Available via license: CC BY 4.0
Content may be subject to copyright.
Webbia. Journal of Plant Taxonomy and Geography 76(1): 65-70, 2021
Firenze University Press
www.fupress.com/webbia
ISSN 0083-7792 (print) | ISSN 2169-4060 (online) | DOI: 10.36253/jopt-10029
WEBBIA
Journal of Plant
Taxonomy
and Geography
Citation: Fernández M., Bogarín D.,
Pupulin F. (2021) A new Muscarella (Orchi-
daceae: Pleurothallidinae) from Tapantí
National Park, Costa Rica. Webbia.
Journal of Plant Taxonomy and Geog-
raphy 76(1): 65-70. doi: 10.36253/jopt-
10029
Received: November 17, 2020
Accepted: January 3, 2021
Published: April 27, 2021
Copyright: © 2021 Fernández M., Bogarín
D., Pupulin F. This is an open access,
peer-reviewed article published by
Firenze University Press (http://www.
fupress.com/webbia) and distributed
under the terms of the Creative Com-
mons Attribution License, which per-
mits unrestricted use, distribution, and
reproduction in any medium, provided
the original author and source are
credited.
Data Availability Statement: All rel-
evant data are within the paper and its
Supporting Information les.
Competing Interests: The Author(s)
declare(s) no conict of interest.
Funding. This paper is part of the
project “Ecología y Biología de las
Orquídeas Epífitas y sus Micorrizas
en el Parque Nacional Tapantí, Cos-
ta Rica”, under number 814-B6-139,
supported by the Vice-Presidency of
Research, University of Costa Rica.
Editor: Riccardo M. Baldini
ORCID
DB: 0000-0002-8408-8841
A new Muscarella (Orchidaceae:
Pleurothallidinae) from Tapantí National Park,
Costa Rica
M F,,,*, D B,, F P,,
1 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Cos-
ta Rica
2 Department of Plant and Soil Science, Texas Tech University, Lubbock, TX 79409, USA
3 Herbarium UCH, Autonomous University of Chiriquí, Chiriquí, Panama
4 Harvard University Herbaria, Cambridge, Massachusetts, U.S.A.
5 e Marie Selby Botanical Gardens, Sarasota, Florida, U.S.A.
*Corresponding author. E-mail: melania.fernandez@ucr.ac.cr
Abstract. A new species of Muscarella from Costa Rica, is described and illustrated.
Muscarella tapantiensis most resembles M. coeloglossa but diers in the longer pedicels
(3-5 vs. 1-2 mm), the shorter (2.5-2.8 vs. 3.75 mm), connate (vs. free) lateral sepals,
and the lip with triangular (vs. broadly rounded) lower lateral lobes. Muscarella xan-
thella also resembles M. tapantiensis; however, it diers in the successively owered
raceme with alternate pedicels, the deeply mbriate petals and the thick, rounded lip
with densely verrucose lateral lobes. We provide illustrations, etymology, notes on
ecology, pictures of the plants and habitat, and a distribution map of the new species.
We discuss the inclusion of this species in the genus Muscarella.
Keywords: oristics, Muscarella coeloglossa, Muscarella xanthella, Neotropics, orchid
taxonomy, Specklinia.
INTRODUCTION
A fundamental principle of any study aimed at understanding the specif-
ic relationships among dierent organisms is to identify them correctly. is
simple rule becomes critical and unpractical when the research is carried out
in a tropical, biologically-rich region (Higgins and Ruokolainen 2004). Biol-
ogists esteem that, in the tropics, only about 86% of life diversity has been
apprehended by science, and the degree of the average knowledge may vary
among living groups (Mora et al. 2011).
With over 50,000 ha, Tapantí-Macizo de la Muerte National Park (TNP)
is one of the largest protected areas in Costa Rica (Sánchez 2002; Bernard et
al. 2009) and one of the species-rich spots for orchids in the country (Pupu-
lin 2003; Pupulin and Bogarín 2012; Crain and Fernández 2020). Because
of this high diversity and the ease to reach the park within less than two
hours from the capital city of San José, ongoing oristic, ecological and bio-
66 Melania Fernández, Diego Bogarín, Franco Pupulin
geographical projects have been carried out by Lankester
Botanical Garden and scientic partners for almost two
decades (Pupulin 2001; 2003, Pupulin et al. 2009; Pupu-
lin and Bogarín 2012). In 2015, a long-term research
project led by Dr. Jyotsna Sharma from Texas Tech Uni-
versity was initiated to understand if mycorrhizal fungi
distribution and specicity inuence the distribution of
epiphytic orchid species in a community. e massive
numbers of co-occurring species there (Figure 1) oer
a unique opportunity to test the limits of the species
co-existence hypothesis. As part of required taxonomic
identication of the epiphytic orchid species subjected to
fungal analyses in the study site, a species of Muscarella
Luer proved impossible to match with any previously
described taxon.
