Content uploaded by A. P. Karremans
Author content
All content in this area was uploaded by A. P. Karremans on Jan 04, 2021
Content may be subject to copyright.
With this work, a rst prodromus (Latin for “precursor”)
to Flora Costaricensis, we are starting a series of papers
that aim to present partial but coherent results obtained
by our group of researchers in the study and critical
appreciation of the orchid ora of Costa Rica. These
prodromi will discuss the systematics of selected genera
or phylogenetically related groups of species in larger
genera, which will be included in forthcoming volumes
of Flora Costaricensis. This magnus opus devoted to the
rich ora of Costa Rica started in 1965 with the visionary
effort of William Burger and has since been led by the Field
Museum of Natural History in Chicago. Thirteen volumes
of Flora Costaricensis have been published to date over a
period of almost 40 years, covering entire plant families
or parts thereof (Burger, 1971; Burger et al., 1977, 1983,
1991, 2000; Pohl, 1980; Durkee and Burger, 1986; Burger
and van der Werff, 1990; Burger and Taylor, 1993; Burger
and Huft, 1995; Atwood and Mora de Retana, 1999; Luteyn
and Wilbur, 2005; Pupulin, 2010).
As the general structure of Flora Costaricensis was
conceived as individual volumes embracing complete
families or at least vast taxonomic groups (to the ranks
of subfamilies, tribes, and subtribes), the completion of
Orchidaceae, with its high internal diversity and the rarity
of certain taxa, proves to be an overwhelming challenge. A
more convenient approach is the publication of individual
papers dealing with the systematics of genera and/or groups
already understood at the specic and intraspecic level, in
anticipation of the nal, comprehensive treatment (see, e.g.,
Pupulin and Bogarín, 2010, 2014b; Pupulin et al., 2010a;
Bogarín, Serracín, and Samudio, 2014; Karremans et al.,
2020). The present prodromus offers the rst systematic
insight into a complete group of Pleurothallidinae orchids
for Flora Costaricensis.
FLORAE COSTARICENSIS SUBTRIBUI PLEUROTHALLIDINIS
PRODROMUS—SYSTEMATICS OF ECHINOSEPALA (ORCHIDACEAE)
Franco PuPulin,1–4 noelia BelFort-oconitrillo,1,5 adam P. Karremans,1,6 and diego Bogarín1,7
Abstract. We provide a general introduction to the prodromi to Flora Costaricensis, and an introduction to the precursors of subtribe
Pleurothallidinae. We present a systematic revision of the genus Echinosepala for the ora of Costa Rica and a detailed discussion
of molecular evidence with regard to most Costa Rican taxa, aimed at phylogenetically framing Echinosepala within the subtribe
Pleurothallidinae, recognizing internal relationships among the species in the genus, and proposing evolutionary hypotheses. We
recognize 10 species of Echinosepala in Costa Rica. Each species is described and illustrated with one or more botanical drawings
and photographs, all based on Costa Rican material. Etymology, distribution, ecology, distinguishing features, and afnities with other
taxa in the genus are discussed, taxonomic notes are provided, and a list of selected material examined for this study is included for
each taxon. Two new species of Echinosepala, E. glenioides and E. isthmica, are described and illustrated, and their relationships are
discussed. New combinations are proposed in Echinosepala for Pleurothallis alexandrae and P. pastacensis, and the former species
is neotypied.
Keywords: ora of Costa Rica, new species, new combination, Pleurothallis alexandrae, Pleurothallis pastacensis, Pleurothallidinae
We would like to thank the staff and curators at AMES, CR, JBL, K, SEL, USJ, and W for their help in retrieving the Echinosepala specimens kept
there and their supportive cooperation. We acknowledge our colleagues at the research department of Lankester Botanical Garden for the unselsh support
they gave throughout the stages of the present work, sharing with us most of the eldwork and providing stimulating ideas, discussion, and materials.
Darha Solano Ulate splendidly rendered most of the illustrations for this paper, proving to be one of the most gifted botanical illustrators active today. The
horticulture staff at Lankester Botanical Garden is thanked for their help in cultivating the living plants documented here. Acknowledgments are extended
to the Costa Rican Ministry of Environment and Energy (MINAE) and its National System of Conservation Areas (SINAC) for issuing the scientic
collection permits under which all the wild specimens for this study were collected. The present paper is part of Project 814-B0-052, “Flora Costaricensis:
taxonomía y logenia de la subtribu Pleurothallidinae, Orchidaceae,ˮ supported by the Vice-Presidency of Research, University of Costa Rica.
1 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Costa Rica
2 Harvard University Herbaria, Cambridge, Massachusetts, U.S.A.
3 The Marie Selby Botanical Gardens, Sarasota, Florida, U.S.A.
4
Corresponding author: franco.pupulin@ucr.ac.cr
5 Orchid Conservation Project, Bosque de Paz Biological Reserve, Bajos del Toro, Costa Rica
6 Naturalis Biodiversity Center, Endless Forms group, Leiden, The Netherlands
7 Herbario UCH, Universidad Autónoma de Chiriquí, David, Panamá
Harvard Papers in Botany, Vol. 25, No. 2, 2020, pp. 155–190.
© President and Fellows of Harvard College, 2020
ISSN: 1938-2944, DOI: 10.3100/hpib.v25iss2.2020.n5, Published online: 31 December 2020
Flora CostariCensis: Pleurothallidinae
When, in 2009, we inscribed a project devoted to the
systematic treatment of the subtribe Pleurothallidinae
Lindl. ex G. Don (Orchidaceae: Epidendreae) for Flora
Costaricensis at the University of Costa Rica (UCR),
we knew that a work of such magnitude had never been
attempted before in a country as highly biodiverse as
Costa Rica. Modern orchid treatments intended for Flora
Costaricensis (see Atwood and Mora de Retana, 1999;
Pupulin, 2010) are aimed at critically discussing the
typication and identity of each taxon to the specic rank,
including subspecic taxa and morphs when required, and
to illustrate from living material all the taxa of a given
group. Subtribe Pleurothallidinae encompasses in Costa
Rica one-third of the total number of orchid species (Pupulin
et al., 2019), making this vast assemblage of taxa one of
the taxonomically more challenging in the country’s ora.
156 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
In the 15 years elapsed since the publication of the senior
author’s checklist of Costa Rican Orchidaceae (Pupulin,
2002), which recorded 427 species of Pleurothallidinae,
and up to the last catalog of pleurothallid orchids given by
Pupulin and coworkers (2019), 134 new species and new
records for the ora of Costa Rica have been added by our
group (Pupulin and Bogarín, 2004, 2007, 2010, 2011a,b,
2012, 2014a,b; Pupulin et al., 2007, 2009, 2010a,b, 2011,
2012; Bogarín et al., 2008, 2013, 2015; Bogarín and
Fernández, 2010; Bogarín and Karremans, 2010, 2016;
Pupulin, Bogarín, and Smith, 2010; Pupulin, Medina, et
al., 2010; Bogarín and Pupulin, 2011; Fernández, 2011;
Fernández and Bogarín, 2011, 2013; Karremans and
Bogarín, 2011, 2013; Karremans and Muñoz García, 2011;
Bogarín, Karremans, et al., 2012; Bogarín, Smith, et al.,
2012; Karremans, 2012, 2014; Karremans and Smith, 2012;
Karremans et al., 2012; Smith and Pupulin, 2012; Fernández,
2013; Smith et al., 2013, 2015; Bogarín and Kisel, 2014;
Bogarín, Serracín, and Samudio, 2014; Fernández et al.,
2014; Bogarín and Jiménez, 2015; Karremans, Bogarín,
et al., 2015; Karremans, Pupulin, et al., 2015; Oses Salas
and Karremans, 2016; Karremans and Díaz-Morales, 2017;
Pupulin, Díaz-Morales, Aguilar, et al., 2017; Pupulin, Díaz-
Morales, Fernández, et al., 2017; Pupulin, Karremans, et al.,
2017). The last available enumeration of Pleurothallidinae
in Costa Rica (Pupulin et al., 2019), which includes data
published until 2017, accounts for 561 species in 30 genera.
After that list, another 20 species were published as new
for science or new records for the ora of the country
(Bogarín et al., 2017, 2020; Karremans and Bogarín,
2017; Rojas-Alvarado and Karremans, 2017; Bogarín,
Fernández, et al., 2018; Bogarín, Karremans, et al., 2018;
Bogarín, Pérez-Escobar, et al., 2018; Karremans and
Jiménez, 2018; Karremans, 2019; Karremans et al., 2019;
Pridgeon, 2020; Pupulin, 2020; Pupulin and Bogarín,
2020; Pupulin and Oses, 2020). Several additional species,
patiently awaiting their turn within the drawers of the
research department at Lankester Botanical Garden, shall
be published in the coming years.
Species richness is not evenly distributed among
Costa Rican Pleurothallids. Traditionally smaller genera
(but please remember that generic denitions and
circumscriptions in the subtribe are still problematic, and
that alternative systems of classication have been proposed
and are sometimes in parallel use) have shown reduced
growth in terms of diversity and number of species, with
increments in overall diversity probably proportional to
the total number of taxa for each genus in the concerned
region. Nonetheless, it is noteworthy that also in most of
the groups that are not centered in Mesoamerica in terms
of species diversity, like Acianthera Scheidw., Anathallis
Barb. Rodr., Dracula Luer, Dresslerella Luer, Myoxanthus
Poepp. & Endl., Octomeria R. Br., Pabstiella Brieger &
Senghas, Restrepiella Garay & Dunst., and Trichosalpinx
Luer, new taxa and new records are continually added to
the checklist of the country’s Pleurothallidinae. The case
is different for the more specious genera like Lepanthes
Sw., Pleurothallis R. Br., and Stelis Sw., each of which
already accounts for around 100–150 species in Costa Rica,
and which together represent more that half of the known
species of Pleurothallidinae in the country. Each of these
genera will include, once their monographic treatments
have been concluded, no less than 150 species.
With the advent of new techniques for the analysis
of molecular data, the supraspecic systematics of the
Pleurothallidinae have been discussed extensively (see,
at least, Pridgeon et al., 2001; Luer, 2002; Karremans,
2016). However, less emphasis has been put into the
effort of clarifying the internal relationships within the
major groups, which would have greatly contributed to a
better understanding not only of the phytogeographic and
phylogenetic patterns of dispersion and speciation of the
largest genera in the different regions of the American
tropics, but also of their true diversity. For almost two
centuries, pleurothallid orchids have been mainly described
within the limits of a strictly oristic approach, with little
or no consideration of either the relationships between
species within any given genus or the delineation of groups
of close relative taxa that could reect the natural history
of the largest genera. Molecular taxonomy mainly focuses
on distinguishing and circumscribing monophyletic genera,
rejecting articial or erroneous systematic frameworks, but
the difculty of obtaining enough samples and molecular
markers for analyses (Bogarín, Pérez-Escobar, et al., 2018)
has in part obscured the true relationships among the
species of the concerned groups. As a result, those genera
whose monophyly received strongest molecular support
were poorly investigated as to their internal relationships,
irrespective of their size.
The ve-year term sponsored by the University of
Costa Rica for our study of the Subtribe Pleurothallidinae
(the maximum period of funding for research projects by
UCR) was highly productive, but it is obvious that the effort
made by the team of Lankester Botanical Garden botanists
to complete the chapter on Pleurothallidinae for Flora
Costaricensis still has to offer novel results. To organize and
present these results, we split Pleurothallidinae into blocks
of genera, and the largest genera into smaller complexes
of related species. Here we introduce the results of our
systematic studies on Echinosepala.
materials and methods
The general methods adopted for this study follow that
presented in Pupulin, Karremans, et al. (2017). Plants of
Echinosepala were collected around the country, from type
localities and other critical areas, and brought to Lankester
Botanical Garden (LBG) for cultivation and subsequent
documentation. At least ve specimens per morphospecies
were collected at any given locality whenever possible.
Field notes were taken during eldwork, including GPS and
political data, elevation, ecological zones, and main types of
vegetation. Plants were cultivated at JBL, and phenological
data recorded.
Morphological Analysis
Over 180 individuals of Echinosepala, belonging to
most of the morphospecies known for the country, were
studied and documented. High-resolution macro- and
microphotographs were taken according to the methods
and with the camera and microscopic equipments detailed
in Pupulin, Karremans, et al. (2017). Drawings of owers
and oral dissection were prepared according to the
methods described therein. Vouchers were conserved in
the liquid collection of JBL and/or in the herbaria of the
National Museum of Costa Rica (CR) and the University of
Costa Rica (USJ). Measurements were mostly taken under
a dissecting stereoscope, or with the aid of the electronic
scale bars inserted in the high-denition images of the
oral details. Collections of the country, as well as relevant
herbaria that allow digital access to their collections, were
revised to study specimens of the taxa in this study, and
specimens annotated when required.
Floral Ecology
Notes on oral ecology were taken from direct
observation of the plants under the semiarticial conditions
of the LBG open greenhouses. Photographs were taken
of individual owers over a lapse of two to three days to
document temporal activity of the perianth.
DNA Extraction and Sequencing
The selection of the material was based on availability
and interspecic variability. Vouchers of the specimens are
kept in the liquid collections at JBL or L, unless specied
otherwise. The list of the vouchers used in the phylogenetic
analyses and their NCBI GenBank accession numbers
are provided in Pupulin, Karremans, et al. (2017: Tab.
1). Methodology for DNA extraction, amplication, and
sequencing of the material followed the steps given therein
under the same heading.
Building the Data Sets
The STADEN (Staden et al., 2003) package was used
for editing the sequences, and the Unicode nomenclature
(IUPAC) was adopted where more than one base pair
was equally probable. In the few cases when Pregap was
unable to build a contig, sequences were merged by lling
in missing positions with N’s. Sequences were aligned
manually in Mesquite v2.72 (Maddison and Maddison,
2007). After the alignments had been edited, additional
sequences were obtained from GenBank, the latter using
nBLAST. Arpophyllum giganteum AF266742 (ITS) was
used as outgroup in all cases, as it is suggested to be the
furthest related of all included species (Pridgeon et al.,
2001; Pupulin, Karremans, et al., 2017).
Phylogenetic Analysis
The Bayesian Evolutionary Analysis Sampling Trees
(BEAST; Drummond and Rambaut, 2007) was used to
analyze the ITS matrix. Substitution and clock models were
unlinked. GTR + Γ model, estimated frequencies, and eight
categories were used. The Lognormal relaxed clock model
was used. The tree prior was speciation-yule birth, and
the number of generations was set to 30,000,000. The rst
20% of the resulting trees were used as burin. Trees were
visualized in FigTree v.1.3.1 (Rambaut, 2009). Posterior
probability (PP) values were added to the branches of the
trees using the labeling option, and branches were reordered
for better visualization as discussed in Pupulin, Karremans,
et al. (2017).
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 157
systematics oF eChinosepala
Echinosepala Pridgeon & M.W. Chase, Lindleyana 17(2):
100–101. 2002.
Synonyms: Echinella Pridgeon & M.W. Chase, Lindleyana
16(4): 253. 2001, nom. illeg. homonym, non Ach.
(1810, Closteriaceae). Type species: Pleurothallis
aspasicensis Rchb.f. ≡ Echinosepala aspasicensis
(Rchb. f.) Pridgeon & M.W. Chase.
Myoxanthus subgen. Satyria Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 44: 5. 1992. Pleurothallis subgen.
Acianthera sect. Satyria (Luer) Luer, Lindleyana 11:
118. 1996. Type species: Pleurothallis pan Luer ≡
Echinosepala pan (Luer) Pridgeon & M.W. Chase.
Myoxanthus subgen. Silenia Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 44: 6. 1992. Pleurothallis subgen.
Acianthera sect. Silenia (Luer) Luer, Lindleyana 11:
118. 1996. Type species: Pleurothallis aspasicensis
Rchb. f. ≡ Echinosepala aspasicensis (Rchb. f.)
Pridgeon & M.W. Chase.
Epiphytic, lithophytic or terrestrial, caespitose to repent,
small to large herbs. Stems erect to curved, terete to slightly
complanate, homoblastic, composed of 2 or more nodes,
enclosed by ancipitous, loose, rarely pubescent sheaths,
commonly disintegrating with age. Leaf coriaceous,
narrowly ovate to linear-elliptic, subacute to acuminate,
cuneate below into a conduplicate, sessile base, with the
veins typically very apparent. Inorescence a fascicle
of single, successive owers from the base or the apex
of the stem, or both; the peduncle terete. Floral bract
loose, usually pubescent. Ovary minutely scaberulous to
tomentose or long-haired. Flowers bilabiate, eshy, with
temporal activity, usually with putrid, reddish-purple colors,
with sepals externally glabrous, scabrous or pubescent.
Dorsal sepal free, subtriangular to lanceolate-elliptic to
oblanceolate, obtuse to acuminate, abaxially glabrous to
verrucose. Lateral sepals connate into an elliptic to obovate
synsepal, subacute-rounded to broadly obtuse, the apices
sometimes free, glabrous to verruculose within. Petals
eshy, lanceolate to obovate, asymmetric, rounded to acute
or truncate. Lip 3-lobed, elliptic from a small claw, the
apical lobe rounded to subtruncate, thickened at apex into
a cushion-like pad, with a pair of intramarginal thin keels,
lateral lobes erect, uncinate; the disc with an erect callus
arising above the base, the lip sometimes with a horseshoe-
158 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
shaped pseudoglenion. Column semiterete, footed, with
narrow wings above the middle. Anther cap globose,
cucullate, with long, stiff hairs on the upper margin, 2-celled.
Pollinia 2, on a short, bilobed caudicle.
Before molecular systematics, Echinosepala species
had been difcult to adequately place into a phylogenetic
context, and only in the present century has its position in
the groups of basal Pleurothallidinae (sensu Karremans,
2016) been fully understood. Of the 14 species currently
recognized as belonging to the genus (Epidendra, 2019;
IPNI, 2020), only 2 were originally described under the
name Echinosepala (Pupulin, Karremans, et al., 2017);
all others have been classied as species of Acianthera,
Myoxanthus, and Pleurothallis.
Carlyle Luer (1992) rst recognized that most species
of the group were closely related and gathered them into
Myoxanthus subgen. Silenia Luer and Satyria Luer. The two
subgenera were mainly distinguished from other groups of
Myoxanthus by the non-scurfy sheaths (except Myoxanthus
pan [Luer] Luer ≡ Echinosepala pan [Luer] Pridgeon &
M.W. Chase) and the owers produced from low on the
ramicaul or terminally at the base of the leaf (instead of
from near the apex of the stem). The long verrucose-ciliate
upper margin of the anther cap is a common synapomorphy
of the two subgenera.
