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Phytotaxa 607 (3): 161–181
https://www.mapress.com/pt/
Copyright © 2023 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Edlley M. Pessoa: 30 Jul. 2023; published: 9 Aug. 2023
https://doi.org/10.11646/phytotaxa.607.3.1
161
Five new species of Pleurothallis (Orchidaceae: Pleurothallidinae) in subsection
Macrophyllae-Fasciculatae from Southeastern Ecuador
MARCO M. JIMÉNEZ1,2,5,8*, LEISBERTH VÉLEZ-ABARCA2,5,9, VIVIANA MASHENDO-JIMBICTI2,10, HENRY
X. GARZÓN-SUÁREZ2,3,5,11, MARCO F. MONTEROS2,4,6,12 & MARK WILSON7,13
1 Grupo de Investigación en Medio Ambiente y Salud (BIOMAS), Ingeniería en Agroindustria, Facultad de Ingeniería y Ciencias
Agropecuarias, Universidad de Las Américas, Vía a Nayón, Quito 170124, Ecuador
2 Grupo Científico Calaway Dodson: Investigación y Conservación de Orquídeas del Ecuador, Quito, 170510, Pichincha, Ecuador
3 Herbario Azuay, Facultad de Ciencia y Tecnología, Universidad del Azuay, Av. 24 de Mayo 7-77 y Hernán Malo, Cuenca, Ecuador
4 Fundación EcoMinga, 270 12 de noviembre and Luis A Martínez, Baños, Tungurahua, Ecuador
5 Herbario HUTPL, Departamento de Ciencias Biológicas, Universidad Técnica Particular de Loja, San Cayetano Alto s/n 11-01-608,
Loja, Ecuador, Ecuador
6 Reserva: The Youth Land Trust, Washington, D.C., USA
7 Department of Organismal Biology and Ecology, Colorado College, Colorado Springs, CO 80903, USA
8
�
marco.jimenez.leon@udla.edu.ec; https://orcid.org/0000-0002-9502-5651
9
�
leis.alexis92@gmail.com; https://orcid.org/0000-0003-3764-9682
10
�
alexa.vivi@outlook.es; https://orcid.org/0000-0003-3857-9129
11
�
hg_palaco14@hotmail.com; https://orcid.org/0000-0003-4049-1652
12
�
marcomonteros24@gmail.com; https://orcid.org/0000-0002-6670-3687
13
�
mwilson@coloradocollege.edu; https://orcid.org/0000-0003-3044-7471
*Author for correspondence:
�
marco.jimenez.leon@udla.edu.ec
Abstract
Five new species of Pleurothallis subsect. Macrophyllae-Fasciculatae from Ecuador are described and illustrated: P. kashi-
menkakarai, P. lapoi, P. marioandresavilae, P. sabanillae and P. tinajillensis. The taxonomic similarities of the new entities
are discussed and information about their distribution, habitat, and conservation status is provided. Additionally, the first
known and confirmed locality for Pleurothallis saueri is provided, a previously described species without collection data, as
well as a lectotype is designated for this name.
Keywords: Amazon, new species, rainforest, Taxonomy
Introduction
The genus Pleurothallis Brown (1813: 211) is the fourth largest genus in the Pleurothallidinae, after Lepanthes Swartz
(1799a: 85), Stelis Swartz (1799b: 239) and Masdevallia Ruiz & Pavón (1794: 122) (Karremans 2016). The species
currently attributed to Pleurothallis are distributed from sea level to more than 3000 m in elevation (Pridgeon 2005),
from Central America and the Caribbean Islands to South America (Luer 1986).
Pridgeon (2005) indicated that members of Pleurothallis are recognized for the mostly epiphytic, caespitose to
repent plants, the stems are erect to rarely pendent, sometimes laterally or apically compressed, enclosed by tubular
sheaths, with or without a conspicuous annulus, the leaves are coriaceous, linear to ovate (rarely semi-terete). The
inflorescences are racemose, fasciculate or solitary-flowered. The flowers are resupinate or non-resupinate. The dorsal
sepal is generally free. The lateral sepals are connate to form a concave synsepal. The petals are linear to obovate,
acuminate to obtuse. The lip is simple or lobed, ligulate to obovate (rarely orbicular), often with a rounded callus
(‘glenion’) at the base and the column is terete or semiterete with the apical anther and two pollinia.
Ecuador is considered one of the most orchid diverse countries in the world, with 4032 species registered of which
1707 are endemic (Endara 2012). After the publication between 2000 and 2004 of the Native Ecuadorian Orchids series,
the most important and comprehensive treatment of the orchid family by Calaway Dodson (Endara 2012), the number
of new species discovered to date has increased the national inventory. Despite extensive taxonomic and collection
JIMÉNEZ ET AL.
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work in recent decades, Ecuador remains under-researched and only within the last 18 months, several species of
orchids were described as new, most of them described in Pleurothallidinae (e.g. Baquero et al. 2022, Doucette et al.
2023a, 2023b, Gutiérrez del Pozo et al. 2022, Monteros et al. 2022a, 2022b, Vélez-Abarca et al. 2022).
When conducting botanical exploration, collection and identification in recent years, we recognize the following
five new species of Pleurothallis subsect. Macrophyllae-Fasciculatae (Lindley 1859: 9), as additions for the orchid
flora of Ecuador which are illustrated and described here.
Materials and methods
All specimens were compared with original descriptions of similar species of Pleurothallis (Luer 1978, Luer 1998,
Luer 2005). Digital images of specimens of Pleurothallis fossulata Luer & Escobar (1998: 91–93), P. complanata
Luer & Hirtz (1996b:154–155), P. ruberrima Lindley (1846: 1) and P. lilijae Foldats (1968: 379) at MO and SEL were
examined online through Tropicos (https://tropicos.org/home) and GBIF (https://www.gbif.org) and for one species,
Pleurothallis fossulata, flowers in alcohol obtained from SEL (although Tropicos lists MO, Luer’s flowers in spirits
are currently housed at SEL).
The measurements of the vegetative and floral parts were made from living material. The fresh flowers were stored
in 70% ethanol and glycerol. Digital images were taken with Panasonic® FZ300, Canon® EOS 1100D and Nikon
D3100 cameras. The plants of the new species were collected under permits No. MAAE-ARSFC-2021-1619, and
MAATE-DBI-CM-2022-0248 granted by the Ministerio del Ambiente y Transición Ecológica de Ecuador (MAATE).
