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LANKESTERIANA 22(1): 37–51. 2022. doi: http://dx.doi.org/10.15517/lank.v22i1.50848
A NEW SPECIES AND A NEW RECORD OF MALAXIS (MALAXIDINAE)
FROM COSTA RICA
Isler F. ChInChIlla1,2,3,6, adam P. Karremans1,4,5 & marIo a. BlanCo1,3,4
1Jardín Botánico Lankester, Universidad de Costa Rica, P.O. Box 302–7050, Cartago, Costa Rica.
2Programa de Posgrado en Biología, Universidad de Costa Rica, Apdo. 11501–2060, San José, Costa Rica.
3Herbario Luis A. Fournier Origgi (USJ), Centro de Investigación en Biodiversidad y Ecología Tropical,
Universidad de Costa Rica, Apdo. 11501–2060, San José, Costa Rica.
4Escuela de Biología, Universidad de Costa Rica, Apdo. 11501–2060, San José, Costa Rica.
5Naturalis Biodiversity Center, Endless Forms Group, Sylviusweg 72, Leiden 2333 BE, The Netherlands.
6Author for correspondence: isler.chinchilla@ucr.ac.cr
ORCID of the Authors: IFC , APK , MAB
Received 14 December 2021; accepted for publication 17 April 2022. First published online: 25 April 2022.
Licensed under a Creative Commons Attribution-NonCommercial-No Derivs 3.0 Costa Rica License.
Introduction. Malaxis Sol. ex Sw. (Epidendroideae:
Malaxideae: Malaxidinae) in its traditional circumscrip-
tion is a cosmopolitan genus comprising ca. 300 species.
It has a broad pantropical distribution and few species in
the temperate regions of the Americas, Asia, and Europe
(Cribb 2005). However, preliminary molecular phylo-
genetic analyses of Malaxidinae suggest that Malaxis in
a broad sense is polyphyletic and that the genus in the
strict sense may rather be restricted to America (Cam-
eron 2005, Radins et al. 2014). In the Americas, there
are 143 species of Malaxis sensu lato currently recog-
nized; Mexico harbors the highest species diversity with
71 published species, 45 of them endemic, followed by
Costa Rica with 22 species, six of them endemic, and
then Panama with 17 species, two of them endemic
(Bernet et al. 2021, Chinchilla 2019, Chinchilla et al.
2020a, Dodson 2002, Dressler 2003, 2009, Espejo Ser-
na et al. 2002, Flora do Brasil 2021, González Tamayo
et al. 2008, Ulloa Ulloa et al. 2018 onwards, Villaseñor
2016). Nonetheless, as detailed taxonomic studies on
Malaxis are carried out in local and regional oras, the
species richness increases in countries where the genus
has previously been underestimated, such as Colombia,
Ecuador, and Peru (Chinchilla 2019).
In Costa Rica, species of Malaxis grow at eleva-
tions from 200 to 3650 m, in tropical wet to subalpine
aBstraCt. We describe and illustrate a new species and a new record of Malaxis from the lower montane and
montane forests of Costa Rica. Taxonomic descriptions, illustrations, distribution maps, and conservation
assessments are provided for each species. Malaxis excentrica sp. nov. is morphologically similar to Malaxis
simillima, from which it can be distinguished by having a single leaf per sympodial unit, shorter, obovate,
erect petals parallel to the column, the lip with triquetrous, acute, convex apical margin and the disc cavity
divided by a costa. Malaxis pittieri is formally recorded for the ora of Costa Rica; Costa Rican specimens
of this species were previously confused with Malaxis majanthemifolia. A lectotype for Microstylis pittieri is
formally designated.
resumen. Describimos e ilustramos una nueva especie y un nuevo registro de Malaxis de los bosques mon-
tano bajo y montano de Costa Rica. Se proporcionan descripciones taxonómicas, ilustraciones, mapas de
distribución y evaluaciones de conservación para cada especie. Malaxis excentrica sp. nov. se asemeja mor-
fológicamente a Malaxis simillima, de la cual se puede distinguir por tener una hoja por unidad simpodial, los
pétalos más cortos, obovados, erectos, paralelos a la columna, el labelo con el margen apical triquetro, agudo,
convexo y la cavidad del disco dividida por una costa. Malaxis pittieri se documenta formalmente para la
ora de Costa Rica; anteriormente, los ejemplares costarricenses de esta especie se confundían con Malaxis
majanthemifolia. Se designa formalmente un lectotipo para Microstylis pittieri.