Because of the strong morphological afnities of
Myoxanthus subgen. Satyria and Silenia with Pleurothallis
subgen. Acianthera (Scheidw.) Luer—a relationship
also recovered by molecular analyses (see particularly
Karremans et al., 2016)—Luer moved most species to
Pleurothallis subgen. Acianthera, creating two new sections
(Luer, 1995; validated in Luer, 1996).
The analyses of DNA data (Pridgeon and Chase,
2001; Pridgeon et al., 2001) conrmed that the species of
Myoxanthus subgen. Silenia and subgen. Satyria formed
a monophyletic group deserving generic recognition, and
Pridgeon and Chase (2001) gave it formal status by creating
Echinella Pridgeon & M.W. Chase (later corrected to
Echinosepala Pridgeon & M.W. Chase [Pridgeon and Chase,
2002]). The phylogenetic analyses also conrmed a close
relationship between Echinosepala and Myoxanthus, as
noted by Luer. Together with Barbosella Schltr., Dresslerella
Luer, Pleurothallopsis Porto & Brade, Restrepia Kunth, and
Restrepiella Garay & Dunst., Echinosepala forms one of
the nine major clades within the Pleurothallidinae, sister to
the basal clade of Octomeria, and consecutively sister to the
Acianthera clade (Karremans, 2016).
Morphologically, species of Echinosepala are
characterized by caespitose to repent plants with
monophyllous stems covered by tubular-ancipitous,
minutely warty to (rarely) pubescent sheaths; the leaf
coriaceous; the inorescence solitary-owered, produced
from the apical or lower nodes of the stem; the ower eshy,
with the sepals frequently hirsute abaxially and variously
verrucose adaxially, the lateral sepals connate into a concave
to cymbiform synsepal; the lip 3-lobed, with the erect
lateral lobes oblong to uncinate, provided with a tall basal
callus; the anther apical incumbent, the anther cap papillose
to ciliate at apex; the 2 pollinia provided with whale-tail-
shaped caudiculae (Luer, 1992; Pridgeon, 2005a).
The point of insertion of the inorescence along the
stem has been used as the main character to distinguish the
two assemblages of species within the group. Luer (1992)
created Myoxanthus subgen. Silenia to separate those
species with a basal inorescence (erroneously reported
to be produced from the rhizome; e.g., Luer, 1992, 2003;
Dressler, 1993; Pridgeon, 2005a) from the rest of the group,
which supposedly have a terminal inorescence. Pupulin,
Karremans, et al. (2017) showed that the inorescences
of Echinosepala species are never rhizomic but are rather
produced along the stem, where it is exerted from the apex
of the second to third internode (see g. 1 in Pupulin,
Karremans, et al., 2017), and they expressed doubts that
owering from the nodes of the rhizome occurs at all in the
Pleurothallidinae. Producing inorescences from the lower
nodes of the secondary stem is apparently an obligate feature
in E. lappiformis (A.H. Heller & L.O. Williams) Pridgeon &
M.W. Chase, E. pan, and E. stonei (Luer) Pridgeon & M.W.
Chase, but it is facultative in E. expolita Pupulin & Belfort,
where both apical and basal inorescences are sometimes
produced simultaneously. This condition has evolved in
other groups of Pleurothallids, especially in Acianthera, and
according to our molecular analyses even in Echinosepala
it could have arisen independently in two different groups
of species.
The “basal” position of the inorescence in species of
Echinosepala convinced Luer (2004) to revive the genus
Brenesia Schltr. (1923), created to accommodate the
anomalous Brenesia costaricensis Schltr., a species with
pendent inorescences produced from the lower nodes of
the stem and hirsute owers. He transferred eight species
previously treated under Echinosepala (Luer, 2004) to
Brenesia, under the belief that they were related to each other.
However, the relationship between species of Brenesia sensu
stricto (i.e., B. costaricensis, B. herrerae [Luer] Luer, and
B. johnsonii [Ames] Luer, plus Acianthera sotoana Solano)
and species of Echinosepala is not supported by DNA
studies. The broad phylogenetic analysis of Acianthera
carried out by Karremans et al. (2016) retrieved that the clade
including the type species of Brenesia is sister to a clade that
includes Antilla and Kraenzlinella, which all together are
highly supported as sisters to the rest of Acianthera.
Florally, species of Echinosepala are mostly characterized
by the putrid, reddish-purple color of the bilabiate perianth,
often spreading only partially, and the complicate abaxial
indumentum of the sepals, which vary from almost
glabrous (E. isthmica, described below), to sparsely warty-
verruculose (E. alexandrae [Schltr.] Pupulin & Bogarín, E.
aspasicensis, E. glenioides [hereafter], E. uncinata [Fawc.]
Pridgeon & M.W. Chase), the warts sometimes grouped
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 159
into small tufts (E. longipedunculata Pupulin & Karremans
and E. vittata [Pupulin & M.A. Blanco] C.O. Morales &
N. Villalobos), to sparsely spiny (E. expolita Pupulin &
Belfort), densely tomentose (E. tomentosa [Luer] Pridgeon
& M.W. Chase), hirsute with soft trichomes (E. lappiformis
[A.H. Heller & L.O. Williams] Pridgeon & M.W. Chase,
E. stonei [Luer] Pridgeon & M.W. Chase), or echinate
with stiff hairs (E. pan). Adaxially, the sepals are almost
glabrous (E. alexandrae, E. aspasicensis, E. uncinata,
E. vittata), sparsely warty (E. tomentosa), to covered with
irregular, low (E. lappiformis) or tall (E. stonei) warts,
or stiff, short, rounded (E. longipedunculata) or pointed
(E. expolita, E. pan) warts. In some species, the base of
the dorsal sepal is semitransparent (e.g. E. aspasicensis,
E. expolita, E. glenioides), forming a window under the dark
purple blotches (Pupulin, Karremans, et al., 2017: g. 3).
The petals are mostly glabrous to sparsely verruculose, but
they may be distinctly warty toward the apex (E. expolita,
E. lappiformis, E. pan, E. stonei). The lip is glabrous in
most species, but it is warty at apex in E. alexandrae and E.
shuarii (Luer) Luer (= Pupulinia shuarii [Luer] Karremans
& Bogarín). It always presents a tall callus at the base,
which can be rounded and broad, massive, slightly concave,
smooth or transversely rugose, occupying all the lip base
(E. alexandrae, E. isthmica, E. lappiformis, E. stonei, E.
uncinata), or provided with a horseshoe-shaped, concave
pseudoglenion (E. aspasicensis, E. expolita, E. glenioides,
E. longipedunculata, E. pan, E. tomentosa, E. vittata). This
produces a gelatinous, non-viscous, shiny exudate, which
is spread through a longitudinal groove to the apex of the
lip, where it apparently dries, becoming matte in two to
three days. In most species, the midline of the lip presents
a pair of intramarginal low lamellae, typically undulate on
the upper margin.
According to our analyses, species of Echinosepala
form a highly supported clade, consecutively sister to
Myoxanthus, Dresslerella, and a clade grouping species
of Pleurothallopsis and Restrepia, and species of
Restrepiella and Barbosella, on two branches (Fig. 1).
Within Echinosepala, E. glenioides (as E. aspasicensis) and
E. longipedunculata are highly supported as sister species,
and in turn they are well supported as sister of E. pan and
E. expolita, which cannot be differentiated by their ITS
sequences. Echinosepala alexandrae (as E. sempergemmata)
and E. isthmica (as E. uncinata) form a well-supported
group, which in turn is sister (with low support) to E. stonei.
In our analysis, the phylogenetic position of E. stonei is
not resolved, but the species is surely not closely related
to the group of E. expolita/E.pan. Echinosepala tomentosa
and E. vittata, both characterized by miniature plant habits
and owers, are sister to all the other species in a highly
supported group.
The actual diversity of this group has still to be fully
understood. Pridgeon (2005a) reported only 8 species for
the genus, but we have recorded 10 species in Costa Rica
alone (Pupulin, Karremans, et al., 2017; herein) (Fig. 2–3).
Including the new taxa described and resurrected in the
present paper, at least 17 species are recognized today in the
genus, distributed from Belize to the West Indies (Jamaica),
to the Guyanas and Brazil, and down to Bolivia, including
Peru (Zelenko and Bermudez, 2009), along the Andes, but
the diversity of Echinosepala is likely much higher.
References: Pridgeon et al. (2001, 2005a,b), Karremans
(2016), Pupulin, Karremans, et al. (2017).
genus discussion For Flora CostariCensis
Echinosepala is a genus of 17 or more species,
distributed from Belize to the West Indies (Jamaica), the
Guyanas and Brazil, and down to Bolivia along the Andes.
According to the number of species recorded in Costa Rica
alone, the diversity of Echinosepala is likely higher. Species
of Echinosepala occur mostly epiphytically (rarely as
lithophytic or terrestrial herbs) in warm tropical, premontane
or cloud, evergreen, wet forests at elevations of 300–2000 m.
Flowering has been mostly recorded from June to December.
Morphologically, species of Echinosepala are characterized
by caespitose to repent plants with monophyllous stems
covered by tubular-ancipitous, minutely warty to (rarely)
pubescent sheaths and coriaceous leaves; the inorescence
solitary-owered, produced from the apical or lower nodes
of the stem; the bilabiate ower eshy, with putrid colors
and complicate indumenta of the sepals, frequently hirsute
abaxially and variously warty adaxially, the lateral sepals
connate into a synsepal; the lip 3-lobed, with the erect lateral
lobes oblong to uncinate, provided with a tall basal callus
occupying all the disc, which can be rounded and broad,
massive, smooth, or provided with a horseshoe-shaped,
concave pseudoglenion, producing a shiny, gelatinous
exudate, and a pair of intramarginal, low keels; the apical
anther incumbent, papillose to ciliate at apex; the 2 pollinia
provided with whale-tail-shaped caudiculae. Echinosepala
is likely to be pollinated by ies that hook the pollinarium to
the rear portion of the scutellum. No pollination studies have
been published to date, but ies of the Drosophilidae family
were recorded visiting the owers. Molecular analyses
carried out by Pridgeon and collaborators (2001) and by
our group (Pupulin, Karremans, et al., 2017) support the
monophyly of Echinosepala, whose species form a highly
supported clade, sister to Myoxanthus, Dresslerella, and a
clade grouping species of Pleurothallopsis and Restrepia,
and species of Restrepiella and Barbosella, on two branches.
Within Echinosepala, the species with a pseudoglenion are
highly supported as sister species, whereas those with a
plate-like, bare callus form another well-supported group,
which in turn is sister to E. stonei (Fig. 4). The species with
miniature plant habits and owers are sister to all the other
species in a highly supported group.
160 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Figure 1. Phylogenetic relationship among the species of Echinosepala (modied from Pupulin, Karremans, et al., 2017). Species names
in red are updated to the nomenclature proposed in the present paper.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 161
Figure 2. Flowers of Echinosepala species from Costa Rica. A–B, E. alexandrae (Bogarín 7137); C–D, E. expolita (Bogarín 1871;
Pupulin 7030); E–F, E. glenioides (Bogarín 3063; Bogarín 8211); G–H, E. isthmica (Bogarín 5855; Bogarín 5871); I, E. lappiformis
(Bogarín 9554). All the vouchers at JBL. Photographs by F. Pupulin.
162 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Figure 3. Flowers of Echinosepala species from Costa Rica. A–B, E. longipedunculata (Bogarín 5449; Karremans 4994); C, E. pan
(Bogarín 10270); D, E. stonei (Bogarín 5713); E, E. tomentosa (Bogarín 5622); F, E. vittata (Blanco 1324). All the vouchers at JBL.
Photographs by F. Pupulin.
Key to the sPecies oF eChinosepala in costa rica
1a. Callus of the lip with an inverted U-shaped callus (pseudoglenion) at the base ................................................. 2
1b. Callus of the lip with the basal callus smooth to rugulose ................................................................. 8
2a. Inorescences produced from the lower nodes of the stem ................................................................ 3
2b. Inorescences produced from the apex of the stem ...................................................................... 4
3a. Sheaths of the ramicauls hirsute; leaves narrowly linear-anceolate, >2.5 cm wide: sepals long-echinate abaxially .................E. pan
3b. Sheaths of the radical glabrous; leaves ligulate, <3 cm wide; sepals shortly tomentose abaxially ...........................E. expolita
4a. Mature plants <20 cm tall; owers small, synsepal >1 mm long, lip >4 mm long ............................................... 5
4b. Mature plants >30 cm tall; owers large, synsepal < 15 mm long, lip <7 mm long .............................................. 6
5a. The dorsal sepal yellow, the synsepal deep purple, adaxially densely tomentose ......................................E. tomentosa
5b. Sepals whitish, striped with red, abaxially with stiff hairs along the veins ..............................................E. vittata
6a. Ovary and abaxial surface of the sepals tomentose; adaxial surface spiny ..............................................E. expolita
6b. Ovary and abaxial surface of the owers warty; adaxial surface verruculose .................................................. 7
7a. Pedicel <3 cm long; owers purple-maroon; lip elliptic, acute, purple-red ...........................................E. glenioides
7b. Pedicel >6 cm long; owers yellow-orange; lip sub rectangular, truncate, yellow ...............................E. longipedunculata
8a. Inorescences produced from the lower nodes of the stem ................................................................ 9
8b. Inorescences produced from the apex of the stem ..................................................................... 10
9a. Dorsal sepal narrowly triangular-lanceolate ..................................................................E. lappiformis
9b. Dorsal sepal broadly oblong ..................................................................................E. stonei
10a. Leaves ligulate; owers autogamous, mostly cleistogamous, adaxially warty, midlobe of lip ovate, narrower than the
lip base ..............................................................................................E. alexandrae
10b. Leaves lanceolate; owers not autogamous, adaxially subglabrous, midlobe of lip transversely rectangular, as wide as the
lip base ................................................................................................ E. isthmica
Figure 4. Labella of Echinosepala species. A–J (left of the bar), species provided with a pseudoglenion; K–Q (right of the bar), species
without pseudoglenion. A–B, Echinosepala expolita; C, E. tomentosa; D, E. pan; E–G, E. glenioides; H–J, E. longipedunculata; K–M,
E. alexandrae; N–P, E. isthmica; Q, E. stonei. Vouchers (all at JBL): Pupulin 7030 (A); Bogarín 1871 (B); Bogarín 5822 (C); Bogarín
10274 (D); Bogarín 1945 (E); JBL-20660 (F); JBL-27660 (G); Bogarín 3863 (H); Bogarín 5449 (I); Karremans 4994 (J); Bogarín 4601
(K); Bogarín 4678 (L–M); Bogarín 5855 (N); Bogarín 5871 (O); Bogarín 5855 (P); Bogarín 7190 (Q). Photographs by F. Pupulin.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 163
taxonomic treatment
1. Echinosepala alexandrae (Schltr.) Pupulin & Bogarín,
comb. nov.
Basionym: Pleurothallis alexandrae Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19: 103. 1923. TYPE: COSTA
RICA. [San José: Cerro el] Tablazo, 1900 m, blühend
im Juli 1909, A. Brade & C. Brade 1159 (Holotype:
B, destroyed; Neotype: designated here, tracings of
Schlechter’s drawings of the habit and oral analysis,
mounted on the same sheet, AMES 00074034,
Fig. 5A–B). Fig. 6–7 (Vouchers: Bogarín 5755, JBL,
and Bogarín 4678, JBL).
Heterotypic synonyms: Pleurothallis sempergemmata Luer,
Selbyana 3(3–4): 386. 1977. TYPE: PANAMA.
Chiriquí: epiphytic in forested valley above
Guadalupe, alt. 2200 m, 17 Dec 1976, C. Luer 1360,
A. Luer, R. Dressler, N. Williams & F. L. Stevenson
(Holotype: SEL).
Myoxanthus sempergemmatus (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 15: 38. 1986.
Echinella sempergemmata (Luer) Pridgeon & M.W.
Chase, Lindleyana 16(4): 253. 2001, nom. illeg.
Echinosepala sempergemmata (Luer) Pridgeon &
M.W. Chase, Lindleyana 17(2): 101. 2002.
Brenesia sempergemmata (Luer) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 255. 2004.
Epiphytic, caespitose, erect herb up to 30 cm tall. Roots
coarse, exuous, 1.5–2.0 mm in diam. Ramicauls stout,
erect, terete, thicker in the distal portion, 5–13 cm long,
homoblastic, composed by 5–8 internodes different in
length, completely enclosed by 6–8 papyraceous, tubular,
ancipitous, apically loose, obliquely truncate, whitish,
glabrous, inated sheaths, increasing in size toward the apex,
1.8–5.2 × 0.8–1.5 cm, the oldest ones breaking longitudinally
and eventually disintegrating with age. Leaf erect, thickly
coriaceous, narrowly elliptic to lanceolate-elliptic, subacute
to acute, minutely emarginate at apex, 12–20 × 2.6–5.0 cm,
the midvein distinctly protruding abaxially. Inorescence a
fascicle of single, successive (or few simultaneous) owers
produced from a quickly degrading spathe at the apex of the
ramicaul; the peduncle terete, suberect to prostrate, glabrous
to minutely puberulous, 2.0–3.2 mm long. Floral bract
papyraceous, inated, lanceolate, acute, ca. 1.0–1.8 cm long.
164 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Pedicel terete, 0.8–1.0 cm long; ovary linear-subclavate, to
7 mm long, papillous-puberolous, the short warts apically
provided with a tuft of stiff, somewhat stellate bristles.
