|
ISSN 1409-3871
LANI E STERIANA
No. 5 SETIEMBRE 2002
Ecce homo, scientia clarus: Luis Fournier Origgi (1935-2002)
CARLOS 0. MORALES --------- --------1
Notes on the natural history of Cyclopogon obliquus
(Orchidaceae: Spiranthinae) in Costa Rica
MARIO A. BLANCO 3
The major sections or groups within Sobralia, with four
new species from Panama and Costa Rica, S. crispissima,
S. gloriana, S. mariannae and S. nutans
ROBERT L. DRESSLER ------- -------- 9
Trichopilia x ramonensis (Orchidaceae),
un hibrido natural de Costa Rica
CARLOS 0. MORALES -------- ---------17
Notas varias sobre Heliconia rodriguezii
(Heliconiaceae) de Costa Rica
CARLOS 0. MORALES --------- 23
A new species of Stellilabium section Taeniorhachis
(Orchidaceae) from Costa Rica
FRANCO PUPULIN and MARIO A. BLANCO --- ---------- 27
LA REVISTA CIENTIFICA DEL JARDIN BOTANICO LANKESTER
UNIVERSIDAD DE COSTA RICA
ESTE 1 I 0 )i. LANKESTERIANA
SE I l CAALA. ICi i 1 I
TI- I ,I I 1. I C-1 LANKESTERIANA
I I I- I CATEDTOTI- I' I -i Cl TCI-
lLlI ''' I'ii'iCfi''
LANKESTERIANA
LA REVISTA CIENTIFICA DEL JARDIN BOTANICO LANKESTER
UNIVERSIDAD DE COSTA RICA
Copyright 2002 Jardin Botanico Lankester, Universidad de Costa Rica
Fecha de publicaci6n valida / Effective publication date: 30 de setiembre del 2002
Diagramaci6n: Jardin Botanico Lankester
Imprenta: Litografia Ediciones Sanabria S.A.
Tiraje: 500 copias
Impreso en Costa Rica / Printed in Costa Rica
R Lankesteriana / La revista cientifica del Jardin Botdnico
Lankester, Universidad de Costa Rica. No. 1
(2001)-- -- San Jos6, Costa Rica: Editorial
Universidad de Costa Rica, 2001--
V.
ISSN-1409-3871
1. Botdnica Publicaciones peri6dicas, 2. Publicaciones
peri6dicas costarricenses
0
LANKESTERIANA 5: 1-2. 2002.
ECCEHOMO, SCIENTIA CLARUS: LUIS FOURNIER ORIGGI (1935-2002)
CARLOS 0. MORALES
Escuela de Biologia, Universidad de Costa Rica. 2060 San Jose, Costa Rica
y Jardin Botanico Lankester. Apdo. 1031-7050 Cartago, Costa Rica
iHe alli al hombre, conocido por su sabiduria! Esta
frase latina del titulo resume mi memorial del Dr. Luis
Alberto Fournier Origgi, eminente ec61logo, botinico
y agr6nomo de la Universidad de Costa Rica (U.C.R.)
fallecido sorpresivamente el pasado 5 de julio. La
gran labor cientifica y agron6mica del Dr. Fournier
pas6 desapercibida para muchos, porque 61 nunca
busc6 imagen o renombre. Fue un cientifico, un
docente y un humanista, que transmitia a los estudi-
antes conocimientos y experiencias, como muy pocas
personas pueden lograrlo. En classes y conferencias,
como un estimulo para los estudiantes, 61 citaba al
poeta latino Virgilio: "Felix qui potuit rerum
cognoscere causes"; es decir, feliz quien ha podido
conocer las causes de las cosas. Don Luis fue, sin
duda, uno de esos series afortunados que lograron
entender el mundo much mis alli de lo comimn.
El se gradu6 de Ingeniero Agr6nomo en la U.C.R.
en 1958. En 1961 obtuvo el titulo de Magister
Agriculturae en el Instituto Interamericano de
Ciencias Agricolas de la O.E.A. La Universidad de
California en Davis le confiri6 el grado de Ph.D. en
Botinica en 1964. De regreso a la U.C.R., en la
Escuela de Biologia hizo grandes contribuciones para
forjar el Herbario de la Universidad de Costa Rica
(USJ), que habia sufrido los estragos de un incendio
el 20 de marzo de 1965. Ademis, la U.C.R. tuvo en 61
a uno de sus mis destacados docentes e investi-
gadores en ciencias biol6gicas desde 1959 hasta
1988. En este afio Don Luis se pension, pero sigui6
colaborando activamente con la Escuela de Biologia
como Profesor Em&rito hasta el dia de su muerte.
Don Luis ha dejado profundas huellas en varias
areas, tales como la ecologia forestal, el pensamiento
conservacionista y la investigaci6n agron6mica (culti-
vo del cafe). D&cadas de studios y observaciones le
permitieron comprender mejor que nadie, en la zona
neotropical, los problems ambientales y el desarrollo
de bosques en areas degradadas por actividades
antrop6genas. A lo largo de 37 afios de persistentes y
rigurosas observaciones en sus fincas de Ciudad
Col6n y Tabarcia, 61 protegi6 areas que se con-
virtieron, con el paso del tiempo, en bosques exube-
rantes con una diversidad comparable a la que
mostraban los antiguos bosques que fueron destrui-
dos. La gran magnitude del legado cientifico y
academico de Don Luis se percibe en sus cerca de
150 publicaciones, que incluyen tesis, articulos cienti-
ficos y various libros. Una de sus publicaciones botini-
cas sobresalientes es Botany of Cocos Island, Costa
Rica, que es un capitulo del libro The Galipagos, edi-
tado por R.I. Bowman y publicado en 1966 por la
Universidad de California. El martes 9 de julio, cuan-
do velamos en la Escuela de Biologia las cenizas de
Don Luis, logramos cubrir todas las mesas de un
extenso laboratorio con muchas de sus publicaciones,
lo que reflej6 claramente cuin fructifera fue su vida
cientifica y academica.
Una s6lida formaci6n academica y cientifica se
complementaba con una profunda cultural y un gran
sentido de humanismo, reflejado en su calidad
humana incomparable. Por su clara conciencia
hist6rica, Don Luis poseia un vastisimo conocimiento
del desarrollo de las ciencias. Una semana antes de su
muerte ofreci6, en la Escuela de Biologia, un colo-
quio sobre los "Antecedentes de la investigaci6n
biol6gica en Costa Rica", exponiendo las raices en la
antigua Grecia y culminando en las d6cadas recientes
de la historic de la biologia en Costa Rica. Nos queda
en el recuerdo, como una lecci6n de rectitud, lo que
Don Luis le dijo al M.Sc. Rodolfo Ortiz cuando &ste
empez6 a trabajar en la U.C.R.: "Rodolfo, sirvale a la
Universidad, pero no se sirva de ella".
2 LANKESTERIANA N 5
Con un apellido frances y otro italiano, Don Luis me del saber, miximos ideales a los que puede aspirar un
cont6, hace various afios, que tambien tenia una abuela espiritu universitario. Como recuerdo imborrable del
alemana. El sinti6 siempre gran admiraci6n y estima Dr. Foumier tenemos una especie pterid6fita de Costa
por la cultural y las ciencias alemanas. Su conciencia Rica y PanamA: Elaphoglossum fournierianum L.D.
preclara siempre estuvo atenta en busca de la verdad y G6mez, Revista Biol. Trop. 20: 33. 1972.
LANKESTERIANA5: 3-8. 2002.
NOTES ON THE NATURAL HISTORY OF CYCLOPOGON OBLIQUUS
(ORCHIDACEAE: SPIRANTHINAE) IN COSTA RICA
MARIO A. BLANCO
Institute Centroamericano de Investigaci6n Biol6gica y Conservaci6n
P.O. Box 2398-250, San Pedro de Montes de Oca, San Jose, Costa Rica
Research Associate, Jardin Botinico Lankester, Universidad de Costa Rica
Present mailing address: University of Florida, Department of Botany, 220 Bartram Hall
P.O. Box 118526 Gainesville, Florida 32611-8526, U.S.A. mblanco@flmnh.ufl.edu
ABSTRACT. The occurrence of Cyclopogon obliquus (J.J. Sm.) Szlach. [= Pelexia obliqua (J. J. Sm.) Garay]
is reported for the first time in Costa Rica. Its taxonomic placement is discussed, and its current geographic
distribution is reviewed. Observations on its habitat, phenology and reproduction are presented.
RESUMEN. Se present el primer informed de la existencia de Cyclopogon obliquus (J.J. Sm.) Szlach.
[= Pelexia obliqua (J. J. Sm.) Garay] en Costa Rica. Se discute su posici6n taxon6mica y se present una
revision de su actual distribuci6n geogrifica. Se presentan observaciones sobre su habitat, fenologia y repro-
ducci6n.
KEY WORDS / PALABRAS CLAVE: Orchidaceae, Cyclopogon, Pelexia, Spiranthinae, Costa Rica, pantropical
species, phenology, deciduousness, autogamy.
In April 2000, I found a leafless, terrestrial
Spiranthine orchid with small whitish flowers at La
Selva Biological Station, in Heredia province, Costa
Rica. It was immediately apparent that it was a novel-
ty for the site's orchid flora (revised by Atwood 1988,
Wilbur 1994), but its identity was a mystery to me.
There were remains of dead leaves at the bases of the
plants, indicating that they were deciduous.
Flowering while leafless is characteristic of a number
of genera in the Spiranthinae, including Sarcoglottis,
Funkiella, Brachystele, Spiranthes, and Schiedeella.
Most species in other genera produce flowers while
their leaves are still present. This vegetative dichoto-
my originally led me to believe that the plant in ques-
tion belonged in the former group. Assuming it to be
a species of Schiedeella Schltr., I concluded it was a
new species, and prepared a description.
Fortunately, Gerardo Salazar who is studying the
generic delimitations within Spiranthinae saw the
description and recognized the plant as a species of
Cyclopogon, and suggested it might be the enigmatic
C. obliquus (J.J. Sm.) Szlach., which I was later able
to confirm. So, the publication of another superfluous
name for this taxon was prevented (see below).
The accompanying illustration (Fig. 1) and following
description are based on the two Costa Rican collec-
tions, which constitute the first report of this species
for the country. In Central America, the species was
previously reported to the north of Costa Rica
(Dressler 1993, and references below). Both Mora-
Retana & Garcia (1992) and Dressler (1993) listed it
as a species of potential occurrence in Costa Rica,
because of its previous report from southern
Nicaragua. Other diagnostic illustrations can be found
in Hamer (1981, as Pelexia hameri, and 1984, 1990,
as Pelexia obliqua) and Hu (1977, as Manniella
hongkongensis).
Cyclopogon obliquus (J. J. Sm.) Szlach., Fragm.
Florist. Geobot. 39(2): 425. 1994. FIG. 1
BASIONYM: Spiranthes obliqua J. J. Sm., Bull. Dep.
Agric. Indes N&erl. 43: 74. 1910. TYPE:
Indonesia, Java, Buitenzorg (Bogor), J. J. Smith
s.n. (BO, not seen).
SYNONYMs: Manniella hongkongensis S. Y. Hu &
G. Barretto, Chung Chi J. 13(2): 6. 1976. TYPE:
China, Hong Kong, S. Y. Hu 13266 (K, not seen).
Pelexia hameri Garay, Bot. Mus. Leafl. Harv.
Univ. 26(1): 22. 1978. TYPE: El Salvador, 0.
Pank in F. Hamer 613 (AMES, photo!).
Pelexia obliqua (J. J. Sm.) Garay, Bot. Mus. Leafl.
Harv. Univ. 28(4): 345. 1980.
