Front Cover

Title: Lankesteriana
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098723/00020
 Material Information
Title: Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Physical Description: v. : ill. (some col.) ; 25 cm.
Language: English
Creator: Jardi´n Bota´nico Lankester
Jardín Botánico Lankester
Publisher: Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Place of Publication: Cartago Costa Rica
Cartago Costa Rica
Publication Date: April 2008
Frequency: three times a year[2002-]
irregular[ former 2001]
three times a year
Subject: Botany -- Periodicals -- Costa Rica   ( lcsh )
Epiphytes -- Periodicals -- Costa Rica   ( lcsh )
Orchids -- Periodicals -- Costa Rica   ( lcsh )
Plantkunde   ( gtt )
Botanische tuinen   ( gtt )
Genre: periodical   ( marcgt )
Spatial Coverage: Costa Rica
Language: In English and Spanish.
Dates or Sequential Designation: No. 1 (mayo 2001)-
Numbering Peculiarities: Issues for May 2001-Oct. 2003 designated no.1-8; issues for Apr. 2004- designated vol. 4, no. 1-
General Note: Latest issue consulted: Vol. 4, no. 1 (abr. 2004).
General Note: International journal on orchidology.
 Record Information
Bibliographic ID: UF00098723
Volume ID: VID00020
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 48491453
lccn - 2001240973
issn - 1409-3871


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Full Text
ISSN 1.409-3871

Vot.. 8, No. I Apim 2008


.T Uri-
d., I



E I\TE RN ATI()NA L J I R N A L R C I i 1 1) L i Y

ISSN 1409-3871

VOL. 8, No. 1 ABRIL 2008

A new addition to the Costa Rican flora: Palmorchis nitida
(Orchidaceae: Neottieae) is documented from the Osa Peninsula

An Asian orchid, Eulophia graminea (Orchidaceae: Cymbidieae),
naturalizes in Florida

Generic realignments in Maxillariinae (Orchidaceae): corrigenda
et addenda

A new species of Masdevallia (Orchidaceae: Pleurothallidinae)
from Peru

Book review. Icones Pleurothallidinarum XXIX. A third century of Stelis
of Ecuador. Systematics of Apoda-Prorepentia. Systematics of miscellaneous
small genera. Addenda: new genera, species, and combinations,
by Carlyle A. Luer 21



Copyright C 2008 Lankester Botanical Garden, University of Costa Rica
Effective publication date: May 5, 2008

Layout: Jardin Botanico Lankester.
Cover: Labella of Palmorchis nitida Dressier, from original scans by C. Bainbridge.
Printer: Litografia Ediciones Sanabria S.A.
Printed copies: 500

Printed in Costa Rica / Impreso en Costa Rica

R Lankesteriana / The International Journal on Orchidology
No. 1 (2001)-- -- San Jose, Costa Rica: Editorial
Universidad de Costa Rica, 2001--

1. Botanica Publicaciones peri6dicas, 2. Publicaciones
peri6dicas costarricenses


IANKESTERIANA 8(1): 1-4. 2008.



Los Charcos de Osa Center for Regional Plant Diversity, Osa Peninsula,
Apdo. # 76-8203, Puerto Jimenez, Golfito, Costa Rica.
'Author for correspondence: cbainbridge@osaresearch.org

ABSTRACT: Palmorchis nitida, previously known from seasonally wet forests of Panama, is reported from
the Osa Peninsula in south-west Costa Rica. Notes and figures are provided regarding its natural history
and phenology. The variable nature of this species is discussed on the basis of recently collected Costa
Rican material.

RESUMEN: Palmorchis nitida, hasta ahora conocida solamente de bosques hfmedos estacionales de Panama,
se report como una especie nueva para la flora de Costa Rica. Se present nueva informaci6n sobre su
historic natural y se discute la naturaleza variable de esta especie con base a los nuevos especimenes
colectados en Costa Rica.

KEY WORD / PALABRAS CLAVE: Orchidaceae, Neottieae, Palmorchis, P. nitida, Costa Rica, Osa Peninsula,

After years of botanical sampling of the species-
rich forests of the Osa Peninsula, located in Costa
Rica's south Pacific, one might assume that nothing
new remained to be discovered. Just how wrong
this is, can be clearly demonstrated by the fact that
new registers and, indeed, new species, continue to
issue from the region, including areas that have long
served as choice collecting sites. One of the latest in
a growing list of surprising plant discoveries to come
out of the Osa Peninsula, is Palmorchis nitida Dressler
(Orchidaceae: Neottieae).

Although a relatively small neotropical genus 22
species are listed by the International Plant Names
Index as Dressler (1983, 1984) explains, spotting the
genus in its natural environment and, subsequently,
distinguishing between species, can be anything but a
straightforward task, as plants look enough like palm
seedlings or broad-leaved forest grasses that they
are easily overlooked when not in flower. With the
exception ofP. powellii, which displays a much larger
growth habit than that of any other species present in
either Costa Rica or Panama, (reaching up to im or

more), members of this genus are similar both in habit
and vegetative appearance. Even in the case ofP. nitida,
where the specific epithet refers to the brilliance of the
leaves, older leaves can appear dull due to the presence
of small mosses growing on their upper surface. Thus,
we depend on the presence of flowers to make anything
more than a generic determination. But, as plants in a
population all flower on the same day, and flowers last
only a few hours, one needs to monitor plants over an
extended time period in order to find flowers.
Dressler was surprised to discover that plants
growing by a well-used trail on Panama's Barro
Colorado Island that he had taken to be P. powellii
were a distinctive new species, P nitida (Dressler
1983, 1984). We were equally surprised to discover
that plants we had taken to be P. silvicola L.O.Williams
(1970) were P. nitida, a species from Panama.

We first observed plants on the Osa Peninsula in
Costa Rica in November 2004, growing in secondary
forest on the slopes of the property Los Charcos de
Osa. However, it was not until July 2006, that we were
lucky enough to finally observe, and photograph, the


FIGURE 1. A A single flower of P. nitida. B -Less commonly P. nitida produces several flowers simultaneously from more
than one inflorescence. C -Ajuvenille plant amongst the leaf litter. Photographs by Reinaldo Aguilar, 2006.

LANKESTERIANA 8(1), abnl 2008. 0 Universidad de Costa Rica, 2008.

BAINBRIDGE & AGUILAR Palmorchid nitida in Costa Rica

flowers of this species (Fig. 1). We then noted that our
plants conformed with the description of P. nitida as
presented in Dressler's (1993) dichotomous key and
with an incomplete description from the Manual de
Plantas de Costa Rica (Dressier, 2004) documented as
Palmorchis Sp. B., which cites a collection made by
Mike Grayum (Grayum et al. 4027, MO), as early as
1984. Probably, P. nitida is the "undescribed species in
the south Pacific lowlands of Costa Rica" that Dressier
(1997) referred to a decade ago.
The apparent leap in geographical distribution -
from eastern Panama to the relatively isolated south
west coast of Costa Rica is perhaps an indication
that P. nitida will prove to have a more continuous
distribution through central Panama and northwards
along the Pacific slopes. Dressier (1983, 1984) faced
a similar geographical disjunct when considering the
possible existence of P silvicola -originally described
from the Osa Peninsula in central Panama. We
anticipate that many more principally Panamanian
species are likely to show up on the Osa Peninsula in
the future.
We report here the first record of P. nitida for
the flora of Costa Rica on the basis of the following

COSTA RICA. Puntarenas: Osa Peninsula, Canton
de Osa, District of Sierpe, Finca Los Charcos
de Osa, 1km before the village of Banegas.
08040'18"N83030'17"W, 30 Nov 2006, C. V
Bainbridge 263 (USJ).

On the basis of fresh material from Costa Rica, we
have observed that the species can be variable. The
more typical form we have observed being compact
plants, of up to 30 cm tall. Less commonly encountered,
is the larger to more elongate form, superficially more
related to P. trilobulata (i.e., as can be seen in A.
Chac6n 931, INB), with a terminal leaf lamina of 25 x
8 cm and floral bracts which are well-spaced along the
rachis (Fig. 2).
Prior to collection, two plants of P. nitida were
monitored by us over a six month period and we
recorded flowers opening on the following dates:

SBainbridge 263, Example A: 23 Jul 2006, 13 Aug
2006, 27 Aug 2006, 17 Sept 2006, 20 Oct 2006, 18
Nov 2006.