Muscarella is a Neotropical genus of the Pleurothal-
lidinae with around 50 species ranging from Mexico to
Brazil and the Antilles (Luer 2006; Pessoa et al. 2014).
e genus is distinguished by the cespitose plants with
loose racemes, f lowers with long-tailed sepals and
fringed, fimbriate or denticulate petals, and a thick,
eshy lip that is somehow ciliate (Luer 2006). Most spe-
cies bear longitudinal calli, and the column-foot is vari-
ously concave (Luer 2006). e species of Muscarella
were initially treated under Pleurothallis R.Br. and later
into a broad concept of Specklinia Lindl. (Pridgeon et
al. 2001). Lindley (1830) described Specklinia to include
minute, herbaceous plants with owers of gibbose sepals,
free petals and a labellum a third length of the sepals,
and membranaceous, winged columns. However, Garay
(1974) transferred it to the subgeneric level under Pleu-
rothallis. en, Luer (1986) built ten sections within
Pleurothallis to group species with similar oral features,
including a section of minute owers with thick, fringed
or verrucose, lobed lips, which he named Pleurothal-
lis section Muscariae Luer (1986). Later, Pridgeon et al.
(2001) proposed a broad concept of Specklinia, sister to
the Platystele-Scaphosepalum clade, for small plants with
an abbreviated stem with an annulus, owers with sepals
connate to dierent degrees, a lip hinged to the column
foot, and a column provided with a toothed apex and
ventral anther and stigma. ey based their ndings on
initial molecular phylogenies of the subtribe. e con-
cept included species of Pleurothallis sections Hymeno-
danthae Barbosa Rodrigues (1882), Muscariae Luer, and
Tribuloides Luer (1986) and species from Acostaea Sch lt r.
(Schlechter 1923), Empusella Luer (Luer 2004) and Pseu-
doctomeria Kraenzl. (Kraenzlin 1925).
Finally, Luer (2006) considered the new circumscrip-
tion of Specklinia as a polyphyletic aggregation of many
taxa; thus, he proposed the transfer of section Muscariae
(Luer 1986) to genus Muscarella. e genus is a mono-
phyletic and morphologically discernible group from the
rest of Specklinia, and it is supported by the most recent
molecular phylogenetic studies in the Pleurothallidinae
(Karremans et al. 2015; Pérez-Escobar et al. 2017)
Figure 1. Landscape at Tapantí National Park, along Rio Grande de Orosi, Cartago, Costa Rica, the type locality of M. tapantiensis. Photo-
graphs by F. Pupulin.
67
A new Muscarella (Orchidaceae: Pleurothallidinae) from Tapantí National Park, Costa Rica
Muscarella comprises 11 species in Costa Rica,
including the species proposed here as new (Luer 2006;
Ossenbach et al. 2007). We describe it hereaer.
MATERIALS AND METHODS
We collected living specimens at TNP, Cartago, Costa
Rica, and cultivated them in the collections at Lankester
Botanical Garden (JBL), University of Costa Rica. Liv-
ing plants were documented with sketches using a Leica®
MZ9.5 stereo-microscope with a drawing tube. e sketch-
es were digitalized, and the images were used for diagram-
ming a dra composite template in Adobe Photoshop® CC.
Photographs were taken with a Nikon D7100, Nikon Bel-
lows PB-6 extension, and a Nikon AF-D 50 mm f/1.8 lens.
Descriptions were prepared from both living specimens
and herbarium material deposited at JBL. Locality data
were obtained with a Garmin eTrex Vista GPS and sup-
porting maps and recorded along with phenology observa-
tions in a computerized database at JBL. Ecological zones
follow the Holdridge Life Zone System (Holdridge 1967,
Holdridge 1987). Distribution maps were made using the
geographic information system soware ArcView GIS 3.3
(ESRI, California, USA). Specimens were preserved in for-
maldehyde: acetic acid: ethanol [FAA (53% ethanol, 37%
water, 5% formaldehyde, and 5% glycerol)].
RESULTS
Muscarella tapantiensis Mel.Fernández & Bogarín, sp.
nov. (Figure 2.).