Keywords/PalaBras Clave: Malaxis acianthoides, Malaxis excentrica, Malaxis fastigiata, Malaxis majanthe-
mifolia, Malaxis pittieri, Malaxis simillima
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LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
pluvial forests. However, the highest species diver-
sity is found between 1000 and 2800 m in the coun-
try (Chinchilla 2019, Chinchilla et al. 2020a). The
montane forests of the main mountain ranges of Costa
Rica, especially the Talamanca mountain range, harbor
the greatest diversity and endemism of Malaxis (Chin-
chilla 2019, Chinchilla et al. 2020a, Dressler 2003).
Despite that numerous botanical expeditions have
been carried out in different locations of Costa Rica,
there are still little explored areas where new or un-
recorded species of orchids may be harbored (Boga-
rín 2011, Bogarín et al. 2020, Chinchilla et al. 2020b,
Fernández et al. 2014, Karremans & Díaz-Morales
2016, Karremans et al. 2012). To update oristic in-
ventories and contribute to the knowledge of Malaxis,
we have carried out botanical explorations in these less
explored areas to collect, document, and identify their
species (Chinchilla 2019). In the Talamanca mountain
range, specimens of Malaxis were collected that did
not match any of the previously reported species in
the country. The examination of herbarium specimens
revealed they coincide with the Panamanian Malaxis
pittieri (Schltr.) Ames, which is here recorded for the
rst time in Costa Rica. Additionally, a Costa Rican
specimen of Malaxis kept at the United States National
Herbarium (US), does not match any of the species de-
scribed for the genus and is described here.
Materials and methods. This study was carried out
between 2015 and 2019. It is based on collections from
Costa Rica and Panama. Living plants of M. pittieri
collected in Costa Rica were photographed using a
Nikon® 7100 camera, and the herbarium specimens
were deposited at JBL and USJ. Sketches of M. pittieri
were prepared from fresh material, while sketches of
M. excentrica were prepared using rehydrated owers
from the holotype specimen. Both were digitized and
diagrammed in a composite plate using Adobe Photo-
shop CS6® and digitally delineated and shaded with an
Apple Pencil® in Procreate application for iPad Pro®
tablet (Apple Inc.). Specimens of Malaxis were exam-
ined in the following herbaria: AMES, CR, F, HLDG,
JBL, MO, SEL, US and USJ (physically), and B, BM,
BRIT, COLO, DAO, G, GH, GM, GOET, IBUG, K, L,
M, NY, P, PMA, UCH and W (through digital images).
A distribution map was prepared using the QGIS
3.8 Zanzibar program (QGIS Development Team
2019), based on satellite imagery updated to 2021
from Microsoft Bing Maps, Microsoft Corpora-
tion©, Earthstar Geographics SIO, 2021© TomTom.
The locations of the examined specimens were
classied according to the Holdridge life zone system
(Holdridge 1967). The phenology was determined
from examination of herbarium specimens and their
labels. The conservation status of each species was
assessed according to the methodology of the Inter-
national Union for Conservation of Nature (IUCN
2019). For M. pittieri, the Extent of Occurrence
(EOO) and Area of Occupancy (AOO) were estimat-
ed based on the studied specimens using geographical
data with the Geospatial Conservation Assessment
Tool (GeoCAT; Royal Botanic Gardens, Kew: http://
geocat.kew.org).
Taxonomic TreaTmenT
Malaxis excentrica Chinchilla, Karremans &
M.A.Blanco, sp. nov. (Fig. 1).
TYPE: Costa Rica. Puntarenas: cantón Coto Brus,
distrito Sabalito, Reserva de la Biosfera de La Amis-
tad, cerca Estación Biológica Las Alturas de Cotón,
08°57’00,3” N, 082°49’56,8” W, 2000 m, 08 julio
1994, W. J. Kress & J. L. Tuxill 94–4775 (holotype:
US!-3314101, barcode 00509021).
Diagnosis: Vegetatively similar to Malaxis simillima
(Rchb.f.) Kuntze, but differs in having one leaf (vs.
two leaves) per sympodial unit, shorter (1.0−1.2 mm
vs. 2.3−3.8 mm) petals that are obovate, erect, paral-
lel to the column (vs. linear, recurved, surrounding the
ovary), the lip with triquetrous, acute, convex (vs. tri-
d, concave) apical margin, and the disc cavity divided
(vs. non-divided) by a costa.