FIGURE 2. Variability in vegetative form in Costa Rican
plants ofP. nitida. A -The more common and compact
form, similar to that ofP. silvicola. B -The less common
more elongated form, similar to that of P. trilobulata.
Photograph by Reinaldo Aguilar. 2006.

* Bainbridge 263, Example B: 29 Jul. 2006, 7 Sept
2006, 21 Oct 2006, 29 Oct 2006, 21 Nov 2006.

Herbarium records show that flowers were
produced in October (Grayum et al. 4027, MO)
and in July (A. Chac6n 931, INB) and buds were
also recorded as present in wild individuals at Los
Charcos on 27 Dec. 2006, 15 Feb. 2007, 15 Apr.
2007, and 15 May 2007. This information suggests
that the species may have an extensive flowering
period. A single mature fruit was collected on 20 Oct
2006. Cited material has been preserved in an alcohol
and glycerine mixture.

The following presents a brief list of associated
plants encountered in the habitat ofP. nitida on the Osa
Peninsula: Cyclanthus bipartitus (C il. illi.ic.... i.
Cryosophila guagara, Asterogyne martiana (Areca-
ceae), Heliconia irrasa, H. latispatha. H. imbricata
(Heliconiaceae), Costus stenophyllus (Costaceae),

LANKESTERIANA 8(1), abnl 2008. O Unversidad de Costa Rica, 2008.


Psychotria acuminata, P. elata (Rubiaceae), Calathea
crotalifera (Marantaceae), Miconia nervosa, M.
osaensis (Melastomataceae), Sarcoglottis hunteriana,
Stenorrynchos lanceolatum, Oeceoclades maculata
(Orchidaceae), Bauhinia bahiachalensis, B.
guianensis, Peltogyne purpurea (Caesalpinaceae),
Williamodendron glaucophyllum (Lauraceae),
Couratari scott-mori, C. guanensis (Lecythidaceae),
Newtonia suaveolens, Acacia alleni (Mimosaceae),
Coccoloba standleyana (Polygonaceae), Sloanea
sulcata (Elaeocarpaceae) and Chaunochiton kappleri

ACKNOWLEDGMENTS: Our thanks to Jorge Lobo, Luis
Diego Gomez, Franco Pupulin and Robert L. Dressier for
their valuable comments during the preparation of this

Dressier, R. L. 1983. Palmorchis in Panama mit einer
neuen Art, Palmorchis nitida, an einem unerwarteten
Standort. Orchidee 34(1): 25-31;
Dressier, R. L. 1984. Palmorchis in Panama: with a new
species where least expected. Orquidea (M6x.) 9(2)
Dressier, R. L. 1993. Field Guide to the Orchids of Costa
Rica and Panama. Cornell University. p.310.
Dressier, R. L. 1997. Orquideologia; Revista de laSociedad
Colombiana de Orquideologia 20(3): 263, f
Dressier, R. L. 2004. Orchidaceae. In: B. Hammel, M. H.
Grayum, C. Herrera & N. Zamora (eds.). Manual de
Plantas de Costa Rica. Missouri Botanical Garden Press.
Vol: 3
Williams, L. O. 1970. Palmorchis silvicola, Fieldiana,
Botany 32(12): 199.
International Plant Names Index (IPNI http://www.ipni.
org, electronic pages consulted 20h March, 2008).

IANKESTERIANA 8(1), abrl 2008. O Universdad de Costa Rica, 200&

LANKESTERIANA 8(1): 5-14. 2008.




'Fairchild Tropical Botanic Garden, 2121 SW 28" Terrace Ft. Lauderdale, Florida 33312
2Department of Biological Sciences, Florida International University, Miami, FL 33199
3Author for correspondence: Robert.Pemberton@ars.usda.gov

ABSTRACT. Eulophia graminea, a terrestrial orchid native to Asia, has naturalized in southern Florida. Orchids
naturalize less often than other flowering plants or ferns, butE. graminea has also recently become naturalized
in Australia. Plants were found growing in five neighborhoods in Miami-Dade County, spanning 35 km from
the most northern to the most southern site, and growing only in woodchip mulch at four of the sites. Plants at
four sites bore flowers, and fruit were observed at two sites. Hand pollination treatments determined that the
flowers are self compatible but fewer fruit were set in selfed flowers (4/10) than in out-crossed flowers (10/10).
No fruit set occurred in plants isolated from pollinators, indicating thatE. graminea is not autogamous. Pollinia
removal was not detected at one site, but was 24.3 % at the other site evaluated for reproductive success. A total
of 26 and 92 fruit were found at these two sites, where an average of 6.5 and 3.4 fruit were produced per plant.
These fruits ripened and dehisced rapidly; some dehiscing while their inflorescences still bore open flowers.
Fruit set averaged 9.2 and 4.5 % at the two sites. No floral visitors were seen during limited (6.5 hr duration)
timed watches of flowers. Individual flowers are open an average 11 days, and the inflorescences may bear
flowers for at least one month. How E. graminea entered Florida is unknown, but capsules, bulbs and plantlets
in flasks are available for sale and/or trade via the interest from Thailand and other places outside the U.S. The
occurrence of the orchid at 900-1000 m elevation its native Sikkim (27-28N) and in Kashmir (above 32N),
suggest that it can live well north of current area of naturalization in southern Florida, which is at sea level
and 26N latitude.

KEY WORDS: breeding system, Eulophia, orchid, mulch, naturalization, pollination

A terrestrial orchid, Eulophia graminea Lindl.,
native to subtropical and tropical Asia, has been detected
in Miami-Dade County in southeastern Florida (Fig.
1-2). Its occurrence is noteworthy because relatively
few orchids naturalize.
In the native region, Eulophia graminea occurs
widely, from South Asia and Southeast Asia to
eastern Asia to subtropical islands in the western
Pacific. More specifically, it has been reported from
Pakistan (Singapur), Kashmir, eastwards to Burma,
Thailand, Malaysia, also Sri Lanka, Nikobar Isl.,
Philippines, Taiwan (Renz 1984). It also occurs in
Singapore (Keng et al. 1989), the Himalayas (Nepal
and Sikkim) from 900-1000 m (Singh 2001), Laos
and Indonesia (GRIN 2008), China (Guangdong,
Guangxi, Guizhou, Hainan, Yunnan; and Hong Kong)

html), the Ryuku Islands (Japan) (Garay & Sweet
1974) and Guam (GRIN 2008).
Within its native range Eulophia graminea occurs
in diverse habitats. In Singapore it is reported to be
common onwasteland, sandy beaches, lawns, roadsides
and other exposed areas (Keng et al. 1998), as well as
open areas in secondary forests and parks (Tan 1993).
In Taiwan, the orchid grows among shrubs in lowlands,
on beaches, coastal grassland (Su 2000). In the Ryukus,
it is uncommon in mountains and open low areas
(Garay and Sweet 1974). The species is threatened in
Sri Lanka (http://www.environmentlanka.com/biodiv/
Eulophia graminea was discovered to be
naturalized in Australia in the Northern Territories
in 2001 il .ip ..1..!! v.au/animal-plant-
health/pests-diseases-weeds/weeds/incursion). In the


Northern Territories the orchid is found in and around
Darwin growing in woodchip mulched garden beds
(Macrae 2002), and also "found pushing its way
through bitumen and concrete" (Hussey & Lloyd
2002). It has also recently reported from Queensland,
Australia (Pier 2008).
This study was undertaken to define and better
understand the orchid's occurrence, its reproduction,
potential to persist and spread in Florida and beyond.


l. i,,, ir.i,. ., The identity of Eulophia graminea
was confirmed with technical descriptions in Flora
of Taiwan (Editorial Committee of the Flora of
Taiwan, 1993, Flora of Pakistan (Renz 1984), and
Flora of the Ryuku Islands (Garay & Sweet 1974),
and by Surangraj Indhamusika, of the Queen
Sirikit Botanical Garden in Chiang Mai Thailand.
To distinguish this orchid from related Florida
orchids, published accounts of the morphological
characteristics of Eulophia graminea and those
orchids were compared. Herbarium specimens
(Pemberton 07-01, 07-02, 08-01, 08-02, 08-03) have
been placed in the herbaria of Fairchild Tropical
Garden (FTG) in Coral Gables, the Florida Museum
of Natural History (FLAS) in Gainesville, and the
University of South Florida (USF) in Tampa.

