantiacum (Schltr.) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium bomboizense (dodson) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium brevilabium (ames & correll) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium burgeri (J. t. atwood) M. a. blanco, comb. nov. 7(3): 519. 2007. 

Camaridium campanulatum (c. Schweinf.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium carinulatum (Rchb. f.) M.a. blanco, comb. nov. 8(1): 15. 2008. 

Camaridium cedralense (J. t. atwood & Mora-Ret.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium cucullatum (Lindl.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium densum (Lindl.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium falcatum (ames & correll) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium fragrans (J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium gomezianum (J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium grisebachianum (nir & dod) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium haberi (J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium hagsaterianum (Soto arenas) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium horichii (Senghas) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium inauditum (Rchb.f.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium insolitum (dressler) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium lankesteri (ames) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium longicolumna (J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium lutheri (J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium meleagris (Lindl.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium micranthum M. a. blanco, nom. nov. 7(3): 520. 2007. 

Camaridium microphyton (Schltr.) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium mombachoense (a. H. Heller ex J. t. atwood) M. a. blanco, comb. nov. 7(3): 520. 2007. 

Camaridium monteverdense (J. t. atwood & G. barboza) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium neglectum (Schltr.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium obscurum (Linden & Rchb.f.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium oestlundianum (L. o. Williams) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium paleatum (Rchb.f.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium praestans (Rchb.f.) M. a. blanco, comb. nov. 7(3): 521. 2007. 


120 

LANKESTERIANA 

Camaridium pygmaeum M. a. blanco, nom. nov. 7(3): 521. 2007. 

Camaridium ramonense (Schltr.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium rhombeum (Lindl.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium scalariforme (J. t. atwood) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium sigmoideum (c. Schweinf.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium soconuscanum (breedlove & d. Mally) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium standleyi M. a. blanco, nom. nov. 7(3): 521. 2007. 

Camaridium stenophyllum (Schltr.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium strumatum (endres & Rchb.f.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium suaveolens (barringer) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium synsepalum (J. t. atwood) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium tigrinum (c. Schweinf.) M. a. blanco, comb. nov. 7(3): 521. 2007. 

Camaridium tricarinatum (J. t. atwood) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Camaridium tuberculare (J. t. atwood) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Camaridium tutae (J. t. atwood) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Camaridium vaginale (Rchb.f.) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Camaridium valerioi (ames & c. Schweinf.) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Camaridium vittariifolium (L. o. Williams) M. a. blanco, comb. nov. 7(3): 522. 2007. 

Chondroscaphe endresii (Schltr.) dressler, comb. nov. 3: 28. 2002. 

Christensonella cepula (Rchb.f.) S. Koehler, comb. nov. 7(3): 522. 2007. 

Christensonella neowiedii (Rchb.f.) S. Koehler, comb. nov. 7(3): 522. 2007. 

Christensonella pacholskii (christenson) S. Koehler, comb. nov. 7(3): 522. 2007. 

Christensonella squamata (barb. Rodr.) carnevali, comb. nov. 7(3): 523. 2007. 

Coryanthes kaiseriana G. Gerlach, sp. nov. 8: 23. 2003. 

Coryanthes maduroana G. Gerlach, sp. nov. 4(1): 70. 2004. 

Cryptocentrum benth. subgenus Anthosiphon (Schltr.) carnevali, comb. et stat. nov. 7(3): 543. 2007. 

Dendrophylax monteverdi (Rchb.f.) ackerman & nir, comb. nov. 4(1): 53. 2004. 

Dichaea elliptica dressler & Folsom, sp. nov. 3: 25. 2002. 

Echinella vittata (pupulin & M.a. blanco) pupulin, comb. nov. 4: 17. 2002. 

Echinorhyncha antonii (ortiz) dressler, comb. nov. 5(2): 94. 2005. 

Echinorhyncha dressler, gen. nov. 5(2): 94. 2005. 

Echinorhyncha ecuadorensis (dodson) dressler, comb. nov. 5(2): 94. 2005. 

Echinorhyncha litensis (dodson) dressler, comb. nov. 5(2): 94. 2005. 

Echinorhyncha vollesii (Gerlach, neudecker & Seeger) dressler, comb. nov. 5(2): 94. 2005. 