Flowers bilabiate, autogamous, mostly cleistogamous and
quickly developing into a fruit before reaching anthesis;
when opening, the owers self-pollinating during the rst
or second day of anthesis, if owering more than one day,
without temporal activity of the perianth; the sepals light
to dull green, the adaxial basal half blotched and spotted
purple-brown, abaxially striped with pale purple; the petals
yellow, blotched with purple-red, sparsely verrucose; the
lip purple-red. Sepals eshy, densely and shortly stellate-
puberolous on the abaxial side; dorsal sepal lanceolate-
oblong, concave, narrowly obtuse, 14–18 × 6–8 mm,
5-veined, the inner surface glabrous, abaxially densely
warty-pubescent, the warts apically provided with a stellate
tuft of short hairs; lateral sepals connate into an ovate,
obtuse to shortly bid, abaxially warty-pubescent synsepal,
the warts terminating with a stellate tuft of short hairs, 15–19
× 10–12 mm, each half 3-veined, the apices sometimes free
to about 1 mm. Petals eshy, elliptic-lanceolate to oblong-
lanceolate, obliquely asymmetric, narrowly obtuse to acute,
7–10 × 2.5–3.0 mm, 3-veined. Lip 3-lobed, ovate-elliptic,
pandurate, from a small, thin, rectangular, white-hyaline
claw, 7.2–8.0 × 2.8–3.1 mm (5.5 mm across the lateral
lobes), the base subtruncate; the apical lobe elliptic to ovate-
elliptic, obtuse, the margins more or less incurved, provided
at the base with a pair of intramarginal, thin, crenulate keels
running inside the lateral lobes toward the disc; lateral
lobes erect, narrowly triangular-uncinate, antrorse; the disc
with an erect, tall, broad, glabrous laminar callus. Column
subarcuate, semiterete, ca. 8 mm long, provided with
thickened, small, elliptic wings above the middle, the foot
ca. 1 mm long. Anther cap ovate, cucullate, with long, stiff
apical hairs, 2-celled. Pollinia 2, ovoid, attened, on a short
bid caudicle. Fig. 2A–B.
Eponymy: named in honor of Rudolf Schlechter’s wife,
Alexandra, née Sobenikoff.
Distribution: Costa Rica to central Panama (Cerro
Campana).
Ecology: in Costa Rica, Echinosepala alexandrae is an
uncommon epiphyte, seemingly restricted to the Caribbean,
Figure 5. Neotype of Pleurothallis alexandrae. A, habit; B, ower analysis. Tracings of Schlechter’s original illustrations based on the
original collection A. Brade & C. Brade 1159 (AMES 00074034). Reproduced with the kind permission of the Curator, Oakes Ames
Orchid Herbarium, Harvard University Herbaria.
Figure 6. Echinosepala alexandrae (Schltr.) Pupulin & Bogarín. A, habit; B, developing ovary and ower remnants; C, dissected perianth;
D, lip, ventral view (the apical lobe in natural position and spread); E, column and lip, lateral view; F, column of self-pollinated ower,
ventral view; G–H, details of the ovary indumentum. Drawn from Bogarin 5775 by F. Pupulin and D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 165
Figure 7. Echinosepala alexandrae (Schltr.) Pupulin & Bogarín. A, habit; B, ower; C, dissected perianth; D, lip in ventral and three-
quarters views; E, column and lip, lateral view; F, column in ventral view (emasculate on the left); G, anther cap; H, pollinarium. Drawn
from Bogarin 4678 by F. Pupulin and D. Solano Ulate.
166 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
submontane wet forests of the Tilarán, Central Volcanic, and
Talamanca mountain ranges, at elevations between 1400
and 2600 m. The species mostly ower during the months
of July–August, but sporadic owering has been recorded
from November to February.
Distinguishing features: the large plants with large,
coriaceous leaves, the owers buds quickly developing
into fruits before or immediately after anthesis, the stellate
appearance of the bristles covering the ovary and the abaxial
surface of the sepals, and the elliptic-pandurate lip with the
apical lobe narrow and with incurved margins distinguish
this species.
The specic epithet alexandrae, created by Schlechter to
honor his wife and conferred to a species of “Pleurothallis”
collected in Costa Rica by the Brade brothers, is the rst
available name for the species that Carl Luer described in
1977 as Pleurothallis sempergemmata on the basis of a
collection from Chiriquí, Panama (Luer, 1977). The type
of P. alexandrae was lost during the bombing of the Berlin
Herbarium in 1943, but fortunately Oakes Ames at Harvard
University had Schlechter’s drawings traced for his own
les before they were destroyed. The two sketches that
Luer (1995) selected as the species lectotype show the large
habit typical of the species, and the pandurate lip with a tall
basal callus—with no pseudoglenion—and a small apical
lobe, which are characteristic of Echinosepala alexandrae
and its synonym P. sempergemmata. Even though Luer
(1992) thought that P. alexandrae was a name for one of
Endrés’s detailed illustrations of Echinosepala (W0021295)
on the basis of material collected in Costa Rica (Endrés
s.n, Restrepia 156, W0021295, W18890142222), which
he treated as Myoxanthus aspasicensis, the two species are
only supercially similar. Endrés’s illustration in Vienna
(reproduced in Ossenbach et al., 2013: 173, g. 171) depicts
a species hereafter treated as E. glenioides, which has a
horseshoe-shaped structure on the top of the basal callus
(pseudoglenion), whereas E. alexandrae has a massive, tall
plate-like callus without a pseudoglenion, similar to that of
E. isthmica, to which it is phylogenetically related.
In his treatment of the Orchidaceae for the Flora
of Panama, Williams (1946) dubitatively included
Pleurothallis alexandrae in synonymy with P. uncinata, and
this induced Dressler (1981) to retain this name, based on
Costa Rican populations, for a Panamanian species distinct
from P. uncinata. However, what Williams considers as
possibly different from Echinosepala uncinata sensu lato—a
collection by Cope from el Valle de Antón (Coclé)—cannot be
E. alexandrae, because Williams describes the callus of this
specimen as “prominent retrorse […] and membranaceous
at the base of the side lobes” (Williams, 1946), that is,
provided with a pseudoglenion, a feature that is lacking in
the true E. alexandrae. Bogarín, Serracín, Samudio, Rincón,
et al. (2014) correctly recognize the afnity of this species,
treating it as E. aspasicensis (or E. glenioides, according
to the present treatment). Also, Luer (1976) interpreted as
P. alexandrae a species locally common in Panama, ranging
south to Venezuela and possibly Colombia. He correctly
stated that P. alexandrae is closely allied to P. uncinata, but
the specimen he illustrated (Luer, 1976: g. 125, likely from
Veraguas), does not match Schlechter’s concept however,
as it clearly presents a lip with a basal pseudoglenion,
and we interpret it as the species described hereafter as
E. glenioides. Luer (1976) also compared the supposed
P. alexandrae with his P. pastacensis, described from an
Ecuadorean collection (Luer, 1976: 156, g. 181), which in
fact also belongs to the group of species provided with a
pseudoglenion and has been treated as conspecic with E.
aspasicensis (e.g., Luer, 1986; Renner et al., 1990; Vásquez
and Ibisch, 2000; Luer, 1992, 1995; Jørgensen and León-
Yánez, 1999; Pupulin, 2002; Bogarín, Serracín, Samudio,
Rincón, et al., 2014; Jørgensen et al., 2014; Kolanowska,
2014). Even though it has been treated under several names,
for example, P. pastacensis (Luer 1976) and Myoxanthus
pastacensis (Luer) Luer (Luer, 1986), and as a synonym of
Brenesia aspasicensis (Jørgensen et al., 2014), Echinella
aspasicensis (Pupulin, 2002), Echinosepala aspasicensis
(Bogarín, Serracín, Samudio, Rincón, et al., 2014), M.
aspasicensis (Luer, 1992; Renner et al., 1990), and P.
aspasicensis (Luer, 1995; Jørgensen et al., 1999; Vásquez
and Ibisch, 2000), we consider it a distinct and good
species. As it lacks specic recognition as a member of
Echinosepala, we propose it here:
Echinosepala pastacensis (Luer) Pupulin, comb. nov.
Basionym: Pleurothallis pastacensis Luer, Selbyana 3(1/2):
156. 1976. TYPE: ECUADOR. Pastaza: epiphytic in
felled tree 20 km east of Puyo, 1000 m, 19 Mar 1976,
C. Luer 949, J. Luer & P. Taylor (Holotype: SEL).
To prevent possible claims of confusion between
Pleurothallis alexandrae Schltr. (1923) and P. alexandri
Schltr. (1922), we would like to emphasize that the names
are different, were created to honor different people, and
refer to different species. Both are therefore legitimate as
they are not likely to be confused under Article 53 of the
International Code of Nomenclature (Turland et al., 2018).
Costa Rican material examined: Cartago: Alvarado,
Cervantes, Parque Nacional Tapantí-Macizo Cerro de la
Muerte, Sendero Arboles Caidos, 9.9016667, -83.7908333,
20 Aug 1992, J. F. Morales 415, V. Nilsson & R. Chacón
Coto (CR). Macizo de la Muerte, southern Panamerican
Highway, km 47, Palo Verde, epiphytic on large trees,
remnants of primary and secondary vegetation, lower
montane wet forest, 2000 m, Nov 2011, F. Pupulin 4284, E.
Salas, H. Léon-Páez & A. C. Rodríguez (JBL). El Guarco,
San Isidro, Casa Mata, Carretera Panamericana Sur, km
47, Palo Verde, 2000 m, 16 Nov 2002, F. Pupulin 4284, E.
Salas, H. Montealegre & E. Salas (JBL). El Guarco, San
Isidro, Casa Mata, Carretera Interamericana, km 40, desvío
a San Cristóbal Norte, epítas en árboles a orillas de la
calle, bosque muy húmedo montano bajo, 9˚46'32.21"N
83˚56'34.40"W, 1869 m, 24 Apr 2008, D. Bogarín 4601, A.
Karremans, Y. Kisel & R. Phillips (JBL). El Guarco, San
Isidro, Madreselva, Tres de Junio, Carretera Interamericana
Sur, km 66, entrada a la nca El Jaular, 9˚40'11.7"N
83˚51'55.9"W, 2609 m, bosque pluvial montano, en bosque
secundario a la entrada de la nca, 24 Apr 2008, D. Bogarín
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 167
168 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
4678, A. Karremans, Y. Kisel & R. Phillips (JBL). El Guarco,
El Empalme, Carretera Interamericana, Cartago to Cerro
de La Muerte, km 66, epiphytic in roadside trees, 9˚42'N
83˚56'W, 2545 m, 23 Jul 2003, M. Whitten 2153, M.A.
Blanco & D. Bogarín (JBL). Jiménez, Pejibaye, Tucurrique,
Bajos del Humo, entre ríos Humo y Vueltas, ladera este
de Cerros Duán, bosque pluvial montano bajo, epítas
en árboles en potreros y borde de bosque, 9˚48'36.7"N
83˚45'16.2"W, 1396 m, 24 Nov 2008, D. Bogarín 5775, R.
L. Dressler, R. Gómez & R. Trejos (JBL). Turrialba, Santa
Cruz–Santa Teresita, Guayabo Arriba, Torito, 50 m después
del Caño Seco, en árboles solitarios del potrero de los Mesén,
10˚00'06.23"N 83˚41'47.85"W, 1470 m, 18 Dec. 2010,
A. P. Karremans 3499 & M. Contreras Fernandez (JBL).
Guanacaste: Monteverde Reserve, 1500 m, 13 Jul 1992, S.
Ingram 1515, K. Ferrell & N. Edmondson (SEL). Heredia:
Cerro Chompipe, 2300 m, 27 Aug 1997, R. L. Dressler s.n.
& D. E. Mora (JBL). San Rafael, Reserva Biológica “El
Chompipe,” 2000–2050 m, 9 Aug 1991, S. Ingram 843 & K.
Ferrell (SEL). San José: Cerro El Tablazo, 1900 m, . Jul
1909, A. Brade & C. Brade 1159 (drawings, AMES).
2. Echinosepala expolita Pupulin & Belfort, Lankesteriana
17(2): 294. 2017. TYPE: Costa Rica. Alajuela: San Ramón,
Piedades, Piedades Norte, road to Bajo La Paz, ca. km 3,
along the Río San Pedro, 10˚08'58.7"N 84˚34'03.3"W,
1300 m, premontane wet forest, secondary and remnants
of primary vegetation, 19 February 2008 F. Pupulin 7030,
R. L. Dressler & A. P. Karremans (Holotype: JBL; Isotype:
JBL). Fig. 8 (Voucher: Pupulin 7030, JBL).
Epiphytic, caespitose, erect herb up to 35 cm tall. Roots
coarse, exuous, 1.5–2.0 mm in diam. Ramicauls stout,
erect, terete, thicker in the distal portion, 6.0–12.5 cm long,
homoblastic, composed of 5 internodes different in length,
completely enclosed by 3–5 papyraceous, tubular, ancipitous,
apically loose, obliquely truncate, whitish, glabrous, inated
sheaths, increasing in size toward the upper one, 1.4–7.7 ×
0.20–0.85 cm, the oldest ones breaking longitudinally and
eventually disintegrating with age. Leaf erect, coriaceous,
linear-elliptic, subacute, minutely emarginate at apex, 16.2–
27.0 × 1.8–2.4 cm, with the midvein strongly protruding
abaxially. Inorescence a fascicle of single, successive
owers produced facultatively at the apex and the base of
the ramicaul at once; the peduncle terete-subclavate, erect,
glabrous, 1.8–2.5 cm long. Floral bract papyraceous, loose,
obliquely truncate-subobtuse, 1.8–2.4 cm long. Pedicel
terete-subclavate, 1.8–2.5 mm long; ovary lanate-hirsute,
completely covered by the bract, linear-subclavate, 3 mm
long. Flowers bilabiate, with temporal activity, fully opening
during the morning and closing in late afternoon; the sepals
light greenish-yellow, with the adaxial basal half mottled
purple-red and the distal half with pointed warts becoming
denser toward the apex, abaxially ushed purple and
densely warty-pubescent; the petals yellow, blotched with
purple-red, sparsely verrucose; the lip dark purple, apically
covered with a translucent wax in fresh owers. Dorsal
sepal elliptic-oblong, 0.4–0.9 × 1.9–2.0 cm, 7- to 9-veined,
showing a semitransparent base, forming a window under
the dark purple-red blotches, tomentose abaxially, adaxially
provided with spiny warts in the apical half. Lateral sepals
connate into an elliptic, abaxially tomentose synsepal, 0.6–
1.7 × 0.4–1.0 cm, each half 5-veined, the apices free about
6 mm, subacute-rounded, with pointed-spiny warts in the
distal half. Petals eshy, rhombic, obliquely asymmetric,
acute, 6–7 × 1–3 mm, 3-veined, apically provided with
spiny warts. Lip 3-lobed, narrowly elliptic from a small,
rounded, thin, dark purple claw, 7.5 × 2.2 mm (5.0 mm
across the lateral lobes), the base subtruncate; the apical
lobe rounded-truncate, thickened at apex into a cushion-
like, elliptic, low pad, provided with a pair of intramarginal,
thin keels running inside the lateral lobes toward the disc,
more prominent to the base of the disk; lateral lobes erect,
narrowly lanceolate, antrorse; the disc with an erect, narrow
callus arising above the base the lip, covered on top by a
horseshoe-shaped, channeled pseudoglenion extending in
front into a low keel ushing into the apical pad. Column
straight to subarcuate, semiterete, 4.4–5.5 mm long,
provided with narrow, uncinate wings above the middle, the
foot ca. 2.3 mm long. Anther cap globose, cucullate, with
long, stiff hairs on the upper margin, 2-celled. Pollinia 2,
obovoid, attened, on a short bid caudicle. Fig. 2C–D.
Etymology: from the Latin expolitus, shining, glossy, in
allusion to the the glossy apex of the lip in the fresh owers.
Distribution: known only from the Caribbean watershed
of the Cordillera de Tilarán in central Costa Rica.
Ecology: epiphytic in tropical, transition to premontane,
and premontane wet forests along the Caribbean watershed
of the Cordillera de Tilarán, at 800–1300 m elevation.
Flowering has been recorded at least from March to June,
and in November, but probably plants may ower at any time
of the year. Flowers show temporal activity, fully opening
during the morning and closing in late afternoon; usually
they are almost to completely closed in the evening. During
anthesis they emit a subtle but pungent smell, reminiscent
of urea. The breakdown of urea to ammonia and carbon
dioxide is known to lure insects, particularly certain ies.
Distinguishing features: the species may be distinguished
by the large plants with narrowly lanceolate leaves longer
than the subtending ramicauls, the inorescences produced
both at the apex and from one of the lower nodes of the
stem, the lanate-hirsute ovary, and the large ower with the
abaxial side of the sepals tomentose.
Florally, Echinosepala expolita is similar to E. pan, to
which it appears related in the phylogenetic reconstruction,
and from which it can be distinguished by the glabrous,
inated sheaths that cover the stem (vs. hirsute, tight), the
distinctly broader ligulate-lanceolate leaves (vs. narrowly
lanceolate), the lanate-hirsute ovary (vs. echinate), the much
larger ower (sepals >15 mm vs. <10 mm long), and the
tomentose abaxial indumentum of the sepals (vs. hirsute-
echinate). At anthesis and in fresh owers, the adaxial
surface of the lip is covered with a translucent wax exudate,
apparently produced within the basal pseudoglenion, which
becomes progressively matte until the lip appears dry during
the last days of anthesis.
The plant architecture of Echinosepala expolita is similar
to E. aspasicensis, but the basal inorescences distinguish
the two species, also in sterile material. The inorescences
Figure 8. Echinosepala expolita Pupulin & Belfort. A, habit; B, ower; C, dissected perianth; D, lip in ventral and three-quarters views;
E, column and lip, lateral view; F, column in ventral view; G–H, pollinarium (two views); I, anther cap. Drawn from Pupulin 7030 by F.
Pupulin and D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 169
170 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
produced from a lower node of the stem may have originated
twice independently (in the E. expolita + E. pan clade and
the clade of E. stonei). Alternatively, it may have evolved
only once, with one or two reversals to the ancestral state
(apical inorescence). Either scenario indicates that species
with basal inorescence do not form a monophyletic group
in Echinosepala.
Costa Rican material examined: Alajuela: San
Ramón, Piedades, Piedades Norte, road to Bajo La Paz, ca.
km 3, along the Río San Pedro, 10˚08'58.7"N 84˚34'03.3"W,
1300 m, premontane wet forest, secondary and remnants
of primary vegetation, 19 February 2008 F. Pupulin 7030,
R. L. Dressler & A. P. Karremans (JBL). San Ramón,
Ángeles, Reserva Biológica Alberto M. Brenes, 10˚13'06"N
84˚36'11"W, 850 m, bosque muy húmedo tropical transición
a premontano, sobre el Sendero La Catarata, epítas en
borde de bosque secundario a orillas del Río San Lorencito,
24 setiembre 2005, D. Bogarín 1871 (JBL). San Ramón,
Piedades S[ur] de San Ramón, Cerros de Laguna, 1190 m,
7 Nov 1922, A. M. Brenes 26219 (425 Herb. Brenes) (CR).
San Ramón, Cordillera de Tilarán, cerca a Balsa, ca. 12 km
NE de San Ramón, 10˚10'30"N 80˚30'30"W, 1100 m, 18
June 1995. Epíta a 2 m. Flores grises con manchas moradas
y rayos blanco-hueso, suculentas, B. Hammel 19870 (CR).