A terrestrial, erect, rhizomatous herb 25.3 cm tall,
including the inorescence. Roots exuous, short, up
to 4.5 cm long × 2 mm in diameter, pubescent. Pseu-
dobulbs 32 × 13 mm, epigeous, heteroblastic, conical,
separated by an ascending rhizome with two internodes
each subtended by one cataphyll, 6 cm long, when im-
mature covered with leaf sheaths becoming scarious
and papyraceous with age, and two ovate, obtuse, cata-
phylls 30–55 × 10–20 mm, the margins free or connate
LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
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ChinChilla et al. — Novelties in Malaxis from Costa Rica
Figure 1. Malaxis excentrica. A. Habit. B. Floral bract, adaxial view. C. Flower, oblique view. D. Dissected perianth (the lip
on the right), attened. E. Apex of ovary, petals and column, lateral view. F. Ovary, lip and column. G. Column, dorsal
(left) and ventral (right) views. H. Pollinia. Drawn from the holotype by Isler F. Chinchilla.
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Figure 2. Distribution of Malaxis excentrica and Malaxis pittieri, based on the specimens examined. Satellite image source:
Microsoft Bing Maps (2021). Prepared by Isler F. Chinchilla.
Figure 3. Flowers of Malaxis simillima. From left to right: front, oblique, lateral and back views (Chinchilla et al. 3183,
JBL-spirit). Scale bar = 3 mm. Photographs by Isler F. Chinchilla.
LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
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ChinChilla et al. — Novelties in Malaxis from Costa Rica
Figure 4. Malaxis pittieri. A. Habit. B. Floral bract, adaxial view. C. Flower, front view. D. Dissected perianth (the lip on
the right), attened. E. Ovary, lip and column. F. Column, dorsal (top) and ventral (bottom) views. G. Pollinia. Based
on Chinchilla 3340 (JBL-spirit). Illustration by Isler F. Chinchilla.
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LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
up to ¾ of their length, shedding as the pseudobulb
matures. Leaves one per sympodial unit, produced
from the base of the pseudobulb, long-petiolate, con-
duplicate, soft-textured; sheaths with the margins con-
nate for ¾ of their length, forming a pseudostem, 8.20
× 0.45 cm; the blade ovate, rounded, acute, the mar-
gins undulate, nine-veined, 12.2 × 6.2 cm, ascending,
green. Inorescence produced from the apex of the
pseudobulb, erect, corymbose, 22.5 cm long; peduncle
21.0 × 0.25 cm, covered near the proximal half with
leaf sheaths, seven-keeled; rachis 9.5 mm long. Floral
bracts ovate, acute to obtuse, concave, green, 1.2–2.5
× 0.6–1.2 mm. Pedicellate ovary terete, inconspicu-
ously keeled, with a 180-degree twist, 6–10 mm long.
Flowers non-resupinate, spreading, green. Dorsal se-
pal elliptic, obtuse, convex, adpressed to the ovary,
2.7–3.5 × 1.0–1.7 mm, three-veined. Lateral sepals
elliptic, obtuse, convex, divergent, margins recurved,
2.5–3.6 × 1.0–1.6 mm, connate at the base 0.5–0.8
mm, three-veined. Petals obovate, rounded, erect,
lightly arcuate, parallel to the column, one-veined,
1.0–1.2 × 0.4–0.5 mm. Lip simple, concave, triangular,
truncate, acute, papillose, eshy, margins entire, thick,
rounded, that apically converge into a triangular rim,
2.0–3.2 × 1.0–1.8 mm; the disc cavity ovate in out-
line, rounded, obtuse, 1.4–2.0 × 0.7–1.3 mm × 0.5–0.6
mm deep, divided by an obcuneate, compressed costa,
adaxially provided with a carina, subcavities narrowly
elliptic, apically delimited by a semilunate emergent
thickening; apical margin triquetrous, convex, porrect,
acute, 0.7–0.9 mm long. Column 0.75–1.0 × 0.6–0.7
mm, rectangular, dorsiventrally compressed, the apex
ventrally four-lobed; rostellum erect, adaxially slightly
convex, emarginate; stigma obcordate, concave, lon-
gitudinally bilobed, ca. 0.4 × 0.5 mm. Pollinia four in
two obovoid, ventrally concave hemipollinaria, 0.50–
0.55 × 0.2–0.25 mm. Fruits unknown.
eTymology: The specic epithet comes from the Latin
excentricus, meaning eccentric or out from the center,
in reference to the shape and position of petals, which
are atypical in Malaxis.