Occurrence After the orchid was first discovered in
a residential garden in South Miami, its occurrence
was determined by surveying the neighborhood
to find plants and counting and describing the
encountered plants. We interviewed people
associated with yards where the plants were found to
learn when the plants were first observed, and what
other information they might have, pertaining to the
possible origin of the orchids. We surveyed parks
and mulched sites within a mile of the South Miami
discovery site for additional plants. Interviews and
discussions with colleagues led to the discovery
of four additional sites with the orchid. Sites in
western Kendall, Little River Miami, and North
Miami were surveyed to confirm the occurrence and
circumstance. Information about the eastern Kendall
occurrence, including photographs of the plant and
site, was provided by the reporting botanist Steve

LANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 2008.

Breeding System and Pollination Eulophia graminea
plants and flowers were observed and manipulated
to gain insights into the orchid's breeding system
and pollination ecology. We examined flowers to
detect rewards and inflorescences to discern odors.
To assess male and female reproductive success, we
measured fruit formation and pollinia removal in the
South Miami and Little River Miami populations
where multiple reproductive plants occurred. Pollinia
removal was scored during single visits in October
and November, 2007 in the South Miami population,
and during one visit during April 2008 to the Little
River Miami population. Fruit set in South Miami
plants was determined by counting the number of fruit
or fruit remnants, and the number of persistent bracts
that subtended the flowers on old inflorescences on
three plants during March, 2008, after the plants had
flowered. Fruit set in the Little River Miami plants
was determined by counting the number of fruit or
fruit remnants, and the number of persistent bracts
that subtended old or fallen flowers on 27 plants
during April, 2008, near the end of flowering period
for most of plants. Timed watches were made during
October, November 2007, January and April 2008 to
observe flower visitors. Flowers at the South Miami
site were hand pollinated with self pollinia and crossed
using pollinia from flowers of other plants at the same
site and date to determine whether the plant is self
compatible, and to compare self and outcross fruit set.
Potted plants raised from bulbs dug at the South Miami
site were placed inside a screen enclosure when their
inflorescences appeared to determine the ability of the
flowers to set fruit without pollinators, which would
indicate either autogamous self pollination or apomixis.
Selected flowers on these inflorescences were tagged
as soon as they opened and followed until wilting to
determine their longevity. Potted flowering plants were
exposed in Ft. Lauderdale and Homestead gardens to
increase observations of potential pollinators and to
access the Eulophia graminea's ability to set fruit at
other locations other than the South Miami and Little
River Miami naturalization sites.

Possible Routes of Introduction Internet searches
were made to find evidence of the plant's cultivation,
presence in commerce, or other human aided
movement. The importation of commodities or other

PEMBERTON et al. Eulophia graminea naturalization

FIGURE 1. Eulophia graminea flowers (ca. 2.5 cm wide). FIGURE 2. Whole flowering plant of the naturalized Eulophia
Note the nearly horizontal position of the lateral sepals graminea growing in woodchip mulch in South Miami,
and the nectar spur. Florida. Note the capsules.

bulk materials that may have carried propagules (bulbs
and/or seed) of the orchid from the native region to the
U.S. was investigated by examining U.S. Department
of Agriculture importation data posted on the Foreign
Agriculture Service website.


Description and distinguishing characteristics
Plants terrestrial and autotrophic. Pseudobulbs ovoid
with 4-5 concentric intemodes, initially covered with
many ovate scales, to 10 cm wide but usually 2.5-
5.0 cm wide, occurring singly or in clusters, usually
partly above the ground (Fig. 3). Leaves 2-5, grass-
like, suberect. Flowers in or out of leaf. Inflorescences
30 to 150 cm tall bearing up to 60 flowers (Fig. 2).
Flowers typically 2.5 cm wide (Fig. 1), with sepals and
petals dull green with brownish purple netted veins;
sepals spreading with lateral sepals held slightly above
horizontal; petals pointing forward above the column;
lip three lobed with two short lateral lobes green with
purple striping and rolled inward to the column to
form a short tube, median lobe white marked with
rose-pink, elongate extending forward and downward,

wavy-crisped along margin, and turning into white and
rose papillae; spur at base of lip long, dilated at tip;
Fruit a capsule ca. 3 cm long by 0.8 cm wide when
fully grown (Fig. 4).
Only one other Eulophia species, the native E.
alta (L.) Fawc. & Rendle, occurs in Florida and the
United States (Romero-Gonzalez 2007). Luer (1972)
included Eulophia ecristata (Femald) Ames in "The
Native orchids of Florida", but this species is now
Pteroglossaspis ecristata (Femald) Rolfe. These three
orchids are readily separated by the symmetry of both
the inflorescences and the individual flowers, as well
as flower color. Pteroglossaspis ecristata flowers are
borne on a raceme and have yellow green petals and
sepals converging over the brown purple lip. Eulophia
alta flowers are in racemes and are green to purplish
(but color can be quite variable), with lateral sepals held
erect and spreading at 45 degree angles. E. graminea
has green flowers with white lips marked with rose
borne on racemes or panicles, with the lateral sepals
spread widely and held almost horizontally. Eulophia
graminea flowers are also smaller than those ofE. alta
(1.4 to 2.6 cm vs. 3.5 to 4.5 cm wide). Although the
plants of all three species can be a meter or more in

LANKESTERIANA 8(1), abnl 2008. O Unversidad de Costa Rica, 2008.


FIGURE 3. Eulophiagraminea plants pulled from a mulch bed. FIGURE 4. Fruit of Eulophia graminea. The capsules (ca.
Note the distinctive nodes on the large primary pseudobulbs 3 cm long x 0.8 cm wide) ripen and dehisce quickly,
(ca. 3.5 cm in diameter), the uppermost nodes with paper allowing the spread of seed while plant is still in flower.
sheaths, and the proliferation of small pseudobulbs from
the bottom and side of the primary pseudobulbs.

height, E. graminea is more slender and has narrower
leaves (1-1.5 cm vs. up to 10 cm wide inE. alta and 3.5
cm in P. ecristata) than the other two species.

Occurrence in Florida Eulophia graminea plants
were found in small numbers in five different areas
of Miami-Dade County in southeastern Florida. The
discovery occurred when three flowering plants of an
unknown orchid growing in the yard of a neighbor were
brought to one of us (SK) during September 2007. We
investigated this South Miami yard on October 5 and
found 17 bulbs of the orchid growing in woodchip
mulch beneath shrubs and small trees. The property
owner first observed a large flowering plant in the yard
"a year or more earlier." He first observed a single large
plant that flowered and produced fruit. Subsequently,
he noticed many bulbs of the orchid, prompting him
to remove "several dozen." He was alarmed at the
proliferation of the plants and suspected that species
could be invasive. Our survey of this South Miami
neighborhood located bulbs in three other yards within
the same block as the reporting neighbor. In the second

IANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 200&

yard with the orchid, a single bulb was growing in
mulch beneath an avocado tree. The third yard had one
plant with three bulbs in mulch beneath a coconut and
two other plants in a mulched, partially shaded, flower
bed. The fourth yard had four large flowering plants and
two other plants with two large non-flowering bulbs.
All were growing in mulch in partial shade of palms
and other trees, except for one flowering plant in an
open mulched bed. The four flowering plants bore tall
(0.7 to 1. 5 m.), slender inflorescences in various stages
of flowering and fruiting. On these four plants, the
number of inflorescences, open flowers and fruits were
as follows: 1)4 inflorescences, 15 flowers, 8 fruits; 2) 10
inflorescences, 82 flowers, 18 fruit; 3) 1 inflorescence,
15 flowers, no fruit; 4) 1 inflorescence, 15 flowers,
no fruit; and an average of 6.5 fruit per plant. This
homeowner with the flowering plants was influenced
by the reporting neighbor to pull and dispose of bulbs
in the garbage, but decided to leave some flowering
individuals. He also obtained mulch from the reporting
neighbor, which could have contained bulbs or seed
of the orchid. Pseudobulb size ranged from 1 to 10