3. Echinosepala glenioides Pupulin, sp. nov.
TYPE: Costa Rica. Cartago: Paraíso, Orosi, Tapantí, Parque
Nacional Tapantí, camino entre el portón del Mirador hacia
el Río Humo, Proyecto Hidroeléctrico Tapantí, 9˚41'32.9"N
83˚47'03.2"W, 1650 m, bosque pluvial premontano “supra
arbores et ad truncos prostratos vetustos ad sylvarum
versuras ad viam umen Humo in Tapantí,” 18 Nov 2010,
D. Bogarín 8211, R. Gómez, A. Karremans, B. Klein, G.
Meza & F. Pupulin (Holotype: JBL). Fig. 9 (Voucher:
Bogarín 8211, JBL).
Species Echinosepala aspasicensis (Rchb. f.) Pridgeon &
M.W. Chase similis sed ovario glabro versus pubescentem,
sepalo postico anguste obovato versus oblongo-lanceum,
petalis obtrullatis versus falcata, labello obtrullato
rotundato versus ligulatum attenuatum retusum, columna
dimidio longa quam labello versus aequilongam recedit.
Epiphytic, caespitose, erect herb up to 50 cm tall.
Roots coarse, exuous, 1.3–2.0 mm in diam. Ramicauls
stout, erect, rarely subpendent, terete, thicker in the distal
portion, 10.0–22.5 cm long, homoblastic, composed of
4–6 internodes different in length, completely enclosed
by 4–6 papyraceous, tubular, ancipitous, loose, obliquely
truncate, whitish, glabrous, inated sheaths, increasing in
size toward the upper one, 1.5–9.0 × 0.3–1.0 cm, the oldest
ones breaking longitudinally and eventually disintegrating
with age. Leaf erect, thickly coriaceous, ligulate to oblong-
elliptic, subacute, minutely emarginate at apex, 16–29 ×
1.9–3.3 cm, with the midvein strongly protruding abaxially
and the lateral veins also faintly visible on both sides.
Inorescence a fascicle of single, successive owers
produced at the apex of the ramicaul; the peduncle terete-
subclavate, erect, microscopically pubescent, 1.0–2.2 mm
long. Floral bract papyraceous, loose, obliquely obtuse,
longer than the pedicel, 2.2–3.2 cm long. Pedicel terete-
subclavate, ca. 5 mm long; ovary terete, linear-subclavate,
glabrous, ca. 5 mm long. Flowers bilabiate, mostly ringent,
with temporal activity, fully opening during the morning
and closing in late afternoon; the sepals light greenish-
yellow, the dorsal semitransparent-fenestrate at the base,
mottled purple-maroon particularly on the basal half, the
dots and blotches more or less aligned with the veins,
abaxially ushed purple and densely warty-pubescent; the
petals dull yellow, heavily blotched or boldly striped with
purple-maroon; the lip purple-red, apically covered with
a translucent wax in fresh owers; the column yellow,
ecked with purple. Dorsal sepal oblanceolate to narrowly
obovate, subobtuse, reclinate, rarely erect, 2.1–2.6 × 0.7–
0.9 cm, 5-veined, with a semitransparent base, forming a
window under the dark purple-red blotches, shortly and
sparsely pubescent abaxially, adaxially provided with low
warts in the apical half. Lateral sepals connate into an
elliptic, obtuse, shortly bid, abaxially shortly and sparsely
pubescent synsepal, 1.80–2.20 × 0.9–1.1 cm, each half
4-veined, the apices free about 1 mm, with low rounded
warts in the distal half. Petals eshy, obtrullate, acute, 6–7
× 1.6–2.0 mm, 3-veined. Lip 3-lobed, elliptic-subrhombic
from a small, rectangular, thin, hyaline claw, 7.0–8.0 × 2.3–
2.6 mm (ca. 5 mm across the lateral lobes), the base truncate
with two short, digitate-conical, retrorse auricles; the apical
lobe ovate, acute, minutely rounded, the apex with sparse
warts, provided with a pair of intramarginal, thin, rounded
keels running inside the lateral lobes toward the disc; lateral
lobes erect, narrowly uncinate-acuminate, antrorse; the disc
with an erect, narrow callus arising above the base the lip,
covered on the top by a horseshoe-shaped pseudoglenion,
channeled in front up to the apical lobe. Column subarcuate,
terete, 5 mm long, provided with narrow, elliptic, stigmatic
wings above the middle, the foot ca. 1 mm long. Anther cap
globose, subspherical, cucullate, with long, stiff hairs on the
upper margin, 2-celled. Pollinia 2, pyriform, on a broad,
obreniform, bid caudicle. Fig. 2E–F.
Etymology: from the words glene, the Greek for “socket
or eyeball,” and the sufx -oides, from the Greek εἶδος
(eîdos, “likeness”), to suggest the imperfect resemblance of
the ornamentation at the base of the callus with the glenion
of other groups of Pleurothallidinae. The pseudoglenion
is apparently an ancestral character in Echinosepala,
as it has been recorded in both the basal E. vittata + E.
tomentosa clade and in the clade including E. glenioides +
E. longipedunculata and E. expolita + E. pan, whereas it is
absent in the derived clades of E. lappiformis + E. stonei
and E. alexandrae + E. uncinata.
Distribution: known at least from central Costa Rica to
the Chiriquí province in western Panama, perhaps ranging
to central Panama.
Ecology: an epiphyte of the large branches and trunks
of trees in pristine and mature vegetation, Echinosepala
glenioides is apparently restricted to the premontane and
lower montane wet forests along the Caribbean watershed
of the Costa Rican cordilleras, ranging from the Tilarán
mountain chain southward to the cordillera of Talamanca,
at elevations between 850 and 2000 m. Flowering mostly
occurs from July to September, but sporadic owering has
been recorded as early as May.
Distinguishing features: the large plant (the largest of
the genus in Costa Rica), with almost linear-ligulate leaves,
Figure 9. Echinosepala glenioides Pupulin. A, habit; B, ower; C, dissected perianth; D, lip in ventral and three-quarters views;
E, column and lip, lateral view; F, column in ventral view; G, pollinarium; H, anther cap. Drawn from Bogarin 8211 by F. Pupulin and
D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 171
172 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
the glabrous ovary, the large owers (with sepals over 2.5
cm long) mostly ringent, the dorsal sepal fenestrate at the
base, the long, elliptic-subrhombic lip provided with narrow
uncinate lateral lobes, and a basal pseudoglenion distinguish
this species among other relatives in Costa Rica.
Populations of Echinosepala glenioides have been
traditionally treated as the northernmost records of E.
aspasicensis, originally described from the Colombian
Andes (Dressler, 1981, 1993; Luer, 1992, 1995, 2003;
Hamer, 2001; Pupulin, 2002; Ossenbach et al., 2010;
Bogarín, Serracín, Samudio, Rincón, et al., 2014; Pupulin
et al., 2019), a poorly known species allegedly distributed
from Costa Rica to Guyana, Brazil, and down to Bolivia
along the Andean chain (Luer, 1992). The plant material
on which the description of Pleurothallis aspasicensis was
based had been sent to Reichenbach by Hermann Wagener
from Caracas, where he was collecting orchids for the
rm of Jean Linden (Reichenbach, 1855). The holotype
specimen, supposedly among Reichenbach’s materials in
Vienna, has not been located. Nevertheless, during one of
his visits to Lindley, Reichenbach gave him a piece of the
original collection, today housed at Kew, which Luer (1992)
selected as lectotype. No vegetative parts are included, and
the crushed ower lacks the lip, so our understanding of
Reichenbach’s concept of P. aspasicensis is largely based
on the original protologue. Several of the features described
by Reichenbach prevent the application of this name to
Costa Rica populations here described as E. glenioides. The
leaves were described as lanceolate, versus oblong-ligulate
in Costa Rican material; the sepals as “oblongo-lanceis,”
versus the dorsal sepal distinctly oblanceolate to obovate;
the lip apically attenuate and retuse, versus acute in E.
glenioides; and the column as long as the lip (“gynostemio
labello aequilongo”), versus barely half the length of the lip
in Costa Rican material.
Florally, Echinosepala glenioides is also similar to
Pleurothallis pastacensis Luer (which has been generally
treated as a synonym of E. aspasicensis), but it differs
from the Ecuadorean taxon by the caespitose habit with
narrow leaves (vs. distinctly repent with wider leaves),
the 1-owered inorescence (vs. simultaneously many-
owered), the distinctly larger owers, the sparse hairs
of the sepals’ indumentum (vs. arranged in tufts), and the
glaborus midlobe of the lip (vs. verucose).
Broadly distributed in Costa Rica, Echinosepala
glenioides is orally variable among populations (Fig. 2E–
F). One of the vouchers used for the molecular analyses does
not group with the other two accessions of E. glenioides
(Fig. 1), leaving an open question about the existence in
Costa Rica on another, cryptic taxon closely related to E.
glenioides. We were unable, however, to characterize this
genetically distinct individual on the basis of distinguishing
morphological features, so given the present status of our
knowledge we favored including it within the variation of
E. glenioides.
Costa Rican material examined: Alajuela: San
Carlos, Fortuna, volcán Chato, sobre el sendero a la laguna
de la cima de volcán, en bosque primario, muy húmedo
premontano, 10.4427778 -84.6808333, 27 Jul 2011, A.
P. Karremans 4407 (CR, JBL). San Ramón, Candelaria,
March–June 1867, A. Endrés s.n. [Restrepia 156] (W). San
Ramón, Candelaria, 1867, A. Endrés s.n. [Restrepia 156]
(W). San Ramón, Piedades, Potrerillos, road to Socorro
de Piedades Sur, slopes of Cerro La Palma, premontane
wet forest, 10˚08'09.9"N 84˚34'47.5"W, 1300–1450 m, 12
Apr 2006, F. Pupulin 6045, R. L. Dressler & A. Carbajal
(JBL). San Ramón, Ángeles, Reserva Biológica Alberto
Brenes, ascenso por el sendero Saíno, bosque muy húmedo
tropical transición a premontano, epíta en bosque
secundario, 10˚13'08"N 84˚35'48"W, 900–1000 m, 25
Sept 2005, D. Bogarín 1945 (JBL). San Ramón, Ángeles,
Reserva Biológica Alberto Brenes, sobre el sendero
Pájaro Sombrilla, 10˚13'N 84˚37'W, 850 m, 3 Oct 2003,
D. Bogarín 436 (JBL). San Ramón, Ángeles, Reserva Río
San Lorenzo de UCR, headwaters of the Río San Lorenzo
below Fila Volcán Muerte. 10- 23 N. 14 Jul 1983, K. A.
Barringer 3783 & B. Pérez (CR). San Ramón, Ángeles,
Balsa, road between San Ramón and La Fortuna de San
Carlos, epiphytic on trees in pastures close to Río Balsa,
premontane rain forest, 10˚10'03.6"N 84˚29'35.7"W, 1150
m, 29 may 2013, F. Pupulin 1150, D. Bogarín, M. Díaz,
& M. Fernández (JBL). San Ramón, Ángeles, Ángeles
Sur, road 141 to La Fortuna, deviation point on the left to
Ángeles Sur, premontane rain forest, epiphytic on scattered
trees in pastures along the border of a secondary, mature
forest, 10˚08'32.78"N 84˚29'28.30"W, 1140 m, 22 Apr 2011,
F. Pupulin 8033 & M. Pupulin (JBL). San Ramón, Santiago,
nca of Jesús Salas Jiménez, mountains toward the towers
of Berlín, lower montane rain forest, epiphytic in old trees
along coffee plantations, 10˚02'21"N 84˚12'02"W, 1300
m, 30 May 2013, F. Pupulin 8497, D. Bogarín, M. Díaz,
& M. Fernández (JBL). Cartago: Orosi, Purisil, Parque
Purisil, 1400–1500 m, 15 Feb 2005, H. León-Páez 92 (JBL).
Heredia: Heredia–San Rafael, Vara Blanca, 2 km noreste de
Alto del Roble, faldas al norte del Cerro Chompipe, Refugio
de Vida Silvestre Cerro Dantas, márgenes de la Quebrada
Grande, epítas en bosque secundario, bosque pluvial
premontano, 10˚05'30,5"N 84˚03'51,4"W, 1981 m, 14 Apr.
2005, D. Bogarín 1521, M. G. Gei & A. Vaughan (JBL).
Limón: Matina, Batán, Parque Nacional Braulio Carrillo,
Carrillo Station, along trail SE of Station, W of Quebrada
González, 10.1625 -83.25, 19 Sept 1990, S. Ingram 530 &
K. Ferrell (CR). San José: Vásquez de Coronado, Jesús,
Bajo La Hondura, Parque Nacional Braulio Carrillo, camino
de la ermita hacia Río Blanco, bosque pluvial premontano,
epítas en árboles a orillas del camino, 10˚03'03"N
83˚59'14"W, 900–1100 m, 21 Jan 2007, D. Bogarín 3063,
R. L. Dressler, F. Pupulin & W. Rossi (JBL).
4. Echinosepala isthmica Pupulin, sp. nov.
TYPE: COSTA RICA. Limón: Siquirres, Pacuarito, 5 km
después de la Estación del Parque Nacional Barbilla,
bosque pluvial premontano, epítas en árbol caído a orillas
del camino, 10˚1'1.5"N 83˚28'29.3"W, 627 m, 3 Dec 2008,
owered in cultivation at Lankester Botanical Garden,
29 Sept. 2102, D. Bogarín 8571, R. L. Dressler, R. Gómez
& R. Trejos (Holotype: JBL). Fig. 10 (Voucher: Bogarín
8571, JBL).
Figure 10. Echinosepala isthmica Pupulin. A, habit; B, ower; C, dissected perianth; D, lip, ventral view; E, column and lip, lateral view;
F, column in ventral view; G, anther cap; I, pollinarium, two views. Drawn from Bogarin 8571 by F. Pupulin and D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 173
174 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
A Echinosepala uncinata (Fawc.) Pridgeon et M.W.
Chase similis, sed caulibus dimidio brevioribus et foliis
parvioribus, labello multo breviore et latiore, lobo apicali
rectangulari vel late obovato integro vs. subrotundatum
minute denticulatum, callo basali insignis massivo
altoque vs. sublaevem, columna longiore distincte curvata
plerumque recedit.
Epiphytic, caespitose to shortly repent, erect herb up to
25 cm tall. Roots coarse, exuous, 1.5–2.0 mm in diam.
Ramicauls stout, erect, terete, 6.0–13.5 cm long, homoblastic,
composed of 5–7 internodes different in length, rarely
producing secondary shoots from the apex, completely
enclosed by 5–8 papyraceous, tubular, ancipitous, apically
loose, obliquely truncate, whitish, glabrous, inated
sheaths, increasing in size toward the upper one, 1.5–4.6 ×
0.3–0.9 cm, eventually disintegrating with age. Leaf erect,
coriaceous, narrowly ovate to lanceolate-elliptic, acute,
minutely emarginate at apex, 13.2–16.0 × 2.2–4.2 cm, the
midvein distinctly protruding abaxially. Inorescence a
fascicle of single, successive owers produced at the apex
of the ramicaul from a narrowly rectangular, papery, loose,
obliquely truncate spathe to about 1.5 cm long; the peduncle
terete, prostrate, furfuraceous to minutely and sparsely
pubescent, 1.8–2.5 cm long. Floral bract papyraceous,
loose, triangular, subacute, 1 cm long. Pedicel terete-
subclavate, 1.8–2.3 mm long; ovary sparsely pubescent,
completely covered by the bract, linear-subclavate, ca. 5
mm long. Flowers bilabiate, not completely spreading,
remaining fully opening after anthesis with no temporal
activity observed; the sepals greenish-cream, striped to
almost solidly mottled purple, subglabrous to sparsely
pubescent abaxially; the petals yellow, blotched with
purple-red; the lip yellow, densely spotted brown apically,
rose-purple, blotched purple at the base, the lateral lobes
yellow; the column white, sparsely dotted purple at the apex,
ushed yellow basally. Dorsal sepal elliptic-lanceolate,
acute, 1.3–1.9 × 0.6–0.9 cm, 5-veined, translucent. Lateral
sepals connate into a narrowly ovate, obtuse to subrounded,
minutely excised synsepal, 0.4–1.7 × 1.6–1.7 cm, each
half 3-veined, glabrous within. Petals eshy, oblanceolate,
obliquely asymmetric, acute, 7–12 × 2–3 mm, 3-veined. Lip
3-lobed, subrectagular-pandurate from a small, transversely
rectangular, thin, hyaline claw, 6.2–7.3 × 2.5–3.0 mm (5.0
mm across the lateral lobes), the base subtruncate, provide
with 2 small, acicular, retrose, pointed auricles; the apical
lobe transversely rectangular to broadly obovate, truncate,
the margins sometimes subundulate, provided with a pair
of intramarginal, thick keels running inside the lateral lobes
toward the disc; lateral lobes erect, narrowly uncinate,
antrorse; the disc with an erect, broad, massive plate-
like callus arising above the base the lip, ushing in front
to the base of the midlobe. Column arcuate, semiterete,
6.3–7.0 mm long, provided with inconspicuous, elliptic
wings above the middle, the foot ca. 1.5 mm long. Anther
cap globose, cucullate, with short, stiff hairs on the upper
margin, 2-celled. Pollinia 2, ovoid, laterally attened, on a
short bid caudicle. Fig. 2G–H.
Etymology: named in reference to the Central American
isthmus, where the species ranges at least from Nicaragua
southward to Panama.
Distribution: Nicaragua, Costa Rica, Panama, and
Colombia.
Ecology: in Costa Rica, Echinosepala isthmica has the
widest ecologic tolerance, spanning from the tropical forests
close to sea level (50 m) to the premontane wet forest at
about 1500 m of elevation. The species has been recorded
from both the Caribbean watersheds of the continental
divide. It mostly owers from July to September, with
sporadic, early owerings recorded in June.