DisTribuTion anD habiTaT: Currently known only from
the type specimen collected at Las Alturas de Cotón
Biological Station in Las Tablas Protective Zone, on
the Pacic slope south of the Talamanca mountain
range in Costa Rica (Fig. 2). It grows in the lower
montane very wet forest at 2000 m in elevation.
Phenology: The only known specimen was collected
in ower in July.
Malaxis excentrica has obovate, erect petals, that
are parallel to the column and barely exceed the length
of the column. This unique feature distinguishes the
species from all other members of the genus. Further-
more, the position of the petals is reminiscent of those
of species belonging to the genus Hippeophyllum
Schltr., another genus of the Malaxidinae endemic to
Melanesia (Cribb 2005).
Due to the ascending rhizome, conical pseudo-
bulb, ovate leaf blade, and corymbose, compact ino-
rescence, M. excentrica is similar to M. simillima, but
the latter has up to three sympodial units with leaves
that are foveate, deeply lustrous, with undulate mar-
gins, the yellowish-green to yellow owers, the petals
incurved, arcuate, the lip deeply concave, with mar-
gins thin (vs. thick), incurved, apically with erect, thin,
diverging lateral teeth, the mid-tooth porrect, thin, at,
exceeding more than twice the length of the teeth lat-
eral, the column robust, eshy, the rostelum deeply
emarginate and the hemipollinaria narrowly (vs. ob-
ovoid) claviform (Fig. 3).
The dried owers of Malaxis excentrica may be
confused with those of Malaxis fastigiata (Rchb.f.)
Kuntze, a species ranging from Mexico and Guate-
mala. The latter can be distinguished from M. ex-
centrica by the longer (ca. 2.5 mm), linear, recurved
petals, the lip subcordate (vs. rounded) at the base,
with disc cavity occupying up to ½ (vs. more than ¾)
of its length, divided by a pandurate (vs. obcuneate)
costa, and the triangular, complanate (vs. triquetrous,
convex) apical margin.
conservaTion sTaTus: Malaxis excentrica is endemic to
Costa Rica, known from a single location in Las Tab-
las Protective Zone, Talamanca mountain range. For-
tunately, the forest where it was found is conserved.
Further studies on the ecology and population size of
this species are required to guarantee its long-term
conservation. Therefore, the species is listed as Least
Concern (LC), following the IUCN (2019: section
10.4) recommendations.
LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
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ChinChilla et al. — Novelties in Malaxis from Costa Rica
FIgure 5. Plants of Malaxis pittieri in situ. A–B. Chinchilla 3338. C. Chinchilla 3340. D. Chinchilla 3341. All the vouchers
at JBL-spirit. Photographs by Isler F. Chinchilla.
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Malaxis pittieri (Schltr.) Ames, Proc. Biol. Soc. Wash-
ington 35: 84. 1922. (Fig. 4, 5, 6, 7). Basionym: Mi-
crostylis pittieri Schltr., Repert. Spec. Nov. Regni Veg.
12: 203. 1913.
TYPE: Panama. Chiriqui: Feuchte Wälder zwischen
Alto de las Palmas und dem Gipfel des Cerro de
Horqueta, 2100−2268 m . d. M., blhend im [11]
März 1911, H. Pittier 3277 (holotype: B, destroyed
[tracing of Schlechter’s oral analysis of the holo-
type: AMES-24153/00101736 (Fig. 8A)]; isotypes:
AMES-00082841, a ower conserved in glycerine;
US-677667/00093447, designated here as lecto-
type (Fig. 8B) [drawings of the lectotype: AMES-
24152/00101737 (Fig. 8C)]; Schlechter’s oral analy-
sis from the holotype, reproduced in Mansfeld (1931:
t. 17, no. 61; Fig. 8D).