Plant a terrestrial herb, geophyte. Roots 3 to 6 in
flowering individuals, tuberous, fusiform, 1-4.5 cm
LANKESTERIANA
long, 6-9 mm thick at the widest point, white-cream,
covered with short hairs when young (early leafing
stage), glabrous when leafless. Leaves petiolate, con-
volute, produced in a basal rosette, 6 or more in num-
ber, held flat against the ground when mature, absent
at flowering; petiole white to light green, 4-14 mm
long, 2-5 mm thick (longer and thinner in the first
smaller leaves), base ,l..ii.ini. lamina elliptic, acute
apically, 1.5-5.8 cm long, 0.8-2.8 cm wide, decurrent
onto the petiole, shiny dark green adaxially, crys-
talline light green abaxially, with a greenish-white
thick (2 mm at base) midvein and two main se-
condary veins diverging from near the base of the
lamina; tertiary venation reticulate, faintly visible
adaxially. Inflorescence terminal, erect, spicate-
pedunculate, 13-32 cm long. Peduncle terete, puberu-
lous, 9-20 cm long, 1-3 mm thick, light green, co-
vered by 4-6 sheathing bracts. Spike dense, 4.5-15 cm
long, 9-40 flowered, with 2-4 flowers open at a time.
Peduncular bracts sheathing, linear-triangular, 2-4
cm long, 3-5 mm wide at the base, pale green. Floral
bracts subulate-caudate, up to 6 mm long and 1.5 mm
wide, scarious. Flowers resupinate, straight to slightly
nodding at the junction of perianth and ovary. Ovary
fusiform, trigonous, 8 mm long, 2.2 mm thick, light
green; spur completely adnate to the ovary, 2.6 mm
long, 0.8 mm wide, covered with glandular hairs to
0.3 mm long. Sepals greenish light-brown, externally
puberulous. Dorsal sepal oblong-lanceolate, obtuse to
rounded, slightly concave at the base, 4.3 mm long, 2
mm wide. Lateral sepals oblong, obtuse apically, sub-
falcate, 4 mm long, 1 mm wide. Petals translucent
white with a central longitudinal reddish-brown line;
oblanceolate, obtuse apically, coherent to the dorsal
sepal, glabrous, 4 mm long, 0.8 mm wide. Labellum
crystalline white with green lateral lobes; pandurate,
auriculate at the base, ecallose, 5.7 mm long (inclu-
ding the basal auricles), 3.3 mm wide; hypochile 2.6
mm long, 3.2 mm wide, margins entire, lateral lobes
enfolding the column; basal auricles subfalcate, 1.5
mm long, 0.5 mm wide, inserted in the spur. Epichile
transversely bilobed, sides slightly incurved to flat, 1.8
mm long, 3.3 mm wide, margin crenulate; apex retuse
with a sinus 0.3 mm deep and 0.4 mm wide. Column
reddish-cream, 4 mm long, 1.3 mm wide, stigma
bilobed; anther brown, 1.2 mm long, 1 mm wide.
Pollinia cuneate-obovate, white, each one 1.7 mm
long, 0.5 mm wide; viscidium dark grey, rhombic, 3
mm wide. Fruit a fusiform capsule, 1.2 cm long, 0.5
cm thick. Most flowers developing into fruits.
COSTA RICAN RECORDS: Heredia: Puerto Viejo de
Sarapiqui, Estaci6n Biol6gica La Selva, Sendero Tres Rios
1550, 1026' N, 8359' W, alt. 50 m, 7 Apr. 2000 (buds,
flowers, fruits, leafless), Blanco, Horvitz, ... Johnson,
& Lange 1488 (USJ, CR, F, MO, SEL, M). Same locality,
18 Jun. 2000 (sterile, 1. .,..1 Blanco 1521 (USJ, CR, F,
MO, SEL).
Cyclopogon obliquus has an interesting historical
record. Even though the genus is otherwise exclusively
Neotropical, it was first found growing spontaneously
in the Bogor Botanic Gardens in western Java, and
described in 1910 by Johannes Jacobus Smith then
director of the Herbarium Bogoriense as a species of
Spiranthes Rich. It was not until 1976 when it was col-
lected again, this time in Hong Kong, and described as
Manniella hongkongensis by Hu and Barretto. It was
collected a few years later in El Salvador, and
described by Garay as Pelexia hameri. A couple of
years later, Garay himself recognized his species as
synonymous with Smith's Spiranthes obliqua, transfer-
ring the latter to Pelexia. In 1994, Szlachetko trans-
ferred it once more, this time to the genus Cyclopogon.
The latter convention is followed here.
Most recent treatments place this taxon under the
genus Pelexia (Hamer 1984, 1990, 2001; Mora-
Retana & Garcia 1992, Dressler 1993, Ackerman
2000, Cribb & Ormerod 2000). However, the combi-
nation of a short column, an oblong, fleshy, blunt ros-
tellum, and a nectary that is totally adnate to the
ovary, suggests a better placement in Cyclopogon (G.
Salazar, pers. commun. 2001). On the other hand,
Ackerman (2000) also used floral morphology to sup-
port a placement in Pelexia. Recent cladistic analyses
based on DNA data (G. Salazar, unpubl.) indicate that
these two genera are not as closely related to each
other as previously assumed (e.g., Bums-Balogh &
Robinson 1983); therefore, molecular data will likely
settle this species' systematic affinities.
Cyclopogon obliquus has also turned up in eastern
Java (Comber 1990), Sri Lanka (Hamer 1984),
Samoa (Cribb & Ormerod 2000), Nicaragua (Hamer
1984, 2001), Guadeloupe, and Cuba (Ackerman
2000). Ackerman (2000) lists vouchers from most of
these locations. According to Comber (1990), it also
occurs in Argentina and Brazil, but no vouchers are
cited. Though the species' geographical origin is
unknown, it is clear it must be Neotropical, as are the
rest of the species in both Cyclopogon and Pelexia.
This case is strikingly similar to that of
Oeceoclades maculata (Lindl.) Lindl., an orchid that
BLANCO Cyclopogon obliquus in Costa Rica
!)kD
U
42
H I J
Figure 1. Cyclopogon obliquus (J.J. Sm.) Szlach. A Flowering plant. B Leafing plant (drawn from dried material;
mature leaves lye flat on the ground in living plants). C Flower, lateral view. D Dorsal sepal and one coherent petal
(the other removed). E Lateral sepal. F Lip. G Floral bract. H Column, lateral view. I Column, dorsal view. J -
Column, ventral view. K Pollinia. ILLUSTRATION VOUCHERS: A, C-K, Blanco et al. 1488; B, Blanco 1521 (both, USJ).
Drawings by the author.
Setiembre 2002
LANKESTERIANA
was described in 1821 from material collected wild in
Brazil, and not found in its original African habitat
until a few decades later (Stem 1988). The range of
0. maculata has expanded very quickly in the
Americas since its presumed introduction less than
two centuries ago, and is presently found in most of
tropical and subtropical America. It also has a long
list of synonyms, mostly due to successive generic
transfers (Stewart 1988). As in Cyclopogon obliquus
(see "Pollination and Fruit Set" below), the success of
0. maculata as an invasive species can be attributed
in part to its self-pollinating capacity (Gonzilez-Diaz
& Ackerman 1988).
HABITAT: The only collection of Cyclopogon
obliquus from what appears to be a natural habitat is
McGillivary 85 (SEL, not found) (Hamer 1984, Cribb
& Ormerod 2000), in shady, wet, lower montane
tropical forest in the Island of Ometepe in Nicaragua.
The Costa Rican collection comes from a lowland
site, even less seasonal as does the original type col-
lection from Bogor. Comber (1990) notes that it has
become more common in Tretes, as "the seasonal cli-
mate in East Java is obviously more to its liking than
the almost everwet climate of Bogor". These observa-
tions, together with its deciduous behavior, support
the idea that this species is better adapted to seasonal-
ly dry habitats.
Most of the collections from both the Americas and
southeast Asia come from man-made habitats, espe-
cially lawns and pots (Garay 1978, Comber 1990,
Cribb & Ormerod 2000), suggesting that the seeds
-or even whole plants- are accidentally transported in
the soil of transplanted crops or ornamentals. This is
the most likely scenario on how it got to Bogor in the
first place; the Botanic Gardens had been actively
importing plants from throughout the tropical world
since the 1840's (Indonesian Institute of Sciences
2002). Cyclopogon obliquus is possibly more widely
distributed in tropical and subtropical areas around
the world than currently recognized, but flowering
plants are easily overlooked due to their inconspicu-
ous flowers and deciduous habit.
This might also explain why this species stayed
undetected at La Selva, where an intensive floristic
inventory program has taken place since 1979
(Hammel & Grayum 1982, Hartshorn & Hammel
1994). The inflorescences look like dry grass stems
from a distance, and they merge with the colors of the
surrounding leaf litter. I noticed them almost acciden-
tally, but a careful inspection of the surrounding area
revealed a small colony of 12 flowering individuals.
Plants were growing in alluvial soil right next to a
paved trail which runs through a late secondary tropi-
cal wet forest, part of an abandoned -and recently
removed- cacao plantation. The area next to the trail
is periodically mowed, which probably facilitated the
establishment of the plants by reducing competition
for light. The leaves form a basal rosette and are held
flat against the ground surface when fully expanded,
so they avoid being damaged by the mowers.
However, some trees with big leaves (e.g. Castilla
elastica, Moraceae) grow in the same area, and their
fallen leaves were observed to totally cover a few
plants at the leafing stage. Light deprivation can be an
important ecological constraint for photosynthesis,
growth, and future reproductive output of individual
plants, as shown by Willems et al. (2001) for a
species of Spiranthes in the Netherlands.
There was a great variation in the size of flowering
individuals. The smallest had an inflorescence 13 cm
tall with nine flowers, and only three roots, the largest
of which was 2.3 cm long. On the other hand, the
tallest individual had an inflorescence of 32 cm with
some 40 flowers and fruits, and with five roots pre-
sent, up to 5 cm in length. There was a whole range
of intermediate individuals (Fig. lA shows one such
"'.% A.y1,L" plant). This suggests that individual plants
may start flowering when quite young. The larger
plants also seemed to have been flowering for a
longer period than the smaller ones, since some of the
fruits lower in the spike had already dehisced; in the
smaller individuals only flowers and developing fruits
were present.
PHENOLOGY: The colony at La Selva was revisited
periodically to gather phenological data. Two months
after the original collection, in June 2000 (early rainy
season, Sandford et al. 1994), numerous leafing
plants were seen at the same spot, even when the
flowering specimens had been previously removed as
vouchers. By October, the plants still had their old
leaves, but were not producing any new leaves. In
late December (early dry season) the leaves had
already disappeared, and no evidence of the presence
of the plants remained at the surface. In mid May
2001 (end of the dry season of that year) most of the
plants had already flowered and were fruiting, still
leafless. Leaves were already present (and the inflo-
rescences gone) by mid July, the last time I checked
BLANCO Cyclopogon obliquus in Costa Rica
the colony.
Contrary to the pattern observed at La Selva,
Comber (1990) reports that populations in Java
flower at the beginning of the rainy season. In any
case, all the available accounts of the species agree in
that inflorescences and leaves never occur simultane-
ously. This deciduous habit is unusual in both
Cyclopogon and Pelexia. Photographs of the flower-
ing and leafing stages are presented by Comber
(1990, as Spiranthes obliqua).
In South Florida, Calvo (1990) found that indivi-
dual plants of Cyclopogon cranichoides (Griseb.)
Schltr. can survive one or two years (and possibly
more) in an underground condition, without produc-
ing any leaves. It is possible that C. obliquus has si-
milar capabilities; if so, point counts on the number
of individuals would likely underestimate the true
population size. However, data from marked indivi-
dual plants are lacking in this species.