Distinguishing features: the distinctly lanceolate
leaves on a medium-sized plant, the comparatively large,
subglabrous to minutely and sparsely tomentose owers,
with the purple blotches of the dorsal sepal arranged in
stripes and the synsepal almost solidly purple-maroon, and
the rectangular-pandurate lip with the apical lobe broader
than long and a massive callus but no pseudoglenion
distinguish this species.
Echinosepala isthmica has been treated as conspecic
with E. uncinata (Williams, 1946; Dressler, 1981, 1993;
Luer, 1992, 1995, 2003; Hamer, 1984, 2001; Pupulin,
2002; Ossenbach et al., 2010; Bogarín, Serracín, Samudio,
Rincón, et al., 2014), originally described from Jamaica by
William Fawcett (1895). Plants of E. uncinata are however
much larger, approaching in size and appearance those of E.
alexandrae, with ramicauls reaching almost 25 cm in length
(vs. up to 13.5 cm in E. isthmica). The owers are also
different. The lip of E. uncinata is comparatively longer,
with minute denticulations on the margin (vs. rectangular
to broadly obovate, entire, in E. isthmica), the basal callus
is low and narrow (vs. massive, as wide as the hypochile
and high), and the column is stout, straight (vs. slender,
distinctly curved), more similar to that of E. alexandrae. The
illustration of Pleurothallis uncinata in Flora of Jamaica
(Fawcett and Rendle, 1910: pl. 10) shows a large plant with
several fruits and, if not for the midlobe of the lip as wide as
the hypochile, we would have been tempted to consider the
native species of Jamaica as the rst available name for the
Costa Rican and Panamanian populations described later as
P. alexandrae and P. sempergemmata.
Echinosepala isthmica may occasionally produce new
vegetative shoots, made up of stem and leaf, and ultimately
roots, at the apex of the pseudobulb (e.g., Bogarín 5855;
Fig. 11). These adventitious plantlets could eventually
detach from the main stem and, on nding the appropriate
conditions, form new individuals independent of the
mother plant. This form of vegetative reproduction is rare
in Echinosepala, and up to now it has been observed only
in E. isthmica, but it is not uncommon in other groups of
Pleurothallidinae.
We are not including Belize at this time in the distribution
of E. isthmica, as the species recorded there by McLeish
and colleagues (1995) could well be conspecic with the
true E. uncinata from the Antilles. Florally, the collection
by Adams (no. 253, K) from the Mountain Pine Ridge in
Cayo District, Belize, is very close to E. isthmica, but the
plant is much larger, with ligulate-oblong leaves, quite
distinct from populations from Nicaragua southward. The
close relationships of the coastal ora of Belize with that
of the West Indies are well known, even in the subtribe
Figure 11. Adventitious plantlets on the apex of the stem in Echinosepala isthmica (Bogarín 5855). Scale bar = 5 cm. Photograph by
F. Pupulin.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 175
176 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Pleurothallidinae, and the species has not been recorded so
far south of Belize. Populations of E. uncinata previously
recorded from Nicaragua (Hamer, 1984, 2001; van den
Berghe and van den Berghe, 2008) and Caribbean Costa
Rica (Atwood, 1987) surely belongs to E. isthmica.
Costa Rican material examined: Alajuela: San
Ramón: Camino a San Carlos, between Potrero Volio
and Legua, July, A. Endrés s.n. [Restrepia 262] (W).
Upala, Bijagua. Parque Nacional Volcán Tenorio, sobre
los senderos principales del parque, bosque primario y
secundario, 10˚42'58.3"N 84˚59'33.1"W, 1121 m, 28 Jul
2011, A. P. Karremans 4446 (JBL). Upala, Bijagua. Parque
Nacional Volcán Tenorio, sobre los senderos principales
del parque. Epítas en bosque primario, bosque húmedo
premontano, 10˚42'54"N 84˚59'14"W, 686 m, 19 May 2016,
A. P. Karremans 7112, N. Davin & J. E. Jiménez (JBL).
Cartago: Jiménez, Pejibaye, entre Taus y Tausito, Selva,
orillas del Río Taus y Quebrada Selva, bosque pluvial
premontano, epítas en bosque secundario a orillas del
camino, 9˚47'10.3"N 83˚45'37.5"W, 1095 m, 24 Nov 2008,
D. Bogarín 5853, R. L. Dressler, R. Gómez & R. Trejos
(JBL). Paraíso, Orosi, Tapantí, sobre el camino a Tausito,
unos 1.5 km del cruce al Parque Nacional Tapantí, epítas y
semi-terrestres sobre el acantilado al lado de la calle, bosque
pluvial premontano, 9˚46'18.48"N 83˚47'34.36"W, 1412
m, A. P. Karremans 6678 & I. Chinchilla (JBL). Turrialba
Santa Cruz–Santa Teresita, Guayabo Arriba, Torito, 50 m
después del Caño Seco, en árboles solitarios del potrero de
los Mesén, 10˚00'06.23"N 83˚41'47.85"W, 1470 m, 18 Dec
2010, A. P. Karremans 3199 & M. Contreras Fernandez
(JBL). Heredia: Sarapiquí, Puerto Viejo, Estación Biológica
La Selva, epítas en ramas caídas, 50 m, 3 Nov 2006, G.
Rojas 144 (JBL). Limón: Pococí, Guápiles, Hacienda La
Cuenca, bosque pluvial premontano, falda norte del Volcán
Turrialba, 10˚8'7.81"N 83˚46'46.2"W, 611 m, 1 Jan 2005,
M. A. Blanco 2762, A.Cháves, L. duToit & C. Ugalde
(JBL). Pococí, Guápiles, Guápiles, Pocora, La Argentina
Sur, márgenes del río Dos Novillos, sendero Las Cataratas,
bosque premontano muy húmedo, vegetación primaria
madura, 500–700 m, 12 Sept 2004, R. Valverde 1303 (JBL).
Siquirres, Pacuarito, 5 km después de la Estación del Parque
Nacional Barbilla, bosque pluvial premontano, epítas en
árbol caído a orillas del camino, 10˚01'1.5"N 83˚28'29.3"W,
627 m, 3 Dec 2008, D. Bogarín 5853, R. L. Dressler, R.
Gómez & R. Trejos (JBL). Same collecting data, D.
Bogarín 5871, R. L. Dressler, R. Gómez & R. Trejos (JBL).
Puntarenas: Buenos Aires, Olán, ascenso por la falda sur
del Cerro Tinuk, bosque pluvial premontano, epíta en
bosque secundario y primario, 9˚17'23.5"N 83˚10'33.5"W,
2164 m, 5 Jul 2012, D. Bogarín 9716, E. Herrera, D.
Jiménez, A. P. Karremans & V. H. Zúñiga (JBL). San José:
Pérez Zeledón, Montecarlo, without collector, 20 Sept 1991,
JBL-02659 (JBL).
5. Echinosepala lappiformis (A. Heller & L.O. Williams)
Pridgeon & M.W. Chase, Lindleyana 17(2): 101. 2002.
Basionym: Pleurothallis lappiformis A. Heller & L.O.
Williams, Fieldiana, Botany 31(2): 42, f. 8. 1964.
TYPE: NICARAGUA. Chontales: epiphytic at
Pistacho Peak near Babilonia Mine, alt. 650 m, Jul
1962, A. Heller 6620 (Holotype: F). Fig. 12 (Voucher,
Bogarín 9554, JBL).
Homotypic synonyms: Myoxanthus lappiformis (A. Heller
& L.O. Williams) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard.15: 38. 1986.
Echinella lappiformis (A. Heller & L.O. Williams)
Pridgeon & M.W. Chase, Lindleyana 16(4): 253.
2001, nom. illeg.
Brenesia lappiformis (A. Heller & L.O. Williams)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 255.
2004.
Epiphytic, shortly repent, erect to subprostrate herb up
to 25 cm tall. Roots coarse, exuous, ca. 2.0 mm in diam.
Ramicauls stout, terete, 15–50 mm long, homoblastic,
composed of 2–3 nodes, the lower one much shorter,
completely enclosed by 3 papyraceous, brous, tubular,
obtuse, whitish sheaths, increasing in size toward the
upper one, to 3 cm long, fragmented with age. Leaf thickly
coriaceous, elliptic, acute, minutely emarginate, 6–12 ×
2–3 cm, cuneate below into a conduplicate, sessile base,
the midvein strongly protruding abaxially. Inorescence a
single ower emerging from the lower nodes of ramicauls;
the peduncle terete, suberect to erect, minutely pubescent,
3–4 mm long, subtended by a brous, brown, acute bract
ca. 3 mm long. Floral bract brous-papyraceous, brownish,
inated, obliquely truncate, 7–8 mm long. Pedicel obconical,
pubescent, 2–3 mm long; ovary clavate, purple, 3 mm long,
densely long-pubescent to echinate. Flowers bilabiate, the
dorsal sepals apically connate to the tip of the synsepal,
eshy, nonresupinate, held almost vertically, neither
temporal activity of the perianth nor scent detected, bright
red-purple, the petals basally white, densely long pubescent-
echinate externally, rugose-verrucose within. Dorsal sepal
narrowly triangular, acute, 17–20 × 4–5 mm, 3-veined.
Lateral sepals connate to the apex into an obovate, obtuse,
concave synsepal, the margins erect, adaxially transversely
verruculose, 20–22 × 14–16 mm. Petals eshy, linear-
oblong, acute, eshy, adaxially verrucose in the distal half,
11–12 × 3 mm, 3-veined. Lip 3-lobed, elliptic from a thin,
rectangular, rose-hyaline claw, 7–8 × 3–4 mm (6 mm across
the lateral lobes), the base truncate; the apical lobe ovate,
minutely truncate, denticulate, verrucose, provided with
a pair of intramarginal, thick, apically verrucolose keels
running inside the lateral lobes toward the disc; lateral lobes
erect, elliptic-subuncinate, antrorse; the disc transversely
rugose-sulcate. Column straight, semiterete, 6–7 mm long,
provided with narrow, rectangular wings above the middle
ending at apex into pointed teeth, the foot ca. 1 mm long.
Anther cap globose, cucullate, with stiff hairs on the upper
margin, 2-celled. Pollinia 2, ovoid, laterally compressed, on
a short bilobed caudicle. Fig. 2I.
Etymology: from the Latin lappa, “bur,” for the
similarity of the ower to a spiny fruit.
Distribution: originally described from Nicaragua
(Williams, 1964), the species is reported to range southward
to northern South America, where it was recorded from
Colombia (Misas Urreta, 2005), Ecuador, and Venezuela
(Luer, 1992). We have examined specimens from Nicaragua,
Figure 12. Echinosepala lappiformis (A. Heller & L.O. Williams) Pridgeon & M.W. Chase. A, habit; B, ower; C, dissected perianth;
D, lip, ventral view; E, column and lip, lateral view; F, column, three-quarters view. Drawn from Bogarín 95541 by F. Pupulin.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 177
178 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Panama, and Colombia, which surely correspond to this
taxon, but we have not had access to materials from Ecuador
and Venezuela to conrm their identities.
Ecology: epiphytic in partial to deep shade on large
branches of trees, in primary and mature secondary
vegetation, Echinosepala lappiformis has been recorded in
Costa Rica exclusively from the warm, wet tropical forests
of the Caribbean plains at elevations between 50 and 650
m. Flowering has been recorded in cultivation from March
to June.
Distinguishing features: Echinosepala lappiformis is
easily recognized by the red-purple, long pubescent-echinate
owers borne at the base of the stems, with the narrowly
triangular dorsal sepal apically connate to the apex of the
synsepal. Echinosepala stonei, supercially similar, has the
dorsal sepal obtuse, free. The Panamanian E. balaeniceps,
not recorded from Costa Rica, has a much larger ower with
a tall, longitudinal callus on the disc.
Costa Rican material examined: Heredia: Sarapiquí,
Horquetas, ca. 80 m, Feb 2005, C. Ossenbach s.n. (JBL).
Limón. Siquirres, Pacuarito, 5 km después de la Estación
del Parque Nacional Barbilla, bosque pluvial premontano,
epítas en árbol caído a orillas del camino, 10˚1'1.5"N
83˚28'29.3"W, 627 m, 3 Dec 2008, D. Bogarín 5876, R. L.
Dressler, R. Gómez & R. Trejos (JBL). Pococí, Guápiles,
Parque Nacional Braulio Carrillo, Sector Quebrada
González, Sendero Las Palmas, detrás de las instalaciones,
10˚9'9.3"N 83˚56'43.5"W, 467 m, bosque muy húmedo
tropical transición a premontano, epítas en bosque
secundario alrededor del sendero, 22 Mar 2012, D. Bogarín
9554, M. Fernández, A. P. Karremans & C. Smith (JBL).
Pococí, Guápiles, 5 km al sur de la carretera, n del camino
paralelo con el Río Blanco, cerca de la Finca Tintoreras,
bosque muy húmedo tropical, 10˚2'16.2"N 83˚53'2.1"W,
350–400 m, 15 Jun 2006, R. L. Dressler 6768, D. Bogarín,
J. Gómez-Laurito & F. Pupulin (JBL). Guápiles, carretera
Braulio Carrillo, ca. 2 km hacia abajo de la entrada del
Teleférico del Bosque Lluvioso, en lomas a orilla de
la carretera, bosque muy húmedo tropical transición a
premontano, epítas en árboles caídos en bosque secundario,
10˚11'32"N 84˚54'27"W, 520 m, 9 Jul 2004, D. Bogarín 890
& F. Pupulin (JBL).
6. Echinosepala longipedunculata Pupulin & Karremans,
Lankesteriana 17(2): 299. 2017. TYPE: COSTA RICA.
Cartago: La Unión, San Rafael, Cerros de La Carpintera,
Campamento Escuela Iztarú, 9˚53'08.2"N 83˚58'15.6"W,
1778 m, bosque húmedo premontano, epítas en potreros
arbolados, 30 octubre 2008, oreció en cultivo en el Jardín
Botánico Lankester, 19 de junio 2012, D. Bogarín 5449, R.
L. Dressler, R. Gómez, F. Pupulin, & R. Trejos (Holotype,
JBL). Fig. 13 (Voucher: Bogarín 5449, JBL).
Epiphytic, caespitose, erect herb up to 30 cm tall. Roots
coarse, exuous, 1.5–2.0 mm in diam. Ramicauls stout, erect,
terete, slightly complanate, thicker in the distal portion, 5.5–
10.0 cm long, homoblastic, composed of 2 nodes different
in length, the lower one much shorter (to 1.5–2.2 cm long),
completely enclosed by 3 papyraceous, tubular, ancipitous,
apically loose, obliquely truncate, whitish sheaths,
increasing in size toward the upper one, 2.0–6.5 × 1.3–2.0
cm, the oldest ones breaking longitudinally into long bers
and eventually disintegrating with age. Leaf erect, thickly
coriaceous, linear-elliptic, subacute, minutely emarginate at
apex, 14–18 × 2.3–3.0 cm, cuneate below into a conduplicate,
sessile base, with the midvein strongly protruding abaxially
and several lateral veins faintly visible on the adaxial blade.
Inorescence a fascicle of single, successive owers at the
apex of ramicaul; the peduncle terete, suberect to gently
arching, sparsely and minutely pubescent, 3–8 cm long,
subtended by a papyraceous, whitish, ancipitous, obliquely
truncate spathe 2.7–4.2 cm long. Floral bract papyraceous,
loose, obliquely truncate-subobtuse, 7–10 mm long. Pedicel
terete-subclavate, 4–5 mm long; ovary completely covered
by the bract, linear-subclavate, 5 mm long, minutely
scaberulous. Flowers bilabiate, with temporal activity,
opening early in the morning and closing in the afternoon
during three or four days; the sepals yellow, mottled dark
purple in the inner side, ushed purple and densely scabrous
to shortly pubescent externally, abaxially verrucose, the
warts scattered at the base and becoming denser toward the
apex, the petals yellow, heavily blotched with purple-red,
the lip orange yellow. Dorsal sepal oblanceolate, obtuse,
19–21 × 6–7 mm, 7- to 9-veined. Lateral sepals connate into
an elliptic synsepal, verruculose in the distal half, 17–19 ×
9–10 mm, each sepal 5-veined, the free apices subacute-
rounded. Petals eshy, narrowly lanceolate, obliquely
asymmetric, acute, 6–7 × 1.5–1.8 mm, 3-veined, the central
vein raised into a rounded keel. Lip 3-lobed, narrowly
elliptic from a small, rectangular, thin, hyaline claw, 7 × 2
mm (4.5 mm across the lateral lobes), the base subtruncate;
the apical lobe rounded to subtruncate, thickened at apex
into a cushion-like, elliptic, low pad, provided with a pair
of intramarginal, thin keels running inside the lateral lobes
toward the disc; lateral lobes erect, narrowly uncinate,
antrorse; the disc with an erect, narrow callus arising at the
base of the lip, the highest part covered by a horseshoe-
shaped, channeled pseudoglenion extending in front into
a low keel ushing into the apical pad. Column straight to
subarcuate, semiterete, 5.0–5.5 mm long, provided with
narrow, elliptic wings above the middle, the foot ca. 1.5 mm
long. Anther cap globose, cucullate, with long, stiff hairs on
the upper margin, 2-celled. Pollinia 2, obovoid, attened,
on a short bilobed caudicle. Fig. 3A–B.
Etymology: from the Latin longipedunculatus, “with
a long peduncle,” in reference to the unusually elongate
peduncle of the ower, which can reach 8 cm in length.
Distribution: known only from Costa Rica, where it
has been recorded from the Caribbean watershed of the
Talamanca, Central Volcanic, and Guanacaste mountain
chains.
Ecology: epiphytic in partial shade on large branches and
trunks of trees, in primary and mature secondary vegetation,
recorded in lower montane wet, premontane rain, and
premontane wet forests at elevations of about 1300–1800 m.
Flowering has been recorded under cultivation from March
to June, September, and November.
Figure 13. Echinosepala longipedunculata Pupulin & Karremans. A, habit; B, ower; C, dissected perianth; D, lip, ventral view; E,
column and lip, lateral view; F, column, ventral view; G, anther cap and pollinarium. Drawn from Bogarin 5449 by F. Pupulin and D.
Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 179
180 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Distinguishing features: Echinosepala longipedunculata
is most similar to E. aspasicensis, with which it shares
the large habit of the plant and the general morphology
of the ower. Nevertheless, E. longipedunculata can be
recognized by the unusually long oral pedicel, which can
reach 8 cm in length (vs. 2.0–3.5 cm in E. aspasicensis), the
greenish-yellow owers (vs. purple-brown), the verrucose-
tuberculate adaxial surface of the sepals (vs. glabrous), and
the shorter, truncate, orange lip (vs. 1.5 × longer, obtuse to
acute, purple).