Type: COSTA RICA, Cartago: Paraíso, Orosi, Tapantí,
Parque Nacional Tapantí, bajando por el sendero Oro-
péndola, epítas a lo largo del borde del río Grande de
Orosi, 9°44’10.15”N 83°46’ 54.80”W, 1330 m, bosque plu-
vial premontano, 25 setiembre 2015, M. Fernández 1080,
L. Oses and A. Orellana (holotype JBL-spirit, J0159).
Diagnosis
Muscarella tapantiensis most resembles M. coeloglos-
sa (Luer and Hirtz) Luer but diers from it by the longer
pedicels (3-5 mm in M. tapantiensis vs. 1-2 mm), the
shorter (2.5-2.8 vs. 3.75 mm), connate (vs. free) lateral
sepals, and the lip with triangular (vs. broadly rounded)
lower lateral lobes.
Description
Plant epiphytic, caespitose, erect or subpendent, very
small, up to 1.8 cm tall. Roots slender, exuous, 0.5 mm
in diameter. Ramicauls slender, erect, 1.8–4.0 mm long,
enclosed by 2 thin, tubular sheaths becoming papyra-
ceous with age. Leaves erect, oblong-elliptic, subacute to
obtuse, thick, coriaceous, conduplicate, abaxially cune-
ate, 0.5–1.3 × 0.3–0.5 cm, the base narrowing into a pet-
iole 3-5 mm long. Inorescence an erect, loose, exuous,
successively owered raceme producing up to 6 owers,
to 3.5 cm long, as long as or exceeding the leaves, devel-
oped laterally from the ramicaul, peduncle liform, to
1.7 cm long, rachis to 2.3 cm long, oral bracts tubular,
acute, to 0.5 mm. Pedicel slender, persistent, to 3-5 mm
long. Ovary terete, widened apically, 0.5 mm long. Flow-
ers inconspicuous, less than 3 mm in diameter, sepals
and petals translucent yellow-greenish to pinkish, hya-
line, the petals with a longitudinal light pink midvein,
the yellow lip turning to light green apically, basally
stained with purple-red. Dorsal sepal elliptic, acute
to acuminate, conduplicate, concave, entire, carinate
beneath, briey connate to the lateral sepals, 2.2–2.5 ×
Figure 2. Illustration of Muscarella tapantiensis: A. Habit. B. Flower.
C. Column and lip, lateral view. D. Dissected perianth. E. Lip, at-
tened, and natural position. F. Anther cap and pollinia. Drawn by
M. Fernández based on M. Fernández 1080 (holotype).
68 Melania Fernández, Diego Bogarín, Franco Pupulin
0.8–1.0 cm. Lateral sepals ovate, slightly acuminate-sub-
falcate, concave, entire, carinate beneath, briey connate
at the base, 2.5–2.8 × 1.0-1.2 mm. Petals elliptic-ovate,
mbriate, aristate, 1.7–2.0 × 0.6–0.8 mm. Lip oblong,
rounded, 3-lobed, 1.5 × 1.0 mm when spread, verrucose
and papillose mostly at base, the lateral lobes triangular,
acute, minutely erose, erect in natural position, shorter
than the midlobe, the midlobe 0.5 mm wide, rounded,
with a longitudinal sulcate callus with two paralell light
keels, the apex slightly revolute, hinged to the column
foot. Column greenish, terete, widened apically, with
two small wings, arcuate, footed, the foot apex concave
with two verrucose, lateral thickenings, 1.2–1.3 × 0.4
mm. Anther and rostellum subapical, the stigma ventral.
Pollinarium with two pyriform pollinia, hooked towards
the base, 0.2 mm, lacking accessory structures. Cap-
sule globose, 6-ridged, ridges prominent, green, purple
between ridges, 3.5–4.0 × 3.0 mm. (Figure 2).
Etymology
Aer Tapantí National Park, the place where plants
of this species were rst collected.
Distribution and habitat
Only known from Costa Rica, growing on mossy
twigs, mostly of Saurauia montana (Actinidiaceae), in
open areas along or close to riparian premontane wet
and rain forests of the Caribbean watershed of the Tala-
manca mountain range (Figure 3).
Flowering time
Plants were seen in ower in the wild from Octo-
ber to April. At TNP, a owering peak occurs between
November and January.