PEMBERTON et al. Eulophia graminea naturalization

cm; we also observed extensive vegetative production
of small bulbs beneath, or adjacent to, the primary
pseudobulbs. The large pseudobulb of one plant, which
grew partly beneath a heavy concrete stepping stone,
lifted the stone ca. 5 cm above the mulch, as it grew
and expanded. Some large inflorescences had unopened
buds on their upper portions and ripe dehiscing fruit on
the lower sections. Surveys of parks and mulched areas
within approximately one mile of this South Miami
occurrence failed to locate additional plants.
The second occurrence of E. graminea was in
a residential neighborhood in west Kendall, ca. 8.5
km southwest of the South Miami population. The
brother of the homeowner (with flowering plants at
the South Miami site) has a landscape maintenance
business, and suggested that we check the yard of
a client because he thought that the orchid might
occur there. A single tight cluster of pseudobulbs
was found in a mulched bed at the house. A survey
of this neighborhood resulted in an additional large
pseudobulb with leaves growing in mulch in a yard
about two blocks to the west of the first house.
The third occurrence of E. graminea was reported
to us by a botanist colleague Steve Woodmansee, who
found a plant growing in mulch in his yard in eastern
Kendall. This site is ca. 2.4 km west of the South Miami
population, and between the South Miami population and
the west Kendall occurrence. He first observed the plant
in 2006. No flowers were observed until an inflorescence
appeared in December, 2007. Flowers opened in mid
January, 2008 on a single inflorescence, 0.38 m tall,
and borne on a bulb 4 cm wide. About 30 flowers were
produced but no fruits formed on the plant.
The fourth occurrence of E. graminea was
reported to us by orchid grower Donald Wallstedt, who
discovered a population of the orchid growing in the
Little River area of Miami. This site is 19.4 km north
of the South Miami population and 31.1 km north of
the southern most site in western Kendall. Wallstedt
first observed the plants during March 2008. The
plants were found growing on adjacent vacant lots
amidst grass and Bidens pilosa L. on shallow sandy
limestone soil scattered with concrete, tile and other
debris. The orchids occurred mostly in the open partly
sunny central area of the two lots, where houses stood
before being torn down during the summer of 2005
(Donald Wilson, pers. comm.). Because the site is due

to be developed, Wallstedt removed 40 of the largest
flowering plants to save them, planting most in his
nearby garden and giving others away. Fifteen of the
transplanted plants in pots in his garden had exposed
pseudobulbs, and these averaged 5.2 cm (sd. = 1.8)
in diameter. One unusually large plant had a mass
of pseudobulbs 35 cm long by 11.3 cm wide, and
bore 12 inflorescences. During a survey of the Little
River Miami naturalization site on April 5, 2008, 29
additional plants including 27 reproductive (flowering
and/or fruiting) plants were found. These plants had
a mean of 1.26 (sd. = 0.447) inflorescences ranging
from 0.3-0.9 m tall with a mean height of 0.45 m (sd.
= 0.141). Twenty one plants had a total of 103 open
flowers and a mean of 5.38 (sd.= 3.38) flowers per
plant. Twenty seven reproductive plants had a total of
92 fruit on 23 different plants, ranging from 0-9 fruit,
and an average of 3.41 (sd. = 2.63) fruit per plant.
The fifth and last site where Eulophia graminea
was detected is in North Miami, which is north of the
Little River Miami site, and 24.3 km north of the South
Miami site and 34.9 km north of the western Kendall
site. Ronald Pereira found the plant and directed us
to the site where we found a single plant growing in
woodchip mulch in a Publix supermarket parking lot
island. The inflorescence of the plant was 0.3 m tall
and bore three old and three fresh flowers but no fruit.

Breeding system and pollination Eulophia graminea
flowers have a nectar spur containing nectar. We found
the nectar to have >2% glucose as indicated by Diastix
R (Bayer) glucose test strips used on three flowers.
No fragrance could be detected in single flowers, but
we were able to detect fragrance from inflorescences
bearing many flowers during the morning. Flower
longevity (determined with 15 flowers on two potted
plants held inside a screen enclosure to exclude
potential pollinators, and tagged the first day they
opened and followed until they wilted) was 5-16 days,
with a mean longevity of 10.9 (sd.= 3.4) days. None of
these tagged 15 flowers, or the other ca. 15 untagged
flowers on these potted plants in the enclosure, set
fruit, indicating the orchid is probably not autogamous.
Hand pollination treatments with self pollinia resulted
in fruit formation in 4/10 flowers, indicating a degree
of self-compatibility. All 10 flowers hand pollinated
with out-cross pollinia set fruit.

IANKESTERIANA8(1), abnl 2008. O Universidadde Costa Rica, 2008.


Flowers on the two flowering plants scored at the
South Miami site for pollinia removal on Oct. 14 had
no pollinia missing (0/15 and 0/82). Two flowers had
pollinia partly out of the anthers, and one of these
almost touched the stigma. The three flowering plants
examined for pollinia removal on Nov. 10 had no
missing pollinia in a total of 28 flowers (0/4, 0/11,
0/13). Dissections of 20 wilted flowers from the site
found two pollinia on the stigmatic surface and pollinia
of many other flowers hanging from below the anther
cap. One hundred thirteen flowers evaluated on 21
plants at the Little River Miami site on April 5, 2008,
had an average pollinia removal of 24.3 % (sd. = 25.6).
Three potted plants with 86 flowers exposed in a garden
in Ft. Lauderdale failed to produce fruit, but two fruits
were produced on a potted plant exposed in a garden
in Homestead that had ca. 15 flowers. The percent fruit
set of the flowers on the 13 inflorescences on the three
plants growing at the South Miami site was: 4.9 % (4
fruit/81 flowers), 2.3% (3 fruit/133 flowers), and 6.0%
(4 fruit/65 flowers), and a mean of 4.45% (sd. = 1.99).
The mean fruit set for 27 reproductive plants growing
at the Little River Miami site was 9.18 % (sd. = 7.14)
per plant.
Aside from ants (Brachymyrmex obscurior
Forel), encountered in some flowers at the South
Miami site, no flower visitors were observed during
limited timed watches (in periods between 8:45am
and 4:20pm on Oct. 14, Nov. 4 and Nov. 10, 2007,
Feb. 2, and April 5, 2008) totaling 390 minutes on
188 flowers on 12 plants.

Possible Introduction Routes The means by which
the orchid entered Florida and the U.S. is unknown.
Although the flowers are not showy compared to
most cultivated orchids, there is some cultivation
of Eulophia graminea and commerce in the plants.
Directions for its cultivation in Singapore are given
by Tan and Sin (1993), but with the comment that the
orchid is not very attractive. Several current postings
on eBay (Feb.-March 2008) offered plants for sale.
A Thai orchid grower (Kasom Orchids in Bangkok)
offered flasks containing 30-40 plantlets for $12/flask.
Another Thai grower (LONUN-Thailand in Bankok)
listed packets of five E. graminea bulbs for $8, and
someone in Scotland offered bulbs for sale. There was
a website posting on the Terrestrial Orchid forum from

LANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 2008.

a man in Germany requesting advice for the cultivation
of E. graminea bulbs that he acquired from Thailand.
A garden website had a posting from someone in
Singapore (part of the native range), offering to trade
an E. graminea capsule (www. Gardenbanter.co.uk).
Also of interest, because of the orchid's naturalization
in Australia, was the selling of the E. graminea by
the Australian eBay Australian store, by another Thai
seller (Thai Tropicals), which currently offers 10 bulbs
for ca. 2.14 Australian dollars.
Another possible introduction route could have been
the importation of bulk raw materials from the orchid's
native region in Asia containing its seed or bulbs. The
bulk material that seemed to have the potential to
carry the orchid material is woodchip component of
the "Logs and woodchips" import category, and these
were imported to the U.S. from Eulophia graminea's
native range (Malaysia, Indonesia, Singapore, China,
Taiwan) (http://www.fas.usda.gov/scriptsw/bico/bico.
asp?Entry=lout&doc=967). It is not known, however,
what portion of this category woodchips represents, if
the woodchips are treated prior to export or after import
in a manner that would kill orchid propagules, or where
these imported woodchips end up in the United States.
No evidence for imported mulch was found except
possibly the woodchips if used for mulch.


Occurrence -It is highly likely that other undetected
plants occur in Miami-Dade County and perhaps
beyond. All five Eulophia graminea occurrences or
populations were found with the assistance of other
people. Our surveys to try to find new occurrences have
been limited, and we have not investigated some of our
local habitats of the same types the orchid occupies in
its native area, such as beach, coastal grasslands, and
secondary forest.