Costa Rican material examined: Cartago: Cartago,
San Francisco, Muñeco, Finca Loma Verde y Jilguero,
camino a Alto Belén, entre Río Sombrero y Quebrada
Patarrá, 9˚46'50.3"N 83˚54'21.1"W, 1430–1620 m, bosque
pluvial premontano, epítas en bosque secundario y árboles
en zonas abiertas, 23 mayo 2007, oreció en cultivo en
el Jardín Botánico Lankester, 30 marzo 2013, D. Bogarín
3863, M. Bonilla, R. Gómez, R. Trejos & J. D. Zúñiga (JBL).
Paraíso, Orosi, Tapantí, Parque Nacional Tapantí, unos 9 km
después de la entrada principal del parque, 500 m después
del puente superior sobre el río Grande de Orosi sobre la
calle que va a la represa, 9˚41'56.24"N 83˚46'53.12"W,
1570 m, epítas en árboles al lado de la calle principal,
bosque pluvial premontano, 10 de febrero 2012, oreció en
cultivo en el Jardín Botánico Lankester, 9 marzo 2013, A.
P. Karremans 4994 (JBL). Guanacaste: Liberia, Liberia.
Parque Nacional Rincón de la Vieja, sendero que recorre el
sector Volcán Santa María, 10˚47'26.00"N, 85˚19'16.00"O,
1304 m, bosque húmedo montano bajo, epíta, 24 Jan. 2015,
owered in cultivation at Lankester Botanical Garden, 29
nov 2016, I. Chinchilla 2216, R. Espinoza, C. Moraga, A.
Guadamuz & A. Rojas-Alvarado (JBL).
7. Echinosepala pan (Luer) Pridgeon & M.W. Chase,
Lindleyana 17(2): 101. 2002.
Basionym: Pleurothallis pan Luer, Selbyana 3(3–4):
360–362, f. 280. 1977. TYPE: PANAMA. Veraguas:
epiphytic in cloud forest northwest of Santa Fé,
alt. ca. 750 m, R. L. Dressler s.n. (Holotype: SEL).
Fig. 14 (Voucher: Bogarín 10274, JBL).
Homotypic synonyms: Myoxanthus pan (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 15: 38. 1986.
Echinella pan (Luer) Pridgeon & M.W. Chase,
Lindleyana 16(4): 253. 2001, nom. illeg.
Brenesia pan (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 255. 2004.
Epiphytic, caespitose to shortly repent herb, up to 25 cm
tall. Roots coarse, exuous, 1.5–2.0 mm in diam. Ramicauls
stout, erect, 4.5–5.7 cm long, homoblastic, composed of
2–3 nodes different in length (increasing from the basal to
the apical internode), each completely enclosed by equally
long, tubular, ancipitous, papyraceous, obliquely truncate,
hirsute, tight, apically loose sheaths, the oldest ones
breaking longitudinally and eventually disintegrating with
age. Leaves erect, coriaceous, narrowly linear-lanceolate,
acute, 1.4–1.8 × 10–22 cm, the base sessile. Inorescence
a fascicle of owers produced singly from the lower nodes
of the ramicauls; the peduncle terete-subclavate, short,
6–10 mm long, pubescent. Floral bract papyraceous,
loose, obliquely truncate-subobtuse, 5–8 mm long, densely
pubescent. Pedicel 2 mm long, pubescent; ovary obconical,
2 mm long, densely long-echinate. Flowers bilabiate, eshy,
short pedunculate, no temporal activity detected, emitting
an urine-like odor, the sepals dull greenish-yellow, spotted
and blotched with dark purple, the petals yellow, dotted with
purple, the lip dark purple-red, the column white to rose-
purple, spotted dark purple. Dorsal sepal elliptical-obovate,
obtuse, eshy, concave, reclined over the column, 11–15 ×
6–9 mm (ca. 4 mm wide at the base), 5-veined, adaxially
provided with spiny warts becoming denser toward the apical
portion, abaxially long-echinate. Lateral sepals connate into
a broadly elliptic-obovate elliptic-ovate, apically excised
for about half their length, 8.3–12.0 × 7.5–9.3 mm, each
half 3-veined, adaxially provided with spiny warts from
the distal half and becoming denser to the apical portion,
abaxially long-echinate. Petals rhombic, eshy, obliquely
asymmetric, subacute, subacutely angled on both margins,
4.3–5.0 × 2.9–3.8 mm, 3-veined, adaxially verrucose in the
thickened apical portion. Lip 3-lobed, narrowly elliptic-
oblong from a small, truncate, dark purple claw, 5.3–6.0
× 2.1–3.6 mm (6.8 mm wide across the expanded lateral
lobes), the base truncate, with two small, conical, retrorse
auricles; the lateral lobes erect, narrowly uncinate; the
apical lobe elliptic to broadly elliptic, rounded-truncate,
thickened at apex into a cushion-like, low pad, covered with
a translucent wax in fresh owers and provided with a pair
of intramarginal, thin keels running inside the lateral lobes
toward the disc; the disc with an erect, narrow callus arising
above the base of the lip, covered on the top by an inverted
U-shaped (horseshoe-shaped), channeled pseudoglenion
ushing into the apical pad. Column stout, straight,
semiterete, to 3.2 mm long, 1.5 mm wide, provided with
narrow stigmatic wings, the foot ca. 2.5 mm long. Anther
cap globose, cucullate, 2-celled, with long stiff hairs in
the upper margin, red-colored abaxially. Pollinia 2, ovoid,
laterally attened, on a short bilobed caudicle. Fig. 3C.
Etymology: named after the Greek god of the forest and
leader of the Satyrs, Pan, whose hair was short-horned and
bristly, resembling the abaxial indumentum of the sepals.
Distribution: known exclusively from Costa Rica
and Panama, ranging from the Tilarán mountain chain in
northern Costa Rica, to the regions of El Valle de Antón and
Santa Fé in central Panama.
Ecology: Echinosepala pan is apparently a rare species,
growing epiphytically in primary and secondary forests,
usually along streams. In Costa Rica it has been recorded in
the premontane wet forests along the Caribbean watershead
of the Cordillera de Tilarán, Cordillera Volcánica Central,
and Cordillera de Talamanca, at elevations ranging between
550 and 1150 m. Flowering has been recorded from June to
September.
Distinguishing features: Echinosepala pan is close
to E. expolita, with which it shares the dark owers and
the lip provided with a narrow, horseshoe-shaped callus.
It can be distinguished from the latter by the narrowly
linear-lanceolate leaves (vs. broader ligulate-lanceolate),
the smaller owers (sepals <10 mm vs. >15 mm long), the
hirsute and tight sheaths clasping the ramicauls (vs. glabrous
and inated), and the abaxial long-echinate indumentum of
the sepals (vs. shortly tomentose).
Figure 14. Echinosepala pan (Luer) Pridgeon & M.W. Chase. A, habit; B, ower; C, dissected perianth; D, lip, three-quarters view; E,
column and lip, lateral view; F, column, ventral view; G, anther cap; H, pollinarium, three views. Drawn from Bogarin 10274 by D.
Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 181
182 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Costa Rican material examined: Alajuela: San Ramón,
1867, A. Endrés s.n. [Pleurothallis No. 315] (W). San
Ramón, Ángeles, Reserva Biológica A.M. Brenes, 30 Dec.
1990, M. Freiberg s.n. (JBL). Same collecting data, owered
in cultivation at Lankester Botanical Garden, Sept 2066, M.
Freiberg s.n. (JBL). San Ramón, Ángeles, Reserva Forestal
de San Ramón, 850–1000 m, Aug. 1991, M. Germani 115
(JBL). San Ramón, Los Ángeles, deviation point to Villa
Blanca, km 1.5, epiphytic in windward premontane forest,
secondary vegetation, 10˚19'11"N 84˚28'28"W, 1120 m, 27
Feb 2003, F. Pupulin 4327, D. Bogarín, H. Montealegre &
A. C. Rodríguez (JBL). Limón: Pococí, Guápiles, Bellavista
600 m oeste de la Escuela La Guaria de Bellavista,
10˚8'35.111"N 83˚48'43.052"W, 569 m, bosque muy
húmedo tropical, epítas en bosque secundario a orillas de
una quebrada, 7 jun 2013, D. Bogarín 10270 (JBL).
8. Echinosepala stonei (Luer) Pridgeon & M.W. Chase,
Lindleyana 17(2): 101. 2002.
Basionym: Pleurothallis stonei Luer, Phytologia 44(3):
171. 1979. TYPE: COSTA RICA. San José: epiphytic
in cloud forest southeast of San José, cultivated in
California, owered in cult. 3 Mar 1978, submitted
to the OIC, no. 2022, C. Luer 2781 (Holotype, SEL).
Fig. 15 (Voucher, Bogarín 5777, JBL).
Homotypic synonyms: Myoxanthus stonei (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard.15: 38. 1986.
Echinella stonei (Luer) Pridgeon & M.W. Chase,
Lindleyana 16(4): 253. 2001, nom. illeg.
Brenesia stonei (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 255. 2004.
Epiphytic, shortly repent, erect herb up to 25 cm tall.
Roots stout, exuous, coarse, 1–2 mm in diam. Ramicauls
erect, stout, terete, 6–8 cm long, homoblastic, composed
of 2–4 internodes different in length (increasing from the
basal to the apical internode), each completely enclosed
by equally long, tubular, papyraceous, loose, obliquely
truncate-subobtuse, glabrous sheath, the oldest breaking
longitudinally and eventually disintegrating with age.
Leaves sessile, erect, coriaceous, broadly elliptic, 9–13 ×
2–4 cm, conduplicate, with a prominent midvein, the apex
asymmetrically retuse, the base cuneate. Inorescence a
fascicle of owers produced singly from the lower nodes
of the ramicauls; the peduncle terete-subclavate, short,
glabrous, ca. 1 cm long. Floral bract papyraceous, inated,
loose, obliquely truncate, acuminate, glabrous, 5–7 mm
long. Pedicel terete, arcuate, glabrous, 5 mm long; ovary
obconical, densely pubescent, not covered by the bract, ca.
4 mm long. Flowers held vertically, nonresupinate, deeply
concave-cucullate, eshy, short pedunculate, with a faint
urine-like scent, the apex of the dorsal sepal not connate to
the synsepal but accommodated inside forming two lateral
entrances; the sepals dark red-purple colored with whitish
hairs abaxially, petals and lip dark red-purple, column dark
red. Dorsal sepal broadly elliptic-oblong, acute, eshy,
14 × 7 mm, 5-veined, the free apex accommodated inside
the synsepal, adaxially covered with irregular, tall warts;
abaxial indumentum hirsute with soft trichomes. Lateral
sepals connate into a broadly oblong, concave, apiculate
synsepal, eshy, to 17 × 13 mm, each half 3-veined,
adaxially covered with irregular, tall warts, abaxially
hirsute with soft trichomes. Petals clavate, eshy, narrowly
rounded, obtuse, transversely rugose, 3 × 8 mm, warty from
the middle portion toward the apex. Lip 3-lobed, eshy,
elliptic-ovate from a small, thin, rectangular, white-hyaline
claw, 4 × 8 mm (ca. 6 mm across the lateral lobes), the
base truncate; the apical lobe elliptic-ovate, obtuse, smooth
becoming densely rugose in the distal portion; the lateral
lobes suberect, narrowly uncinate, antrorse, acuminate;
the disc provided with a massive, tall, rounded and broad,
slightly concave callus, occupying all the lip base and
extending to the disc; pseudoglenion absent. Column stout,
straight, semiterete, 5–6 mm long, 1–2 mm wide, provided
with narrow, uncinate wings above the middle, the foot ca. 3
mm long. Anther cap globose, 2-celled, cucullate, with stiff
hairs in the upper margin. Pollinia 2, ovoid, on a granulose,
short, bid caudicule. Fig. 2I.
Eponymy: named for Richard L. Stone of Los Altos
Hills, California, who discovered the species and owered
it in the United States.
Distribution: known only from Costa Rica.
Ecology: a rare epiphyte on trees, paddocks, and
secondary forest edges. Echinosepala stonei is apparently
restricted to the lower montane and premontane rain forests
along the Caribbean watershed of the Talamanca mountain
chain (and perhaps the Central Volcanic Cordillera), where
it has been recorded from 800 to 1900 m in elevation.
Flowering has been recorded from January to March.
Distinguishing features: the purple-red, hirsute
owers borne at the base of the ramicaul, with the dorsal
sepal broad, elliptic, obtuse, apically free, and the clavate
petals with tall warts distinguish Echinosepala stonei from
the closely related E. lappiformis, which has a narrowly
triangular, acute dorsal sepal apically connate to the apex
of the synsepal, and narrowly lanceolate, low-tuberculate
petals.
Echinosepala stonei is apparently a very rare species,
for a long time exclusively known from the type collection,
owered in cultivation in California, and allegedly collected
“southeast of San José” (Luer, 1979), an imprecise locality.
Here we can substantiate the presence of E. stonei in Costa
Rica, on the basis of three collections from the Caribbean
slopes of Cerro Duán, in the northern region of the
Talamanca cordillera.
Costa Rican material examined: Cartago: Jiménez,
Pejibaye, Tucurrique, Bajos del Humo, entre ríos Humo
y Vueltas, ladera este de Cerros Duán, 9˚48'36.7"N
83˚45'16.2"W, 1396 m, bosque pluvial montano bajo,
epítas en árboles en potreros y borde de bosque, 24 Nov
2008, D. Bogarín 5714, R. L. Dressler, R. Gómez & R.
Trejos (CR). Jiménez, Pejibaye, Tucurrique, Bajos del
Humo, entre ríos Humo y Vueltas, ladera este de Cerros
Duán, bosque pluvial montano bajo, epítas en árboles en
potreros y borde de bosque. 9˚48'36.7"N 83˚45'16.2"W,
1396 m, 24 Sept 2008, D. Bogarín 5740, R. L. Dressler, R.
Gómez & R. Trejos (JBL). Same locality data, D. Bogarín
5777, R. L. Dressler, R. Gómez & R. Trejos (JBL). Paraíso,
Figure 15. Echinosepala stonei (Luer) Pridgeon & M.W. Chase. A, habit; B, ower; C, dissected perianth; D, lip, ventral view; E, column
and lip, lateral view; F, column, ventral view; G, anther cap; H, pollinarium. Drawn from Bogarin 5777 by D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 183
184 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Cachí, Peñas Blancas, entre Cerros Duán y Alto Velo de
Novia, 9˚48'43.76"N 83˚46'36.61"W, 1829 m, bosque muy
húmedo premontano, epítas en árboles en potreros y borde
de bosque, 11 Mayo 2009, D. Bogarín 7190, R. Gómez, Y.
Kisel, P. Renshaw & R. Trejos (JBL).
9. Echinosepala tomentosa (Luer) Pridgeon & M.W. Chase,
Lindleyana 17: 101. 2002.
Basionym: Pleurothallis tomentosa Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 76: 177. 1999. TYPE: COSTA
RICA. Without locality, [San José: Candelaria]
ca. 1867, A. R. Endres 630 (Holotype: W). Fig. 16
(Voucher: Bogarín 5622, JBL).
Homotypic synonyms: Pleurothallis tomentosa Luer,
Orquideología 21: 337. 2000, nom. illeg.
Echinella tomentosa (Luer) Pridgeon & M.W. Chase,
Lindleyana 16: 253. 2001.
Myoxanthus tomentosus (Luer) Pupulin & M.A.
Blanco, Lankesteriana 2: 18. 2001.
Brenesia tomentosa (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 255. 2004.
Epiphytic, caespitose, erect herb up to 2 cm tall. Roots
coarse, exuous, 1 mm in diam. Ramicauls stout, erect,
terete, slightly complanate, thicker in the distal portion,
2.5–12.5 long, homoblastic, composed of 2 internodes
different in length, the lower one much shorter (to 1.0–
1.5 cm long), completely enclosed by four papyraceous,
tubular, ancipitous, apically loose, obliquely truncate,
greenish sheaths, increasing in size toward the upper one,
1.5–5.0 × 0.5 cm, the youngest with purple spots, the oldest
ones breaking longitudinally into long bers and eventually
disintegrating with age. Leaf erect, thickly coriaceous,
narrowly elliptic, subacute, minutely emarginate at apex,
8.2–11.2 × 1.5–2.4 cm, cuneate below into a conduplicate,
sessile base, with the midvein strongly protruding abaxially
and several lateral veins faintly visible on the adaxial blade.
Inorescence a fascicle of single, successive owers at the
apex of ramicaul; the peduncle terete, suberect, sparsely
and minutely pubescent, 3 mm long, subtended by a
papyraceous, whitish, ancipitous, obliquely truncate spathe
7 mm long. Floral bract papyraceous, loose, obliquely
truncate-subobtuse, 7 mm long. Pedicel terete-subclavate,
4 mm long, pubescent; ovary completely covered by the
bract, linear-subclavate, 1.5 mm long, pubescent. Flowers
bilabiate, no temporal activity of the perianth observed; the
dorsal sepal yellow, blotched with dark purple in the inner
side along the veins, the synsepal deep purple, adaxially
densely tomentose, the lip dark purple, column yellow with
purple blotches. Dorsal sepal linear-elliptic, obtuse, 11.5–
13.0 × 3.3–4.0 mm, 5-veined, adaxially tomentose. Lateral
sepals connate into an elliptic, synsepal, verruculose in
the distal half, 11–13 × 5–6 mm, each half 8-veined, the
free apices subacute-rounded, adaxially tomentose. Petals
eshy, narrowly lanceolate, obliquely asymmetric, acute,
4.5–5.0 × 1.3–2.0 mm, 3-veined, the central vein raised into
a rounded keel. Lip 3-lobed, ovate to oblong from a small,
rectangular, thin, hyaline claw, 6 × 2 mm (3.2 mm across the
lateral lobes), the base subtruncate; the apical lobe obtuse,
narrowly uncinate, antrorse; with a pair of intramarginal,
thin keels running toward the apex and canaliculate along
the middle; lateral lobes suberect, with a horseshoe-shaped,
channeled pseudoglenion at the base. Column straight to
subarcuate, semiterete, clavate, 4.3 × 1.2 mm long, the
foot ca. 1.3 mm long. Anther cap globose, cucullate, with a
mbriate crest, 2-celled. Pollinia 2, obovoid, attened, on a
short bilobed caudicle. Fig. 3E.