POLLINATION AND FRUIT SET: Orchids in the
"Pelexia-alliance" are adapted to bee pollination; the
dorsal viscidium attaches to the underside of the
labrum, and the fragile, friable pollinia assume a pro-
tected position when the proboscis is retracted
(Singer & Sazima 1999). Species of Cyclopogon
appear to be specialized for pollination by halictid
bees (Singer & Cocucci 1999, Singer & Sazima
1999). However, autogamy is reported for several
species (Ackerman 1995, Singer & Sazima 1999, van
der Cingel 2001). The plants of C. obliquus at La
Selva appear to be autogamous (at least facultatively),
since virtually all the flowers develop into fruits (but
see below). As in C. elatus (Sw.) Schltr. in Puerto
Rico (Ackerman 1995), the simultaneous presence of
buds, open flowers and dehisced fruits in some inflo-
rescences indicate an ephemeral anthesis and rapid
fruit ripening. Self-pollination becomes advantageous
in conditions where suitable pollinators are absent, as
might be the case in southeast Asia.
Both Catling (1982) and Singer & Sazima (1999)
report enlargement of ovaries without seed produc-
tion in unpollinated flowers in several Spiranthine
genera, including Cyclopogon. Although seeds were
observed in some dehisced fruits in C. obliquus at La
Selva, I cannot rule out the possibility that the plants
require pollinator services in order to set seed, and the
unpollinated flowers simply enlarge their ovaries
without producing any seeds.
In May 2001, an inspection of the inflorescences
revealed some indehisced fruits with no seeds within.
Most of these had round exit holes, but a few had
hymenopteran pupae inside (one per fruit), about 3
mm long. Unfortunately, no adults could be reared
when inflorescences were put in clear plastic bags for
a few days. Dissection of the pupae revealed they are
Eurytomid wasps, possibly belonging to the large
genus Eurytoma. Species of this genus have been
found parasitizing fruits in several orchid species,
including Pelexia adnata (Sw.) Spreng. in Belize
(Catling & Greenwood 1988). The absence of such
seed predators in Indonesia might partly account for
the apparent higher population densities of
Cyclopogon obliquus there than in the Neotropics.
ACKNOWLEDGMENTS: I thank Carol Horvitz (ii --1.
of Miami) for funding my stay at La Selva during field-
work for a project assessing alluvial forest regeneration
following removal of cacao trees, when flowering material
was collected. Additional collecting was possible due to a
research grant by the Andrew W. Mellon Foundation and
the Organization for Tropical Studies. Gerardo Salazar
(Jodrell Laboratory, Royal Botanic Gardens, Kew) provid-
ed much useful information and interesting discussions.
Paul Hanson (Universidad de Costa Rica) identified the
wasps. Carlos Ossenbach kindly provided several critical
references. Robert Dressler, Carlos 0. Morales, and
Franco Pupulin made useful suggestions for the improve-
ment of the manuscript.
LITERATURE CITED
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the Virgin Islands. Mem. New York Bot. Gard. 73.
Ackerman, J. D. 2000. Notes on the Caribbean orchid
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Atwood, J. T. 1988. The vascular flora of La Selva
Biological Station, Costa Rica: Orchidaceae. Selbyana
10: 76-145.
Burns-Balogh, P. & Robinson, H. 1983. Evolution and
phylogeny of the Pelexia Alliance (Orchidaceae:
Spiranthoideae: Spiranthinae). Syst. Bot. 8: 263-268.
Calvo, R. 1990. Four-year growth and reproduction of
Cyclopogon cranichoides (Orchidaceae) in South
Florida. Amer. J. Bot. 77: 736-741.
Catling, P. M. 1982. Breeding systems of northeastern
North American Spiranthes (Orchidaceae). Can. J. Bot.
60: 3017-3039.
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Setiembre 2002
LANKESTERIANA
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LANKESTERIANA5: 9-15. 2002.
THE MAJOR SECTIONS OR GROUPS WITHIN SOBRALIA,
WITH FOUR NEW SPECIES FROM PANAMA AND COSTA RICA,
S. CRISPISSIMA, S. GLORIANA, S. MARIANNAE AND S. NUTANS
ROBERT L. DRESSLER
Missouri Botanical Garden, Florida Museum of Natural History, Marie Selby Botanical Gardens
Mailing address: 21305 NW 86th Ave., Micanopy, Florida 32667, U.S.A.
RESUMEN. Se present una discusi6n de las secciones o grupos de Sobralia, basada en las secciones nom-
bradas por Brieger. Se described cuatro species nuevas de Panama y Costa Rica: S. gloriana, una especie
escasa y muy llamativa del area de El Valle de Ant6n, pariente de S. atropubescens pero de flores mas
grandes y de coloraci6n amarilla; S. mariannae, una plant pequefia del bosque nuboso de Cerro Jefe, con
flores blancas y pelos gruesos en el labelo; S. crispissima, pariente cercana de S. lindleyana, que se extiende
desde Panama hasta el sur de Mexico, y S. nutans, una especie distintiva de la Seccion Globosae, de
Panama.
ABSTRACT. The sectional classification of Sobralia is discussed, based on a system proposed by Brieger, and
four new species of Sobralia are described from Panama and Costa Rica, S. crispissima, S. gloriana, S.
nutans and S. mariannae.
Sobralia is a conspicuous element in the orchid
flora of the American tropics that remains poorly
known. The delicate, ephemeral flowers are often
showy, but unless they are prepared carefully and
dried quickly they make very poor pressed speci-
mens. Even flowers in alcohol may disintegrate with
time. Since botanical classification is traditionally
based on dried material, Sobralia has been much
neglected. To achieve a better understanding of this
group, we should, ideally, study living material. The
sectional classification is here discussed, and four new
species from Panama and Costa Rica are described.
THE SECTIONS AND GROUPS WITHIN SOBRALIA
Lindley (1854) divided Sobralia into 3 sections, A,
B and C. Reichenbach (1873) used the Sections
Eusobralia and Brasolia for two South American
groups, but the selection of S. dichotoma Ruiz &
Pav6n as lectotype of the genus by Angely (1973: p.
1268) nullified this and made Section Brasolia
Rchb.f. a synonym of Section Sobralia. Now, Brieger
(1983) has published some additional sectional
names, and names are now available at least for the
three main groups listed by Lindley. Brieger's names
are validly published and clearly typified, but there
are several species and complexes that do not fit well
in Brieger's classification. It might be premature to
publish new sectional names at this time. That can
wait until we have information on phylogeny from
molecular studies.
Brieger separates Sections Sobralia and Racemo-
sae by the position of the inflorescence, lateral in
Sect. Sobralia and terminal in Racemosae. However,
one may sometimes find both terminal and upper la-
teral inflorescences on the same stem. Dr. Dodson
suggests the relative bract length as a much better and
clearer way to distinguish what appear to be two dis-
tinct groups (pers. comm.).
Brieger (1983) creates a section Intermediae for S.
fragrans Lindl., but that species seems closely allied
to S. bletiae Rchb.f., S. mucronata Ames & C.
Schweinf. and others, so I tentatively apply the name
Intermediae to a complex of species with smaller
flowers and inflorescences. Brieger includes S. cro-
cea (Poepp. & Endl.) Rchb.f. in Sect. Globosae, but it
is very unlike S. candida, so I place it with the
Intermediae for now.
It should be noted that the photograph labelled as
S. fragrans in Brieger (1983), is actually S. callosa
L.O. Williams, treated by Brieger as Lindsayella
amabilis, while the photograph labelled as S. candida
could well be S. fragrans.
The Section Abbreviatae is much the largest group,
with perhaps 50 known species.
As here delimited, the members of Sobralia
Section Globosae have narrow, acuminate leaves, the
column is very narrow basally, and the bract cluster
increases in length as flowers are produced, often
LANKESTERIANA
reaching 3-4 cm in length. This group includes only a
few species, treated in more detail below.
Sobralia luteola Rolfe, with a very condensed
raceme, may be allied to Section Intermediae, though
the inflorescence is strikingly different. Sobralia
macrophylla Rchb.f. and its allies, with the bract
cluster largely hidden, may not be a phylogenetic
group, though the plants are easily recognized, with
or without flowers. Sobralia undatocarinata C.
Schweinf. and its allies, with loose bract clusters and
2-5 relatively durable flowers produced simultane-
ously, is quite distinctive, and a few close allies
remain to be named. Both S. amabilis (Rchb.f.) L.O.
Williams and S. callosa (Lindsayella amabilis) are
superficially similar and probably pollinated by hum-
mingbirds, but they are very distinct in their details.
Indeed, S. callosa has pollinia very like those of
Elleanthus, and may have the best claim to generic
status of any of the anomalous groups.
Ames and Schweinfurth (1937) emphasized the
basal callus as being similar to the basal calli of
Elleanthus, but nearly all species of Sobralia have a
callus or a pair of calli at the base of the lip, on which
the base of the column appears to rest; of course, the
basal calli of Sobralia may well be homologous with
those of Elleanthus.
SUMMARY OF SECTIONS, ANOMALOUS GROUPS AND SPECIES
1. Inflorescence racemose, with prominent intemodes
2. Floral bracts shorter than pedicel and ovary .............................................. .........................................................
........................................................ ........ Section Sobralia [Type: S. dichotoma Ruiz & Pav6n, aprox. 13 species]
2 Floral bracts longer than pedicel and ovary .................................................. ......................................................
....................................................... Section Racemosae Brieger [Type: S. rosea Poepp. & Endl., aprox. 8 species]
1. Inflorescence condensed, with very short intemodes
3. Rachis of inflorescence conic, exposed, with flowers and buds exposed simultaneously .................... S. luteola
3. Rachis of inflorescence concealed by bracts, the bracts forming a cone-like structure
4. Bract clusters continuing growth, flowering over long period
5. B ract clusters grow ing in length .......................................................... ......................................................
................................... Section Globosae Brieger [Type: S. candida (Poepp. & Endl.) Rchb.f., 3-4 species]
5. B ract clusters grow ing in girth ...................................................................................... ............... S. valida
4. Bract clusters flowering only one season, not enlarging appreciably after flowering starts
6. Bract clusters basally concealed by sheaths of inflorescence bracts (upper leaves); the exposed apices of
floral bracts wide, obtuse or retuse .................................................. ................. S. macrophylla complex
6. Bract clusters basally concealed or not; apices of floral bracts narrow, acute or acuminate
7. Bract clusters 3-6 cm
8. Bract clusters compact, usually producing 1 flower at a time; tube of lip subequal to blade or longer
........................... Section Abbreviatae Brieger [Type: S. fimbriata Poepp. & Endl., 45-50 species]
8. Bract clusters loose, often producing 2-several flowers simultaneously; tube of lip much shorter
than blade
9. Lip with conspicuous undulate keels; all flowers at once, lasting 3-4 days .. ..................
...................................................................................... S. undatocarinata com plex (3-5 species)
9. Lip without conspicuous, undulate keels; flowers ephemeral, 1-2 at a time ............ S. amabilis
7. Bract clusters less than 2 cm, usually producing 1 or 2 flowers at a time
10. Pollinia flattened, w without distinct caudicles .................................. ................................................
................................................ Section Intermediae Brieger [Type S. fragrans Lindl., 10-15 species]
10.Pollinia obovoid, attached to distinct caudicles .......................... S. callosa (Lindsayella amabilis)
NEW SPECIES
Sobralia crispissima Dressler, sp. nov. FIG. 1
HOLOTYPE: PANAMA. Chiriqui: al oeste de Volcin,
carretera a Rio Sereno; alt. 1200 m; 25 ago. 2001,
s6palos y p6talos blancos, labelo amarillo-anaranjado
con manchas rojas. A. Maduro & E. Olmos 223 (MO,
Isotypes, PMA, FLAS).