Additional specimens examined (paratypes)
COSTA R ICA. Cartago: Paraíso, Orosi, Tapantí,
Parque Nacional Tapantí, Sendero Oropéndola, orillas
del Río Grande de Orosi, 9°44’37.86”N 83°47’04.28”W,
1296 m, epítas en árbol de Saurauia sp. (Actinidiaceae)
a orillas del río, bosque pluvial premontano, 30 octu-
bre 2011, D. Bogarín 9362 and M. Bonilla (JBL-spirit,
D4865) (Figure 4). Cartago: Turrialba, Tayutic, Platanil-
lo, siguiendo el camino paralelo al río Platanillo, que va
hacia el origen de este, unos 3.5 km del centro de Pla-
tanillo, sobre árboles solitarios en potreros, 9º47’52.90”N
83º33’44.51”W, 1049 m, 4 febrero 2012, A. Karremans
4934 (JBL-spirit, D5464).
DISCUSSION
e new species is recognized by the small, less
than 3 mm in diameter, translucent yellow-greenish
to pinkish owers with concave sepals, mbriate and
aristate petals, and a lip with two short lateral lobes,
characters that agree with the concept of Muscarella.
e new species most resembles the Ecuadorian Mus-
carella coeloglossa (Luer & Hirtz) Luer, but diers from
it by the longer pedicels (3-5 mm in M. tapantiensis
vs. 1-2 mm), the shorter (2.5-2.8 vs. 3.75 mm), con-
nate (vs. free), apically acuminate (vs. rounded) lateral
sepals, and the lip with triangular (vs. broadly round-
ed) lower lateral lobes. e Ecuadorian Muscarella xan-
thella (Luer) Luer also resembles M. tapantiensis, mostly
because of the more or less elliptic leaves and the small
pinkish owers with concave, translucid sepals. Never-
theless, Muscarella xanthella produces a fasciculous-
like inorescence with pedicels congested at the apex of
the peduncle, dierent from the successively owered
raceme with alternate pedicels of M. tapantiensis. When
looking closer to the ower, the lacerate petals of Mus-
carella xanthella dier from the deeply mbriate ones
of M. tapantiensis, and the lip is thick, rounded with
densely verrucose lateral lobes. In contrast, Muscarella
tapantiensis bears thinner, triangular lobes at the base
and is papillose mostly at the base.
Figure 3. Distribution of Muscarella tapantiensis in Costa Rica
(Central America), based on M. Fernández 1080 (holotype), D.
Bogarín 9362 & A. Karremans 4934 (paratypes). Only two marks
are shown since the holotype and one paratype were collected in
the same area.
69
A new Muscarella (Orchidaceae: Pleurothallidinae) from Tapantí National Park, Costa Rica
ACKNOWLEDGMENTS
To Dr. Jyotsna Sharma, for her support during the
Ph.D. studies of the rst author and initial comments
on this manuscript. To Adam Karremans for previous
comments on the phylogenetic position of Muscarella,
and to Lizbeth Oses and Andrea Morales for eld sup-
port. Meryll Arias, Fabricio Carbonell, Mahyar Shirazin-
ia, Lisandro Zúñiga, and all the sta at TNP for their
constant help with logistics and permits. anks to the
National System of Conservation Areas (SINAC) for the
collecting permits.
REFERENCES
Barbosa Rodrigues J. 1882. Genera et Species Orchide-
arum Novarum quas collegit, descripsit et iconibus
illustravit. II: 9-10. Sebastianopolis: Imprimerie de C.
& H. Fleiuss.
Bernard F, de Groot RS, Campos JJ. 2009. Valuation of
tropical forest services and mechanisms to nance
their conservation and sustainable use: A case study
of Tapantí National Park, Costa Rica. For Policy
Econ, 11, 174–183. URL:https://www.sciencedirect.
com/science/article/abs/pii/S1389934109000264
Crain BJ, Fernández M. 2020. Biogeographical analyses
to facilitate targeted conservation of orchid diversity
hotspots in Costa Rica. Diversity and Distribution.
26: 853–866. URL: https://onlinelibrary.wiley.com/
doi/full/10.1111/ddi.13062
Garay LA. 1974. Acostaea y los géneros del complejo
Pleurothallis. Orquideología. 9(2): 121–123.
Higgins MA, Ruokolainen, K. 2004. Rapid tropical for-
est inventory: a comparison of techniques based on
inventory data from western Amazonia. Conserva-
tion Biology. 18(3): 799–811. URL: https://www.jstor.
org/stable/3589090?seq=1
Holdridge LR. 1967. Life Zone Ecology. San José, Costa
Rica: Tropical Science Center.
Holdridge LR. 1987. Ecología basada en zonas de vida.
San José, Costa Rica: IICA.
