A new Lycaste (Orchidaceae: Maxillariinae) from Costa Rica

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After its introduction to Europe by the Spanish
expedition to Peru and Chile by H. Ruiz and J.
Pavón in the eigthteeth century, and the description
of the first specimens referable to the genus Lycaste
Lindl., by E. Poeppig & L. Endlicher in Nova
Genera and Species Plantarum (1833-1835), plants
of this genus became one of the most famous and
popular orchids in cultivation (Oakeley 1993). The
genus was created by J. Lindley in 1843 based on
morphologic characters, separating this group of
orchids from the genus Maxillaria Ruiz & Pav.
Molecular studies used to assess the phylogenetic
relationships between the species of Lycaste and
Anguloa Ruiz & Pav. showed that the mainly South
American group of Lycaste section Fimbriatae
Fowlie, deserves a generic status (Ryan et al. 2000).
Ryan & Oakeley (2003) proposed the genus Ida A.
Ryan & Oakeley to include the former group, reduc-
ing Lycaste to species mainly Mesoamerican in distri-
bution.
In spite of this generic separation, both genera have
been recognized for their beauty and horticultural
value. Species pertaining to groups of orchids popular
in cultivation are often well known taxonomically,
and it is not surprising that in his Essential Guide of
Lycaste species, Oakeley (1993) opened his introduc-
tion saying “if it is not in here it has not been
described”. However, when a good taxonomic system
is developed in countries rich in plants species, sup-
ported by an efficient system of information, taxo-
nomic novelties often appear also in previously
monographed and botanically well-sampled genera
(Pupulin & Bogarín 2005, Dressler & Pupulin 2006,
Pupulin 2007).
Hereafter, Lycaste bruncana is presented as a new
species to the science.
Lycaste bruncana Bogarín, sp. nov.
TYPE: COSTA RICA. Puntarenas: Golfito, Golfito,
Esquinas, La Gamba, Bosque de Los Austriacos,
LANKESTERIANA 7(3) : 543-549. 2007.
A NEW LYCASTE (ORCHIDACEAE: MAXILLARIINAE) FROM COSTA RICA
DIEGO BOGARÍN
Jardín Botánico Lankester, Universidad de Costa Rica. P.O. Box 1031-7050 Cartago, Costa Rica, A.C.;
Centro de Investigación en Orquídeas de los Andes “Ángel Andreetta”, Universidad Alfredo Pérez Guerrero,
Extensión Gualaceo, Ecuador.
dbogarin@cariari.ucr.ac.cr
ABSTRACT: Lycaste bruncana from Costa Rica is described and illustrated. This species is sister to L. tricol-
or, its closest relative, from which is easily distinguished by the lip shape, with a narrow isthmus and
spathulate midlobe, the lanate to lanulose infrastigmatic surface, the pollinarium with a longer stipe, the
subrectangular pollinia and the tomentose anther cap. However, among Costa Rican species of Lycaste, the
most remarkable character of L. bruncana is the twisted column, which may be indicative of a reproductive
barrier. Information related to biogeography, distribution, morphology, molecular data and pollination is
provided and discussed.
RESUMEN: Se describe e ilustra Lycaste bruncana de Costa Rica. Esta especie es hermana de L. tricolor, su
pariente más cercana, de la cual se distingue fácilmente por la forma del labelo, con un istmo estrecho y el
lóbulo medio espatulado, la superficie infraestigmática lanada a lanulosa, el polinario con un estípite más
largo, los polinios subrectangulares y la capa de la antera tomentosa. Sin embargo, entre las especies
costarricenses de Lycaste, el carácter más notable de L. bruncana es la columna torcida, la cual puede ser
indicativa de una barrera reproductiva. Se proporciona y discute información relacionada con biogeografía,
distribución, morfología, datos moleculares y polinización.
KEY WORDS: Orchidaceae, Lycaste, Lycaste bruncana, new species, Costa Rica, DNA barcoding, orchid
pollination.

Tropenstation La Gamba, 8°42’40”N 83°13’00”W,
100-400 m, 3-4 Julio 2004, M. Blanco 2610 & E.