Sobraliae lindleyanae Rchb.f. similis, foliis
majoribus et floribus brevioribus, labello reflexo mar-
gine valde undulato (crispo), marginibus falcatis
appendicibus columnae integris dignoscenda.
Roots 6-8 mm in diameter; stems 20-40 cm, ca. 5
mm wide, slightly compressed; sheaths striate; leaves
5 or 6, 9-14 x 3.5-5.4 cm, ovate, slightly attenuate,
acute; inflorescence terminal, with floral bracts con-
DRESSLER Sobralia
cealed in infundibuliform sheaths of inflorescence
bracts (upper leaves with reduced blades) with 1 or 2
additional reduced blades of floral bracts exposed,
blade of lower inflorescence bract 7-11.5 x 2.2-3.5
cm, blade of upper inflorescence bract ca. 2.5 x 2 cm;
ovary and pedicel ca. 20 mm. Sepals and petals white,
lip orange-yellow with red spots; sepals 3.5-4.5 x 0.9-
1.2 cm, oblong or oblong-oblanceolate, apiculate;
petals 3.5-4.5 x 1-1.5 cm, oblong-oblanceolate,
obtuse or apiculate; lip 3-4 x 3-3.8 cm, broadly obo-
vate, emarginate, blade strongly reflexed to near mid-
dle of tube, with a single rounded basal callus ca. 4.5
x 4 mm, pilose distally, with 3 low ventral keels
basally, margin of blade strongly crisped-undulate;
column ca. 1.8 cm, with porrect, falcate arms (or
"wings").
Sobralia lindleyana Rchb.f. is common as terrestri-
al or lithophyte in the Llanos del Volcin, north of
Hato del Volcin. This new and related species was
recently found to the west of Volcan, in forested
areas, and very similar plants without flowers were
found to the east, usually growing epiphytically.
When I noted that this species resembled S. lin-
t.ii,' ......1 Erick Olmos said "No, S. lindleyana is the
species in the Llanos, its leaves are smaller and the
flowers larger," which nicely summarizes the differ-
ences between these species. Reichenbach's drawing
of S. lindleyana is suggestive of S. crispissima, but
the type specimen is clearly the familiar species of
the Llanos. The flowers of S. crispissima are some-
what smaller than those of S. ',.i I ...... and the tube
is proportionately shorter. These plants may be distin-
guished from S. lindleyana by the following key.
1. Leaves 5-6 cm long; sepals and petals 5.5-6 cm long;
tube of lip subequal to sepals and petals; blade of lip
spreading, but not reflexed; column arm serrate-
dentate dorsally .................................... S. lindleyana
1. Leaves 10-14 cm long; sepals and petals 3.5-5 cm
long; tube of lip distinctly shorter than sepals and
petals; blade of lip reflexed, reaching the middle of the
tube; column arm entire dorsally ............ S. crispissima
The epithet crispissima refers to the strongly wavy
margins of the lip.
PARATYPES: MEXIco. Chiapas: Ocosingo, ca. 8.5
km del camino Monte Libano-NahA, tomando el
camino a Villa las Rosas y El Pozo Ocotal
(PEMEX), aprox. 87.2 km al E de Ocosingo, ca. 940
m, 1655'N, 9133'W, Selva median perennifolia, de
montafia con Talauma, Quercus y Terminalia, sepa-
los y p6talos blancos, labelo amarillo con manchas
caf6-rojizo, fecha de colecta 7 ago 1992, prensado de
material cultivado 24 sept. 1997, M. A. Soto, R.
Solano y L. Izquierda 6958 (AMO, FLAS).
NICARAGUA. Jinotega: Cerro Grande, elev. 3950 ft.,
Nov., A. A. Heller 1314 (SEL); Matagalpa: Finca
Bavaria, 4100 ft., A. A. Heller 4980 (SEL). COSTA
RICA. Cartago: below Pavones on road from
Turrialba to Siquirres, 25 Aug. 1963, C. H. Dodson
2516 (SEL).
Sobralia (Abbreviatae) gloriana Dressler, sp. nov.
FIG. 2, 3
HOLOTYPE: PANAMA. Cocld: Area de El Valle de
Ant6n, Alt. 800-1000 m. 23 julio 2001, s6palos y
p6talos amarillo palido, bafiados por rojo-castafio,
bordes y vena media amarillos, labelo rojo-rosado
con borde rosado palido y quillas amarillas. A.
Maduro & E. Olmos 212 (MO, Isotype PMA).
Herba epiphytica foliis ellipticis, floribus ab glo-
mere bractearum conico-ellipsoideo products, luteis,
sepalis petalisque castaneo-ferrugineo aspersis, sepa-
lis ellipticis vel lanceolato-ellipticis, petalis oblongo-
lanceolatis, labello obovato quinque carinis humilis
ornato.
Roots 3-5 mm in diameter; stems 20-80 cm, 2.5-4
mm wide, slightly compressed; sheaths striate, with
very short bristles to ca. 0.5 mm on both sheaths and
stem; leaves several, 8.5-11 x 3.7-4.3 cm, elliptic,
short-acuminate, strongly plicate; sheaths verrucu-
lose; inflorescence terminal, bract cluster ellipsoid,
ca. 3 x 0.8 cm, largely concealed by sheaths of upper
leaves (or foliar bracts); ovary and pedicel 20-25 mm;
Sepals and petals pale yellow, flushed with purplish
brick red, sepals with yellow median stripe, petals
with broader median stripe and yellow margins, lip
dull purple with yellow keels in throat; sepals 5-6 x
1.4-1.8 cm, elliptic or lance-elliptic, apiculate; petals
4-5 x 1.6-1.8 cm, oblong-oblanceolate; lip 5.2-6.2 x
3.5-4 cm, cuneate, obovate, basal ridges 9 x 2 mm,
with 5 low ventral keels, 3-4 of these reaching nearly
to apex, margin distally undulate; column 2.6-3.2 cm.
When I first saw this attractive Sobralia, I thought
it might be a natural hybrid between S. atropubescens
and some other species with yellow or white flowers.
Now several plants of this kind have appeared, all
from the area of El Valle de Ant6n, and it is surely a
distinct species, though not very common. It is evi-
dently related to S. atropubescens, but the flowers are
larger and much more attractive with a distinct color
Setiembre 2002
LANKESTERIANA
FIGURE 1. Sobralia crispissima Dressier. Flower, from
the type locality. Photograph by K. Dresser.
FIGURE 3. Sobralia gloriana Dressler. A flower cultivat-
ed in Florida. Photograph by K. Dresser.
PIGURE 3. ~ooralta nutans Dressier. severall intlores-
cences, some now form part of Maduro y Olmos 236,
Bocas del Toro, Panama. Photograph by K. Dresser.
PIGURE 2. oralia gloriana Dressier. A Ilower, mnca
Dracula, Panama. Photograph by K. Dresser.
FIGURE 4. Sobralia mariannae Dressier. A flower,
somewhat faded, Cerro Jefe, Panama. Photograph by K.
Dresser.
PIGURE b. mo0ralia nutans Dressier. tower, El valle
Chiquito, near El Valle de Ant6n, Panama. Photograph by
E. Olmos.
DRESSLER Sobralia
pattern. One hopes that it will be propagated from
seed, and become more widely available for orchid
collections and gardens.
The epithet gloriana honors Licenciada Gloria
Maduro, the charming wife of Andres Maduro.
Since she was foolish enough to marry a rabid orchid
nut, she deserves, at the very least, that there be an
attractive new species named in her honor.
Sobralia (Intermediae) mariannae Dressler, sp. nov.
FIG. 4
HOLOTYPE: PANAMA. Panama, Cerro Jefe, flow-
ered in cult. 24 May 2002, epiphyte, flower white,
callus yellow, brown at base, fragrant, R. L. Dresser
6352, (MO).
Herba epiphytica caule supra medium foliato, foliis
ovatis acuminatis, inflorescentia pedunculo super-
posita, sepalis oblongis, petalis ellipticis, labello obo-
vato sex carinis trichomatibus crassiusculis quasi
digitiformibus ornatis.
Epiphytic or terrestrial, caespitose, roots 1.5-2.5
mm, stems 28-60 x 0.15-0.2 cm. Leaves 4.5-7 x 1.5-2
cm, elliptic-ovate, acuminate, closely spaced at mid-
stem and more widely spaced above; sheaths verru-
cose. Inflorescence terminal, with a reduced foliar
bract (sometimes two) 2-3 x 0.8-1.6 cm whose inflat-
ed sheath covers much of the bract cluster; bract clus-
ter 2-2.5 x 0.4-0.5 cm, about half concealed by sheath
of foliar bract, bracts glabrous; ovary and pedicel 15-
22 mm; flower white, callus yellow, brown at base;
dorsal sepal 2.7 x 0.67 cm, oblong, apiculate; lateral
sepals 2.5 x 0.85 cm, oblong-elliptic; petals 2.3-2.4 x
0.8 cm, elliptic, acute or apiculate; lip 2.5 x 2.1 cm,
broadly obovate, 3-lobed distally, with 1 rounded
basal callus, 6 yellow keels from base, these short-
pilose, reaching near apex, midlobe 6 x 11 mm,
retuse; column c. 8 x 3 mm.
This species is frequent in the cloud forest on
Cerro Jefe. Until recently, we had only a field-collect-
ed plant in which the flowers are poorly preserved,
but we have found a single flower in situ (photo) that
was preserved in alcohol, permitting a much more
complete description. Even without flowers, the
plants are very distinctive, with the leaves closely
spaced at mid-stem (1-3 cm apart), with a much
longer internode (6-10 cm) above. This species, then,
has a distinct peduncle, much like that of S. fragrans.
This distinctive species may well occur in other areas
of cloud forest in eastern Panama. The epithet honors
Marianne Akers, an enthusiastic orchid student in
Panama, who has collected the only other pressed
specimen I have seen.
OTHER SPECIMEN SEEN: from the type locality, M.
Akers 488-28 (SEL).
Sobralia (Globosae) nutans Dressler, sp. nov.
FIG. 5-7
HOLOTYPE: PANAMA. Bocas del Toro: Km. 63, al
norte de Fortuna, alt. 1050 m, 9 sep. 2001, terrestre;
flores p6ndulas, blancas o crema, s6palos amarillentos
por fuera, labelo crema con manchas rojo-castafio por
fuera, por dentro amarillo en las quillas, con manchi-
tas rojo-castafio. A. Maduro & E. Olmos 236 (MO,
Isotypes PMA, SEL).
Herba terrestris foliis ellipticis acuminatis, floribus
pendulis ab glomere bractearum elongato in tempo
crescenti products, sepalis petalisque ellipticis, label-
lo ovato obscure trilobato septem carinis ornato,
tribus internis basi humili lobulis altis in apice, exter-
nis omnino crenato-undulatis.
Terrestrial, roots 3-5 mm in diameter; stems 60-
130 cm, 3-4 mm wide, slightly compressed; sheaths
striate; leaves several, 20-36 x 2.5-4.7 cm, elliptic,
acute to long-acuminate, strongly plicate; inflore-
scence terminal, gradually elongating, producing 1-3
pendent flowers at a time, to 4 cm. long; ovary and
pedicel 20-25 mm; floral bracts c. 18 x 5-6 mm,
lance-ovate, closely spaced; sepals at first yellowish-
green, petals and lip cream, lip with red-brown spots
without, keels yellow with red-brown spots near mid-
line; sepals 28-31 x 8.6-10.5 mm, elliptic, apiculate,
lateral sepals carinate; petals 26-36 x 8-10.5 mm,
elliptic, apiculate; lip 22-23 x 18 mm, ovate, weakly
3-lobed, midlobe markedly undulate, with incurved
margins; basal callus ca. 2 x 2 mm, 2-parted; keels 7,
the 3 inner keels are low in the basal half and then
form erect fin-like lobes 2-2.5 mm tall, to 4 mm long,
there is then a crenate-undulate keel on each side, fol-
lowed by a shorter, similar keel on each side; column
c. 13 mm, very narrow basally, c. 4 x 4 mm distally,
wing-like teeth ca. 2 x 1.3 mm, porrect; capsule 10.5
x 1 cm.