A terrestrial or epiphytic, erect, subcaespitose to
rhizomatous herb 4.5–20.0 cm tall, including the ino-
rescence. Roots exuous, short, up to 3.5 cm long × 2.0
mm in diameter, pubescent. Pseudobulbs 7−20 × 3−10
mm, epigeous, heteroblastic, conical, separated by an
ascending rhizome with two internodes, subtended by
one cataphyll, 0.3–6.0 cm long, pale greenish, when
immature, covered with leaf sheaths becoming scarious
and papyraceous with age, and one greenish, ovate, ob-
tuse cataphyll with inconspicuous tufts of hairs at the
base, 12–40 × 5–10 mm, the margins connate, shed-
ding as the pseudobulb matures. Leaves one per sym-
podial unit, produced from the base of the pseudobulb,
long-petiolate, conduplicate, soft-textured; sheaths
whitish-green, four-angled, the margins connate form-
ing a pseudostem, 3.5−9.5 × 0.15−0.30 cm; the blade
ovate, cordate-amplexicaul, acute, the margins undu-
late, ve-seven-veined, 1.7−4.6 × 1.8−3.8 cm, hori-
zontal, adaxially dark green, lustrous, abaxially light
green to whitish-green, dull. Inorescence produced
from the apex of the pseudobulb, erect, racemose, lax,
4–18 cm long; peduncle 3.5−13.5 × 0.1−0.2 cm, cov-
ered up to on lower 3/4 with leaf sheaths, sometimes
with a 180 degree twist, ve-seven-keeled; rachis
5–65 mm long. Floral bracts ovate, acuminate, pap-
illose, concave, greenish, pellucid, 0.5–1.5 × 0.7–1.0
mm. Pedicellate ovary terete, inconspicuously keeled,
with a 360-degree twist, 3–7 mm long, white, pellu-
cid. Flowers non-resupinate, spreading, lustrous, pel-
lucid; sepals and petals white, lip white with a light
to dark green stripe along the costa that extends to the
apex of the lip, the column greenish with the opercu-
lum and rostellum white; becoming orangish with age.
Dorsal sepal elliptic, obtuse, convex, adpressed to the
ovary, one-veined, 2.2–2.5 × 0.8–1.0 mm. Lateral se-
pals asymmetrically elliptic, acute to obtuse, convex,
convergent, the outer margins becoming recurved as
anthesis progresses, 1.8–2.3 × 0.7–1.0 mm, the inner
margins connate along 1.3−1.7 mm, one-veined. Petals
narrowly lanceolate, obtuse to acute, margins revolute,
recurved, arcuate, surrounding the ovary, one-veined,
2.0–2.3 × 0.3–0.5 mm. Lip trilobed, concave, sagittate,
papillose, membranaceus, margins entire, thin, 1.7–2.4
× 1.2–1.6 mm; lateral lobes triangular, obtuse to acute,
recurved, spreading, 0.5–0.8 × 0.4–0.5 mm; midlobe
triangular, 1.4–1.7 × 0.9–1.1 mm, with the disc cavity
ovate in outline, subcordate, obtuse, 0.7–1.0 × 0.4–0.8
mm × 0.3–0.5 mm deep, divided by a faint, compressed
costa, basally wider, subcavities narrowly lanceolate,
pellucid, apically delimited by a semilunate emergent
thickening, shortly raised into a rounded margin; api-
cal margin triangular, obtuse, concave, porrect, 0.4–0.6
mm long. Column 0.5–0.7 × 0.5–0.7 mm, rectangular,
dorsiventrally compressed, the apex ventrally four-
lobed; rostellum erect, truncate, bearing two viscaria
at the apex; stigma transversally oblong, subcordate,
concave, longitudinally bilobed, 0.15–0.25× 0.35–0.40
mm. Pollinia four in two obovoid, ventrally concave
hemipollinaria, 0.30–0.50 × 0.15–0.30 mm. Fruits el-
lipsoid capsules, 3.0–8.0 × 1.5–3.0 mm.
eTymology: The specic epithet honors the Swiss bot-
anist Dr. Henri François Pittier (1857–1950), who col-
lected the type specimen. Pittier was one of the most
inuential scientists for Costa Rican botany; his work,
leadership, and academic knowledge contributed to the
formation of what is now the National Herbarium of
Costa Rica and the development of scientic research
on the ora of the country.
DisTribuTion anD habiTaT: From Costa Rica and Pan-
ama, where it grows at 1500−3036 m of elevation.
Malaxis pittieri is recorded here for the rst time for
the ora of Costa Rica on the Pacic slope of the Cen-
LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
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ChinChilla et al. — Novelties in Malaxis from Costa Rica
Figure 6. Inorescences of Malaxis pittieri bearing recently opened owers, showing the lip with the entire apex. A. From
Costa Rica (Chinchilla et al. 2341, JBL-spirit). B. From Panama (Bogarín et al. 11179, UCH). Photographs by Isler F.
Chinchilla (A) and Diego Bogarín (B).
FIgure 7. Flowers of Malaxis pittieri. A–C. From Costa Rica: A. A recently opened ower (Chinchilla et al. 2341, JBL-
spirit). B–C. A ower in an early stage of dehydration showing the partially contracted lip (Chinchilla 3340, JBL-spirit).