Possible routes of introduction Although no evidence
for cultivation or commerce of the orchid in the U.S.
was discovered, we found that the orchid is cultivated
overseas and is available as seed, bulbs, and plantlets
in flasks by web-based businesses and private
individuals. This suggests that importation to Florida
could easily occur, although CITES restrictions on
orchid importation might restrain and limit its legal
importation. But both legal and illegal importation could

PEMBERTON et al. Eulophia graminea naturalization

have occurred. The availability of Eulophia graminea
is coupled with considerable enthusiasm for orchid
growing in southern Florida. The American Orchid
Society alone has ca. 25 affiliated chapters in the three
highly populated counties (Broward, Miami-Dade, and
Palm Beach) in southeastern Florida. The importation
of bulk materials from the native area, which could
introduce E. graminea seed or pseudobulbs, is another
possible introduction route, but less plausible than
purposeful importation for horticulture.

Breeding System and Pollination Although we
determined that Eulophia graminea needs a pollinator
for pollination, we did not identify the pollinator(s).
Fruit set was observed at the South Miami and Little
River Miami naturalization sites, and in an exposed
potted plant in Homestead. No fruit was seen in the
flowering plants at the east Kendall and North Miami
naturalization sites, or potted plants exposed in Ft.
Lauderdale in Broward County, just to the north of
Miami-Dade County. At the South Miami site, pollinia
removal was not observed during the two scoring
periods during October and November, but a low level
of fruit was present during and after those assessments;
with a 4.5 % fruit set detected after flowering finished.
At the Little River Miami site, pollinia removal was
documented (with a mean of 24 %), and a mean fruit
set was 9%. Perhaps the greater number of flowering
plants at the Little River Miami site, ca. 70 compared
to 4 at the South Miami site, may have resulted in
both more flower visitation and increased chances for
reproductive success. In addition, the much greater
presence of flowering weeds such as B. pilosa at the
Little River Miami site could have also increased
pollinator presence and activity. In other studies of
tropical orchid pollination (Tremblay et al. 2005),
pollinia removal rates were usually higher than fruit
set rates. Pollinia removal rates were, however, higher
in deceptive tropical orchids, but not in rewarding
tropical orchids (Tremblay et al. 2005) such as E.
graminea. Because our visits and observation periods
were few, we could have missed the activity periods
of pollinators responsible for the fruit set. It might
also be possible, despite the absence of fruit set in
enclosed plants or in exposed plants in Ft Lauderdale,
that some low level autogamy occurs. The presence of
nectar and fragrance in the daytime, the prominent lip

with coloration contrasting coloration, papillae, and a
gullet (formed by the lateral lobes of the lip) suggest
pollination by bees. More observations are needed to
detect the pollinator(s) and additional treatments to
determine whether a limited amount autogamy might
be possible in plants in the field.

Potential to persist The underground (or partially
underground) pseudobulbs, particularly those buried
in mulch, probably provide a means to survive the
dry season and drought. Mulch is widely utilized in
southern Florida, in part, to conserve moisture for
garden plantings. Part of the plant's native range (South
and Southeast Asia) experiences an intense dry season
with higher temperatures than occur in Florida's dry
season. During these times the leaves dry and the plant
become dormant (Tan & Sin 1993). The pseudobulb,
particularly those that are underground, may be better
able to survive the rare freeze events southern Florida
experiences, and may allow it to persist farther north in
Florida, and perhaps beyond.

Potential to spread Eulophia graminea has capsules
which ripen and dehisce quickly, suggesting that its
spread by seed is likely. The occurrence of multiple
plants in relatively small areas of three neighborhoods
also suggests local spread by air-bome seed. The
occurrence in five separated areas of Miami-Dade
County might also be due to wind blown seed, but
spread by the movement of seeds or pseudobulbs
in mulch is also possible. We know that mulch was
shared from the first yard in the South Miami (where
the orchid was found) with another neighbor where
we also found the plants. The ability of the orchid to
persist as dormant pseudobulbs can probably facilitate
their survival during movement with mulch.
Eulophia graminea occurs beyond the tropics in
some parts of its native range. The northeast part of
its range in Okinawa (Garay & Sweet 1974) is just
above 260N, which is the same latitude as Miami-Dade
County. The orchid lives in the Himalayas between
900-1000 m in Nepal and Sikkim) (Singh 2001). Nepal
lies between 26-320N, but areas at or below 1000 m
lie in the lower latitudes of the country. Sikkim lies
between 27 and 280N. The orchid's occurrence in
Kashmir (Renz 1984), an elevated region, whose
southern border lies above 32oN, is the region with
the most temperate climate in the plant's native range.

LANKESTERIANA 8(1), abnl 2008. 0 Unversidad de Costa Rica, 2008.


One inland Chinese location for the plant is in Anhui
province is ca. 320N (Chinese distribution map for the
species, Tropicos, Missouri Botanical Garden), but
this locality is unconfirmed. The occurrence of the E.
graminea at higher latitudes and higher elevations than
the 26N and near sea level occurrence in southern
Florida, suggests that the plant could survive well north
of its current area of naturalization. If the orchid is able
to live at 320N in the southeastern U.S., it could occur
from southern Georgia to eastern Texas. The source
area of the plants that have naturalized in southern
Florida is, however, unknown, and the naturalized
plants may not necessarily have the cold tolerance of
plants occurring in the more northern parts of the native
range. The large native geographic range, including its
occurrence on isolated islands, suggests an ability to
disperse and to spread widely. Eulophia graminea was
one of the first orchids to colonize Krakatau Islands
after the 1883 massive eruption, being detected in
1908 (Partomihardjo 2003).
This combination of the orchid's more northern
occurrence in its native range, its ability to seed
quickly and disperse over great distances, vegetative
proliferation of its pseudobulbs, and terrestrial
pseudobulbs which can be dormant, all suggest that
Eulophia graminea will probably spread in Florida and
possibly north of Florida to Georgia the Carolinas. The
orchid could spread to the West Indies by windborme
seed. It could also be moved accidentally (as
pseudobulbs or seed) as contaminants of horticultural
potted plants exported from Florida to warmer parts
of the US and to the West Indies. Eulophia graminea
could also be moved intentionally by people for
We found the orchid growing in mulch at four of the
five sites in southern Florida; it has also naturalized in
mulch in Australia. All of the detected plants in mulch
were found in woodchip mulch, although the roots of
some plants may have been growing into soil beneath
the mulch. Mulch might have the mycorrhizae needed
by the orchid for seed germination and establishment.
Mulch harbors many fungi, and alien fungi have been
first detected in ornamental woodchip mulch (Shaw et
al. 2004). The orchid is reported to have a wide tolerance
of environmental conditions (Tan & Sin 1993), and its
occurrence on sandy beaches, grasslands, and other
shrubby, open areas, and secondary forests in its native

LANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 2008.

area, indicate that it could colonize these habitat types
in Florida. The habitats the orchid colonizes will
probably depend on whether suitable mycorrhizae
occur in them.

Orchid naturalization Only 11 species of orchids
are naturalized in Florida (Wunderlin and Hansen
2004). With the addition of Eulophia graminea, this
is around 10% of the total orchid species in the state.
This indicates that naturalization of orchids in Florida
occurs less frequently than either other flowering
plants (ca. 33% naturalization) or ferns (ca. 33%)
(Pemberton 2003). Lower naturalization rates in
orchids appears to be a general occurrence (Daehler
1998). Eleven of 12 of the orchids naturalized in
Florida are terrestrial species, suggesting that terrestrial
species may naturalize more easily than epiphytes.
This is noteworthy because epiphytic orchids
dominate horticulture and home cultivation in Florida,
although a couple of terrestrial species, Spathoglottis
plicata Blume and Phaius tancarvilleae (Banks ex
L'Her.) Blume, are commonly sold. Both orchids are
naturalized in Florida to a limited extent (Wunderlin
& Hansen 2008). The lack of specialist pollinators
and appropriate mycorrhizae are thought to limit
orchid naturalization (Daehler 1998). Terrestrial
environments, particularly mulch, may provide more
potential mycorrhizae needed by orchid seed to
germinate than do the arboreal habitats of epiphytic
orchids. An invasive orchid, Disa bracteata Sw., and a
weedy native orchid, Microtis media R.Br., inAustralia
form associations with a greater array of mycorrhizal
fungi of more world-wide occurrence than do non-
weedy native orchids (Bonnardeaux et al. 2007).
While the breeding system of colonizing orchids
may be diverse (Sun 1997), the only wide-spread
naturalized orchids in Florida, Oeceoclades maculata
(Lindl.) Lindl. and Zeuxine strateumatica (L.) Schltr.),
are thought to be autogamous (Sun 1997, Gonzalez-
Diaz &Ackerman 1988). This supports, in part, the idea
that pollination limitation may be a barrier to orchid
naturalization. The recent naturalization of two tropical
bees in southern Florida, an orchid bee (Pemberton &
Wheeler 2006), and an oil-collecting bee (Pemberton
& Liu 2008), may reduce this pollinator limitation
and both promote some already naturalized orchids
and facilitate the naturalization of some ornamental