Echinosepala vittata (pupulin & M.a. blanco) c.o. Murales & n. Villal., comb. nov. 4(3): 202. 2004. 

Elleanthus ligularis dressler & bogarín, sp. nov. 7(3): 539. 2007. 

Encyclia cajalbanensis Mújica, bocourt & pupulin, sp. nov. 4(3): 211. 2004. 

Encyclia monteverdensis M. a. díaz & ackerman, sp. nov. 4(1): 50. 2004. 

Epidendrum cancanae (p.ortiz) Hágsater, comb. nov. 5(1): 73. 2005. 

Epidendrum fuscinum (dressler) Hágsater, comb. nov. 5(1): 73. 2005. 

Epidendrum macdougalli (Hágsater) Hágsater, comb. nov. 5(1): 74. 2005. 

Epidendrum misasii Hágsater, nom. nov. 5(1): 74. 2005. 

Epidendrum montis-narae pupulin & L. Sánchez S., sp. nov. 1: 7. 2001. 



Epidendrum paruimense G. a. Romero & carnevali, sp. nov. 4(3): 229. 2004. 

Epidendrum parviexasperatum (Hágsater) Hágsater, comb. nov. 5(1): 74. 2005. 

Epidendrum stolidium Hágsater, nom. nov. 5(1): 74. 2005. 

Epidendrum x monteverdense (pupulin & Hágsater) Hágsater, comb. nov. 5(1): 74. 2005. 

Epidendrum zunigae Hágsater, Karremans & bogarín, sp. nov. 8(2): 63. 2008. 

Euryblema dressler, gen. nov. 5(2): 94. 2005. 

Euryblema anatonum (dressler) dressler, comb. nov. 5(2): 94. 2005. 

Euryblema andreae (ortiz) dressler, comb. nov. 5(2): 94. 2005. 

Govenia viaria dressler, sp. nov. 3: 26. 2002. 

Guarianthe dressler & W.e. Higgins, gen. nov. 7: 37. 2003. 

Guarianthe aurantiaca (bateman ex Lindl.) dressler & W.e. Higgins, comb. nov. 7: 38. 2003. 

Guarianthe bowringiana (Veitch) dressler & W.e. Higgins, comb. nov. 7: 38. 2003. 

Guarianthe patinii (cogn.) dressler & W.e. Higgins, comb. nov. 7: 38. 2003. 

Guarianthe skinneri (bateman) dressler & W.e. Higgins, comb. nov. 7: 38. 2003. 

Inti M. a. blanco, gen. nov. 7(3): 524. 2007. 

Inti bicallosa (Rchb.f.) M. a. blanco, comb. nov. 7(3): 524. 2007. 

Inti chartacifolia (ames & c. Schweinf.) M. a. blanco, comb. nov. 7(3): 524. 2007. 

Ixyophora dressler, gen. nov. 5(2): 95. 2005. 

Ixyophora aurantiaca (Senghas & Gerlach) dressler, comb. nov. 5(2): 95. 2005. 

Ixyophora carinata (ortiz) dressler, comb. nov. 5(2): 95. 2005. 

Ixyophora viridisepala (Senghas) dressler, comb. nov. 5(2): 95. 2005. 

Lankesterella glandula ackerman, sp. nov. 4(1): 49. 2004. 

Lepanthes gerardensis M.a. blanco, sp. nov. 8: 19. 2003. 

Lycaste bruncana bogarín, sp. nov. 7(3): 543. 2007. 

Malaxis brevis dressler, sp. nov. 4(1): 97. 2004. 

Malaxis insperata dressler, sp. nov. 4(1): 97. 2004. 

Malaxis rostratula dressler, sp. nov. 4(1): 97. 2004. 

Malaxis triangularis dressler, sp. nov. 4(1): 97. 2004. 

Mapinguari carnevali & R. Singer, gen. nov. 7(3): 525. 2007. 

Mapinguari auyantepuiensis (Foldats) carnevali & R. Singer, comb. nov. 7(3): 525. 2007. 

Mapinguari desvauxianus (Rchb.f.) carnevali & R. Singer, comb. nov. 7(3): 525. 2007. 

Mapinguari foldatsianus (carnevali & i. Ramírez) carnevali & R. Singer, comb. nov. 7(3): 525. 2007. 

Mapinguari longipetiolatus (ames & c. Schweinf.) carnevali & R. Singer, comb. nov. 7(3): 525. 2007. 