In addition to the features given below in the key, S.
nutans is rather larger than the other species of the sec-
tion. The margins of the midlobe are somewhat folded
in over the terminal lobes of the keels. The epithet,
Setiembre 2002
LANKESTERIANA
2 cm \
I 1/ 5mm
FIGURE 7. Sobralia nutans Dressier. A Flower, natural position. B Flower, front view. C Flower parts, spread. D -
Lip, spread. E Column, lateral and ventral views. F Anther.
nutans, refers to the nodding, or pendent, flowers. Mesa, 4 mi. E of El Valle, 836'N 80007'W, 25 Mar.
PARATYPES: PANAMA. Bocas del Toro: same area as 1993, T Croat 74806 (MO).
type, 850 m, 27 Oct. 1999, R. L. Dresser & J. T. Better material from Costa Rica may show that
Atwood 6256 (MO, PMA). Cocld: Vicinity of La Ingram & Ferrell 813 and other Costa Rican speci-
Mesa, beyond El Valle, on northern slope of Cerro mens of the Section Globosae represent a species dis-
Gaital, 837'N 80007'W, 850-950 m, 12 July 1978, G. tinct from S. lancea, or that S. pardalina is a syn-
McPherson 11227 (MO); El Valle de Ant6n, La onym of S. lancea.
KEY TO THE SPECIES OF SECTION GLOBOSAE
1. Lip distinctly pandurate, with 2 crenate or undulate keels
2. Leaves subcoriaceous; lip ecallose basally ................... S. lancea Garay [Costa Rica, Colombia, Ecuador]
2. Leaves thin, membranous; lip with "V"-like basal callus ............................. S. pardalina Garay [Ecuador]
1. Lip ovate or subovate, with 5-7 keels
3. Lip with 7 keels, outer 2 keels on each side prominent, crenate-dentate throughout, 3 inner keels low in
basal halves with prominent, erect apical lobules ........................................ S. nutans Dressler [Panama]
3. Lip w ith 5 low keels, all keels sim ilar, dentate distally .................................................................................
................................ S. candida (Poepp. & Endl.) Rchb.f. [Venezuela, Colombia, Ecuador, Peru, Bolivia]
Setiembre 2002 DRESSLER Sobralia 15
ACKNOWLEDGMENTS. I wish to thank Andres Maduro and Brieger, F. G. 1983. Subtribus Sobraliinae. In R.
Erick Olmos for their help in finding material of these Schlechter, Die Orchideen, I, 13. Lieferung, eds. F. G.
species, Franco Pupulin for his help with the Latin diag- Brieger, R. Maatsch, K. Senghas. Verlag Paul Parey,
noses, and Stig Dalstr6m for the drawings of S. nutans. Berlin & Hamburg. p. 780-800.
Lindley, J. 1854. Folia Orchidacea, Sobralia. London, J.
LITERATURE CITED Matthews.
Angely, J. 1973 ("1970"). Flora Analitica e Fitogeografica Reichenbach, H. G. 1862-1874. Xenia Orchidacea vol. 2.
do Estado de Sio Paulo, tomo 6. Sio Paulo. Leipzig, F. A., Brockhaus.
Page
Missing
or
Unavailable
LANKESTERIANA5: 17-21. 2002.
TRICHOPILIA X RAMONENSIS (ORCHIDACEAE),
UN HIBRIDO NATURAL DE COSTA RICA
CARLOS 0. MORALES
Escuela de Biologia, Universidad de Costa Rica. 2060 San Jose, Costa Rica
Jardin Botanico Lankester. Apdo. 1031-7050 Cartago, Costa Rica
ABSTRACT. The natural hybrid Trichopilia x ramonensis (T. marginata x T suavis) from the San Ram6n
region of Costa Rica is here described and illustrated. The name of this hybrid was used for the first time
in 1992, nevertheless it has so far not been validly published.
RESUMEN. Se describe e ilustra el hibrido natural Trichopilia x ramonensis (T. marginata x T. suavis) de
la region de San Ram6n, Costa Rica. El nombre de este hibrido se us6 por primera vez en 1992; sin
embargo, hasta ahora no se ha publicado vilidamente.
PALABRAS CLAVE / KEY WORDS: Trichopilia marginata, Trichopilia suavis, Trichopilia x ramonensis,
hibridos naturales, Orchidaceae, Costa Rica.
El g6nero neotropical Trichopilia Lindl.
(Orchidaceae) se distribuye desde Mexico y Cuba
hasta el sur de Brasil y posee unas 30 species
(Horich 1996, Mora-Retana 1999, Bock & GruB
2000). Garay (1972) ubica el g6nero en la subtribu
Trichopiliinae, mientras que Dressler & Williams
(1970) y Dressler (1993b) lo hacen en Oncidiinae.
Estudios recientes de ADN (Williams et al. 2001)
sugieren que este g6nero conforma un grupo aislado
entire los taxones basales de Oncidiinae. En
Mesoam&rica se conocen entire 12 y 15 species; seis
de &stas habitan en Costa Rica (Mora-Retana 1999,
Pupulin 2002). No obstante, ya se tienen evidencias de
que en este pais existen otros taxones de Trichopilia
que rara vez han sido vistos o se han confundido con
species ya descritas (R.L. Dresser, F. Pupulin, com.
pers. 2001). En general, este taxon es dificil; asi lo
reconocen distinguidos estudiosos de las orquideas,
como Horich (1996) y Dressler (1993a, 2001).
Trichopilia es, sin duda, uno de los g6neros mis
atractivos de las orquideas centroamericanas. Son
plants mayormente epifitas que habitan en bosques
lluviosos de bajuras, asi como en bosques himedos
de altitudes medias. Los pseudobulbos crecen densa-
mente agrupados, son aplanados, subovados a oblon-
gos y cada uno tennrmina en una sola hoja. Las flores
son a menudo aromiticas, de coloraci6n variada, con
morfologia en general similar a la de Cattleya. El
labelo es tubiforme o infundibuliforme, en algunas
species abierto y extendido en la parte distal.
En bosques del canton de San Ram6n, Alajuela, en
la Cordillera de Tilarin, Costa Rica, se han hallado
plants con caracteristicas intermedias entire las de T.
marginata Henfr. (s.l.) y T suavis Lindl., con flores
relativamente grandes, de labelo roseo-purpura,
extendido y ondulado. Seghn comunicaci6n del fina-
do Joaquin Garcia con Clarence K. Horich (1996), ya
en el siglo XIX, hace 105 a 110 afios, Richard Pfau
observ6 plants similares cerca del poblado de San
Ram6n, y el mismo Horich (loc. cit.: 179) lo hizo por
ahi de 1973 cerca de La Balsa de San Ram6n. La
existencia de un hibrido natural de Trichopilia fue
comunicada por Dressler (1993a) y Mora-Retana
(1999). En enero de 1993 Horich envi6 semillas
maduras a Alemania; &stas germinaron en tres sema-
nas y las plants florecieron siete afios despues en
Bad Gandersheim, Baja Sajonia (Lucke 2000).
Mora-Retana & Garcia (1992) anotan un nombre
inedito para este taxon: "Trichopilia x ramonensis J.
Garcia & Mora-Retana (nat. hyb.), ined.". Sin embar-
go, no existe una publicaci6n vilida, aun cuando
plants de este hibrido se han reproducido por semi-
llas y han florecido al menos en Costa Rica,
Alemania y Estados Unidos de Norteam&rica. El
hibrido artificial ha sido llamado Trichopilia x
Charles (Dressler, com. pers. 2001).
LANKESTERIANA
Trichopilia x ramonensis J. Garcia & Mora-Ret. ex
C.O. Morales, nothosp. nova FIG. 1
HOLOTIPO: COSTA RICA. Alajuela; cant6n San
Ram6n, distrito San Rafael. Berlin. 1003'N,
8428'0. Leg. Luis Acosta s.n. Floreci6 en marzo de
1991. USJ-57879 (en liquido.
Nothospecies nova hic descripta, sed multos annos
ab egregiis orchideologis nota, apud J. Garcia et
Mora-Retana in anno 1992 iam citata sed adhuc non
descripta. Epiphyta parva, pseudobulbis dense aggre-
gatis, elongatis, 6-11 cm longis, 1,7-2,1 cm latis, plus
minusve rectangulatis, complanatis, interdum falcatis.
Foliis elliptico-lanceolatis, petiolo 0,5-2,0 cm long,
lamina 12-24 cm longa, 3,5-6,9 cm lata.
Inflorescentia brevis, saepe 1 vel 3 (5) floribus.
Sepalis petalisque rubriusculis, sepalis lateralibus ad
basim 1,0-1,5 cm connatis; labello roseo-purpureo,
inter Trichopiliam marginatam et T. suavem interme-
die structurato, 6,1-7,5 cm long, 4,7-5,8 cm lato
[complanato disposito], conspicue venato, ad apicem
ampliato, profunde emarginato et obscure tetralobato,
marginibus valde undulatis simul irregulatis; column
erecta, 2,8-3,1 cm longa, ad basim 1,0-1,5 cm cum
labello connata; clinandrio supra anthera conspicue
laciniato. Fructum non vidi.
Epifita con pseudobulbos densamente agrupados,
alargados, 6-11 x 1,7-2,1 (2,5) cm, un poco mis
anchos cerca de la base, de forma mis o menos rec-
tangular, con bordes laterales casi paralelos, aplana-
dos, a veces falcados. Hojas eliptico-lanceoladas,
peciolo 0,5-2 cm de largo, limina 12-24 cm de largo,
3,5-6,9 cm de ancho maximo. Inflorescencia un raci-
mo corto, generalmente arqueado, a veces colgante,
surgiendo de la base de un pseudobulbo nuevo; con el
raquis liso, glabro, levemente aplanado dorsiventral-
mente, de 3 a 5 mm de diametro, entrenudos de 0,6 a
2,5 (3) cm de largo, entrenudo terminal reducido, a
veces con una brictea est&ril. Brdcteas florales
ovado-lanceoladas, agudas, mis anchas en la base,
1,8 cm largo, 1,2-1,4 cm de ancho maximo, grises,
papiriceas, con puntos y manchas marr6n claro.
Pedicelo levemente curvo hasta doblado en hoz, 1,3 a
1,5 cm de largo. Flores generalmente 1 a 3 (5); sepa-
los y p6talos pardo-rojizos, labelo roseo-puirpura;
ovario 2,7-2,8 cm de largo, sulcado; sepalo dorsal
5,7-6,6 x 1,1-1,6 cm, lanceolado; sepalos laterales
5,3-7,1 x 1,0-1,3 cm, lanceolados, connados 1,0-1,5
cm en la base, a menudo retrorsos en la parte media,
con el haz ligeramente acanalado en el tercio proxi-
mal; petalos 5,2-6,7 x 1,1-1,5 cm, oblongos, ligera-
mente fusiformes, agudos a apiculados; labelo 6,4-7,5
cm de largo, 5,3-5,8 cm de ancho maximo [siendo
extendido y ligeramente aplanado], formando un tubo
ensanchado, abierto y extendido hacia el apice, de
forma intermedia entire T. marginata y T. suavis,
envolviendo la column, profundamente emarginado
en el apice y oscura e irregularmente tetralobado en la
parte distal, con bordes muy ondulados e irregulares,
16bulos distales 2,2-2,8 x 1,1-2,3 cm, venaci6n longi-
tudinal conspicua, callo simple: una quilla estrecha
elevada en el centro del labelo hasta ca. 0,5 cm mis
arriba de la column; column 2,8-3,1 cm de largo
(hasta la antera), 0,4-0,5 cm de diimetro, recta, para-
lela a la base estrecha del labelo, ventralmente fusio-
nada 1,0-1,5 cm con la base del labelo; clinandrio
prominent, laminar, finamente laciniado, de hasta
0,8 cm sobre la antera y rodeindola; antera 0,5 cm de
largo, 0,35 cm de ancho, ca. 0,30 cm de alto; polinios
2, cada uno 2,5 x ca. 1,5 mm.