PEMBERTON et al. Eulophia graminea naturalization

orchids. The orchid bee, Euglossa viridissima Friese,
visits many different ornamental orchids (Pemberton
2007a), and is pollinating and causing considerable
fruit set in Guarianthe skinneri (Bateman) Dressler
& W.E. Higgins (Pemberton 2007b). The oil-
collecting bee, Centris nitida Smith, is pollinating
and inducing fruit set in the ornamental Oncidium
sphacelatum Lindl. (Pemberton, in press), and is the
only known pollinator of the invasive cowhom orchid,
Cyrtopodium polyphyllum (Vell.) Pabst ex F. Barrios,
(Liu and Pemberton, unpubl. data). It is doubtful that
either of these naturalized bees visits the flowers of
Eulophia graminea because they are quite unlike the
flowers used by them.
Eulophia graminea is probably destined to be a
permanent part of Florida's flora, given its multi-site
and relatively wide occurrence during the early phase
of its colonization, as well as its ability to produce
ample fruit and persist during the dry season via its
dormant pseudobulbs. It will be interesting to observe
the extent of this alien orchid's colonization and spread
in the New World and to try to understand the reasons
for its successful naturalization.

ACKNOWLEDGEMENTS. Surangraj Indhamusika, of the
Queen Sirikit Botanical Garden in Chiang Mai Thailand,
confirmed the identity of Eulophia graminea, Stig Dalstrom
of the Orchid Identification Center at Marie Selby Gardens
and Mario Blanco of University of Florida assisted in
the identification of the orchid. Harvey Bernstein first
brought plants of an unknown orchid to SK and shared his
observations of the plants with us. Tom Dickinson suggested
the possible occurrence of the orchid at the west Kendal
site. Steve Woodmansee shared his discovery of a plant
in east Kendall site. Donald Wallstedt and Ronald Pereira
shared their discoveries of plants at Little River Miami and
North Miami sites, respectively. We appreciate the Harvey
Bernstein and the Gordon Lee Dickinson families for
allowing us to study E. graminea in their yards, and realtor
Donald Wilson for permission to study the plants at the
Little River Miami site and for information about the site.

Bonnardeaux, Y, M. Brundrett, A. Batty, K. Dixon, J. Koch
& K. Sivasithamparam. 2007. Diversity of mycorrhizal
fungi of terrestrial orchids: compatibility webs, brief
encounters, lasting relationships, and alien invasions.
Mycolog. Res. 111: 51-61.

Daehler, C.C. 1998. The taxonomic distribution of invasive
plants: ecological insights and comparison to agricultural
weeds. Biol. Conserv. 84: 167-180.
Garay, L.A. & H.R. Sweet. 1974. Orchids of southern Ryuku
Islands. Harvard Univ. Press.
Gonzalez-Diaz, N. & J.D. Ackerman.1988. Pollination and
seed production in the orchid Oeceoclades maculata.
Lindleyana 3:150-155.
GRIN (Germplasm Resources Information Network)
Taxonomy of plants. 2008. http://www.ars-grin.gov/
cgi-bin/npgs/html/taxgenform.pl (last accessed March,
Hussey, B.M.J. & S.G. Lloyd. 2002.Western weeds, additions,
deletions and name changes. Western Australian
Government. http://members.iinet.net.au/-weeds/ww
update.pdf (last accessed February, 2008).
Keng, H., S.C. Chin & H.T.W. Tan. 1998. The concise floral
of Singapore, Vol. II Monocots. National University
Singapore Press. 234 p.
Luer, C. 1972. The native orchids of Florida. The New York
Botanical Garden, New York.
Macrae, C. 2002. New Weed Found in Darwin. Department
of Business, Industry and Resource Development,
Primary Industry and Fisheries, Northern Territory
Government, Australia. (last accessed 31 January
Partomihardjo, T. 2003. Colonisation of orchids on the
Krakatau Islands. Telopea 10: 299-310.
Pemberton, R.W. 2003. The common staghorn fern,
Platycerium bifurcatum, naturalizes in southern Florida.
Amer. Fern J. 93: 204-207.
Pemberton, R.W. & G.S. Wheeler. 2006. Orchid bees don't
need orchids, evidence from the naturalization of an
orchid bee in Florida. Ecology 87: 1995-2001.
Pemberton, R.W. 2007a. An orchid bee naturalizes in
Florida; implications for orchids and other plants.
Orchids 76: 446-448.
Pemberton, R.W. 2007b. Invasive orchid bee, Euglossa
viridissima, pollinates the ornamental orchid (Gua-
rianthe skinneri) in Florida. Lankesteriana 7: 461-468.
Pemberton, R.W. & H. Liu. 2008. The naturalization of
an oil collecting bee Centris nitida in Florida and the
eastern United States, with notes on the Centris species
native to Florida. Florida Entomol. 91: 101-109.
Pemberton, R.W. in press. Pollination of the ornamental
Oncidium sphacelatum by the naturalized oil-collecting
bee (Centris nitida) in Florida. Selbyana.
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http://www.hear.org/pier/prospective.htm (last accessed
in March 2008).
Renz, J. 1984. Orchidaceae. Pp. 1-63 in: E. Nasir & S.I.
Ali (eds.), Flora of Pakistan 164. PanGraphics Ltd.,

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Gonzalez, G.A., G.C. Femrndez-Concha, R.L. Dressier,
L.K. Magrath & G.W. Argus. 2007. Orchidaceae, Flora
of North America Vol. 26, p. 640. Online version http://
www.efloras.org (last accessed Feb., 2008)
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woodchips in Surrey. Mycologist 18: 12-15.
Singh, A.P 2001. Flowering plants of Nepal (Phanerogams).
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with contrasting mating systems. Amer. J. Bot. 84: 224-
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LANKESTERIANA 8(1), abnl 2008. 0 Umversidad de Costa Rica, 2008.

IANKESTERIANA 8(1): 15. 2008.





1 Department of Botany, University of Florida, 214 Bartram Hall, Gainesville, FL 32611-8526, U.S.A.
(mablanco@ufl.edu, I i..i'i. ..i .1.. lendara@flmnh.ufl.edu)
2 Jardin Botanico Lankester, Universidad de Costa Rica, Apdo. 1031-7050, Cartago, Costa Rica.
3Herbario CICY, Centro de Investigaci6n Cientifica de Yucatan (CICY), calle 43 No. 130,
Col. Chuburn de Hidalgo, 97200 Merida, Yucatan, Mexico. (careval@cicy.mx)
4 Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL 32611-7800, U.S.A.
(whitten@flmnh.ufl.edu, orchid@flmnh.ufl.edu)
5 Departamento de Botanica, Instituto de Biociencias, Universidade Federal de Rio Grande do Sul,
CEP 91501-970, Porto Alegre, RS, Brazil. (rbsingerl@yahoo.com)
6 Departamento de Gen6tica, Escola Superior de Agricultura "Luiz de Queir6z", Universidade de Sao Paulo,
C.P. 83, Piracicaba, SP, Brazil 13400-970. (samantha.koehler@gmail.com)
7Center for Plant Research, University of British Columbia, #302-Macmillan Building, 2357 Main Mall,
Vancouver, B.C., Canada V6T 1Z4. (isidro@interchange.ubc.ca)

1) Two new combinations forthe basionymMaxillaria
bomboizensis Dodson were made in Blanco et al.,
(2007), one in Camaridium (page 519) and another
in Sauvetrea (page 535). The first was accidentally
left undeleted after we confirmed the correct
placement of this taxon in Sauvetrea, and therefore
should be considered a synonym:

Sauvetrea bomboizensis (Dodson) M. A. Blanco,
Lankesteriana7: 535.2007. Camaridium bomboizense
(Dodson) M. A. Blanco, Lankesteriana 7: 519. 2007,
syn. nov.