D. From Panama: A dehydrated ower with a contracted lip that simulates being apically trilobed (Woodson & Schery
474, MO). Photographs by Isler. F. Chinchilla.
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LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
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ChinChilla et al. — Novelties in Malaxis from Costa Rica
tral Volcanic and Talamanca mountain ranges (García
Castro et al. 1993 included it in an unvouchered list of
orchid species common to Costa Rica and Panama, but
their report must be considered speculative). In Pana-
ma, M. pittieri it is known from Chiriqui province (Fig.
2). It inhabits in lower montane and montane pluvial
forests, in understory clearings, on very moist, mossy,
and well-drained soils. It usually grows as a terrestrial,
but sometimes as an epiphyte up to 1.5 m in height on
moss-covered tree trunks (Fig. 5).
Phenology: Plants ower from March, June to Sep-
tember and bear fruits from June to August and De-
cember in the eld.
oTher sPecimens sTuDieD: COSTA RICA. Heredia:
Parque Nacional Braulio Carrillo, Volcán Barva Sta-
tion, path and forest between park ofce and crater,
10°07’20” N, 84°06’00” W, 2700−2900 m, 13 Jul
1991, H. Gay et al. 1507 (CR); Barva, San José de la
Montaña, Área de Conservación Central, Parque Na-
cional Braulio Carrillo, sector Volcán Barva, cerca del
sendero que conduce a la laguna del Volcán Barva,
aprox. 230 m de la Estación del MINAE, 10°08’05” N,
84°07’10” W, 2711 m, 15 Dic 2018, I. Chinchilla et al.
4031 (JBL-spirit); Braulio Carrillo Nat. Park, canton of
Barva, Barva Station, in pastures around station, near
stream, 10°07’20” N, 84°06’00” W, 2580 m, 10 Sep
1990, S. Ingram et al. 507 (CR). Puntarenas: Buenos
Aires, Cuenca Térraba-Sierpe. Buenos Aires, Potrero
Grande, Tres Colinas, ca. 800 m, bajando hacia He-
lechales, 09°06’16,80” N, 83°03’53,60” W, 1600 m,
14 Jun 2006, D. Santamaría et al. 4504 (CR); Bue-
nos Aires, P. N. La Amistad. Cuenca Térraba-Sierpe,
Sabanas Esperanza, entre 1º y 3º Sabana, 09°04’33”
N, 83°01’55” W, 1600 a 1800 m, 01 Jun 2006, J. F.
Morales 13929 (CR). San José: Pérez Zeledón, Al-
bergue Cuericí, mirador trail above the Albergue
Cuericí, 09°33’30” N, 83°39’42” W, 2600 m, 23 Jun
2002, B. Boyle et al. 6331 (USJ); [Valle del El Gen-
eral, San Isidro] Vale Gen. S. Isidro, Jun, A. R. Endrés
s.n. (W-1889−0030705); Pérez Zeledón, Rivas, Parque
Nacional Chirripó, aprox. 200 m después del km 10, a
la orilla del sendero, 09°26’35,30” N, 83°31’48,20” W,
3021 m, 23 Jul 2015, I. Chinchilla et al. 2305 (USJ);
same data, I. Chinchilla et al. 2341 (JBL-spirit); same
data, I. Chinchilla et al. 2342 (JBL-spirit); same lo-
cality, aprox. 300 m después del kilómetro 10, a la
orilla del sendero, 09°26’33,67” N, 83°31’47,15” W,
3036 m, 13 Ago 2017, I. Chinchilla 3338 (JBL-spir-
it); same data, I. Chinchilla 3340 (JBL-spirit); same
data, I. Chinchilla 3341 (JBL-spirit); [Pérez Zeledón]
Fila Ventisqueros, Chirripó Gde., en el humus de la
montaña alta [09°27’52” N, 83°31’21” W], 2000–
3000 m, Ago 1971, L. D. Gómez-Pignataro & W. C.