PARATIPOS: COSTA RICA. Alajuela: San Ram6n. Cataratas.
Leg. Luis Acosta s.n. 19 de marzo de 1992. USJ-44878
(material seco, piezas florales extendidas). "Trichopilia cf.
ramonensis (Hibrido natural). Tiene caracteres intermedios
entire T. marginata y T. suavis, sobre todo en el labelo"
(det. Joaquin Garcia & Dora E. Mora). Alajuela: San
Ram6n [sin localidad exacta]. Leg. Luis Acosta s.n.
Febrero de 1987. Floreci6 en cultivo, marzo de 1990. USJ-
31985 (material seco, un pseudobulbo, una hoja y una
inflorescencia con tres flores). "Hibrido natural. Plantas
con pseudobulbos alargados, con hojas eliptico-lanceo-
ladas, anchas. Flores en racimo de 3-5, blanco-rosadas, con
puntos y manchas rosado oscuro".
MATERIAL ADICIONAL VISTO: COSTA RICA. Alajuela: San
Ram6n [sin localidad exacta]. Leg. desconocido. Floreci6
en enero de 2002, Jardin Botanico Lankester, Cartago,
1350 m (JBL # 33). Planta dibujada, FIG. lA.
"Hibrido artificial. Cruce hecho por Claudio Salas.
Floreci6 en marzo de 1991". USJ-57881 (en liquido.
Este hibrido natural se reconoce por sus pseudobul-
bos alargados y estrechos, inflorescencias cortas,
arqueadas, con 1 a 5 flores, con el labelo roseo-pur-
pura, ensanchado y abierto hacia el apice, profunda-
mente emarginado, fuerte e irregularmente ondulado
y oscuramente tetralobado en la parte distal. Una va-
riante de T. marginata s.l. con labelo pfirpura ha sido
confundida con el hibrido (por ejemplo, USJ-57902,
en liquid, de la Reserva Biol6gica Alberto Brenes,
San Ram6n, Alajuela), pero las flores del hibrido son
MORALES Trichopilia x ramonensis
4 cm
2 cm
Figura 1. Trichopilia x ramonensis J. Garcia & Mora-Ret. ex C.O. Morales. A. Habito; plant con restos de dos inflo-
rescencias. B. Sepalo dorsal. C. Sepalos laterales. D. PNtalo. E. Labelo (algo extendido y aplanado). F. Columna con
pedicelo (N.B. clinandrio incomplete). G. Antera, vista dorsal. H. Antera, vista ventral. I. Polinios. A: Dibujo del autor;
plant del Jardin Botdnico Lankester. B-I: Dibujos del holotipo (USJ), por Marcia GonzAlez Garay.
Setiembre 2002
3mm
LANKESTERIANA
much mis grandes y voluminosas, con el labelo mis
extendido y ondulado en la parte distal. Los pseudo-
bulbos son morfol6gicamente muy similares en el
hibrido y en T. marginata, pero en esta especie a
menudo son much mis largos que en el hibrido.
ILUSTRACIONES: Lucke (2000) y Bock & GruB (2000)
ofrecen excelentes fotos en colors del hibrido y de
los progenitores. En Horich (1996) se halla una
fotografia del hibrido en blanco y negro.
ETIMOLOGIA: El epiteto latinizado ramonensis
(espafiol: ramonense) es el gentilicio que se refiere a
San Ram6n, el cant6n donde se ha hallado el hibrido
natural.
Las flores observadas y medidas muestran notables
variaciones de tamafio. Una flor de un cruce artificial
(C. Salas s.n., USJ, en liquido, que tambi6n fue
media detalladamente, exhibe un tamafio de piezas
florales menor que en el holotipo; sin embargo, es
ffcil predecir que en hibridos artificiales mantenidos
en 6ptimas condiciones de cultivo las flores seran
mis grandes que en plants silvestres. Es probable
que la variaci6n en tamafio depend no s6lo del am-
biente, sino tambien de la especie en la que se form
la capsula. Probablemente, las flores son mis grandes
cuando la capsula se form en T suavis, cuyo labelo
es mis voluminoso que el de T. marginata. En el
hibrido, a menudo el labelo posee el borde blanco, lo
cual tambi6n, al parecer, depend del origen de la
capsula (R.L. Dresser, comun. pers. 2002). En el
campo, es probable la aparici6n de especimenes con
s6palos laterales libres o unidos s6lo por unos
milimetros, tal como se observa en el hibrido artifi-
cial mencionado. Ademis, el taxon que, en sentido
amplio, conocemos como T. marginata es notable-
mente variable, sobre todo en el color de las flores, lo
que seguramente determine cierto grado de variabili-
dad en el hibrido. Los ejemplares secos de herbario
muestran piezas florales notablemente reducidas de
tamafio respect a ejemplares preservados en alcohol
o formalina. Por ello, todas las mediciones de 6rganos
vegetativos y piezas florales de la descripci6n corres-
ponden a ejemplares vivos o preservados en liquid.
DISTRIBUCI6N Y ECOLOGiA: Horich (1996: 182) sitiua
el hibrido entire Angeles Norte y el rio San Lorenzo,
cant6n de San Ram6n, a altitudes entire 900 y 1100 m.
Ahora sabemos que la distribuci6n es mis amplia.
Trichopilia x ramonensis habitat en areas de bosque
tropical huimedo y premontano humedo, entire 800 y
1100 m de altitude, en la Cordillera de Tilarin,
incluyendo la Reserva Biol6gica Alberto Brenes, en
San Ram6n, puesto que alli se han recolectado las dos
species progenitoras (ejemplares en CR y USJ).
Tambien es probable la presencia de este hibrido en la
zona montafiosa de Monteverde, Puntarenas, que co-
linda con la Reserva Biol6gica Alberto Brenes.
Ciertas plants procedentes de algun lugar del can-
t6n de Perez Zeled6n, zona sur de Costa Rica, culti-
vadas en el Jardin Botinico Lankester, son muy simi-
lares a las de T x ramonensis, tanto vegetativamente
como respect a color y morfologia floral. Sin embar-
go, las pocas flores de esas plants que he visto son
algo mis pequefias, el labelo es menos ondulado y
tiene l6bulos mejor definidos que en las plants de
San Ram6n. En Bock & GruB (2000: 758) la foto 2,
determinada como Trichopilia marginata 'W6ssen',
muestra flores con labelo puirpura muy similares a las
de P&rez Zeled6n. Por cortesia de R.L. Dressler
(comun. pers., abril de 2002) he visto la foto de una
plant florecida de Panama que, casi con seguridad,
es T. x ramonensis. En Die Orchidee 43(6): 52
(Orchideenbewertung 1991), 1992, aparece una
fotografia en colors determinada como "Trichopilia
elegans 'Mariechen' SM(B) / D.O.G.", que muestra
una similitud extraordinaria con el hibrido aqui
descrito. El nombre "T. elegans" no aparece vilida-
mente publicado. Es necesario revisar detalladamente
el material del sur de Costa Rica y de Panama. Muy
probablemente, pronto se revelar6 que el ambito de
distribuci6n del hibrido natural es much mayor que
el conocido hasta ahora. Te6ricamente, el hibrido
podria hallarse en todas las zonas donde coinciden T.
marginata y T. suavis (Costa Rica y Panama, entire
800 y 1200 m de elevaci6n, aproximadamente; cf.
Mora-Retana & Atwood 1992, pero aparentemente no
existen recolectas de ninguna de las dos species en
Colombia). No obstante, las poblaciones silvestres de
los progenitores han desaparecido o han sido fuerte-
mente diezmadas debido a la destrucci6n de grandes
areas boscosas y a la recolecta illegal de orquideas con
fines comerciales. En la actualidad, es casi impossible
hallar estas species y el hibrido de ambas en los mis-
mos sitios sin protecci6n estatal donde fueron obser-
vados por Pfau, Brenes, Horich y Mora-Retana.
Aunque la &poca de floraci6n de ambas species se
traslapa (Mora-Retana 1999), no sabemos casi nada
sobre las condiciones microambientales que favore-
cen la hibridaci6n natural y el crecimiento de las
plantulas.
MORALES Trichopilia x ramonensis
AGRADECIMIENTOS: Agradezco sinceramente las sugeren-
cias y adiciones de Mario A. Blanco (USJ, FLAS), Franco
Pupulin (Jard. Bot. Lankester) y Robert L. Dresser
(FLAS), que mejoraron sustancialmente el texto. Tambien
agradezco a Marcia Gonzalez las ilustraciones del
holotipo. Aqui se honra la memorial de Joaquin B. Garcia
(1944-2001, vide Lankesteriana 3), quien queda ademas
inmortalizado en Prosthechea joaquingarciana Pupulin
[Selbyana 22(1): 19, fig. 3. 2001] y de Dora Emilia Mora
(1940-2001, vide Lankesteriana 2). Tambien recordamos a
Clarence Klaus Horich (1930-1994), cuyo esfuerzo, asi
como el de otros naturalistas y horticultores alemanes, nos
permit hoy conocer mejor la flora orquidacea de Costa
Rica.
LITERATURE CITADA
Bock, Irene & GruB, 0. 2000. Trichopilia Lindley, Introd.
Nat. Syst. Ed. II: 446, 1836. Orchidee (Hamburg) 51(6):
758-759.
Dresser, R.L. 1993a. Field guide to the orchids of Costa
Rica and Panama. Ithaca and London, Comstock
Publishing Associates/Cornell University Press. 374 p.
Dresser, R.L. 1993b. Phylogeny and classification of the
Orchid Family. Portland, Oregon, Dioscorides Press.
314 p.
Dresser, R.L. 2001. Mesoamerican orchid novelties: 5.
Oncidiinae. Selbyana 22(1): 9-13.
Dresser, R.L. & Williams, N.H. 1970. An overlooked
genus in the Oncidiinae. Amer. Orch. Soc. Bull. 39(11):
988.
Garay, L.A. 1972. Notas sobre la Alianza Trichopilia.
Orquideologia 7(4): 191.
Horich, C.K. 1996. Sch6ne und seltene Trichopilia-Arten
von Zentralamerika. Orchidee (Hamburg) 47(4): 177-
184.
Lucke, E. 2000. Aussaat und Bliite von Trichopilia x
ramonensis. Orchidee (Hamburg) 51(4): 443-445.
Mora-Retana, Dora E. 1999. Trichopilia Lindl. In:
Atwood, J.T. & Mora-Retana, Dora E. Orchidaceae:
Tribe Maxillarieae: Subtribes Maxillariinae and
Oncidiinae. In: Burger, W. (ed.). Flora Costaricensis.
Fieldiana, Bot. n.s. 40: 169-172.
Mora-Retana, Dora E. & Atwood, J.T. 1992. Orchids of
Costa Rica. Part 2. Icon. Pl. Trop. 15: pl. 1497 & 1498.
Mora-Retana, Dora E. & Garcia, J.B. 1992. Lista actua-
lizada de las orquideas de Costa Rica (Orchidaceae).
Brenesia 37: 79-124.
Pupulin, F. 2002. Catalogo revisado y anotado de las
Orchidaceae de Costa Rica. Lankesteriana 4: 1-88.