Taxonomy
1. Pleurothallis kashi-menkakarai Mashendo-Jimbicti, Vélez-Abarca & M.M.Jiménez, sp. nov. (Figures 1–2).
Type:—ECUADOR. Zamora Chinchipe: El Pangui, Cordillera del Cóndor flank, 3°38’32’’ S, 78°30’37” W, 1302 m,
23 October 2021, L. Vélez LV-0069 (holotype: HUTPL 14771!)
Similar to Pleurothallis fossulata Luer & Escobar from which it differs by its shorter, elliptic dorsal sepal (7.5–8
mm vs. 11 mm long, oblong), shorter, translucent green synsepal (6.5–7.0 mm vs. 10 mm long, red brown), crenulate,
verrucose petals, equal in length to the lip (vs. entire, glabrous, shorter than the lip), and involute, thickened, verrucose,
crenulate lip margins (vs. revolute, thin, smooth, entire).
Description:—Plant epiphytic, caespitose, to 30 cm tall. Roots slender, white, flexuous, 1 mm in diameter.
Ramicauls erect, green, 8.9–20.1 long and 0.1 cm in diameter enclosed by two papyraceous sheaths at the base and
one below the middle. Leaves 6.6–11.0 × 2.1–4.5 cm, sessile, coriaceous, erect to suberect, ovate, long acuminate,
shiny, cordate at the base, with faintly raised lateral nerves, margin revolute, purplish. Inflorescence single-flowered,
resupinate, produced in a successive fascicle from a reclining spathaceous bract to 1.4 cm long; peduncle abbreviate
ca. 1 mm long, concealed within a spathaceous bract, floral bract tubular ca. 4 mm long, pedicel cylindrical, verrucose,
straight, 3.9–5.0 mm long. Sepals glandulous-cellular, translucent yellow to reddish brown with red-brown and yellow
veins, rarely in almost purely translucent yellow; dorsal sepal 7.5–8 × 4.5–5.0 mm, broadly elliptic, obtuse, concave,
3-veined; lateral sepals connate into a broadly ovate, subacute, synsepal, 6.5–7.0 × 4.5–5.0 mm, 4-veined, slightly
concave at the base; Petals 4.5–5.0 × 0.9–1.0 mm, narrowly oblong, slightly falcate at the base, acute, inconspicuously
1-veined, papillose-verrucose, reddish brown to red purple, rarely tan yellow, margins yellow and crenulate-erose. Lip
5 × 2.5–2.7 mm, shortly unguiculate, oblong-ovate, obtuse and incurved at the apex, slightly convex above the middle,
3-veined abaxially, papillose-verrucose, reddish brown to red-purple, rarely tan yellow, margins crenulate, thickened,
purple; the base concave, hinged to the column-foot; a small, narrowly elliptic glenion in the middle. Column purple,
stout, complanate, papillose, 2.0 mm long, 1.5–1.8 mm wide, 2-channeled dorsally, widening to the base, column-foot
glandulous, paler. Anther narrowly obovate, purplish with a touch of yellow, papillose, stigma apical, bilobed, 0.7 mm
long. Pollinia 2, clavate, 0.6 mm long. Ovary cylindrical, subverrucose, brownish purple, slightly curved at the joint
with the pedicel, 3.5–4.6 mm long, 0.8 mm wide.
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 163
FIGURE 1. Pleurothallis kashi-menkakarai A. Habit. B. Flower. C. Dissected perianth. D. Ovary, column and lip in lateral view. E. Lip
adaxial and abaxial views. F. Column in lateral view. Illustration by Vélez-Abarca based on the type.
Distribution and ecology:—Pleurothallis kashi-menkakarai is known from two locations within the Cordillera
del Cóndor in south-eastern Ecuador (Figure 3). The species is found around 1400 m in elevation in lower montane
forests near watercourses of sandstone plateaus. The holotype of this species was found growing sympatrically with
Maxillaria grayi Dodson (1994a: 69), P. cordata (Ruiz & Pavón 1798: 234) Lindley (1830: 5) and P. adeleae Luer
(1981: 200), between the lower and middle strata of the forest. This forest is characterized by a mixture of Andean and
lowland trees of the Amazon, where the canopy can grow up to 30 m in height and the understory is dense. It was also
found growing together with Dichaea histrio Reichenbach (1859: 330), Epilyna embreei Dodson (1994b: 146) and
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Maxillaria bicallosa (Reichenbach 1878: 9) Garay (1962: 527), over the roots of Socratea exorrhiza (Martius 1824:
36) Wendland (1860: 103) palms, in a nearby location. This species has been only recorded in the province of Zamora
Chinchipe and is considered as endemic.
FIGURE 2. Floral comparison of A. Pleurothallis kashi-menkakarai and B. P. fossulata C. Different color forms of Pleurothallis kashi-
menkakarai D. Column with ovary in lateral view (uppermost), lip in adaxial view (left) and lateral view (right) of Pleurothallis kashi-
menkarai. Elaborated by L. Vélez-Abarca from photos: A, C (left) and D by M.M. Jiménez based on the type and M. Jiménez 1190B by
Mark Wilson based on a cultivated plant, C (right) by L. Vélez-Abarca based on M. Jiménez 1191.
Etymology:—From the words kashi (night), and menkakarai (lost), meaning “lost at night” in the Shuar language,
because the new species was discovered at night when researchers were lost in the forest of a Shuar community.
Taxonomic discussion:—Pleurothallis kashi-menkakarai is similar to Pleurothallis angusta Ames & C.
Schweinfurth (1925: 23–24) from Costa Rica and P. fossulata from Colombia with which it shares flowers with a
maroon-burgundy colored lip, a triangular, cream-colored area between the apex and edges of the lip and glenion and
two calli flanking the deep glenion. The new species is distinguished from Pleurothallis fossulata (Figure 2) by its
smaller plants (30 cm vs. 42 cm tall), the narrower (3.0–4.5 cm vs. 5–7 cm wide), cordate, attenuate leaves (vs. deeply
cordate, acuminate), the shorter pedicel (3.9–5.0 mm vs. 10 mm long), the shorter ovary (3.5–4.6 vs. 6 mm long), the
one flower produced per fascicle (vs. to 2–3 often simultaneous), the non-dilated petals (vs. dilated at the base), the
subacute synsepal (vs. obtuse) and the strongly incurved apex of the lip (vs. convex).