Masdevallia vilcabambensis L. Valenz. & Suclli, sp. nov. 8(1): 17. 2008. 

Maxillariella M. a. blanco & carnevali, gen. nov. 7(3): 527. 2007. 

Maxillariella acervata (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella alba (Hook.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella anceps (ames & c. Schweinf.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella appendiculoides (c. Schweinf.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella arbuscula (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella brevifolia (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella caespitifica (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella cassapensis (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 


122 

LANKESTERIANA 

Maxillariella caucana (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella cobanensis (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella costaricensis (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella curtipes (Hook.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella densifolia (poepp. & endl.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella diuturna (ames & c. Schweinf.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella elatior (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella estradae (dodson) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella funicaulis (c. Schweinf.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella graminifolia (Kunth) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella guareimensis (Rchb.f) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella houtteana (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 528. 2007. 

Maxillariella infausta (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella lawrenceana (Rolfe) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella linearifolia (ames & c. Schweinf.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella longibracteata (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella luteorubra (F. Lehm. & Kraenzl.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella mexicana (J. t. atwood) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella microdendron (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella nitidula (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella oreocharis (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella pardalina (Garay) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella pastensis (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella ponerantha (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella procurrens (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella prolifera (Sw.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella purpurata (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella robusta (barb. Rodr.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella sanguinea (Rolfe) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella spilotantha (Rchb.f.) M. a. blanco & carnevali, comb. nov. 7(3): 529. 2007. 

Maxillariella stenophylla (Rchb.f. ) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella stictantha (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella tenuifolia (Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella tuerckheimii (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella variabilis (bateman ex Lindl.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella vinosa (Rolfe) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella vulcanica (F. Lehm. & Kraenzl.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella x yucatanensis (carnevali & R. Jiménez) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Maxillariella xanthorhoda (Schltr.) M. a. blanco & carnevali, comb. nov. 7(3): 530. 2007. 

Mormolyca acutifolia (Lindl.) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca aureoglobula (christenson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca chacoensis (dodson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca cleistogama (brieger & illg) M. a. blanco, comb. nov. 7(3): 531. 2007. 



Mormolyca dressleriana (carnevali & J. t. atwood) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca hedwigiae (Hamer & dodson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca lehmanii (Rolfe) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca moralesii (carnevali & J. t. atwood) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca pudica (carnevali & tapia-Muñoz) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca richii (dodson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca rufescens (Lindl.) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca sanantonioensis (christenson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca schlimii (Linden & Rchb.f.) M. a.blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca sotoana (carnevali & Gómez-Juárez) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca suarezorum (dodson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Mormolyca tenuibulba (christenson) M. a. blanco, comb. nov. 7(3): 531. 2007. 

Myoxanthus vittatus pupulin & M.a. blanco, sp. nov. 2: 16. 2001. 

Oerstedella x monteverdensis pupulin & Hágsater, sp. nov. 8: 32. 2003. 

Oncidium zelenkoanum dressler & pupulin, sp. nov. 8: 37. 2003. 

Ornithidium adendrobium (Rchb.f.) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium affine (poepp. & endl.) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium cachacoense (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium canarense (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium condorense (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium fasciculatum (c. Schweinf.) M.a. blanco & i. ojeda, comb. nov. 8(1): 15. 2008. 

Ornithidium fimbriatilobum (carnevali & G. a. Romero) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium gualaquizense (dodson) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium haemathodes (Ruiz & pav.) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium lasallei (Foldats) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium machinazense (d. e. benn. & christenson) M. a. blanco, comb. nov. 7(3): 532. 2007. 

Ornithidium maldonadoense (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 532. 2007. 

Ornithidium minutiflorum (d. e. benn. & christenson) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium nicaraguense (Hamer & Garay) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium oxapampense (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium patellum (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium patulum (c. Schweinf.) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium pseudonubigenum (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium pustulosum (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium rauhii (d. e. benn. & christenson) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium repens (L. o. Williams) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium rigidum (barb. Rodr.) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium scandens (d. e. benn. & christenson) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium scullianum (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium sillarense (dodson & Vásquez) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Ornithidium simplex (J. t. atwood) M. a. blanco & ojeda, comb. nov. 7(3): 533. 2007. 