Burger 3418 (CR). PANAMA. Chiriquí: Boquerón,
Cordillera, Paso Ancho, Aguacate, Finca Aguacatal,
camino a la naciente del Río Breque, 08°47’28,6” N,
82°34’39,7” W, 2126 m, 04 Ago 2014, D. Bogarín et
al. 11179 (UCH); Boquete, [08°48’18” N, 82°26’16”
W], 5000 ft [1524 m], Jun. 27, 1938, M. Davidson 801
(AMES, F, US); Potrero Muleto to summit, Volcán
de Chiriquí, [08°48’40” N, 82°31’35” W], 13−15 Jul
1940, R. E. Woodson & R. Schery 474 (MO); Vicini-
ty of Casita Alta, Volcán de Chiriqué [08°49’01” N,
82°29’35” W], ca. 1500−2000 m, 28 Jun-02 Jul 1938,
R. E. Woodson et al. 830 (AMES, BM, US).
Malaxis pittieri is distinguished by having a single
leaf per sympodial unit with the margins of the sheaths
connate, forming a pseudostem and the blade cordate-
amplexicaul at the base, the inorescence racemose,
lax, bearing small, white owers, the lateral sepals
connate for more than half their length forming a ga-
lea behind the lip, and the lip trilobed, sagittate, with a
green stripe along the costa that extends to the apex of
the lip. Historically, it has been confused with Malaxis
majanthemifolia Schltdl. & Cham., a species native
from Mexico to Costa Rica (Fig. 9). However, M. pit-
tieri differs in having white pedicels with a 360-degree
twist (vs. greenish, not twisted), the sepals, petals, and
lip white (vs. green to yellowish-green), the lip with a
Left: FIgure 8. Type material of the Microstylis pittieri. A. Tracing of the original drawings of the oral analysis of the
holotype of M. pittieri (AMES-24153 / 00101736), excluding the habit of a plant of a different species. Courtesy of the
Orchid Herbarium of Oakes Ames, Harvard University Herbaria. B. Lectotype of M. pittieri (US-677667 / 00093447).
Courtesy of the United States National Herbarium (US). C. Drawings of the lectotype of M. pittieri (AMES-24152 /
00101737). Courtesy of the Orchid Herbarium of Oakes Ames, Harvard University Herbaria. D. Schechter’s drawings
of the oral analysis from the holotype of M. pittieri, reproduced in Mansfeld (1931: t. 17, no. 61).
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LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
vertical green stripe (vs. dark brownish-green on the
distal half), the disc cavity divided (vs. non-divided)
by a costa into two subcavities, ending in a raised (vs.
non-raised), rounded, concave (vs. at) apical margin.
Malaxis pittieri is vegetatively similar to M. acian-
thoides (Schltr.) Ames from Mexico and Guatemala,
but the latter has green to yellowish-green owers, the
petals unguiculate, obliquely and transversely rhombic
(vs. simple, narrowly lanceolate), the lip with lateral
lobes of similar length to the mid lobe (vs. up to half
the length of the midlobe), with ciliate-papillose (vs.
entire) margins, and the disc cavity non-divided (vs. di-
vided by a costa). Malaxis cobanensis Archila, Szlach.
& Chiron, a Guatemalan endemic, is also vegetatively
similar. But the latter has green sepals, petals and lip,
the lip with disc cavity occupying up to ⅓ (vs. more
than ¾) of its length, non-divided (vs. divided by a
costa), and basally delimited by a transverse thicken-
ing (vs. not with a transverse thickening).
The rst collection of M. pittieri in Costa Rica was
made by the French botanist and orchidologist Au-
guste R. Endrés between 1866 and 1874 (Endrés s.n.,
W-1889−0030705!; see Ossenbach et al. 2010). Sub-
sequently, it has been collected multiple times in the
country, but specimens have been misidentied as M.
majanthemifolia and its presence has gone unnoticed
for over 147 years. For example, the specimen cited as
a Costa Rican voucher of M. majanthemifolia by both
Pupulin (2002) and Dressler (2003) (Ingram et al. 507,
CR) is a plant of M. pittieri.
Similarly, the specimens cited as a Panamanian
vouchers of M. majanthemifolia (see Bogarín et al.
FIgure 9. Plant habit (A) and inorescence (B) of Malaxis majanthemifolia in situ (J. F. Morales 21193, CR). Photographs
by Juan. F. Morales.
LANKESTERIANA 22(1). 2022. © Universidad de Costa Rica, 2022.
49
ChinChilla et al. — Novelties in Malaxis from Costa Rica
2014, Dressler 2009, Williams 1946) are indeed M.
pittieri. Malaxis majanthemifolia to date has not been
recorded in Panama.
nomenclaTural noTes: Schlechter (1913) described
Malaxis pittieri (as Microstylis pittieri Schltr.) with
the lip trilobed at the apex as shown in the drawings
of the type (Fig. 8C, D). The tracing of Schlechter’s
drawings of the holotype of M. pittieri at AMES
(24153/00101736; Fig. 8A) show a oral analysis that
coincides with the protologue of M. pittieri (and also
with the original drawings of Schlechter, published
posthumously by Mansfeld, 1931: t. 17, no. 61; Fig.