It is also similar to Pleurothallis angusta but is distinguished by the smaller plants (30 cm vs. 50 cm tall), the
shorter (6.6–11.0 cm vs. 8.0–16.5 cm wide), ovate, long-acuminate leaves (vs. lanceolate, acute), the shorter pedicel
(3.9–5.0 mm vs. 14 mm long), the shorter ovary (3.5–4.6 mm vs. 5.0–7.0 mm long), the translucent yellow or reddish
brown flowers (vs. dark purple), the shorter (8 mm vs. 11 mm long), broadly elliptic, obtuse dorsal sepal (vs. ovate,
acute), the 4-veined synsepal (vs. 5-veined), the smaller (4.5–5.0 × 0.9–1.0 mm vs. 6.3–7.0 × 1.5–2.0 mm), narrowly
oblong, acute petals (vs. falcate, obtuse), the lip with crenulate margins (vs. minutely ciliate) and the narrowly obovate
anther cap (vs. obtriangular) (Pupulin 2021).
Conservation status:—Pleurothallis kashi-menkakarai has not been reported within the Ecuadorian National
System of Protected Areas. The species is known from two abundant populations, where a population density of 3 to
4 plants per phorophyte can be found. However, the range in which the species occurs is under threat by gold mining
and grazing since P. kashi-menkakarai has only been observed within an area of 9 km2 (Figure 3).
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 165
FIGURE 3. Distribution map of Pleurothallis kashi-menkakarai (black circle), P. lapoi (black square), P. marioandresavilae (black star),
P. sabanillae (black cross), P. saueri (also black square), P. tinajillensis (black diamond) in Ecuador and mining threats (red squares). Map
by Henry X. Garzón.
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166 • Phytotaxa 607 (3) © 2023 Magnolia Press
2. Pleurothallis lapoi M.M.Jiménez & Vélez-Abarca, sp. nov. (Figures. 4–5)
Type:—ECUADOR. Zamora Chinchipe: Near El Pangui, 781253.08 E, 9597652.43 S, 1560 m, M. Jiménez 1424
(holotype: HUTPL 14640!)
FIGURE 4. Pleurothallis lapoi. A. Habit. B. Flower. C. Dissected perianth. D. Ovary, column and lip in lateral view. E. Lip adaxial and
abaxial views. Illustration by L. Vélez-Abarca based on the type.
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 167
FIGURE 5. Floral structure comparison of Pleurothallis lapoi (above) and P. erythrium (down). A–D. Flower, lip and sepals of P. lapoi
based on the type. E–G. Flower, lip and sepals of P. erythrium based on M. Jiménez 318. Elaborated by L. Vélez-Abarca from photos by
Marco M. Jiménez.
Similar to Pleurothallis erythrium Luer (1978: 386) from which differs by its ovate-triangular dorsal sepal (vs.
oblong-ovate), the oblong-triangular petals (vs. narrowly linear-oblong) and the yellow, triangular-ovate and shallow
lip (vs. red-purple, oblong-ovate, rounded at the apex and convex) with two calli (vs. concave at the end).
Description:—Plant epiphytic, up to 19 cm tall, roots slender, white, flexuous, 0.6–1.0 mm in diameter.
Ramicauls slender, suberect, terete, 14.0–16.6 long and 0.1 cm in diameter, enclosed by papyraceous, tubular sheaths,
upper sheath 24.6–31.0 mm long, basal sheath 12.6–15.7 mm long. Leaves green, slightly falcate to falcate, ovate,
shortly attenuate, glossy, centrally channeled, 5.1–6.3 × 2.1–2.6 cm, base cordate, margins revolute. Inflorescence a
fascicle of successive flowers produced from a reclined, papyraceous, spathaceous bract, 6.6–8.0 mm long; peduncle
6.3–6.8 mm long, floral bract infundibuliform, 3.6–4.0 mm long; pedicel verrucose, dilated at the junction with
the ovary, 6.2 mm long. Flowers glossy, sepals papillose, subverrucose below the apical third. Dorsal sepal purple,
ovate-triangular, 3-veined, 6.1–6.6 × 3.2 mm, acute, shallow, margins slightly involute, callose at the base. Lateral
sepals united into an ovate, purple infused with yellow to the apex and veins, 4-veined, shallow, acute, 5.5–5.7 × 3.7
mm synsepal, apex minutely incurved. Petals purple with yellow borders, oblong-triangular, deflexed, lanceolate, 1-
veined, microscopically papillose-verruculose, acute, carinate abaxially at the midvein, 3.8–3.9 × 0.7–0.8 mm, margin
minutely erose. Lip dark yellow, tan and purple to the apex, triangular-ovate, 3-veined, shallow, verrucose-papillose,
2.6–2.9 × 2.0 mm, with a Y-shaped cream-colored depression and two triangular calli around the glenion that reaches
up to the apical third, margins minutely erose, incurved, base cordate with a small, ovate glenion, apex acute, center of
the base of the lip with two calli. Column stout, pale green suffused with purple dorsally and to the stigmatic margin,
complanate, papillose, 1.2–1.6 × 1.1 mm, stigma reniform, rostellar flap minute, 0.1 mm long, column foot 0.6 mm
long, with longer papillae. Anther broadly ovate, microscopically papillose, orange-purple, 0.5 mm long. Pollinarium
with 2, ovoid, yellow pollinia, 0.4 mm long. Ovary green-purple, lightly verrucose, subterete, slightly arcuate, 3.8–4.0
× 0.9 mm.
Distribution and ecology:—Pleurothallis lapoi was found in southeastern Ecuador growing in an evergreen
lower montane forest of the eastern Andes of southern Ecuador, Zamora Chinchipe province, El Pangui canton (Figure
3). Populations have been observed within the Cayamatza micro-basin at 1560 m elevation. This area is characterized
JIMÉNEZ ET AL.
168 • Phytotaxa 607 (3) © 2023 Magnolia Press
by very steep slopes with rapidly accelerating deforestation due to expansion of the agricultural frontier. These forests
are dominated by Erythrina amazonica Krukoff (1939: 270) (Fabaceae), Cedrelinga cateniformis (Ducke 1915: 17)
Ducke (1922: 70) (Fabaceae), Jacaranda copaia (Aublet 1775: 650) Don (1823: 267) (Bignoniaceae) and Inga sp.