Serrano, floreció en cultivo en el Jardín Botánico
Lankester, 19 Julio 2007, D. Bogarín 3987 (holotype:
CR; isotype: JBL-spirit). FIG. 1.
Lycaste tricolori (Klotzsch.) Rchb.f. affinis, a qua
lobulo medio labelli spathulato, isthmo angustiore
longioreque, anthera tomentosa, superficie infrastig-
matica lanato-lanulosa, stipite pollinarii longiore et
apice columnae leviter torto imprimis differt.
Plant epiphytic, cespitose, up to 60 cm tall. Roots
slender, up to 3 mm in diameter, covered by absorp-
tive hairs. Pseudobulbs 5-7 x3-4.5 cm, ovoid, sulcate,
ancipitous, with inconspicuous spines when deciduous
and subtended by scarious sheats. Leaves two (rarely
three) apical leaves and 2-3 lateral leaves adpressed to
each young pseudobulb, articulate, persistent during
the blooming period, deciduous with age. Leaf blade
petiolate, 5.5 x51 cm, plicate, obovate, acute to
acuminate, with 5 conspicuous veins. Inflorescence,
one to six, pedunculate, to 9 cm long, 1-flowered, con-
cealed by 2-3 persistent, imbricate, ovate, acute,
bracts, flowering usually with a new growth. Floral
bract 2.5-3 x2-2.5 cm, ovate, acute, enclosing the
ovary. Ovary cylindrical, sulcate, smooth, 1.7 cm
long, 3.5 mm in diameter. Flowers showy, from 1 to
several open at the same time, sepals pale green to
pinkish, petals white, spotted with pink, lip white with
purple dots around the callus and pink dots in the mid-
lobe. Sepals equal, rectangular-obovate, acute, 4.5 x
1.7 cm, basally tomentulose, margin entire, straight in
natural position. Petals ovate, acute, slightly crenate at
apex, 3.5-4.2 x1.5-1.7 cm, recurved in natural posi-
tion. Lip three-lobed, 2.7 x1.2 cm, attached to the col-
umn foot, with a rectangular thick, emarginate callus
up to 1.3 cm long, lateral lobes rectangular, obtuse,
thick, perpendicular to the callus, 1.2 x0.5 cm, mid-
lobe spathulate, thin, the isthmus narrow, emarginate,
erose at apex, 1.5-1.7 x0.3 cm. Column with a purple-
spotted foot, twisted to the left, basally tomentulose,
silky at apex, infrastigmatic surface lanulose, the stig-
ma ventral; anther apical, up to 2.5 long. Pollinarium
6 mm long, with four pollinia, two rectangular-ovoid
and two subequal, smaller; stipe 3.5 mm long. Anther
cap cucullate, tomentose. Capsule not seen.
PARATYPES: San José: Pérez Zeledón, Pejibaye, Fila
Costeña, Zapote de Pejivalle, 9°07’37.3”N
83°33’22.7”W, 740-810 m, along the margins of a
small river, remnants of primary forest and secondary
forest, 10 December 2000, F. Pupulin 2654, D.
Castelfranco & J. Cambronero (JBL-Spirit). Pérez
Zeledón, San Isidro de El General, carretera a
Dominical, Fila Tinamastes, La Alfombra,
9°19’19.0” N 83°45’58.1” W, 1000 m, bosque húme-
do premontano, bosque secundario alterado y árboles
a orillas de la carretera, 23 Octubre 2005, A.
Karremans 1076, J. Cambronero, R.L. Dressler, R.
Gómez, D. Bogarín, F. Pupulin, A. & S. Rambelli
(JBL-Spirit). Puntarenas: Aguirre, Naranjillo, road
between Quepos and San Marcos de Tarrazú, 750 m,
9°33’01”N 84°05’01”W, wet premontane transition
to lower montane forest remnants of primary and sec-
ondary mature vegetation, 1 January 2001, F. Pupulin
2780, D. Castelfranco & E. Salas (JBL-Spirit).