2) The wrong year (1964) was given for the basionym
Ponera adendrobium Rchb.f. [Ornithidium
adendrobium (Rchb.f.) M. A. Blanco & Ojeda]
(page 532). The correct year of publication for
Ponera adendrobium is 1865. The rest of the
citation is correct and therefore the transfer is valid
(see article 33.4 of the Code; McNeill et al., 2006).

3) The following combinations were not included in
the article, and are thus validated here:

IANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 200&

Camaridium carinulatum (Rchb.f.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria carinulata Rchb.f, Flora 41:
6. 1877.

Ornithidium fasciculatum (C. Schweinf.) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria fasciculata C. Schweinf.,
Bot. Mus. Leafl. 15: 162. 1952.

ACKNOWLEDGEMENTS. We would like to thank Eric Hagsater
(AMO) and James D. Ackerman (UPRRP) for calling our
attention to errors 1 and 2 respectively.

Blanco, M.A., G. Carnevali, W. M. Whitten, R. B. Singer,
S. Koehler, N. H. Williams, I. Ojeda, K. M. Neubig, L.
Endara. 2007. Generic realignments in Maxillariinae
(Orchidaceae). Lankesteriana 7: 515-537.
McNeill, J., F. R. Barrie, H. M. Burdet, V. Demoulin, D. L.
Hawksworth, K. Marhold, D. H. Nicholson, J. Prado, P.
C. Silva, J. E. Skog, W. J. Wiersema, & N. J. Turland.
2006. International Code of Botanical Nomenclature
(Vienna Code). Reg. Veg. 146.


LANKESTERIANA 8(1): 17-19. 2008.



Jardin Botanico de Missouri Peru
Prolongaci6n Bolognesi Mz. E Lote 6, Oxapampa, Cerro de Pasco, Peru
luis_gin@yahoo.es; 1ii '.l. i. i.iil ,.in

ABSTRACT. Masdevallia vilcabambensis, a new species from the department of Cuzco in Peru, is described
and illustrated. It is similar to M. minute, but it has much smaller plants and flowers, the flowers are
yellowish and blotched with purple, and the sepals abruptly reflexed at the middle. Also, M. vilcabambensis
is exclusively know from around 2500 meters elevation, whilst M. minute is restricted to warm regions
below 350 meters of altitude.

KEY WORDS : Orchidaceae, Pleurothallidinae, Masdevallia vilcabambensis, Peru, new species

Introduction. The cloud forest of the region of
Vilcabamba in Peru hosts an astonishing variety in
plant species, and the efforts by botanists and concerned
institutions to enlighten this floristic mega diversity
promote the continue discovery of taxa new to the
science. The characteristic climate of the cloud forest,
with its high humidity and the constant presence of
fog, favors the establishment of a rich epiphytic flora,
among which the family Orchidaceae is outstanding
for the great number of species (Schweinfurth 1944,
1951, 1958-1961, 1970, Bennett & Christenson
1993, 1995, 1998, 1999, 2001, Rodriguez 1999).
The orchids of the subtribe Pleurothallidinae (Tribu
Epidendreae) account for approximately 3000 species,
which means more o less 10% of the world orchids.
Among them, Masdevallia Ruiz & Pav. has nearly 500
species, 150 of which are endemic to Peru, but it is easy
to estimate that this number will quickly increase as
new difficult-to-reach areas still covered with pristine
vegetation will be explored.
The genus Masdevallia is distributed from
southern Mexico to the South of Brazil, with the
greatest species diversity in the central mountain
ranges of Colombia, Ecuador, Peru, and Bolivia. It is
characterized by terrestrial, epiphytic or lithophytic
plants provided with non-pseudobulbous stems and a
single leaf, fleshy to coriaceous, cuneate at the base.
The basal inflorescences can be single- or many-
flowered, with the peduncle usually cylindrical.
The flowers are showy and frequently with brilliant

colors. The sepals are always adnate, with the free
apexes ending in more or less elongate tails. Species
of Masdevallia are most commonly found in wet and
cold climates, with some taxa spanning toward more
temperate regions.

Taxonomic treatment. During one of the explorations
done by the team of the Missouri Botanical Garden
to collect material from the Vilcanota River basin,
Efrain Mountafies Suclli, one of the team members,
discovered a tiny orchid that is described here as

Masdevallia vilcabambensis L. Valenzuela & E.
Suclli, sp. nov.

TYPE: Peru. Convenci6n: Vilcabamba, Oyara, 2500
m, 1301'29"S 7250'08"W, 22 Febrero 2007,
L. Valenzuela 8896, E. Suclli, I Huamantupa, G.
Calatayud, F Zamora & N. Su6rez holotypee, CUZ;
isotype MO). FIG. 1.

Species Masdevallia minute Lindl. similis, sed statura
multo minore, floribus minoribus, sepalis reflexis,
habitat divers recedit.

Herb epiphytic, small, up to 25 mm tall. Ramicaul
to 23 mm long. Leaves obovate-elliptic, glabrous,
10-25 x 5-8 mm, narrowed toward the base into a
conduplicate petiole 8-12 mm long. Inflorescences
single-flowered, emerging from the basal sheaths,
2 -3 per ramicaul, to 20 mm long, green. Flowers


1 mm

1 mm

3 mm

FIGURE 1. Masdevallia vilcabambensis L. Valenzuela & E. Suclli. A Habit. B Apex of the leaf, abaxial view. C Flower.
D Flower, spread (the petals removed), the reflexed tails not visible. E Column and lip, lateral view. F Lip. G Petal.
H Anther cap. Drawn by L. Valenzuela from the holotype.

LANKESTERIANA 8(1), abnl 2008. 0 Universidad de Costa Rica, 2008.

VALENZUELA GAMARRA A new Masdevallia from Peru

with yellowish sepals, blotched vivid purple at the
base, the tails yellow; petals and lip purple; column
green with the apex cream-hyaline. Sepals narrowly
triangular-ovate, connate at the base ca. 1 mm, ending
into reflexed, conduplicate tails, 3 x 1 mm excluding
the tail. Petals obcuneate-subfalcate, truncate, 1.5 x
1 mm. Lip narrowly ovate-subpandurate, abruptly
reflexed-folded at the middle, acute, 2 x 1.5 mm.
Column terete, slender, subclavate, with shallow
clinandrium, 1.6 mm long. Anther cap cucullate,
helmet-shaped, cream.

DISTRIBUTION AND ECOLOGY: Epiphytic in cloud forest
at an altitude of between 2400-2500 m. The plants
grow in shady places, on the stems of vines (up to
2 cm in diameter) covered with mosses, at medium
height (up to 3 meters from the ground), forming small
populations usually of 4-8 individuals.

ETYMOLOGY: Named from the district to which belongs
the locality where the species was found, in the district
of Vilcabamba, the historical place where the Incas
directed by Manco Inca offered their last resistance to
the Spanish invasion in 1532.

The only species similar to M. vilcabambensis is
M. minute Luer, but the former has much smaller plants
and flowers, the flowers are yellowish, blotched with
purple (vs. white in M. minute), the disposition of the
column and the shape of the sepals, and sepals abruptly
reflexed at the middle (vs. straight). Furthermore, M.
minute is restricted to warm regions below 350 meters,
while M. vilcabambensis is exclusively know from a
cold region at around 2500 meters elevation.

ACKNOWLEDGEMENTS. The authors wishes to thank Stig
Dalstr6m for help with the translation of the manuscript,
Franco Pupulin for help in any addition to the manuscript,

and the Missouri Botanical Garden for support of this and
other floristic projects in Peru, allowing new plant taxa to be
discovered and described.