(Fabaceae) trees. In this forest it is found on tree trunks and branches of the lower canopy stratum, together with other
orchid species such as Masdevallia don-quijote Luer & Andreetta (1985: 63), Myoxanthus exasperatus (Lindley 1859:
15) Luer (1982: 36), Pleurothallis saueri (Luer 2012: 355) Shaw (2016: 39) and Stelis spp.
Etymology:—Named after Lester Lapo, outstanding orchid grower from El Pangui, Zamora Chinchipe Province,
who discovered the species.
Taxonomic discussion:—The new species is distinguished from other species of Pleurothallis subsect.
Macrophyllae-Fasciculatae by its small purple flowers, acute sepals, and verrucose-papillose, triangular-ovate, yellow
lip. It is most similar to Pleurothallis erythrium. Pleurothallis lapoi can be distinguished from P. erythrium by its larger
plants (up to 19 vs. 12 cm tall), convex leaves with revolute margins (vs. concave with flat margins) and the flowers
with acute, verrucose sepals and lip (vs. papillose, subacute).
Conservation status:—Pleurothallis lapoi has not been found in any formally protected area in Ecuador. So far,
no more than five individuals have been sighted within the Cayamatza micro-basin. Deforestation is ongoing in the
area for the establishment of cattle pastures and subsistence crops by local residents.
3. Pleurothallis marioandresavilae Vélez-Abarca & M.M.Jiménez sp. nov. (Figures 6–8).
Type:—ECUADOR. Zamora Chinchipe: Next to the El Quimi reserve, 820 m, 19 December 2021, L. Vélez LV-0072
(holotype: ECUAMZ 08648!)
Similar to Pleurothallis complanata Luer & Hirtz from differs by its yellowish-brown, verrucose, acute dorsal
sepal (vs. light yellow green, glabrous, obtuse to rounded at the apex), 4–veined, elliptical-oblong, obtuse lateral
sepals (vs. faintly 6–veined, ovate, acute) and smaller lip (2.1–2.3 × 1.6–1.8 mm vs. 4.0 × 2.5 mm), elliptical, obtuse,
verrucose-striated (vs. oblong-ovate, obtuse, verrucose), and a glenion bilobed (vs. obovoid).
Description:—Plant epiphytic, up to 26 cm tall, erect; roots slender, flexuous, 0.5–0.7 mm in diameter. Ramicauls
very slender, suberect, terete, 12.5–23.3 long and 0.1 cm in diameter, enclosed by papyraceous, tubular sheaths, upper
sheath 2.8 mm long, basal sheath 1.25 mm long. Leaves erect, green suffused with purple at the margins and the
underside, slightly convex, coriaceous, microscopically papillate, ovate, channeled in the middle, 5.5–7.4 × 1.5–1.8
cm, acuminate, slightly falcate at the apex, the base sessile, cordate. Inflorescence a fascicle of successive flowers
produced from a suberect, spathaceous bract, ca. 0.9 mm long; floral bract infundibuliform, 3 mm long; pedicel
tubular, 2.5 mm long. Flowers glossy, dorsal sepal yellowish-brown, slightly suffused with purple at the base, edges
and veins, elliptical-obovate, 7.0–8.5 × 3.8–4.0 mm, 3-veined, subverrucose, convex in the apical third; lateral sepals
connate into a 4-veined, elliptical-oblong, obtuse, minutely bifid pale purple with yellowish green base, 4.6–5.0 ×
3.7–4.0 mm. Petals pale purple apex and base yellowish green, narrowly oblong, slightly constricted in the first apical
third, acute, 3.8–4.0 × 0.6–0.8 mm, 1-veined, papillose, margins entire. Lip red purple combined with orange in the
base and margins, elliptical, obtuse, verrucose-striated, convex, 2.1–2.3 x 1.6–1.8 mm, 3-veined, margin irregular; the
base truncate, centrally channeled towards the apex of the lip, hinged to the column foot. Column yellowish green,
transversely subrectangular, dorsally complanate, 1.3–1.5 × 1.0–1.2 mm, glabrous, stout, 3-ribbed dorsally, stigma
transversely bilobed with entire margins, rostellar flap 0.8 mm long. Anther cordiform, yellowish green, 0.3 mm long.
Pollinarium with 2 obovoid, yellow pollinia, 0.2 mm long. Ovary subverrucose, curved in the middle, 2.0–2.3 × 1.0
mm.
Distribution and ecology:—The new species has been recorded in Zamora Chinchipe, Ecuador. It apparently
also occurs in Piura department, northern Peru, where it was previously recorded photographically by Benavente
et al. (2020) without any herbarium voucher, as Pleurothallis compressa Luer (1996a: 75), a Costa Rican species
with different habit and flower morphology. The species has been found growing as an epiphyte on Pouteria caimito
(Ruiz & Pavón 1802: 18) Radlkofer (1882: 333) (Sapotaceae) (Figure 9) and Andesanthus lepidotus (Bonpland 1808:
38) Guimarães & Michelangeli (2019: 948) (Melastomataceae). The population of Pleurothallis marioandresavilae
occurs in unprotected areas where selective timber extraction is common. The specimens were found growing together
with other small orchids such as Brachionidium sp., Macroclinium sp., Pleurothallis ariana-dayanae Vélez-Abarca,
Jiménez & Gutiérrez del Pozo (2022: 109), Maxillaria splendens Poeppig & Endlicher (1836: 66) and Stelis spp.
Additional specimens examined:—ECUADOR. Zamora Chinchipe: San Andrés, close to a river, 2075 m, 29
November 2022, M. Jiménez & G. Iturralde 1619 (HUTPL 14756!).
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 169
FIGURE 6. Pleurothallis marioandresavilae. A. Habit. B. Flower. C. Dissected perianth. D. Ovary, column and lip in lateral view. E. Lip
adaxial and abaxial views. Illustration by L. Vélez-Abarca based on the type.
Etymology:—Named after Mario Andrés Ávila, professor at the Universidad Estatal Amazónica, in Ecuador, in
gratitude for his unconditional help and educational support.