Golfito, Golfito, Esquinas, La Gamba, Bosque de Los
Austriacos, Tropenstation La Gamba, 8°42’40”N
83°13’00”W, 100-400 m, 3-4 Julio 2004, M. Blanco
2610 & E. Serrano (JBL-Spirit).
DISTRIBUTION: known only from the southern Pacific
watershed of Cordillera de Talamanca and Cordillera
Costeña, Costa Rica, between 700 and 1000 m of ele-
vation. Southernmost collections suggest that this
species may occur around the Chiriquí river valley in
Panama, which represents the southernmost extension
of the Cordillera Costeña. Plants of L. tricolor have
similar elevation range but are restricted to the
Atlantic slopes of Costa Rica and Panama.
ETYMOLOGY: From the emblematic name “Brunca”,
which comes from the indigenous people who live in
the southern Pacific lands of Cordillera de
Talamanca. Brunca also names the Región Brunca,
one of the socioeconomic regions of Costa Rica that
occupies most of the distribution range of this
species.
HABITAT: Plants of this species grew epiphytically in
primary and secondary vegetation in premontane rain
forest, tropical wet forest-premontane belt transition,
tropical moist forest-premontane belt transition and
tropical wet forest. All these life zones strictly corre-
spond to the southern Pacific watershed of Cordillera
de Talamanca and Fila (Cordillera) Costeña.
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BOGARÍN- A NEW Lycaste 545
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
FIGURE 1. A-G. Lycaste bruncana Bogarín. A. Habit. B. Flower in natural position. C. Dissected perianth. D. Lip. E.
Column and lip, lateral view. F. Column. lateral and frontal view. G. Pollinarium (front view, dorsal view and lateral
view) and anther cap. Drawn from the holotype by Diego Bogarín.

CONSERVATION STATUS: The species has been record-
ed in protected areas. The type collection comes from
The Field Station La Gamba, situated in the south of
Puntarenas province, in Golfito. It constitutes the first
record for this research center, where no Lycaste
species were previously recorded (Weissenhofer et al.
2001). Biogeographical similitudes and proximity
between this area, Piedras Blancas National Park and
Corcovado National Park, also suggest that L. brun-
cana may well occur in those protected areas. An
unvouchered specimen in cultivation (J. Clarke 314)
from Boracayán Wildlife Refuge around Fila Alivio
in the upper Río Barucito basin, Cordillera Costeña,
is also recorded from a protected area. Apparently,
this species is not rare and some specimens were col-
lected in disturbed areas. This species should not be
regarded as endangered.
PHENOLOGY: Plants in cultivation flower from May to
July. Some plants can extend the flowering season up
to October.
This species is closely related to L. tricolor, a wide-
spread species from the Atlantic lands of Costa Rica
and Panama. In a survey of Lycaste species in Costa
Rica, Horich (1977) cited several specimens of L. tri-
color from the Atlantic side of both Tilarán and
Central Volcanic chains. Also, he stated that L. tricolor
has no close affinities or intermediate relatives in the
genus. It seems that he never had access to material
from the Pacifc watershed of Cordillera de Talamanca
(Horich1966). Oakeley (199 1, 1993) discussed L. tri-
color as a tiny pink to white flower species with the lip
provided with a marked isthmus, and a projecting dis-
coid callus. All geographic data and pictures presented
in his work are correctly attributed to L. tricolor.
The only record that may eventually correspond to
a first collection of L. bruncana is the “small dwarf
race” (Fowlie 1970: 7) of L. tricolor that was appar-
ently collected in San Vito de Java, a region that is
included in the current distribution area of L. brun-
cana. However, no herbarium specimens or vouchers
of this species were apparently prepared by the author
(Fowlie 1970). Recent treatments of Costa Rican and
Mesoamerican orchidaceae do not mention any col-
lections of Lycaste tricolor from the Pacific water-
shed of Cordillera de Talamanca (Dressler 2003,
Ossenbachet al. 2007).