Palmorchis eidae dressler, sp. nov. 3: 26. 2002. 

Paphinia vermiculifera G. Gerlach & dressler, sp. nov. 8: 27. 2003. 



Pescatorea ecuadorana (dodson) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea hemixantha (Rchb.f.) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea hirtzii (Waldvogel) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea lalindei (Linden) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea lawrenceana (Rchb.f.) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea pulvinaris (Rchb.f.) dressler, comb. nov. 5(2): 95. 2005. 

Pescatorea violacea (Lindl.) dressler, comb. nov. 5(2): 95. 2005. 

Phragmipedium andreettae p.J.cribb & pupulin, sp. nov. 6(1): 1. 2006. 

Phragmipedium manzurii W.e. Higgins & p. Viveros, sp. nov. 8(3): 89. 2008. 

Pleurothallis grammata dressler, nom. nov. 3: 28. 2002. 

Polycycnis blancoi G. Gerlach, sp. nov. 4(1): 67. 2004. 

Prosthechea micropus (Rchb.f.) W.e. Higgins, comb. nov. 4(3): 223. 2004. 

Prosthechea tardiflora Mora-Retana ex pupulin, sp. nov. 3: 23. 2002. 

Rhetinantha M. a. blanco, gen. nov. 7(3): 534. 2007. 

Rhetinantha aciantha (Rchb.f.) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha acuminata (Lindl.) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha cerifera (barb. Rodr.) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha divaricata (barb. Rodr.) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha encyclioides (J. t. atwood & dodson) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha friedrichsthalii (Rchb.f) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha mariaisabeliae (J. t. atwood) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha monacensis (Kraenzl.) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha neilii (dodson) M. a. blanco, comb. nov. 7(3): 534. 2007. 

Rhetinantha notylioglossa (Rchb.f.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Rhetinantha ophiodens (J. t. atwood) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Rhetinantha pastorellii (d. e. benn. & christenson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Rhetinantha schistostele (Schltr.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Rhetinantha scorpioidea (Kraenzl.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Rhetinantha witsenioides (Schltr.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea bomboizensis (dodson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea bomboizensis (dodson) M.a. blanco, comb. nov. 8(1): 15. 2008. 

Sauvetrea chicana (dodson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea cornuta (c. Schweinf.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea laevilabris (Lindl.) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea machupicchuensis (christenson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea napoensis (dodson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Sauvetrea sessilis (Lindl.) M. a. blanco, comb. nov. comb. nov. 7(3): 535. 2007. 

Sauvetrea trigona subsp. amaroensis (d. e. benn. & christenson) M. a. blanco, comb. nov. 7(3): 535. 2007. 

Scaphyglottis bidentata (Lindl.) dressler, comb. nov. 3: 28. 2002. 

Scaphyglottis cuniculata (Schltr.) dressler, comb. nov. 3: 28. 2002. 

Scaphyglottis imbricata (Lindl.) dressler, comb. nov. 3: 28. 2002. 

Sobralia gloriana dressler, sp. nov. 5: 11. 2002. 

Sobralia nutans dressler, sp. nov. 5: 13. 2002. 



Sobralia mariannae dressler, sp. nov. 5: 13. 2002. 

Sobralia crispissima dressler, sp. nov. 5: 10. 2002. 

Sobralia quinata dressler, sp. nov. 6: 27. 2003. 

Solenidium portillae dalström & Whitten, sp. nov. 6: 1. 2003. 

Stanhopea confusa G.Gerlach & beeche, sp. nov. 4(3): 217. 2004. 

Stanhopea manriquei Jenny & nauray, sp. nov. 4(2): 109. 2004. 

Stanhopea naurayi Jenny, sp. nov. 5(1): 77. 2005. 

Stelis megachlamys (Schltr.) pupulin, nom. nov. 4: 74. 2002. 

Stellilabium erratum dressler, sp. nov. 2: 11. 2001. 

Stellilabium smaragdinum pupulin & M.a. blanco, sp. nov. 5: 28. 2002. 

Stenia falcata (ackerman) dressler, comb. nov. 5(2): 93. 2005. 

Stenotyla dressler, gen. nov. 5(2): 96. 2005. 

Stenotyla lankesteriana (pupulin) dressler, comb. nov. 5(2): 96. 2005. 

Stenotyla lendyana (Rchb.f.) dressler, comb. nov. 5(2): 96. 2005. 