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FIGURE 7. A. Pleurothallis marioandresavilae in situ attached to a Pouteria caimito tree. B. Leisberth Vélez-Abarca, taking in situ
photograph. Elaborated by L. Vélez-Abarca from photos of the type by A. by L. Vélez-Abarca and B. by Jimmy Zari.
FIGURE 8. Pleurothallis marioandresavilae. A. Close-up of the lip and apex of the synsepal. B. Lateral view of the ovary, column, and
lip. C. Floral comparison of P. marioandresavilae. D. Flower of P. complanata. Elaborated by L. Vélez-Abarca from photos by A, B, C
(left) by Marco M. Jiménez based on the type D. (right) by Ulli Lorimer based on a cultivated plant.
Taxonomic discussion:—Pleurothallis marioandresavilae (Figure 10C) has been confused in cultivated
specimens with P. complanata, (Figure 10D) which is the most similar species. But the new species differs by shorter
leaves (5.5–7.4 × 1.5–1.8 cm vs. 8.5–10.5 × 2.5–3), shorter pedicel (2.5 vs. 6.0–7.0 mm long), acute, yellowish-brown,
verrucose dorsal sepal (vs. obtuse to rounded, yellow-green, glabrous), 4-veined, elliptical-oblong, obtuse lateral sepals
(vs. 6–veined, ovate, acute), narrowly oblong petals (vs. narrowly linear-triangular), elliptical, verrucose-striated lip
(vs. oblong-ovate, minutely verrucose), and bilobed glenion (vs. obovoid) (Figure 10).
Conservation status:—Pleurothallis marioandresavilae is known from only two localities, one near the protected
forests of the El Quimi Biological Reserve, the other near the border with Peru, both in the Province of Zamora
Chinchipe, in the southeast of Ecuador (Fig. 3), and apparently it also occurs in northern Peru. Both localities have
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 171
been influenced by the strong anthropogenic activities (mining, logging and cattle grazing); however, it is not ruled out
that the species may be present within the aforementioned reserve.
4. Pleurothallis sabanillae M.M.Jiménez & Vélez-Abarca, sp. nov. (Figures 9–10).
Type:—ECUADOR. Zamora Chinchipe: Loja-Zamora road, near El Tambo, 3°56’40.08’’ S, 79°02’45.00’’ W, 2102 m,
4 August 2022, M. Jiménez & L. Vélez-Abarca 1253 (holotype: HUTPL 14636!).
FIGURE 9. Pleurothallis sabanillae. A. Habit. B. Flower. C. Dissected perianth. D. Ovary, column and lip in lateral view. E. Lip adaxial
and abaxial views. Illustration by L. Vélez-Abarca based on the type.
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FIGURE 10. Photographs of Pleurothallis lilijae A. and B. P. sabanillae. Elaborated by L. Vélez-Abarca from photos by A. Sebastián
Moreno and B. Marco M. Jiménez based on the type.
Similar to Pleurothallis lilijae Foldats (1968: 379), from which it differs by its broadly ovate, 7–veined dorsal
sepal (vs. broadly elliptic, 5–veined), ovate, 3-veined petals (vs. triangular ovate, 1-veined), the obtuse, convex lip (vs.
triangular, concave) with a deep, broad glenion and an apical sulcus (vs. shallowly concave, small and no sulcus).
Description:—Plant epiphytic, up to 36 cm tall, descendent to pendent; roots slender, flexuous, 0.6–1.3 mm in
diameter. Ramicauls slender, suberect, terete, 11.1–19.4 cm long and 0.1 cm in diameter, enclosed by papyraceous,
tubular sheaths, upper sheath 12.9–18.9 mm long, basal sheath 6.2–7.7 mm long. Leaves deflexed, coriaceous,
microscopically papillate, lanceolate to narrowly ovate, channeled in the middle, 2.4–3.7 × 12.0–14.5 cm, acuminate at
the apex, the base sessile, deeply cordate with the lobes unequal up to 1.6 cm long. Inflorescence a fascicle of successive
flowers produced from a suberect, scabrous spathaceous bract, 1.1–1.3 mm long; floral bract infundibuliform, 4.4–
4.6 mm long; pedicel subflexuous, 2.4–3.5 mm long. Flowers resupinate or not, glossy, dorsal sepal purple with
yellow borders to pale yellow suffused with purple at the base and veins, broadly ovate, subacute to obtuse, strongly
deflexed at the apex, 9.8–11.5 × 8.9–9.4 mm, 7-veined, subverrucose-papillose, minutely pubescent, convex below the
middle; lateral sepals connate into a 6-veined, elliptical-oblong, subverrucose-papillose, strongly revolute margins,
purple with yellow borders synsepal, 6.1–6.5 × 2.9–4.6 mm. Petals purple with yellow borders, ovate above the third
quarter, oblique, narrowed below the middle, shortly attenuate, strongly deflexed near the apex, acute, 8.1–10.1 ×
4.1–5.0 mm, 3-veined, pubescent-verruculose, margins ciliate-denticulate, revolute. Lip dark purple with paler or
whitish border, broadly ovate, obtuse, diffusely glandulous-verrucose, concave, 2.7–2.9 × 3.3–3.5, 3-veined, margins
thickened, involute apically; the base subtruncate, hinged to the column foot, with very short obtuse angles and a large,
deep, obovoid glenion, occupying the middle third of the lip, concave and deeply sulcate towards the apex of the lip.
Column dark purple, transversely subrectangular, dorsally complanate, 2.8 × 3.0–3.2 mm, papillose, glandulous on the
edges, stout, 3-ribbed dorsally, stigma transversely bilobed with minutely ciliate margins, rostellar flap 0.4 mm long.
Anther narrowly triangular-obovate, white to pale yellow, papillose, 1.1 mm long, bilobed at the base. Pollinarium
with 2 obovoid, yellow pollinia, 0.9 mm long, brought together by a drop-like viscidium. Capsules 1.6 × 0.3 cm. Ovary
subverrucose-glandulous, straight, 2.6–3.0 × 1.0 mm.