Karremans AP, Albertazzi FJ, Bakker FT, Eurlings MCM,
Pridgeon A, Pupulin F, Gravendeel B. 2016. Phyloge-
Figure 4. Flowers of Muscarella tapantiensis from Tapantí National Park showing two variations of sepal color: A. Variation with reddish
sepals, from the holotype (M. Fernández 1080). B. Variation with yellow sepals, based on D. Bogarín 9362.
70 Melania Fernández, Diego Bogarín, Franco Pupulin
netic reassessment of Specklinia and its allied genera
in the Pleurothallidinae (Orchidaceae). Phytotaxa.
272: 1–36. URL: https://www.biotaxa.org/Phytotaxa/
article/view/phytotaxa.272.1.1
Kraenzlin FWL. 1925. New species of Masdevallia and
allied genera. Bull Misc Inform Kew. 3: 116.
Lindley J. 1830. e genera and species of orchidaceous
plants 8. London: J. Ridgway.
Luer CA. 1986. Icones Pleurothallidinarum III. Systemat-
ics of Pleurothallis (Orchidaceae). Monogr Syst Bot
Missouri Bot. 20: 1–89.
Luer CA. 2003. Pleurothallis. In: B.E. Hammel, M.H.
Grayum, C. Herrera, N. Zamora, editors. Manual de
Plantas de Costa Rica. Vol. III. Monogr Syst Bot Mis-
souri Bot. 95; p. 386–452.
Luer CA. 2004. Icones Pleurothallidinarum XXVI. Sys-
tematics of Pleurothallis subgenus Acianthera (Orchi-
daceae): a second century of new species of Stelis,
Epibator, Ophidion, Zootrophion. Monogr Syst Bot
Missouri Bot. 95: 1–258.
Luer CA. 2006. Icones Pleurothallidinarum XXVIII. A
reconsideration of Masdevallia. Systematics of Speck-
linia and vegetatively similar taxa (Orchidaceae).
Monogr Syst Bot Missouri Bot. 105: 1–94.
Mora C, Tittensor DP, Adl S, Simpson AGB, Worm B.
2011. How Many Species Are ere on Earth and in
the Ocean? PLoS Biol. 9(8): e1001127. URL: https://
journals.plos.org/plosbiology/article?id=10.1371/
journal.pbio.1001127
Ossenbach C, Pupulin F, Dressler RL. 2007. Orquídeas
del Istmo Centroamericano catálogo y estado de con-
servación. Orchids of the Central American Isthmus.
San José, Costa Rica: Editorial 25 de Mayo.
Pérez-Escobar OA, Chomicki G, Condamine FL, Karre-
mans AP, Bogarín D, NJ Matzke, Silvestro D, Antonelli
A. 2017. Recent origin and rapid speciation of Neo-
tropical orchids in the world’s richest plant biodiver-
sity hotspot. New Phytologist. 215(2): 891–905. URL:
https://nph.onlinelibrary.wiley.com/doi/full/10.1111/
nph.14629
Pessoa E, De Barros F, Alves M. 2014. Specklinia integrip-
etala and S. ianthina spp. nov. (Orchidaceae: Pleu-
rothallidinae) from northeastern Brazil. Nord J Bot.
32: 129–132. URL: https://onlinelibrary.wiley.com/
doi/abs/10.1111/j.1756-1051.2013.00183.x
Pridgeon AM, Solano R, Chase MW. 2001. Phylogenetic
relationships in Pleurothallidinae (Orchidaceae): com-
bined evidence from nuclear and plastid DNA sequenc-
es. American Journal of Botany. 88(12): 2286–2308.
URL: https://pubmed.ncbi.nlm.nih.gov/21669661/
Pupulin F. 2001. Extending the range - A new Dracula
from Costa Rica. Orchids. 70(6): 564-567.
Pupulin F. 2003. Tapantí - Orquídeas – Orchids. Cartago,
Costa Rica: Jardín Botánico Lankester, Universidad
de Costa Rica.
Pupulin F, Bogarín D, Jiménez D. 2009. New species and
records in Mesoamerican Lepanthes. Orch Dig. 73:
136–145.
Pupulin F, Bogarín D. 2012. Lepanthes novae tapantiensis.
Orch Dig. 76(1): 20–29.
Sánchez J. 2002. Aves del Parque Nacional Tapantí / Birds
of Tapantí National Park. Heredia, Costa Rica: Ed.
INBIO.
Schlechter FRR. 1923. Beiträge zur Orchideenkunde von
Zentralamerika. II: Additamenta ad Orchideologiam
Costaricensem. Repert Spec Nov Regni Veg. Beih 19:
22–23.