According to Fowlie (1970) and following his clas-
sification of the genus, both species should be placed
in the section Deciduosae subsection Paradeciduosae
Fowlie. Although the similarities between the two
species (i.e., the three-lobed lip, the deciduous
pseudobulbs without conspicuous spines, the similar
size of the plant, and the gross flower morphology)
could have confused taxonomists in the past, Lycaste
borucana may be easily distinguished on the basis of
a consistent set of morphological features.
Lycaste bruncana is distinguished from L. tricolor
by the lip shape with a narrow isthmus and spathulate
midlobe (vs. ovate), the lanate to lanulose infrastig-
matic surface (vs. smooth), the pollinarium with a
longer stipe (up to 3.5 mm long), the subrectangular
pollinia (vs. ovoid) and the tomentose anther cap (vs.
smooth) (Figs. 2-5). However, the most remarkable
character to recognize L. bruncana is the twisted col-
umn, which may be indicative of a reproductive barri-
er with respect of its relative (Fig. 5). Among the
species of Costa Rican Lycaste, this is the only
species that presents a twisted column (Dressler
2003).
All the former characters are diagnostic and show
no variation among individuals. The length of lip is
variable, ranging from 1.5 to 2.7 cm, however the
midlobe is spathulate in all collections studied (Fig.
2). Small plants cultivated at Lankester Botanical
Garden produced smaller flowers than the robust
plants. The number of flowers produced by a plant
varies from one to six in each growth and it depends
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LANKESTERIANA
546
FIGURE 2. Variation in the lip length among individuals of
L. bruncana. A. from A. Karremans 1076. B. from F.
Pupulin 2654. C. from F. Pupulin 2780. D. from D.
Bogarín 3987. All vouchers in JBL-spirit collection.
Scale bar = 1 cm.

on both the robustness of the plant and the number of
pseudobulbs present. Flower coloration is rather con-
stant, with concolor sepals and petals with pink spots,
however, slightly paler flowers have been observed
among the different collections. The distinctive col-
umn, twisted to the left, showed no variation among
individuals and the individual flowers of the same
plant. All the studied flowers showed twisted
columns.
After the submission of this paper, an anonymous
reviewer suggested commenting on two names not
validly published that supposedly have been used
for this species. Although the International Code of
Botanical Nomenclature (McNeill et al. 2006)
expressly recommends (Recomm. 34A) that authors
should avoid mentioning previously unpublished
names, I found two citations of the name Lycaste
crystallina, both of them nomina nuda, with differ-
BOGARÍN- A NEW Lycaste 547
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FIGURE 5. Comparison of the column shape in L. bruncana
and L. tricolor. A-A’. L. bruncana. A. Upper view,
showing the twisted column. A’. Frontal view, showing
the twistened column and the lanulose infraestigmatic
surface. From A. Karremans 1076 (JBL-spirit). B-B’.
Lycaste tricolor. B. Upper view, showing the straight
column. B’. Frontal view, showing the straight column
and smooth infraestigmatic surface. From D. Bogarín
733 (JBL-spirit). Scale bar = 1 cm.
FIGURE 3. Comparison of the pollinarium shape between
L. bruncana and L. tricolor. A-C, L. bruncana. A. from
D. Bogarín 3987. B. from F. Pupulin 2780. C. from A.
Karremans 1076. A’-C’ L. tricolor. A’. from JBL-
02995. B’. from D. Bogarín 733. C’. from JBL-06408.
All vouchers in JBL-pollinaria collection. Scale bar = 5
mm.
FIGURE 4. Comparison between anther cap shape in L.
bruncana and L. tricolor. A-C. L. bruncana. A. from D.
Bogarín 3987. B. from F. Pupulin 2780. C. from A.
Karremans 1076. A’-B’ L. tricolor. A’. from D.
Bogarín 733. B’. from JBL-06408. All vouchers in JBL-
spirit collection. Scale bar = 5 mm. MAP 1. Distribution of L. bruncana and L. tricolor.