Distribution and ecology:—Up to now Pleurothallis sabanillae is only known from the steep hillsides near
the village of Sabanilla in the Zamora Chinchipe province of Ecuador (Figure 3). The species is found growing as an
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 173
epiphyte on understory lianas and tree trunks, together with other orchid species such as Elleanthus vernicosus Garay
(1978: 105), Octomeria colombiana Schlechter (1920: 121), O. hirtzii Luer (2002: 29), Pleurothallis adonis Luer
(1976: 40) and P. omoglossa Luer (1976: 152). The upper montane forests of this region are dominated by Purdiaea
nutans Planchon (1846: 251), Myrica pubescens Humboldt & Bonpland ex Willdenow (1806: 746), Myrsine andina
(Mez 1902: 378) Pipoly (1992: 7) and pioneer species such as Andesanthus lepidotus and Graffenrieda harlingii
Wurdack (1976: 7) (Bussman 2005).
Etymology:—Named after Sabanilla, known also as El Tambo, an eminent farming village in the Zamora
Chinchipe Province of Ecuador close to the type locality.
Taxonomic discussion:—Pleurothallis sabanillae belongs to the Pleurothallis cardiostola-lilijae complex, a
group within the subsection Macrophyllae-Fasciculatae according to Luer (2005) and Wilson et al. (2022). The new
species is most similar to Pleurothallis lilijae, but is distinguished by the shorter plants (up to 36 cm vs. 45 cm tall) and
flowers (1.1 × 1.6 vs. 1.6 × 2.2 cm), the purple flowers (vs. pale yellowish brown), the strongly deflexed dorsal sepal
(vs. erect), strongly recurved petals (vs. slightly deflexed), the broadly ovate lip (vs. ovate-triangular) and the purple
column (vs. purple and white).
Conservation status:—To date this species is known from one locality with a few dozens of individuals we
recommend to consider it as a restricted endemic. Pleurothallis sabanillae is present near forests threatened by human
activities (frequent forest fires, works close to high voltage electrical lines, livestock breeding and deforestation).
Fortunately, this species is expected to be found within the Podocarpus National Park, where its conservation would be
assured.
5. Pleurothallis tinajillensis M.M.Jiménez, H. Garzón & Vélez-Abarca, sp. nov. (Figures 11–12)
Type:—ECUADOR. Morona Santiago: Municipal Conservation Ecological Area Tinajillas–Río Gualaceño, between
Limón and Gualaceo, 3°00’45.10’’ S, 78°31’29.06” W, 26 May 2022, 2336 m, M. Jiménez & H. Garzón 1410 (holotype:
HUTPL 14634!).
Similar to Pleurothallis ruberrima, from which it differs by its lanceolate, attenuate leaves (vs. narrowly ovate,
acute), flowers lying on the leaf surface (vs. dangling over the sides of the leaf), elliptic-ovate dorsal sepal (vs. narrowly
ovate), the base of the petals unlobed, separated from each other (vs. with an obtuse lobe, very close to each other), the
lip triangular-ovate, sulcate without a callus (vs. broadly ovate, convex with a rounded callus).
Description:—Plant epiphytic, up to 23 cm tall, roots slender, flexuous, 0.6–0.9 mm in diameter. Ramicauls
slender, suberect, terete, 13.5–21.6 long and 1.3–1.6 mm in diameter, enclosed by papyraceous, tubular sheaths, upper
sheath 33.4–37.2 mm long, basal sheath 22.6–24.8 mm long. Leaves green adaxially, suffused with purple abaxially,
lanceolate, attenuate, dull, centrally channeled, 9.0–13.0 × 2.5–3.2 cm, concave with slightly raised lateral nerves,
base cordate, margins purple. Inflorescence a fascicle of successive flowers produced from a reclined, papyraceous,
spathaceous bract, 14.0–17.7 mm long; peduncle short, 3.6 mm long, floral bract infundibuliform, 8.4 mm long;
pedicel very long, verrucose, dilated at the junction with the ovary, 12.1 mm long. Flowers glossy, delicate, lying
against the leaf, sepals and petals pale garnet green toward the ends, with purple minute pubescence. Dorsal sepal
elliptic-ovate, 7-veined, 8.0 × 11.7 mm, convex, margins revolute. Lateral sepals united into an ovate, 6-veined,
slightly concave, obtuse, 14.0 × 8.2 mm synsepal. Petals narrowly oblong, slightly falcate, lanceolate, 3-veined, ribbed
at the midvein, 10.1–10.4 × 1.9–2.0 mm, base thickened, whitish, verrucose, recurved, separated at the upper margin
from the other petal, apex acute, minutely incurved. Lip greenish white, tan and purple to the apex, triangular-ovate,
3-veined, verrucose-papillose, 4.3 × 3.4 mm, adaxially sulcate in the middle with two lateral depressions at each side,
margins minutely erose, base truncate with a small glenion, apex apiculate, center of the base of the lip without a
callus. Column stout, white, complanate, papillose, 3.0 mm long including the rostellar flap, 1.7 × 1.2 mm, apex flared,
thickened. Anther narrowly obovate, whitish brown, papillose, 1.1 mm long, bilobed at the base, pollinia 2, yellow,
clavate, 1.1 mm long. Ovary subverrucose, subclavate, almost straight, 5.1 × 1.3 mm.
Distribution and ecology:—Up to now Pleurothallis tinajillensis is only known from the type locality on the
steep slopes into the AECMTRG in the Morona Santiago Province (Figure 3). The species grows at the foot of the hills
in the montane rainforest at around 2300 m in elevation, on the Eastern slope of the Cordillera Oriental of the Andes. It
grows on tree trunks, together with other orchid species such as Elleanthus vernicosus Garay (1978: 105), Maxillaria
meridensis Lindley (1846: 19), Oncidium cruentoides Chase & Williams (2008: 24), Pleurothallis eccentrica Luer &
Hirtz (1988: 140) and Pleurothallis amplectens Luer (1980: 73). The local vegetation is characterized by Cecropia
andina (Cuatrecasas 1945: 286), Oreopanax andreanus Marchal (1880: 90), Andesanthus lepidotus and Weinmannia
fagaroides Kunth (1823: 54–55).
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FIGURE 11. Pleurothallis tinajillensis. A. Habit. B. Flower. C. Dissected perianth. D. Ovary, column and lip in lateral view. E. Lip
adaxial and abaxial views. Illustration by L. Vélez-Abarca based on the type.