Williams, N.H., Chase, M.W., Fulcher, T., & Whitten,
W.M. 2001. Molecular systematics of the Oncidiinae
based on evidence from four DNA sequence regions:
expanded circumscriptions of Cyrtochilum, Erycina,
Otoglossum, and Trichocentrum and a new genus
(Orchidaceae). Lindleyana 16(2): 113-139.
Setiembre 2002
Page
Missing
or
Unavailable
LANKESTERIANA5: 23-25. 2002.
NOTAS VARIAS SOBRE HELICONIA RODRIGUEZII
(HELICONIACEAE) DE COSTA RICA
CARLOS 0. MORALES
Escuela de Biologia, Universidad de Costa Rica. 2060 San Jose, Costa Rica,
y Jardin Botanico Lankester. Apdo. 1031-7050 Cartago, Costa Rica
ABSTRACT. Heliconia rodriguezii was described in 1982, but all the Costa Rican specimens were at the
USJ Herbarium. Isotypes were distributed to other herbaria lately in May and June 2002. Moreover the
herbarium acronyms for the isotypes are erroneously cited in the original publication. Notes on distribu-
tion, ecology, and morphology of this rare, endemic, and poorly known species are given.
RESUMEN. Heliconia rodriguezii fue descrita en 1982, pero todos los especimenes de Costa Rica se halla-
ban en el Herbario USJ. Los isotipos se enviaron a otros herbarios hasta mayo y junio de 2002. Ademis,
los acr6nimos de herbarios para los isotipos se citan mal en la publicaci6n original. Se anotan datos
sobre distribuci6n, ecologia y morfologia de esta especie endemica, escasa y poco conocida.
PALABRAS CLAVE / KEY WORDS: Heliconiaceae, Heliconia rodriguezii, especimenes tipo, Costa Rica
En Costa Rica se conocen unas 42 species nativas
de Heliconia. De estas, unas once species (ca. 26 %)
son endemicas de este pais y es probable que afin se
describan algunas species nuevas. Aunque en la lite-
ratura aparecen unos 450 nombres de species, va-
riedades e hibridos, Berry & Kress (1991) situian
entire 200 y 250 el niamero total de species de
Heliconia. Asi, la diversidad registrada en Costa Rica
represent entire 15 y 20 % de las species del g6nero.
Stiles (1985) sefiala que el centro de diversificaci6n
se halla en la Sudamerica tropical. En Costa Rica,
seguin las observaciones del mismo autor, la mayor
diversidad de Heliconia se concentra en la vertiente
atlantica al pie de las cordilleras Central y de
Talamanca, entire 100 y 700 m de altitude, y en la
Peninsula de Osa y areas vecinas.
En lugar de estudiar material de herbario, Daniels
& Stiles (1979) y Stiles (1975, 1979, 1980, 1982,
1985) observaron y describieron las heliconias in
vivo. De este modo, su trabajo ha sido critic para
entender tanto la taxonomia heliconiacea costarri-
cense como la estrecha relaci6n ecol6gica y evolutiva
existente entire estas plants y dos grupos de aves:
colibries, que polinizan todas las species de
Heliconia de Costa Rica, y aves frugivoras, que dis-
persan frutos y semillas.
Una de las species menos conocidas, Heliconia
rodriguezii F. G. Stiles (1982: 222), dedicada al
insigne botinico, artist y humanista costarricense
Rafael Lucas Rodriguez, fue descrita de La Montura,
en el Parque Nacional Braulio Carrillo, donde, segun
el autor, la especie es comfin en laderas y barrancas
boscosas y se halla a menudo en claros ocasionados
por caida de arboles. Los datos del tipo indican lo
siguiente: "Stiles 80-27, forest understory at La
Montura, el. 1050 m, Parque Nacional Braulio
Carrillo, Provincia San Jose, Costa Rica, 27 July
1980; UCR holotypee), MNCR, FMNH, USNM"
Tenemos dos problems con estos datos:
1) Los acr6nimos de los herbarios citados son
incorrectos. Sin duda, "UCR" se refiere al Herbario
de la Universidad de Costa Rica (USJ), "MNCR"
alude al Herbario Nacional (CR), en el Museo
Nacional de Costa Rica, "FMNH" es el Herbario del
Field Museum of Natural History (F) en Chicago,
EE.UU., y "USNM" tiene que ser el United States
National Herbarium (US), en Washington, EE.UU.
2) En USJ observe siete ejemplares completes del
tipo de Heliconia rodriguezii, lo que claramente reve-
la que los isotipos citados por Stiles nunca fueron
enviados a los respectivos herbarios. Esto ya fue con-
firmado por curadores de los herbarios CR, F y US.
En mayo y junio de 2002 han sido enviados ejem-
plares isotipos a los herbarios CR, F, M, ULM y US,
LANKESTERIANA
Figura. 1. Heliconia -- ....: F.G. Stiles. Isotipo (USJ). Dibujo del autor.
mientras que el holotipo (asi anotado por Stiles) y un La primera se diferencia de &stas dos por el hibito
isotipo permanecen en USJ. mis musoide, vistago (pseudotallo) mis grande y
mis robusto, hojas mis grandes e inflorescencias mis
Seguin Stiles, las species mis emparentadas con anchas y voluminosas. Stiles consider que, mor-
H. rodriguezii (Fig. 1) son H. gracilis G.S. Daniels & fol6gicamente, H. rodriguezii es un eslab6n entire las
F.G. Stiles y H. ignescens G.S. Daniels & F.G. Stiles. dos species mencionadas. Uno de los isotipos (F)
MORALES Notas sobre Heliconia rodriguezii
indica: "Planta de 1.5 m de altura. Inflorescencia roja,
flor amarillenta." Kress (in Hammel et al., ined.)
sefiala que las bricteas medias poseen una superficie
externa glabra, roja a rojo-anaranjada. Ademas, indica
en su clave de las heliconias de Costa Rica que H.
ignescens tiene liminas foliares menores que 50 cm
de largo y bracteas cincinales anaranjadas, mientras
que H. rodriguezii muestra liminas foliares mayores
que 50 cm de largo y bricteas cincinales rojas. Esta
especie, que aparentemente florece la mayor parte del
afio, se consider rara y es endemica en el Parque
Nacional Braulio Carrillo, en las laderas atlAnticas de
la Cordillera Central, entire 800 y 1200 m de ele-
vaci6n, justamente en el Area de mayor diversidad del
genero Heliconia en Costa Rica. Stiles (1982) incluye
una fotografia de la inflorescencia de H. rodriguezii
en blanco y negro. En Berry & Kress (1991: 167)
aparece una fotografia en colors. En esta filtima
obra, asi como en Daniels & Stiles (1979), aparecen
tambien fotografias en colors de las dos species
emparentadas, mencionadas anteriormente.
OTROS ESPECIMENES VISTOS:
F.G. Stiles 80-32 (USJ). Localidad tipica. 27 de julio de
1980. "... flor amarilla."
F.G. Stiles 83-68 (USJ). "Braulio Carrillo". 8 de abril de
1983.
F.G. Stiles s.n. (USJ-30864). Localidad tipica, 1020 m de
altitude. 23 de abril de 1989. "Br6cteas rojas; flores amari-
llas."
Es indudable que H. rodriguezii ha permanecido
desconocida para la mayoria de botanicos que traba-
jan en Costa Rica, en parte debido a que los isotipos
no se distribuyeron. Ademas, muy pocos botinicos
han recolectado esta especie despues de Stiles. Asi,
por ejemplo, antes de mayo de 2002 no habia ningfin
ejemplar en CR. En la base de datos TROPICOS, del
Jardin Botinico de Missouri, se cita un solo ejemplar:
B.A. Loiselle 224 (MO) [Heredia, 1530 m, 10 14' N,
84 05' W, January 1987, 1-1.5 m; red upright bracts;
inflorescence and unripe fruit in January]. La base de
datos ATTA, del Instituto Nacional de Biodiversidad,
incluye asimismo un solo ejemplar: P.J.M. Maas
7809 (INB) [Area del Rio Sucio, Parque Nacional
Braulio Carrillo]. Durante muchos afios esta especie
ni siquiera apareci6 en algunas listas de plants de
Costa Rica.
LITERATURE CITADA
Berry, F. & Kress, W.J. 1991. Heliconia, an identification
guide. Washington/London. Smithsonian Institution
Press. 334 p.
Daniels, G.S. & Stiles, F.G. 1979. The Heliconia taxa of
Costa Rica. Keys and descriptions. Brenesia 15 (Supl.):
1-150.
Institute Nacional de Biodiversidad (INBio). 2002. Sistema
ATTA. http://atta.inbio.ac.cr/
Kress, W.J. Heliconiaceae. In: Hammel, B.E., Zamora, N.
& Grayum, M.H. (eds.). Manual de Plantas de Costa
Rica. Vol. 1: Monocotiled6neas. Heredia, INBio, y St.
Louis, Missouri Bot. Gard. (manuscrito).
Missouri Botanical Garden. 2002. TROPICOS.
http://mobot.mobot.org/W3T/Search/vast.html
Stiles, F.G. 1975. Ecology, flowering phenology, and
hummingbird pollination of some Costa Rican
Heliconia species. Ecology 52: 285-301.
Stiles, F.G. 1979. Notes on the natural history of Heliconia
(Musaceae) in Costa Rica. Brenesia 15 (Supl.): 151-
180.
Stiles, F.G. 1980. Further data on the genus Heliconia
(Musaceae) in northern Costa Rica. Brenesia 18: 147-
154.
Stiles, F.G. 1982. Taxonomic and distributional notes on
Costa Rican Heliconia (Musaceae), II: Parque Nacional
Braulio Carrillo, Cordillera Central. Brenesia 19/20:
221-230.
Stiles, F.G. 1985. Aspectos de la ecologia, distribuci6n y
evoluci6n del genero Heliconia en Costa Rica. In:
D'Arcy, W.G. & Correa, Mireya D. (eds.). The botany
and natural history of Panama / La botanica e historic
natural de Panama. p. 273-285. Missouri Botanical
Garden.
Setiembre 2002
Page
Missing
or
Unavailable
LANKESTERIANA 5: 27-30. 2002.
A NEW SPECIES OF STELLILABIUM SECTION TAENIORHACHIS
(ORCHIDACEAE) FROM COSTA RICA
FRANCO PUPULIN' and MARIO A. BLANCO2
'Jardin Botinico Lankester, Universidad de Costa Rica
P.O. Box 1031-7050 Cartago, Costa Rica, A.C. fpupulin@cariari.ucr.ac.cr
Research Associate, Marie Selby Botanical Gardens, Sarasota, USA
2 nstituto Centroamericano de Investigaci6n Biol6gica y Conservaci6n
P.O. Box 2398-250, San Pedro de Montes de Oca, San Jose, Costa Rica
Research Associate, Jardin Botinico Lankester, Universidad de Costa Rica
Present mailing address: University of Florida, Department of Botany, 220 Bartram Hall
P.O. Box 118526 Gainesville, Florida 32611-8526, U.S.A. mblanco@flmnh.ufl.edu
RESUMEN. Se describe e ilustra Stellilabium smaragdinum de los bosques montanos de encino de la
Cordillera de Talamanca. S. smaragdinum se distingue de otras species de la secci6n Taeniorhachis por
su inflorescencia con 1 a 3 (4) flores simultineas, los 16bulos basales del labelo oblongos a ovoides y
redondeados en el apice y la peculiar coloraci6n de la setas en los 16bulos laterales de la column, marfil
con bandas puirpura.
ABSTRACT. The new species Stellilabium smaragdinum is described and illustrated from the montane oak
forests of the Cordillera de Talamanca, Costa Rica. Among the species of Sect. Taeniorhachis, S.
smaragdinum may be recognized for the inflorescence bearing 1-3 (4) simultaneous flowers, the oblong
to ovoid, rounded basal lobules of the lip, the non-ciliate margins of the lip midlobe and the peculiar
colour of the column setae, cream banded with purple.