ent authorship. Lycaste crystallina M.A. Dix, nom.
nud., (Missouri Botanical Garden 2007) is ostensi-
bly based on Blanco 2610, designated here as a
paratype of L. bruncana. The other citation, as
Lycaste crystallina Wubben ex H.F. Oakeley, nom.
nud., was found at BIBLIORCHIDEA (Swiss Orchid
Fundation 2007) and refers to a recent article by
Oakeley (2007) where the author anticipated the
publication of L. crystallina as a new species, how-
ever it does not constitute a valid name (see Code,
art. 34.1). No information at all was retreived about
the other unpublished name suggested by the anony-
mous reviewer.
BIOGEOGRAPHY - According to the locality data avail-
able for both species, they are geographically separat-
ed (Map 1). All the collections of L. tricolor corre-
spond to the Atlantic watershed of the Cordillera de
Tilarán, the Central Volcanic chain and the
Talamanca range. On the other side, all collections of
L. bruncana come from the Pacific watershed of the
Cordillera de Talamanca and the Cordillera Costeña.
Analogous examples of sister orchid species in the
Atlantic and Pacific regions of Talamanca range have
been documented in the genus Kefersteinia (Pupulin
2001). These examples support the role of the
Cordillera de Talamanca as a geographic isolation
barrier, where these sister species have evolved sepa-
rately.
Though the Cordillera Costeña and nearby areas
have been explored botanically since several years
ago, new species have been recently revealed from
the region (Gómez-Laurito 2006). A well defined
floristic pattern is present in Cordillera Costeña and
its neighboring areas. Its geographic position, geolog-
ic history and the inherent climatological characteris-
tics given by the rain patterns and humid winds that
come from the Pacific, make this area rich in
endemisms.
MOLECULAR DATA - Sequences using matK chloro-
plast region obtained to assess the candidate barcode
for flowering plants support the clustering of the
accessions of L. tricolor. This species is placed sepa-
rately from the accessions of L. bruncana. The plant
of L. bruncana from which the type specimen was
prepared (Gene Bank accession EU214496) was used
to obtain the matK sequence analysed for barcoding
pourposes (Lahaye et al. in prep). In the former work,
authors concluded that L. borucana (cited there as
Lycaste cf. tricolor) is a cryptic species supported by
DNA barcoding. Lycaste bruncana does not cluster
with the three accessions of L. tricolor assessed in the
study. In conclusion, molecular evidence supports the
identity of L. bruncana, as a species distinct from its
relative, L. tricolor.
POLLINATION ECOLOGY -Lycaste is a typical bee-pol-
linated flower (Roubik & Hanson 2004). The dis-
tinctive twisted column present in L. bruncana
might also be indicative of a reproductive barrier
whereby L. bruncana would deposit the pollinarium
on the shoulder of the pollinating bees and L. tricol-
or, with its straight column, would deposit the polli-
narium on the scutellum. Pollination studies in
orchids around Golfito area did not reveal any
potential bee pollinator of Lycaste species (Lobo &
Bolaños 2005). More observations and comparisons
between pollination biology of both species may
confirm the reproductive isolation barrier inferred
by flower morphology. This acquires special interest
since the two species overlap their flowering season
so, differences in both, flower morphology and the
way in which these species use the pollinator to
carry the pollinarium, ensure a reproductive isola-
tion. This isolation barrier supports the distinction
between L. bruncana and L. tricolor.
ACKNOWLEDGEMENTS.I want to thank Franco Pupulin
and Robert L. Dressler for their invaluable support and
their useful comments on the paper. To my colleagues José
Daniel Zúñiga and Mario Blanco who kindly provided use-
ful information. The scientific services of Costa Rican
Ministery of Environment and Energy (MINAE) and its
National System of Conservation Areas (SINAC) for issu-
ing the collecting permits under which wild species treated
in this study were collected. The staff of Tropenstation La
Gamba for granting access to the installations and for the
research facilities. The staff of Jodrell Laboratory of Royal
Botanic Gardens, Kew, specially Guillaume Gigot, Martyn
Powell and Vincent Savolainen, for their help with the
DNA barcoding work and to Renaud Lahaye who analysed
the molecular data. The UK Darwin Initiative for the
Survival of Species sponsored field work activities under
the project “Conservation and Monitoring of Meso-
American Orchids” (Ref.14001).
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LANKESTERIANA
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