Etymology:—Named after Tinajillas, which is the Municipal Conservation Ecological Area Tinajillas–Río
Gualaceño (AECMTRG), where the type locality is found. It is an important local protected area in southeastern
Ecuador, home to a considerable area of natural vegetation that includes montane and premontane forest ecosystems.
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 175
FIGURE 12. Flower and frontal view of the lip. A. Pleurothallis ruberrima B. P. tinajillensis. Close-up of the lips and columns. C.
Pleurothallis ruberrima. D. P. tinajillensis. Elaborated by Henry Garzón from photos by A. Henry X. Garzón based on a in situ specimen
and B. Marco M. Jiménez based on the type.
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FIGURE 13. Lectotype of Lindleyalis saueri Luer. Reproduced in Harvard Papers in Botany 17:333–368. 2012. C. Luer illustr. 21701.
Fig. 27.
Taxonomic discussion:—Pleurothallis tinajillensis is most similar to P. ruberrima but it is distinguished by its
smaller plants, up to 23 cm tall (vs. 35 cm), concave, centrally channeled, suffused with purple at the abaxial surface
of the leaf (vs. convex, sulcate, green at the abaxial surface of the leaf ), shorter pedicel (12 mm vs. 20–60 mm
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 177
long), minutely pubescent sepals and petals (vs. glabrous), shorter dorsal sepal (11.7 vs. 15–23 mm long), extended
horizontally petals (vs. reflexed), pulvinate, apiculate, basally truncate lip (vs. convex, broadly obtuse to rounded,
basally subcordate), and whitish, complanate column (vs. green and white or purple, ancipitous) with a longer rostellum
(1.2 mm long vs. 0.4 mm).
FIGURE 14. Pleurothallis saueri. A. Habit. B. Flower. C. Perianth dissected and close-up to margin of the petal. D. Ovary, column,
and lip, lateral view and close-up of the margin of the lateral lobe and the midlobe. E. Lip adaxial and abaxial views with the basal lobes
expanded. F. Column in lateral, ¾, and frontal views, with a close-up to the papillae of the clinandrium. G. Anther cap adaxial and abaxial
views and pollinia. LCDP by L. Vélez-Abarca elaborated with photographs by Marco M. Jiménez from L. Vélez 0073.
JIMÉNEZ ET AL.
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Conservation status:—This species has not been reported within the Ecuadorian National System of Protected
Areas but the type locality is located within the AECMTRG, an area of ~32,928 hectares protected since 2002.
Pleurothallis tinajillensis appears to have a very restricted distribution and may be endemic to the province of Morona
Santiago. Additional data is required about distribution and abundance of this species before a valid conservation
assessment can be made.
Taxonomic notes on Pleurothallis saueri (Luer) J.M.S. Shaw
Pleurothallis saueri was originally described as Lindleyalis saueri Luer in Luer & Thoerle (2012: 355) from a wild
specimen from Ecuador but without a known locality, which was imported by Tropical Orchid Farm and grown
by E. Sauer in the United States (Luer & Thoerle (2012). It is now confirmed that this species grows in Zamora
Chinchipe Province, southeast Ecuador, in the same location of P. lapoi (Figure 3). A specimen of P. saueri was
found, photographed, and collected without flowers in the confirmed locality in November 2020, it was grown at the
Orquideario La Paphinia, where it bloomed 18 months later (Figure 14).
6. Pleurothallis saueri (Luer) J.M.S. Shaw (2016: 39).
Basionym: Lindleyalis saueri Luer (2012: 355).
Type:—ECUADOR. Without collection data, obtained from Tropical Orchid Farm in Hawaii, flowered in cultivation
in Centerville, OH, December 2011, E. Sauer s.n. (holotype: MO, not found, lectotype here designated), C. Luer illustr.
21701, in Harvard Papers in Botany 17: 333–368. 2012. (Fig. 13). Epitype (designated here):—ECUADOR. Zamora
Chinchipe: cerca de El Pangui, 1560 m, 17 January 2022, L. Vélez-Abarca, LV 0073 (ECUAMZ!) (Figure 14).
While searching for the original material of the species, the type material was sought at Missouri Botanical
Garden (MO) to verify if the specimen is deposited there. It was known that the holotype of Pleurothallis saueri is not
found at the herbarium, therefore, based on the original illustration a lectotype is selected here.
Acknowledgments
We acknowledge Luis Ocupa Horna for his review of the previous draft of this manuscript and Gabriel A. Iturralde
for his help with this investigation. MMJ thanks Universidad de Las Américas (UDLA) for funding orchid research in
Ecuador, grant No. AGR.LBR.22.03. We would like to thank the Universidad Técnica Particular de Loja herbarium
(HUTPL) for their support in the development of this research. To the Ministerio del Ambiente, Agua y Transición
Ecológica (MAATE) of Ecuador for granting the research permits No. MAAE-ARSFC-2021-1619 and MAATE-
DBI-CM-2022-0248. We thank Sebastián Moreno and Ulli Lorimer for their photos used in this article. To Mary
Merello, Assistant Curator of the MO, for the information about Pleurothallis saueri. The authors also acknowledge
the reviewers of this manuscript for helping with comments and corrections of this work.
References
Aublet, J.B.C.F. (1775) Histoire des Plantes de la Guiane Françoise 2: 650–653.
Ames, O. & Schweinfurth, C. (1925) New or noteworthy species of orchids from the American tropics. Schedulae Orchidianae 8: 1–91.
Baquero, R.L.E., Iturralde, G. & Martel, C. (2022) Telipogon crisariasae (Orchidaceae) a new species from northern Ecuador. Phytotaxa
564: 248–256.
https://doi.org/10.11646/phytotaxa.564.2.8
Benavente, L., Ocupa-Horna, L., Ugaz, A., Charcape-Ravelo, M. & Saldaña, I.S. (2020) Orquídeas CITES del Caserío El Hormiguero,
distrito de El Carmen de la Frontera, provincia de Huancabamba, región Piura, Noroeste del Perú. Arnaldoa 27: e1–e12.
Bonpland, A.J.A. (1808) Monographie des Melastomatacées. Rhexies. Monographia Melastomacearum 2: 38.
Brown, R. (1813) Clasis XX. Gynandria-Monandria. Hortus Kewensis; or, a Catalogue of the Plants Cultivated in the Royal Botanic
Garden at Kew 5: 188–220.