8D), but includes a plant habit drawing from a differ-
ent species of Malaxis, because it has two leaves in the
same sympodial unit, with opposite blades rounded at
the base, and a corymbose inorescence; inorescence;
Fig. 8A, center). This indicates a confusion with part of
the type material when tracing the ower analysis and
the plant habit of the holotype of M. pittieri in the Ber-
lin herbarium (many of the ower analyses and plant
habit of the types of orchid species that Schlechter de-
scribed were traced after their publication upon Ames’s
request, for his own herbarium; Ames 1944). Oakes
Ames, in a herbarium annotation on the specimen here
designated as lectotype (US-677667/00093447, visible
in the upper left corner in Fig. 8B) suggested that this
drawing by the morphology of the leaves and inores-
cence resembles Microstylis carpinterae Schltr. [Mal-
axis carpinterae (Schltr) Ames]. But it is impossible
to determine with certainty from this drawing alone.
However, M. pittieri consistently has one leaf cordate-
amplexicaul at the base, and the lip with the entire apex
(Fig. 6A–B); what happens is that when dehydrated,
the lip apex undergoes a strong contraction that cleaves
the apical margin, simulating a trilobed apex (Fig.
7A−D). The artifact described above was also veried
with fresh material collected near the type locality of
M. pittieri (D. Bogarín et al. 11179, UCH; Fig. 6B).
The isotype of the Microstylis pittieri designated
here as the lectotype was previously indicated by
Dressler (2009) as the holotype, and he also wrote it
in an annotation on the specimen sheet. However, the
holotype specimen that Schlechter used to describe M.
pittieri was destroyed in 1943 in the bombing of the
Berlin-Dahlem Botanical Museum (Ames 1944). In
his annotation on the lectotype specimen, Ames also
indicated that there was a duplicate in Berlin, which
was undoubtedly the holotype.
conservaTion sTaTus: Malaxis pittieri is a native of
Costa Rica and western Panama. It is known from 11
locations (six in Costa Rica, ve in Panama), in the
Central Volcanic (Costa Rica) and Talamanca moun-
tain ranges (Costa Rica and Panama). Its extent of oc-
currence (EOO) was estimated at 3562.297 km2 with
an area of occupancy (AOO) of 48 km2. This species is
known from ve protected areas: Braulio Carrillo Na-
tional Park and Chirripó National Park in Costa Rica;
Volcán Barú National Park in Panama; and Talamanca
Range-La Amistad Reserves / La Amistad National
Park in both Costa Rica and Panama. In Costa Rica,
ve locations are found in protected areas, and one
location is also in a forest near a protected area; simi-
larly, in Panama, four locations are in protected areas,
and one location near a protected area. Therefore, the
species is listed on a global level and national level as
Least Concern (LC).
acknowleDgmenTs. We thank the Vicerrectoría de Inves-
tigación of the Universidad de Costa Rica by its nancial
support to the research project 814–C1–055 Flora Costari-
censis: Taxonomía y logenia de la subtribu Malaxidinae
(Orchidaceae) en Costa Rica. To the Ministerio del Am-
biente y Energía of Costa Rica (MINAE) and the Sistema
Nacional de Áreas de Conservación (SINAC) for provi-
ding the scientic permits 138–16-ACLA-P and SINAC-
ACC-309–2018. To the staff and curators at the CR, JBL,
MO, and USJ herbaria, who allowed access to their equip-
ment, libraries, and collections, and the staff of AMES, F,
SEL, and US that loaned Mexican and Central American
specimens of Malaxis to MO for examination by the rst
author. To Juan F. Morales, Diego Bogarín, and José E. Ji-
ménez, for providing us with photographs of living plants.
The rst author thanks the Missouri Botanical Garden for a
Shirley A. Graham Research Fellowship in Systematic Bo-
tany and Biogeography, and its researchers Barry Hammel,
Michael Grayum, James Solomon, Mary Merello, Alba Ar-
belaez, John Pruski, Rosa Ortiz, Shirley Graham, and Alan
Graham. To Carlos O. Morales for the constructive sugge-
stions at the beginning of this investigation. We thank the
editors of Lankesteriana and the anonymous reviewers for
their constructive comments to improve the manuscript.
This contribution represents part of the rst author’s MSc
thesis, in the Programa de Posgrado en Biología of the Uni-
versidad de Costa Rica.
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