KEY WORDS / PALABRAS CLAVE: Stellilabium smaragdinum, Stellilabium sect. Taeniorhachis,
Orchidaceae, Costa Rica.
Mesoamerican species of the genus Stellilabium
Schltr. (Orchidaceae) are not easy plants to spot in the
field. In many cases they lack leaves and also when
the leaves are present they are usually very small,
rarely reaching 3 cm in lenght, so one needs to rely
on inflorescences to distinguish them among the rich
epiphytic vegetation of Neotropical forests. It is likely
this is the main reason why two thirds of
Mesoamerican species of the genus were only descri-
bed in the last twelve years (Atwood 1989; Salazar-
Chavez & Higsater 1991, Atwood & Dressler 1995,
Dressler 1999, 2001), thanks to the help of resident
botanists carrying out systematic collections and stu-
dies intended for large floristic projects.
Generic circumscription of Stellilabium was revi-
sed by Braas & Liickel (1982b), Garay & Romero-
Gonzalez (1998) and Dressler (1999). Stellilabium in
the strict sense, characterized by flowers with simple
column, is a group eminently Andean in distribution,
perhaps closely allied to Trichoceros Kunth and
Telipogon Kunth, which includes Cordanthera L.O.
Williams, Darwiniella Braas & Liickel, and
Sodiroella ecuadorensis Schltr. A formal reinstate-
ment of Dipterostele Schltr. was proposed by Braas
& Liickel (1982a, 1982b) to include those species
presenting narrowly triangular-acuminate lip and bri-
stles of the column short. Garay & Romero-Gonzilez
(1998) adopted a much wider circumscription of the
genus Dipterostele Schltr. and formally assigned to it
all the Stellilabium species with distinct lateral lobes
on the column and a cucullate clinandrium. Dressler
(1999) interpreted Stellilabium in a broad sense,
including Dipterostele Schltr., and identified four
distinctive groups within the genus giving them sec-
tional status. Section Stellilabium and Sect.
Dipterostele (Schltr.) Dressler, both with persistent
leaves, mainly terete or triangular rhachis, and mar-
gins of floral bracts not decurrent on rhachis, are
exclusively South American. Species of Stellilabium
from Mesoamerica, on the contrary, present epheme-
LANKESTERIANA
ral leaves and margins of floral bracts decurrent on
the rhachis, so that the flowers are borne on face
rather than on the edges of the flattened rhachis.
Although their relationships are yet unclear, among
species from Central America two main groups may
be distinguished. In Section Taeniorhachis Dressler
the column is 3-lobed, and the fleshy stigma is adnate
to the base of the lip, whereas species of Sect.
Ramphostele Dressler have simple column and a por-
rect stigma free from the lip. With the notable excep-
tion of S. lankesteri Ames, flowers of the latter sec-
tion are completely glabrous (Dressler 1999).
Samples of DNA sequences for the genus are still
scanty, but they support distinction between sections
Stellilabium, Dipterostele and Taeniorhachis (com.
by N.H. Williams, in Dressler 2001), though the
three groups may eventually prove to be subgroups
of Telipogon. It should be noted that if Stellilabium
Sect. Taeniorhachis is ever elevated to the generic
rank it would have to be renamed, since that name is
already occupied by one species in Poaceae (Cope
1993).
A species pertaining to Stellilabium section
Taeniorhachis from the Talamanca range in Central
Costa Rica is here described as new to science:
Stellilabium smaragdinum Pupulin & M.A. Blanco,
sp. nov. FIG. 1.
TYPE. COSTA RICA. Cartago. El Guarco, La Chonta,
Turbera, 0942'00"N 8356'20"W, 2400 m, epifita
sobre Quercus sp., en ramas bajas, 17 julio 2001, M.
Blanco 1965, F. Rizo-Patr6n, A. Vasco & 0. Vargas
holotypee, USJ!, USJ-Spirit!; isotypes, CR!, SEL!).
Inter species sections Taeniorhachidi floribus 1 vel 4
simultaneis lobulis lateralibus labelli oblongis vel
ovoideis, rotundatis, marginibus lobi intermedi labelli
integris, setis columnae eburneis purpureo striatis
dignoscenda.
Plant epiphytic, small, nearly acaulescent, the
abbreviated stem about 5 mm long. Roots thick,
flexuous, rounded in section, ca. 1 mm wide. Leaves
3-4, distichous, narrowly elliptic to lanceolate, acute,
minutely apiculate, 15-18 mm long, 2-3 mm wide, the
base enclosing the subterete stem, sometimes absent
at flowering. Inflorescence a primarily simple raceme
10-33 cm long, producing secondary branches in suc-
cession from the lower bracts of the previous axis;
peduncle terete, 5-9 cm long, with 3-4 triangular,
acute bracts about 4 mm long; rhachis flattened, 1-1.3
mm wide, successively many-flowered (to 27), with
1-3 (4) flowers simultaneously opened; floral bracts
triangular, acute, from the margin of the flattened rha-
chis, 2 mm long, 1.2 mm wide. Ovary linear-subcla-
vate, 3.6-4.6 mm long including the pedicel. Flowers
small, with olive-green sepals and petals, the lip and
column emerald green, the dendroid hairs of the
column cream, banded purple. Dorsal sepal widely
elliptic, obtuse, with a rounded apicule, 2.9 mm long,
2.4 mm wide. Lateral sepals obliquely widely ellip-
tic-suborbicular, rounded, provided with a rounded
apicule, the apical margins somewhat involute in
natural position, 2.7 mm long, 2.4 mm wide. Petals
narrowly elliptic, slightly falcate, subobtuse, apicula-
te, the margins ciliate, 3.7 mm long, 1.7 mm wide.
Lip sagittate, 3.4 mm long, 2.3 mm wide, covered
with sparse, stiff hairs, the basal lobules oblong to
ovoid, rounded, flattened, the median lobe ligulate,
velutine, with a prominent, acute apicule, the margins
minutely ciliate. Column 3-lobed, the lateral lobes
elliptic, flattened, 1.5 mm long, 1 mm wide, the dor-
sal margins covered with dendroid-stellate setae ca.
0.6 mm long, the median lobe helmet-shaped, roun-
ded, 0.8 mm long, 0.7 mm wide, covered with simple
setae 0.2-0.3 mm long; stigma globose, rounded.
Anther cap cucullate, deeply cordate, 2-celled.
Pollinia 4 in two pairs of different size, obovate, late-
rally complanate-concave, on a narrow, ligulate stipe;
viscidium uncinate. Fruit an elliptic-globose capsule,
4 mm long.
ETIMOLOGY: From the Latin smaragdinus, emerald-
green, in reference to the bright green color of lip and
column.
ECOLOGY: Epiphytic on shady, lower branchlets
amidst a dense cover of mosses and other epiphytes
in wet montane oak forest at 2400 m altitude. More
plants possibly grew higher up the tree, where they
would be virtually impossible to detect from the
ground.
Among the species of Stellilabium Sect.
Taeniorhachis, S. smaragdinum may be recognized
Figure 1. Stellilabium smaragdinum Pupulin & M.A. Blanco. A, B Habit. C Plant. D Flower. E Dissected
perianth. F Column and lip, lateral view. G Setae of lateral lobes of column. H Setae of midlobe of colum. L -
Anther cap and pollinarium.
Setiembre 2002 PUPULIN and BLANCO A new Stellilabium 29
,/" 5 mm
3 mm
1 mm
II
,E B ',\,
S1 cm
'/ L( /
5 mm
LANKESTERIANA
for the inflorescence bearing 1-3 (4) simultaneous
flowers, the elliptic, rounded lateral lobes of lip, the
non-ciliate margins of the lip midlobe and the pecu-
liar colour of the column setae, cream banded with
purple. Stellilabium smaragdinum is closely related to
other species of Sect. Taeniorhachis presenting den-
droid-stellate bristles on the lateral lobes of column
and simple setae on the midlobe (which sometimes is
only glandular-papillose), i.e. S. butcheri Dressler, S.
erratum Dressler, S. helleri L.O. Williams, and S.
minutiflorum (Kraenzl.) Garay. However, plants of
the latter species are much smaller (the type, Endrds
s.n., W-R 2006!, 2007!, 2011!) and the flower pre-
sents a smaller lip with short, narrowly triangular
basal lobes (illustration of type, W!. See discussion
on S. minutiflorum in Dressler 1999: 471).
Stellilabium helleri has a fringe of hairs in front of the
column that are absent in S. smaragdinum, and the
column midlobe is obscurely muricate. Stellilabium
erratum has dark wine-purple flowers with compara-
tively narrower petals and a markedly hastate lip. The
Panamanian S. butcheri presents a convex lip with
papillose-hispid margins and narrow basal lobules.
ACKNOWLEDGMENTS: The type was collected during the
field course "Sistematica de Plantas Tropicales" (OET 01-
18), under the auspices of the Organization for Tropical
Studies, funded in part by the Andrew W. Mellon
Foundation.
LITERATURE CITED
Atwood, J.T. 1989. Orchids of Costa Rica, part 1. Icon. Pl. Trop. 14: pl. 1392-1395.
Atwood, J.T. & R.L. Dresser. 1995. Another new species of Stellilabium from Monteverde, Costa Rica. Selbyana 16:
239-241.
Braas, L.A. & E. Liickel. 1982a. Darwiniella, eine neue Gattung der Subtribus Telipogoninae Schltr. Orchidee
(Hamburg) 33: 167-170.
1982b. Die Gattungen mit Bestimmungsschliissel der T /ip',,'u-Verwandtschaft (Subtribus Telipogoninae
Schltr.): T lipqtowi H.B.K., Trichoceros H.B.K., Stellilabium Schltr, Dipterostele Schltr., Darwiniella Braas &
Liickel. Orchidee (Hamburg) 33: 170-176.
Cope, T.A. 1993. Taeniorhachis: a new genus of Gramineae from Somalia. Kew Bull. 48: 403-405.
Dresser, R.L. 1999. A reconsideration of Stellilabium and Dipterostele. Harvard Pap. Bot. 4: 469-473.
2001. Stellilabium erratum, a comedy of blunders. Lankesteriana 2: 11-13.
Garay, L.A. & G.A. Romero-Gonzalez. 1998. Schedulae Orchidum. Harvard Pap. Bot. 3: 53-62.
Salazar Chavez, G.A. & E. Hagsater. 1991. Stellilabium standleyi, unico representante mexicano de la subtribu
Telipogoninae (Orchidaceae). Orquidea (Mex.) 12: 231-236.
El JARDIN BOTANIC LANKESTER
Universidad de Costa Rica
anuncia el
1ER CONGRESS INTERNATIONAL DE ORQUIDEOLOGIA NEOTROPICAL
continuacion del SEMINARIO MESOAMERICANO DE ORQUIDEOLOGIA Y CONSERVACION
y
3 ENCUENTRO DEL GRUPO REGIONAL MESOAMERICANO DE ESPECIALISTAS EN ORQUIDEAS, UICN/CSE
The JARDIN BOTANICO LANKESTER
Universidad de Costa Rica
announces the
1"s INTERNATIONAL CONFERENCE OF NEOTROPICAL ORCHIDOLOGY
continuation of the MESOAMER1CAN SEMINAR ON ORCHIDOLOGY AND CONSERVATION
and
MEETING OF THE REGIONAL MESOAMERICAN ORCHID SPECIALIST GROUP, IUCN/Ssc
San Jos'e, Costa Rica
Ciudad de la Investigacibn de la Universidad de Costa Rica
23 al 26 de mayo del 2003
May 23-26, 2003
Un event cienfifico patrocinado por
A scientific event sponsored by
Fundad6n
Lankester
|