Etymology: from the Latin tomentosus, “covered by
short, rigid hairs,” in reference to the adaxial surface of
sepals.
Distribution: endemic to the central Pacic watershed
of the Cordillera de Talamanca, Costa Rica.
Ecology: epiphytic in partial shade on large branches
and trunks of trees, in primary and mature secondary
vegetation, in premontane rain, and premontane wet forests
at elevations of about 1200–1400 m. Flowering has been
recorded in cultivation in April and March.
Distinguishing features: Echinosepala tomentosa is
distinguished by the inorescences developed at the apex of
the stem, the leaves of fertile stems less than 12 cm long and
the adaxially densely tomentose sepals, the yellow dorsal
sepal and the purple synsepal. It is similar to E. vittata, but
that species has sepals abaxially with stiff hairs along the
veins and they are whitish, striped with red.
Luer (1999) described Pleurothallis tomentosa based on
a collection by A. R. Endrés in 1867, apparently without
locality data. However, the description made by A. R.
Endrés (Restrepia 630, W0020266) states “Candelaria,
March,” a place comprising the drainage basins of the
Tarrazú, Alumbre, and Santa Elena rivers owing into the
Pacic watershed of Cordillera de Talamanca (Ossenbach
et al., 2010). Luer suspected the species to be extinct, as
no collections after 1867 were known. However, we found
populations of this species in the Central Pacic region
(Tarrazú) of Costa Rica, where the species is locally
relatively frequent.
Additional Costa Rican material examined: Puntarenas:
Parrita, Parrita, 9 km suroeste de San Carlos de Tarrazú,
La Virgen, potreros hacia la cumbre de Fila Chonta,
9˚34'33.92"N 84˚9'24.49"W, 1240.2 m, bosque pluvial
montano bajo, epítas en bosque secundario remanente y
potreros, 25 Febrero 2020, D. Bogarín 12945, S. Abarca, I.
Chinchilla, G. Parra & R. Parra (JBL). San José: Tarrazú,
San Lorenzo, ca. 4 km al sureste de Santa Marta, camino
a Bajo Reyes, 9˚36'38.2"N 84˚00'52.1"W, 1475 m, bosque
muy húmedo premontano, en parche de bosque secundario
maduro, 20 Noviembre 2008, D. Bogarín 5622, R. L.
Dressler, M. Fernández, R. Gómez & R. Trejos (JBL).
10. Echinosepala vittata (Pupulin & M.A. Blanco) C.O.
Morales & N. Villalobos, Lankesteriana 4(3): 203. 2004.
Basionym: Myoxanthus vittatus Pupulin & M.A. Blanco,
Lankesteriana 2: 16. 2001. TYPE: COSTA RICA.
San José: Pérez Zeledón, El Brujo, near Río División,
450 m, 30 Jan. 2000, owered in cultivation at Jardín
Botánico Lankester, 3 June 2001, M. A. Blanco 1324
(Holotype: USJ, Isotype: JBL). Fig. 17 (Voucher:
Blanco 1324, USJ).
Figure 16. Echinosepala tomentosa (Luer) Pridgeon & M.W. Chase. A, habit; B, ower; C, dissected perianth; D, lip, ventral and three-
quarters views; E, column and lip, lateral view; F, column, three-quarters view; G, anther cap; H, pollinarium. Drawn from Pupulin 5622
by F. Pupulin and D. Solano Ulate.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 185
186 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Figure 17. Echinosepala vittata (Pupulin & M.A. Blanco) C.O. Morales & N. Villalobos. A, habit; B, ower; C, dissected perianth; D, lip,
ventral view; E, column and lip, lateral view; F, pollinarium; G, anther cap. Drawn from Blanco 1324 by F. Pupulin.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 187
Homotypic synoyms: Echinella vittata (Pupulin & M.A.
Blanco) Pupulin, Lankesteriana 4: 17. 2002, nom.
illeg.
Pleurothallis grammata Dressler, Lankesteriana 3:
28. 2002, nom. subst.
Echinosepala vittata (Pupulin & M.A. Blanco) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 119.
2007, nom. illeg., nom. super.
Epiphytic, caespitose herb up to 17 cm tall. Roots coarse,
ca. 2 mm in diam. Ramicauls stout, erect, 4–7 cm long,
enclosed by 3–4 loose, tubular sheaths, fragmented with age.
Leaf erect, thickly coriaceous, narrowly lanceolate-elliptic,
minutely emarginate, 7–10 × 1.3–1.5 cm, cuneate below
into a conduplicate, sessile base. Inorescence a fascicle of
single, successive owers, produced at the apex of ramicaul,
the peduncle terete, sparsely pubescent, 0.8–1.3 cm long,
subtended by a papyraceous, ancipitous spathe 1.4–1.6 cm
long. Pedicel terete-subclavate, 2–3 mm long; ovary less
than 1 mm long, densely pubescent. Flowers small, eshy,
whitish, longitudinally striped with purple, densely short-
pubescent externally, glabrous within. Dorsal sepal linear-
oblanceolate, obtuse, 1.0–1.2 × 0.3 cm, 5-veined. Lateral
sepals connate into a lanceolate, shortly emarginate, concave
synsepal, 1.0–1.3 × 0.4–0.5 mm, each half 4-veined. Petals
eshy, linear-oblong, obtuse, minutely apiculate, adaxially
provided with a tuft of short papillae near the apex, 4 × 1
mm, 3-veined. Lip 3-lobed, ligulate, subacute, 4 mm long,
2.5 mm wide between lateral lobes; the apical lobe smooth,
slightly undulate along the margins; the lateral lobes erect,
narrowly uncinate, antrorse; the disc with an erect, narrow,
low, horseshoe-shaped, channeled callus above the base,
extending in front into a low keel just to near the lip apex,
laterally provided with a pair of low lamellae extending to
the apex, hinged on the end. Column arcuate, semiterete, 1.6
mm long, provided with narrow wings above the middle,
the foot less than 1 mm long. Anther cap globose, cucullate,
with long hairs on the upper margin, 2-celled. Pollinia 2,
obovoid, attened, on a short, bid caudicle. Fig. 3F.
Etymology: from the Latin vittatus, “longitudinally
striped,” in reference to the stripes on sepals and petals.
Distribution: known only from northern and central
Costa Rica, where it has been recorded on both watersheds
of the continental divide.
Ecology: epiphytic in tropical wet forest, premontane
belt transition, at 450–750 m in elevation. Flowering has
been recorded from November to June.
Distinguishing features: Echinosepala vittata is closely
related to E. glenioides, E. longipedunculata, and particularly
to E. tomentosa, all provided with a pseudoglenion at the
base of the lip, but it is distinguished by the small size, the
densely pubescent ovary, the whitish ower striped with
purple, the sepals adaxially smooth, and the ligulate, obtuse
lip. Echinosepala tomentosa may be distinguished from
E. vittata by the purple-black owers with an oblong lip,
rounded at apex.
Together with Echinosepala tomentosa, the other species
with a short vegetative habit and small ower, E. vittata
groups into the basal-most clade of Echinosepala, sister to
all the other species in the genus.
Additional Costa Rican material examined: Alajuela:
Upala, Bijagua. En la calle nueva desde el Celeste Moutain
Lodge al parque Volcán Tenorio, orillas del bosque y
potrero al lado del camino, 10˚43'8.46"N 85˚0'2.07"W,
729 m, 14 Dec 2016, A. P. Karremans 7524, M. Cedeño, I.
Chinchilla, M. Díaz y G. Rojas-Alvarado (JBL). San José:
León Cortés, San Pablo, R.F. Los Santos, El Abejonal, 7 Jan
1935, E. Azoefa 286 (CR). Pérez Zeledón, El Brujo, near
Río División, 450 m, 30 Jan. 2000, owered in cultivation
at Jardín Botánico Lankester, 3 June 2001, M. A. Blanco
1324 (JBL). Pérez Zeledón, Viento Fresco de El Brujo,
9˚25'13"N 83˚56'27"W, 450 m, 21 Jan 2001, F. Pupulin
2878, D. Castelfranco & E. Elizondo (JBL).
literature cited
atwood, J. T. 1987. The vascular ora of la Selva Biological
Station, Costa Rica. Orchidaceae. Selbyana 10(1): 76–145.
atwood, J. T., and D. E. mora de retana. 1999. Family #39,
Orchidaceae: Tribe Maxillarieae: Subtribes Maxillariinae and
Oncidiinae. In W. Burger, ed., Flora Costaricensis. Fieldiana,
Bot. n.s. 40.
Bogarín, D., I. F. chinchilla and M. cedeño Fonseca. 2020. Two
new species of Lepanthes (Orchidaceae: Pleurothalldinae) from
Costa Rica and their phylogenetic afnity. Plant Systematics and
Evolution 306(2): 20.
Bogarín, D., and M. Fernández. 2010. Lepanthes arenasiana
(Pleurothallidinae: Orchidaceae), a new species from Costa
Rica. Lankesteriana 9(3): 487–489.
Bogarín, D., M. Fernández, and z. serracín. 2018. Lepanthes
aures-ursinae and L. vertebrata spp. nov. (Orchidaceae: Pleuro-
thallidinae) from Panama. Nordic J. Bot., 36(1–2), njb-01292.
Bogarín, D., and d. Jiménez. 2015. Lepanthes novae Durikaensis.
Lindleyena [in Orchids (Bull. Amer. Orchd. Soc.)] 84: 430–440.
Bogarín, D., and a. P. Karremans. 2010. A new Platystele
(Orchidaceae: Pleutothallidinae) from central Costa Rica.
Orquideología 27: 208–220.
––––––. 2016. A New Brachionidium (Orchidaceae: Pleuro-
thallidinae) from the First Botanical Expedition to the Volcán
Cacho Negro, Costa Rica. Syst Bot. 41(4): 919–923(5).
Bogarín, D., a. P. Karremans, and m. Fernández. 2018.
Genus-level taxonomical changes in the Lepanthes afnity
(Orchidaceae, Pleurothallidinae). Phytotaxa 340(2): 128–136.
Bogarín, D., a. P. Karremans, and m. muñoz garcía. 2015.
Brachionidium kirbyi, eine neue Art zu Ehren des Gründers
des Orchideenprojektes “Bosque de Paz” in Costa Rica. Die
Orchidee 66: 404–409.
Bogarín, D., a. P. Karremans, and F. PuPulin. 2008. New species
and records of Orchidaceae from Costa Rica. Lankesteriana
8(2): 53–74.
Bogarín, D., a. P. Karremans, r. rincón, and B. gravendeel.
2013. A new Specklinia (Orchidaceae: Pleurothallidinae) from
Costa Rica and Panama. Phytotaxa 115(2): 31–41.
Bogarín, D., and y. Kisel. 2014. A new Lepanthes from Costa
Rica. Orchid Rev. 122(1305): 28–31.
Bogarín, D., l. oses, and c. m. smith. 2017. Masdevallia
luerorum (Orchidaceae: Pleurothallidinae), a new species from
Costa Rica. Lankesteriana 17(2): 235–244.
188 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
Bogarín, D., o. a. Pérez-escoBar, d. groenenBerg, s. d. holland,
a. P. Karremans, e. m. lemmon, a. r. lemmon, F. PuPulin, e.
smets, and B. gravendeel. 2018. Anchored hybrid enrichment
generated nuclear, plastid and mitochondrial T markers resolve
the Lepanthes horrida (Orchidaceae: Pleurothallidinae) species
complex. Molec. Phylogen. Evol. 129(2018): 27–47.
Bogarín, D., and F. PuPulin. 2011. Lepanthes daniel-jimenezii
Bogarín & Pupulin, spec. nov. (Pleurothallidinae: Orchidaceae),
a new species from Costa Rica close to Lepanthes guardiana.
Die Orchidee 62(2): 111–115.
––––––. 2012. Three new Lepanthes (Orchidaceae: Pleurothallid-
inae) from South-East Costa Rica. Lankesteriana 12(2): 107–114.
Bogarín, D., z. serracín, and z. samudio. 2014. Illustrations
and studies in Neotropical Orchidaceae, The Speccklinia
condylata group (Pleurothallidinae) in Costa Rica and Panama.
Lankesteriana 13(3): 185–206.
Bogarín, D., z. serracín, z. samudio, r. rincón, and F. PuPulin.
2014. An updated checklist of the Orchidaceae of Panama.
Lankesteriana 14(3): 135–364.
Bogarín, D., c. m. smith, and d. Jiménez. 2012. The Lepanthes
ovalis group (Pleurothallidinae) with three new species from
Costa Rica. J. Bot. Res. Inst. Texas 6(2): 361–373.
Burger, w. c. 1971. Family #40, Casuarinaceae. Family #41,
Piperaceae. In W. Burger, ed., Flora Costaricensis. Fieldiana,
Bot. 35.
Burger, w. c., K. Barringer, a. gentry, and l. d. gómez. 2000.
Family #193, Scrophulariaceae; Family #193a, Schlegeliaceae;
Family #194, Bignoniaceae; Family #195, Pedaliaceae; Family
#196, Martyniaceae; Family #197, Orobanchaceae. In W.
Burger, ed., Flora Costaricensis. Fieldiana, Bot. n.s. 41.
Burger, w. c., J. g. Furlow, l. d. gómez P., and d. e. stone.1977.
Family #42, Chlorantaceae; Family #43, Lacistemaceae;
Family #44, Salicaceae; Family #45, Garryaceae; Family #46,
Myricaceae; Family #47, Juglandaceae; Family #48, Bataceae;
Family #49, Betulaceae; Family #50, Fagaceae; Family #51,
Ulmaceae; Family #52, Moraceae; Family #52a, Cannabaceae;
Family#53, Urticaceae. In W. Burger, ed., Flora Costaricensis.
Fieldiana, Bot. 40.
Burger, w. c., and m. huFt. 1995. Family #113, Euphorbiaceae.
In W. Burger, ed., Flora Costaricensis. Fieldiana, Bot. n.s. 36.
Burger, w. c., J. KuiJt, K. Barringer, l. d. gómez, and r.
BaKer. 1983. Family #54, Podostemaceae; Family #55,
Proteaceae; Family #56, Olacaeae; Family #57, Opiliaceae;
Family #58, Loranthaceae; Family #59, Aristolochiaceae;
Family #60a, Hydnoraceae; Family #60b, Rafesiaceae; Family
#61, Balanophoraceae; Family #62, Polygonaceae; Family
#63, Chenopodiaceae; Family #64, Amaranthaceae; Family
#65, Nyctaginaceae; Family #66, Phytolaccaceae; Family
#67, Aizoaceae; Family #68, Portulacaceae; Family #69,
Basellaceae; Family #70, Caryophyllaceae. In W. Burger, ed.,
Flora Costaricensis. Fieldiana, Bot. n.s. 13.
Burger, w. c., t. Plowman, B. B. simPson, and n. zamora.
1991. Family #97, Krameriaceae; Family #98, Oxalidaceae;
Family #99, Geraniaceae; Family #100, Tropaeolaceae; Family
#101, Linaceae; Family #101a, Humiriaceae; Family #102,
Erythroxylaceae; Family #103, Zygophyllaceae. In W. Burger,
ed., Flora Costaricensis. Fieldiana, Bot. n.s. 28.
Burger, w. c., and c. m. taylor. 1993. Family #202, Rubiaceae.
In W. Burger, ed., Flora Costaricensis. Fieldiana, Bot. n.s. 33.
Burger, w. c., and h. van der werFF. 1990. Family #80,
Lauraceae; Family #81, Hernandiaceae. In W. Burger, ed.,
Flora Costaricensis. Fieldiana, Bot. n.s. 23.
dressler, r. l. 1981. The Orchids: Natural History and
Classication. Harvard University Press, Cambridge,
Massachusetts, U.S.A.
Pleurothallidinae1993. Phylogeny and Classication of the Orchid
Family. Cambridge University Press, Cambridge.
drummond, a. J., and a. ramBaut. 2007. BEAST: Bayesian
evolutionary analysis by sampling trees. BMC Evol. Biol. 7:
214.
durKee, l. h., and w. Burger. 1986. Family #200, Acanthaceae;
Family #201, Plantaginaceae. In W. Burger, ed., Flora
Costaricensis. Fieldiana, Bot. n.s. 18.
ePidendra. 2019. The Global Orchid Taxonomic Network. Jardín
Botánico Lankester, Universidad de Costa Rica (accessed June
2019). Available from: http://epidendra.org/taxones/index.html
Fawcett, w. 1895. Two new orchids from Jamaica. J. Bot. 33(385):
12.
Fawcett, w., and a. B. rendle. 1910. Flora of Jamaica:
Orchidaceae. British Museum, London.
Fernández, m. 2011. A reconsideration of Trichosalpinx
minutipetala (Ames & C.Schweinf.) Luer (Orchidaceae:
Pleurothallidinae). Orquideología 28(2): 119–134.
––––––. 2013. Tubella—those other Trichosalpinx. Die Orchidee
64(4): 310–317.
Fernández, m., and d. Bogarín. 2011. A new Trichosalpinx
(Orchidaceae: Pleurothallidinae) from the northern Pacic
lowlands of Costa Rica. Phytotaxa 38: 41–48.
––––––. 2013. A new species of Trichosalpinx (Orchidaceae:
Pleurothallidinae) from Costa Rica. Brittonia 65(1): 96–101.
Fernández, m., d. Bogarín, a. P. Karremans, and d. Jiménez.
2014. New species and records of Orchidaceae from Costa Rica.
III. Lankesteriana 13(3): 259–282.
hamer, F. 1984. Orchids of Nicaragua, Part 5. Icones Plantarum
Tropicarum 12: pl. 1101–1200.
––––––. 2001. Orchidaceae. Pages 1612–1853 in w. d. stevens,
c. ulloa ulloa, a. Pool, and o. m. montiel, eds., Flora de
Nicaragua. Monogr. Syst. Bot. Missouri Bot. Gard. 85(2).
IPNI. 2020. International Plant Names Index (accessed May 2020.
http://www.ipni.org/
Jørgensen, P. m. and s. león-yánez, eds. 1999. Catalogue of the
vascular plants of Ecuador. Monographs in Systematic Botany
from the Missouri Botanical Garden 75: 1-1181.