Bennett, D. E. & E. A. Christenson. 1993. Icones Orchida-
cearum peruvianum. Part 1, t. 1-200. A. Pastorelli de
Bennett ed., Lima.
Bennett, D. E. & E. A. Christenson. 1995. Icones Orchida-
cearum Peruvianum. Part 2, t. 201-400. A. Pastorelli
de Bennett ed., Lima.
Bennett, D. E. & E. A. Christenson. 1998. Icones Orchida-
cearum Peruvianum. Part 3, t. 401-600. A. Pastorelli
de Bennett ed.
Bennet, D. E. & E. A. Christenson. 1999. A new species
of Telipogon Kunth (Orchidaceae: Telipogoninae) del
Peru. Araldoa 6 (1): 61-64.
Bennett D. E. & E. A. Christenson. 2001. Icones
Orchidacearum Peruviarum. Part 4, t. 601-800. A.
Pastorelli de Bennett ed., Sarasota, USA.
Rodriguez, A. 1999. Orquideas en Machupicchu. Egemsa &
CBC, Cusco, Peru.
Rolando, I. 1999. Orchids of the Pueblo Hotel, Machupicchu,
Urubamba, Cusco, Pern. Presented by J. Koechlin Von
stein and J. Purisaca.
Schweinfurth, C. 1944. The Orchids of Cusco. Revista
Universitaria, UNSAAC 87: 35-46.
Schweinfurth, C. 1951. Orchidaceae peruvianae VIII. Bot.
Mus. Leafl. 15(3): 79-109.
Schweinfurth, C. 1958. Orchids of Peru, part 1. Fieldiana,
Bot. 30: 1-260.
Schweinfurth, C. 1959. Orchids of Peru, part 2. Fieldiana,
Bot. 30:261-351.
Schweinfurth, C. 1960.Orchids of Peru, part 3. Fieldiana,
Bot. 30: 352-786.
Schweinfurth, C. 1961. Orchids of Peru, part 4. Fieldiana,
Bot. 30: 787-1026.
Schweinfurth, C. 1970. First supplement to the Orchids of
Peru. Fieldiana, Bot. 33: 1-8.

LANKESTERIANA 8(1), abnl 2008. O Unversidad de Costa Rica, 2008.


IANKESTERIANA 8(1): 21-22. 2008.


Luer, C.A. 2007. Icones Pleurothallidinarum XXIX. A third century of Stelis of Ecuador.
Systematics of Apoda-Prorepentia. Systematics of miscellaneous small genera. Addenda:
new genera, species, and combinations. Monogr. Syst. Bot. Missouri Bot. Gard. 112: 1-130.

With this recent delivery of a third century of
new species of Stelis Sw. from Ecuador (pp. 1--82),
Carlyle A. Luer raised to three hundreds the number
of Stelis species he and his co-workers (notably
Alex Hirtz and Lorena Endara) revealed to science
in the last five years (Luer 2002, 2004). Taking in
account a forthcoming lot of 130 previously known
and new species (announced in the Introduction),
plus some 50 names among those recorded by
Schlechter (1921), the total number of accepted
species known from Ecuador is close to 500, or over
the half of the number of species presently known in
the genus. However, according to Luer's estimate,
Ecuadorean leadership in Stelis diversity is destined
to fade as soon as a similar amount of time and
effort in studying this genus will be expanded to the
neighboring regions of Colombia and Peru, both of
which are more than four times larger in geographic
area. This let us easily envision a genus Stelis in the
strict sense (as it is treated by Luer in his series)
encompassing more than 1500 valid species, which
will convert it into the largest Neotropical orchid
genus and perhaps into the largest natural genus of
the Orchidaceae as a whole.
As it is usual in Luer's "green books" series, each
species is fully described, and paragraphs are provided
on etymology and a short discussion for each taxon.
With the exceptions of S. lynniana and S. paulula,
which are known only from cultivated specimens
without exact provenience, locality data are provided
for all the species. All the new species are illustrated
in composite ink plates by the well-trained hands by
Luer himself and Stig Dalstrom, and it is unfortunate
that, for editorial reasons, the 101 plates had to be
strongly reduced in size from the original full-page
format, someway impeding a clearer view of the often-
intricate floral details of Stelis.
Unlike the two previous parts of the treatment of
Stelis of Ecuador (Luer 2002, 2004), section Labiatae

IANKESTERIANA 8(1), abrl 2008. O Universdad de Costa Rica, 200&

is combined here with sect. Stelis, the diagnostic
character of sect. Labiatae (the degree of connation of
lateral sepals) proving to be inconsistent. The species
of the new century are accordingly arranged into three
Sections, namely Nexipous (8 species), Humboldtia (8
species) and Stelis (87 species), the latter including sect.
Labiatae. Because of the comparatively stable flower
morphology and the mostly stereotyped vegetative
architecture of Stelis, which makes difficult to manage
such a large group for identification purposes, a key to
the Ecuadorian species will be a welcome addition to
part four of the series, where a systematic treatment is
Following the third century of new Stelis from
Ecuador, the fascicle includes, systematic monographs
of Apoda-Prorepentia (Luer) Luer, and other
miscellaneous pleurothallid genera not previously
treated in the "green" series.
Apoda-Prorepentia, based on Pleurothallis sect.
Apoda-Prorepentes Lindl. and elevated at generic
rank by Luer (2004), includes in the present treatment
8 species, mostly characterized by the repent and
often pendent habit, but with highly diverse floral
morphology. According to the author, the genus ranges
from Mexico to Brazil, Ecuador, and the West Indies,
with a main center in northwestern Andes. A key to the
species, full descriptions and notes, and a composite
plate for each species are provided in the acclaimed
style of previous Icones Pleurothallidinarum
systematic treatments.
Six monotypic genera [namely Cucumeria Luer,
Empusella (Luer) Luer, Mirandopsis Szlach. & Marg.,
Mixis Luer, Mystacorchis Szlach. & Marg., and
Psedoctomeria Kraenzl.] are monographed; references
are given to the species illustrations that appeared
in previous volumes, with the exception of Mixis
incongrua Luer, which is depicted at page 104 of the
present volume. Also monographed are Physosiphon
Lindl. (2 species from Mexico to Peru, both illustrated,


P. emarginatum in two different morphs), and
Physothallis Garay (2 species endemic to Ecuador,
illustrated on page 105).
In the Addenda section (pp. 106-130), devoted to
miscellaneous new genera, species, and combinations,
Luer describes Eitti'lla.. a genus based on
Pleurothallis subgen. Etttiia, Luer, and Niphantha,
with 2 species segregated from the same subgenus
(all treated a Stelis sensu lato by Pridgeon & Chase,
2001). The first new genus is admittedly polymorphic
and scarcely distinct from Pabstiella Brieger &
Senghas, at the point that generic assignments should
rely "on interpretation of subtle floral characters and
distribution". Although noting that Ettilitill, and
Pabstiella might constitute a single genus, 40 new
combinations are published here under Ettliui'lla
(pp. 106-107, plus E. vellozoana amid Pabstiellas
on p. 121), while 67 species are formally transferred
to Pabstiella (pp. 119-121). Two new species of
Ettulitill, are described and illustrated from Peru
and Ecuador. Nyphantha is mainly distinguished
from Eltlittill, by a comparative large spathe and the
lateral sepals free nearly to the base, and allegedly
supported by an unpublished DNA analysis.
Additionally, new species are described and
illustrated in the genera Alaticaulia (a recent segregate

from Masdevallia, of which the author describes 3 new
species from Ecuador and Peru) Lepanthes (4 species
from Colombia and Ecuador), Pleurothallis (P. davisii
from Ecuador), Restrepia (the Colombian R. fritillina),
Spilotantha (another genus split from Masdevallia,
of which a new species is described from Ecuador),
Stelis (S. maduroi from Panama), Trichosalpinx (T
sipapoensis from Venezuela), and Trisetella (the
Ecuadorean T klingeri). Among nomenclatural
changes, the volume offers 4 new combinations in
Acianthera, 8 in Alaticaulia, 12 in Anathallis, 2 in
Echinosepala [but E. vittata (Pupulin & M.A. Blanco)
Luer is a superfluous name predated by E. vittata
(Pupulin & M.A. Blanco) C.O. Murales & N. Villal.
2004), 1 each in Reichantha, Specklinia, and Tigivesta
(replacing name for Vestigium Luer, predated by a
fungal genus), and 4 in Teagueia.
The plates of the four new species referable to
genera allied to Masdevallia (Alaticaulia inamoena, A.
neukermansii, A. rojohnii and Spilotantha nigricans,
pp. 122-130) are numbered according to the format
of Systematics of Masdevallia, bringing the total of
numbered illustrations to 695.

Franco Pupulin
Lankester Botanical Garden

LANKESTERIANA 8(1), abrl 2008. 0 Universdad de Costa Rica, 2008.

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