Stenotyla picta (Rchb.f.) dressler, comb. nov. 5(2): 96. 2005. 

Telipogon acicularis (dressler) n.H. Williams & dressler, comb. nov. 5(3): 168. 2005. 

Telipogon alexii n.H. Williams & dressler, nom. nov. 5(3): 170. 2005. 

Telipogon alticola (dodson & R. escobar) n.H. Williams & dressler, comb. nov. 5(3): 168. 2005. 

Telipogon anacristinae (pupulin) n.H. Williams & dressler, comb. nov. 5(3): 168. 2005. 

Telipogon andinus (L.o. Williams) n.H. Williams & dressler, comb. nov. 5(3): 168. 2005. 

Telipogon barbozae (J.t. atwood & dressler) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon bennettii (dodson & R. escobar) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon bergoldii (Garay & dunst.) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon boliviensis (R. Vásquez & dodson) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon boylei (J.t. atwood) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon bullpenensis (J.t. atwood) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon butchii n.H. Williams & dressler, nom. nov. 5(3): 169. 2005. 

Telipogon calueri n.H. Williams & dressler, nom. nov. 5(3): 170. 2005. 

Telipogon campbelliorum (J.t. atwood) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon distantiflorus (ames & c. Schweinf.) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon embreei n.H. Williams & dressler, nom. nov. 5(3): 170. 2005. 

Telipogon erratus (dressler) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon fortunae (dressler) n.H. Williams & dressler, comb. nov. 5(3): 169. 2005. 

Telipogon helleri (L.o. Williams) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon hystrix (dodson) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon ibischii (R. Vásquez) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon jostii (dodson) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon microglossus (Schltr.) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon monteverdensis (J.t. atwood) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon morganiae (dodson) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon morii (dressler) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon niri ackerman, sp. nov. 4(1): 48. 2004. 

Telipogon ortizii n.H. Williams & dressler, nom. nov. 5(3): 169. 2005. 



Telipogon pampatamboensis (dodson & R. Vásquez) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon perlobatus (Senghas) n.H. Williams & dressler, comb. nov. 5(3): 170. 2005. 

Telipogon pseudobulbosus (d.e. benn. & christenson) n.H. Williams & dressler, comb. nov. 5(3): 71. 2005. 

Telipogon reventadorensis n.H. Williams & dressler, nom. nov. 5(3): 171. 2005. 

Telipogon roberti n.H. Williams & dressler, nom. nov. 5(3): 171. 2005. 

Telipogon selbyanus n.H. Williams & dressler, nom. nov. 5(3): 171. 2005. 

Telipogon smaragdinus (pupulin & M.a. blanco) n.H. Williams & dressler, comb. nov. 5(3): 171. 2005. 

Telipogon tanii (dodson) n.H. Williams & dressler, comb. nov. 5(3): 171. 2005. 

Telipogon tsipiriensis (pupulin) n.H. Williams & dressler, comb. nov. 5(3): 171. 2005. 

Trichopilia x ramonensis J. García & Mora-Ret. ex c.o. Murales, nothosp. nov. 5: 18. 2002. 

Warczewiczella guianensis (Lafontaine, Gerlach & Senghas) dressler, comb. nov. 5(2): 96. 2005. 

Warczewiczella lobata (Garay) dressler, comb. nov. 5(2): 96. 2005. 

Warczewiczella palatina (Senghas) dressler, comb. nov. 5(2): 96. 2005. 

RHaMnaceae 

Krugiodendron acuminatum J. a. González & poveda, sp. nov. 8: 16. 2003. 

RUbiaceae 

Hoffmannia stephaniae L.a. González & poveda, sp. nov. 4(3): 183. 2004. 

RUtaceae 

Amyris magnifolia Gómez-Laur. & Q. Jiménez, sp. nov. 6: 5. 2003. 

ScRopHULaRiaceae 

Gibsoniothamnus ficticius J. F. Morales, sp. nov. 4(1): 2. 2004. 

SteRcULiaceae 

Byttneria osaënsis cristóbal, sp. nov. 4(3): 175. 2004. 

SyMpLocaceae 

Symplocos retusa Kriebel, sp. nov. 4(1): 57. 2004. 

Symplocos striata Kriebel & n. zamora, sp. nov. 4(3): 171. 2004.

VOL. 8, No. 3 - Lankesteriana - Universidad de Costa Rica

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