Jørgensen, P. m., m. h. nee and s. g. BecK, eds. 2014. Catálogo
de las plantas vasculares de Bolivia. Monographs in Systematic
Botany from the Missouri Botanical Garden 127(1-2): 1-1744.
Karremans, a. P. 2012. Illustrations and studies on Neotropical
Orchidaceae. 3. On the identity of Dracontia pachyglossa and D.
ramonensis; with a new species, D. lueriana (Pleurothallidinae).
Ann. Naturhist. Mus. Wien, B. 113: 119–132.
––––––. 2014. Lankesteriana, a new genus in the Pleurothallidinae
(Orchidaceae). Lankesteriana 13(3): 319–332.
––––––. 2016. Genera Pleurothallidinarum: An updated
phylogenetic overview of Pleurothallidinae. Lankesteriana
16(2): 219–241.
––––––. 2019. To be or not to be a Stelis. Lankesteriana 19(3):
281–343.
Karremans, a. P., d. aguilar sandí, m. artavia-solís, m.
cedeño-Fonseca, i. F. chinchilla, K. gil, g. roJas-alvarado,
n. solano-guindon, and J. villegas-murillo. 2019.
Nomenclatural notes in the Pleurothallidinae (Orchidaceae):
Miscellaneous. Phytotaxa 406(5): 259–270.
Karremans, a. P., and d. Bogarín. 2011. Pleurothallis adventurae
(Orchidaceae: Pleurothallidinae), eine neue Art aus einer
unerforschten Region in Costa Rica/Pleurothallis adventurae
(Orchidaceae: Pleurothallidinae), una nueva especie de una
región inexplorada en Costa Rica. Orchideen Journal 18(3):
111–114.
––––––. 2013. Three new species of Dracontia (Pleurothallidinae,
Orchidaceae) from Costa Rica. Syst. Bot. 38(2): 307–315.
––––––. 2017. Two novelties in Platystele (Ochidaceae:
Pleurothallidinae) from Costa Rica. Lankesteriana 17(2): 215–
221.
Karremans, a. P., d. Bogarín, m. díaz-morales, m. Fernández,
l. oses, and F. PuPulin. 2016. Phylogenetic reassessment of
Acianthera (Pleurothallidinae: Orchidaceae). Harvard Pap. Bot.
21(2): 171–187.
Karremans, a. P., d. Bogarín, m. Fernández, c. m. smith, and
m. a. Blanco. 2012. New species and records of Orchidaceae
from Costa Rica. II. Lankesteriana 12(1): 19–51.
Karremans, a. P., d. Bogarín, and B. gravendeel. 2015.
Specklinia lugduno-batavae (Pleurothallidinae: Orchidaceae), a
new species in the S. digitalis group. Blumea 59(3): 180–184(5).
Karremans, a. P., and m. díaz-morales. 2017. Novelties in Costa
Rican Stelis (Orchidaceae: Pleurothallidinae): Two new species
and a new record in the “Dracontia Group.” Lankesteriana
17(2): 193–202.
Karremans, a. P., and J. e. Jiménez. 2018. Pleurothallis hawkingii
and Pleurothallis vide-vallis (Orchidaceae; Epidendroideae),
two new species from Cordillera de Guanacaste in Costa Rica.
Phytotaxa 349(2): 185–191.
Karremans, a. P., and m. muñoz garcía. 2011. Pleurothallis
silvae-pacis, a new species. Orchid Rev. 119(1295): 155–159.
Karremans, a. P., F. PuPulin, and B. gravendeel. 2015. Specklinia
dunstervillei, a new species long confused with Specklinia
endotrachys (Orchidaceae: Pleurothallidinae). PLoS One 10(7):
e0131971.
Karremans, a. P., g. salguero, d. Bogarín, l. oses, and m.
cedeño-Fonseca. 2020. Illustrations and studies in Neotropical
Orchidaceae: The Specklinia brighamii group (Pleurothallidinae)
in Costa Rica. Phytotaxa 447(1): 16–30.
Karremans, a. P., and c. m. smith. 2012. A note on genus
Dracontia (Orchidaceae: Pleurothallidinae), with a new species.
Harvard Pap. Bot. 17(1): 13–17.
KolanowKa, m. 2014. Orchids of the Darien gap. Koeltz Scientic
Books, Koenigstein, Germany.
luer, C. A. 1976. Icones Pleurothallidinarum. Miscellaneous
species of Pleurothallis. Selbyana 3: 38–201.
––––––. 1977. Icones Pleurothallidinarum (Orchidaceae).
Miscellaneous species in the Pleurothallidinae. Selbyana 3(3–4):
386–387, f. 293.
––––––. 1979. Miscellaneous new species in the Pleurothallidinae.
Phytologia 44: 164.
––––––. 1986. Icones Pleurothallidinarum. 1. Systematics of
Pleurothallidinae. Monogr. Syst. Bot. Missouri Bot. Gard. 15:
65–69.
––––––. 1992. Icones Pleurothallidinarum. 9. Systematics of
Myoxanthus. Addenda to Platystele, Pleurothallis subgenus
Scopula, and Scaphosepalum (Orchidaceae). Monogr. Syst. Bot.
Missouri Bot. Gard. 44: 1–128.
––––––. 1995. A re evaluation of the Pleurothallid subgenera
Satyria and Silenia (Orchidaceae). Page 146 in Icones
Pleurothallidinarum. 12. Systematics of Brachionidium.
Addenda to Dresslerella, Platystele and Porroglossum. Monogr.
Syst. Bot. Missouri Bot. Gard. 57.
––––––. 1996. Validation of taxa in the Pleurothallidinae.
Lindleyana 1(2): 118.
––––––. 1999. Icones Pleurothallidinarum. 18. Systematics of
Pleurothallis subgen. Pleurothallis sect. Pleurothallis subsect.
Antenniferae, subsect. Longiracemosae, subsect. Macrophyllae-
Racemosae, subsect. Perplexae, subgen. Pseudostelis, subgen.
Acuminatia. Monogr. Syst. Bot. Missouri Bot. Gard. 76: 177,
f. 33a.
––––––. 2000. Miscellaneous new species in the Pleurothallidinae.
Orquideologia 21(3): 318–340.
––––––. 2002. A systematic method of classication of the
Pleurothallidinae versus a strictly phylogenetic method.
Selbyana 23(1): 57–110.
––––––. 2003. Pleurothallis. Pages 386–452 in B. e. hammel, m. h.
grayum, c. herrera, and n. zamora, eds., Monocotiledóneas
(Orchidaceae–Zingiberaceae). Vol. 3 in Manual de Plantas de
Costa Rica. Monogr. Syst. Bot. Missouri Bot. Gard. 93.
––––––. 2004. Icones Pleurothallidinarum. 26. New genera and
combinations in the Pleurothallidinae. Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 253–265.
––––––. 2007. Icones Pleurothallidinarum. 29. A third century
of Stelis of Ecuador. Systematics of Apoda–Prorepentia.
Systematics of miscellaneous small genera. Addenda: New
genera, species, and combinations (Orchidaceae). Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 119.
luteyn J. l., and r. l. wilBur. 2005. Family #172, Ericaceae.
In W. Burger, ed., Flora Costaricensis. Fieldiana, Bot. n.s. 45.
maddison, w. P., and d. r. maddison. 2007. Mesquite: A modular
system for evolutionary analysis. Mesquite v. 2.72. Available at
http://mesquiteproject.org
mcleish, i., n. r. Pearce, B. r. adams, and J. s. Briggs. 1995.
Native Orchids of Belize. A. A. Balkema, Rotterdam, The
Netherlands.
misas urreta, G. 2005. Orquídeas de la Serranía del Baudó,
Chocó, Colombia. Corporacion Capitalina de Orquideología,
Bogotá, Colombia.
oses salas, l., and a. P. Karremans. 2016. A note on Masdevallia
zahlbruckneri and M. utriculata (Orchidaceae). Harvard Pap.
Bot. 21(2): 325–336.
ossenBach, c., F. PuPulin, and r. Jenny. 2010. Orchid itineraries
of Augustus R. Endrés in Central America: A biographic and
geographic sketch. Lankesteriana 10(1): 19–47.
––––––. 2013. The Illustrations and the Catalogue. Vol. 2 of
Orchids in the Life and Work of Auguste R. Endrés. Verlag des
Naturhistorischen Museums, Vienna.
Pohl, r. w. 1980. Family #15, Gramineae. In W. Burger, ed.,
Flora Costaricensis. Fieldiana, Bot. n.s. 4.
Pridgeon, A. M. 2005a. Echinosepala. Pages 356–358 in A. m.
Pridgeon, P. J. criBB, m. w. chase, and F. n. rasmussen,
eds., Epidendroideae. Part 1.Vol. 4 of Genera Orchidacearum.
Oxford University Press, Oxford.
––––––. 2005b. Echinosepala. Pages 246–249 in F. PuPulin, ed.,
Acianthera—Kegeleilla. Vol. 1 of Vanishing Beauty. Native
Costa Rican Orchids. Editorial de la Universidad de Costa Rica,
San José.
––––––. 2020. Pleurothallis. Pages 850–909 in F. PuPulin, ed.,
Lacaena—Pteroglossa. Vol. 2 of Vanishing Beauty. Native
Costa Rican Orchids. Koeltz Botanical Books, Oberreifenberg,
Germany.
Pridgeon, A. M., and m. w. chase. 2001. A phylogenetic
reclassication of Pleurothallidinae (Orchidaceae). Lindleyena
16(4): 235–271.
––––––. 2002. Nomenclatural notes on Pleurothallidinae
(Orchidaceae). Lindleyana 17(2): 98–101.
Pridgeon, A. M., r. solano, and m. w. chase. 2001. Phylogenetic
relationships in Pleurothallidinae (Orchidaceae): Combined
evidence from nuclear and plastid DNA sequences. Amer. J. Bot.
88(12): 2286–2308.
PuPulin, F. 2002. Catálogo revisado y anotado de las Orchidaceae
de Costa Rica. Lankesteriana 4: 1–88.
––––––. 2010. Family #39, Orchidaceae: Tribe Cymbidieae:
Subtribe Zygopetalinae. In W. Burger, ed., Flora Costaricensis.
Fieldiana, Bot. n.s. 49.
––––––. 2020. Pabstiella. Pages 809–811 in F. PuPulin, ed., Lacaena–
Pteroglossa. Vol. 2 of Vanishing Beauty. Native Costa Rican
Orchids. Koeltz Botanical Books, Oberreifenberg, Germany.
2020 PUPULIN ET AL., FLORAE COSTARICENSIS (ORCHIDACEAE) PRODROMUS 189
190 HARVARD PAPERS IN BOTANY VOL. 25, NO. 2
PuPulin, F., and d. Bogarín. 2004. Two new species of Lepanthes
(Orchidaceae: Pleurothallidinae) from Costa Rica. Kew Bull.
59: 559–563.
––––––. 2007. A second species of Restrepiella (Orchidaceae:
Pleurothallidinae). Willdenowia 37(1): 323–329.
––––––. 2010. Illustrations and studies in Neotropical Orchidaceae—
The Lepanthes jimenezii group (Pleurothallidinae). Harvard
Pap. Bot. 15(1): 111–121.
––––––. 2011a. Two Lepanthes (Orchidaceae: Pleurothallidinae)
with strongly reduced corolla. OrchidsJournal 1(1): 31–35.
––––––. 2011b. Two New Lepanthes from Costa Rica. Lindleyana
[in Orchids (Bull. Amer. Orch. Soc.)] 80(3): 178–181.
––––––. 2012. Lepanthes novae Tapantienses. Orch. Digest 76(1):
20–29.
––––––. 2014a. Aliae Lepanthes Machogaffenses (Orchidaceae:
Pleurothallidinae). Harvard Pap. Bot. 19(2): 193–200.
––––––. 2014b. Illustrations and studies in Neotropical Orchidaceae.
6. The “Lepanthes guatemalensis” group (Pleurothallidinae) in
Costa Rica. An. Jard. Bot. Madrid 71(1): 1–17.
––––––. 2020. Lepanthes. Pages 444–511 in F. PuPulin, ed., Lacaena–
Pteroglossa. Vol. 2 of Vanishing Beauty. Native Costa Rican
Orchids. Koeltz Botanical Books, Oberreifenberg, Germany.
PuPulin, F., d. Bogarín, and m. Fernández. 2010a. Illustrations and
studies in Neotropical Orchidaceae. 2. A note on Pleurothallis
luctuosa (Orchidaceae: Pleurothallidinae), with a new species.
Ann. Naturhist. Mus. Wien. B.112: 239–252.
––––––. 2010b. On the identity of Myoxanthus scandens
(Orchidaceae: Pleurothallidinae), with a new species.
Lankesteriana 9(3): 467–473.
PuPulin, F., d. Bogarín, and d. Jiménez. 2009. New species and
records in Mesoamerican Lepanthes. Orchid Digest 73: 136–
145.
PuPulin, F., d. Bogarín, and a. P. Karremans. 2019. How many
species of Pleurothallidinae are there? Pages 301–362 in a.
Pridgeon, ed., Proceedings of the Sixth Scientic Conference on
Andean Orchids, 2017. Guayaquil, Ecuador.
PuPulin, F., d. Bogarín, and c. m. smith. 2010. Two new
species of Lepanthes from Costa Rica close to L. schizocardia
(Orchidaceae: Pleurothallidinae). Lankesteriana 9(3): 423–430.
PuPulin, F., m. díaz-morales, J. aguilar, and m. Fernández.
2017. Two new species of Pleurothallis (Orchidaceae:
Pleurothallidinae) allied to P. cardiothallis, with a note on ower
activity. Lankesteriana 17(2): 329–356.
PuPulin, F., m. díaz-morales, m. Fernández, and J. aguilar.
2017. Two new species of Pleurothallis (Orchidaceae: Pleuro-
thallidinae) from Costa Rica in the P. phyllocardia group.
Lankesteriana 17(2): 153–164.
PuPulin, F., a. P. Karremans, and n. BelFort oconitrillo.
2017. Two new species of Echinosepala. Lankesteriana 17(2):
285–304.
PuPulin, F., a. P. Karremans, and B. gravendeel. 2012. A recon-
sideration of the empusellous species of Specklinia (Orchidaceae:
Pleurothallidinae) in Costa Rica. Phytotaxa 63: 1–20.
PuPulin, F., h. medina, and d. Bogarín. 2010. Two Lepanthes
(Orchidaceae: Pleurothallidinae) with a reduced corolla.
Orchideen J. 17(3): 117–121.
PuPulin, F., and l. oses. 2020. Masdevallia. Pages 572–609 in
F. PuPulin, ed., Lacaena–Pteroglossa. Vol. 2 in Vanishing
Beauty. Native Costa Rican Orchids. Koeltz Botanical Books,
Oberreifenberg, Germany.
PuPulin, F., c. ossenBach, r. Jenny, and e. viteK. 2011. Typi
Orchidacearum ab Augusto R. Endresio in Costa Rica lecti.
Annalen Naturhistorisches Museums Wien, Ser. B, Botanik.
Zoologie. 112: 265–313.
PuPulin, F., g. roJas, and J. d. zúñiga. 2007. Three new species
of Acianthera (Orchidaceae: Pleurothallidinae) from Costa Rica.
Harvard Pap. Bot. 12(1): 163–170.
ramBaut, A. 2009. FigTree v1.3.1. Available at http://tree.bio.
ed.ac.uk/ software/
reichenBach, h. g. 1855. Wagener’s Orchideen aus Ocaña.
Bonplandia 3(5): 65–73.
renner. s. s., h. Balslev and l. B. Jolm-nielsen. 1990. Flowering
plants of Amazonian Ecuador–A checklist. AAU Reports 24:
1–241.
roJas-alvarado, g., and a. P. Karremans. 2017. Additions to
the Costa Rican Myoxanthus (Orchidaceae: Pleurothallidinae).
Lankesteriana 17(2): 203–214.
schlechter, R. 1923. Beiträge zur Orchideenkunde von
Zentralamerika. II. Additamenta ad Orchideologiam
Costaricensem. Repert. Spec. Nov. Regni Veg. Beih. 19: 200.
smith, c. m., d. Bogarín, and F. PuPulin. 2015. A new Reichantha
(Orchidaceae: Pleurothallidinae) from Nicaragua and Costa
Rica. Syst. Bot. 40(1): 83–87.
smith, c. m., d. Jiménez, and F. PuPulin. 2013. Lepanthes
variabilis (Orchidaceae: Pleurothallidinae), a new endemic
species from Costa Rica. Brittonia 65: 469–476.
smith, c. m., and F. PuPulin. 2012. Contributions to a taxonomic
revision of Masdevallia (Orchidaceae: Pleurothallidinae) in
Costa Rica. A note on Masdevallia attenuata. Harvard Pap. Bot.
17(1): 25–38.
staden, r., d. P. Judge, and J. K. BonField. 2003. Analyzing
sequences using the Staden Package and EMBOSS. In s. a.
Krawetz and d. d. womBle, eds., Introduction to Bioinformatics.
A Theoretical and Practical Approach. Humana Press, Totawa,
New Jersey.
turland, n. J., J. h. wiersema, F. r. Barrie, w. greuter, d. l.
hawKsworth, P. s. herendeen, s. KnaPP, w.-h. KusBer, d.-z.
li, K. marhold, t. w. may, J. mcneill, a. m. monro, J. Prado,
m. J. Price, and g. F. smith, eds. 2018. International Code of
Nomenclature for Algae,Fungi, and Plants (Shenzhen Code)
Adopted by the Nineteenth International Botanical Congress
Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz
Botanical Books, Glashütten.
van den Berghe, e. P., and i. g. van den Berghe. 2008. Las
orquídeas de Nicaragua, una guía de campo (accessed July 17,
2019). http:// www.caftadr-environment.org/spanish/outreach/
publications/37_cafta_Orquideas_nal.pdf.
vásquez, r. and P. l. iBisch. 2000. Orchids of Bolivia, diversity
and conservation status, Subtribu Pleurothallidinae. Editorial
F.A.N., Santa Cruz de la Sierra, Bolivia.
williams, l. o. 1946. Orchidaceae. Pages 107–246 in w. g.
d’arcy, ed., Flora of Panama. Part 3. Fascicle 2. Ann. Missouri
Bot. Gard. 33: 1–140.
––––––. 1964. Tropical American plants. 6. Orchidaceae. Fieldiana,
Bot. 31(2): 17–44.
zelenKo, h., and P. Bermúdez. 2009. Orchids. Species of Peru.
ZAI Publications, Quito, Ecuador.