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 179
Bussman, R.W. (2005) Bosques andinos del sur de Ecuador, clasificación, regeneración y uso. Revista Peruana de Biología 12: 203–
216.
https://doi.org/10.15381/rpb.v12i2.2394
Chase, M.W., Williams, N.H., Neubig, K.M. & Whitten, W.M. (2008) Taxonomic transfers in Oncidiinae to accord with Genera
Orchidacearum, Volume 5. Orchids 77: 27.
Cuatrecasas, J. (1945) Notas a la Flora de Colombia—VII. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales
6: 286.
Dodson, C.H. (1994a) New orchid species and combinations from Ecuador 3. Orquideología 19: 51–96.
Dodson, C.H. (1994b) New orchid species and combinations from Ecuador 2. Orquideología 19: 123–149.
Don, D. (1823) Description of five new genera of plants, belonging to the natural order Bignoniaceae. Edinburgh Philosophical Journal
9: 267.
Doucette, A., Wilson, M., Portilla, J., Kay, A., Moreno, J.S. & Cameron, K.M. (2016) Dos especies nuevas de Pleurothallis y un nuevo
nombre para Acronia rinkei. Orquideología 33: 135–139.
Doucette, A., Medina, H. & Portilla, J. (2023a) New Records of Lepanthes Sw. from Ecuador. Part 1: Three new species. Orchids 92:
294–299.
Doucette, A., Medina, H. & Portilla, J. (2023b) New Records of Lepanthes Sw. from Ecuador. Part 2: Two new species, two new records
and a heterotypic synonym. Orchids 92: 372–378.
Ducke, W.A. (1915) Plantes nouvelles ou peu connues de la région amazonienne (Avec 19 planches). Archivos do Jardim Botânico do Rio
de Janeiro 1: 17.
Ducke, W.A. (1922) Plantes nouvelles ou peu connues de la région amazonienne (IIe Partie). Archivos do Jardim Botânico do Rio de
Janeiro 3: 70
Endara, L. (2012) Orchidaceae. En: León-Yánez, S., Valencia, R., Pitman, N., Endara, L., Ulloa Ulloa, C. & Navarrete, H. (Eds.) Libro
Rojo de Plantas Endémicas del Ecuador. Publicaciones del Herbario QCA, Pontificia Universidad Católica del Ecuador, Quito.
Foldats, E. (1968) Contribución a la orquideoflora de Venezuela IV. Acta Botánica Venezuélica 3: 305–426.
Garay, L.A. (1962) Studies in American Orchids, V. Caldasia: 517–529.
Garay, L.A. (1978) Orchidaceae (Cypripedioideae, Orchidoideae, Neottioideae). In: Harling, G. & Sparre, B. (Eds.) Flora of Ecuador, vol.
9. University of Göteborg and Swedish Museum of Natural History, Göteborg and Stockholm, pp. 105.
Guimarães, P.J.F., Michelangeli, F.A., Sosa, K. & de Santiago Gómez, J.R. (2019) Systematics of Tibouchina and allies (Melastomataceae:
Melastomateae): A new taxonomic classification. Taxon 68: 937–1002.
https://doi.org/10.1002/tax.12151
Gutiérrez del Pozo, D., Jiménez, M.M., Vélez-Abarca, L. & Baquero, L.E. (2022) A new Masdevallia (Orchidaceae: Pleurothallidinae)
from the Ecuadorian Amazon foothills of the Llanganates mountains. Phytotaxa 522: 191–200.
https://doi.org/10.11646/phytotaxa.552.3.3
Karremans, A. (2016) Genera Pleurothallidinarum: An updated phylogenetic overview of Pleurothallidinae. Lankesteriana 16: 219–241.
https://doi.org/10.15517/lank.v16i2.26008
Krukoff, B.A. (1939) The American species of Erythrina. Brittonia 3: 205–337.
https://doi.org/10.2307/2804812
Humboldt, A, Bonpland, A.J. & Kunth, K. (1823) Nova Genera et Species Plantarum (quarto ed.), vol. 6. Paris, Sumtibus Libreriae
Graeco-Latino-Germanicae, 600 pp.
Lindley, J. (1830) Pleurothallis cordata. In: The Genera and Species of Orchideaceous Plants. London, Ridgways, pp. 5.
https://doi.org/10.5962/bhl.title.120492
Lindley, J. (1846) Orchidaceae Lindenianae. pp. 1–19.
Lindley, J. (1859) Folia Orchidacea, an enumeration of the known species of orchids, Pleurothallis, part 2. J. Matthews, London.
Luer, C.A. (1976) Miscellaneous species of Pleurothallis. Selbyana 1: 38–201.
Luer, C.A. (1978) Icones Pleurothallidinarum (Orchidaceae). Miscellaneous new species and combinations in the Pleurothallidinae.
Selbyana 2: 367–390.
Luer, C.A. (1981) Miscellaneous new species and combinations in the Pleurothallidinae (Orchidaceae). Phytologia 49: 197–240.
https://doi.org/10.5962/bhl.part.15137
Luer, C.A. (1982) A reevaluation of the genus Myoxanthus (Orchidaceae). Selbyana 7: 34–54.
https://doi.org/10.5962/bhl.part.14425
Luer, C.A. (1985) Miscellaneous new species in the Pleurothallidinae (Orchidaceae). Phytologia 57: 63.
Luer, C.A. (1986) Icones Pleurothallidinarum III. Systematics of Pleurothallis. Monographs in Systematic Botany from the Missouri
Botanical Garden 20: 1–109.
https://doi.org/10.5962/bhl.title.149317
JIMÉNEZ ET AL.
180 • Phytotaxa 607 (3) © 2023 Magnolia Press
Luer, C.A. (1988) A revision of some sections of subgenus Pleurothallis. Lindleyana 3: 133–149.
Luer, C.A. (1996a) New species of Pleurothallis (Orchidaceae) from Ecuador. Lindleyana 11: 54–113.
Luer, C.A. (1996b) New species in the Pleurothallidinae from Costa Rica. Lindleyana 11: 141–197.
Luer, C.A. & Escobar, R.R. (1998) Nuevas especies de Pleurothallis de Colombia. Orquideología 21: 72–106.
Luer, C.A. (2002) Miscellaneous new species in the Pleurothallidinae (Orchidaceae). Selbyana 23: 1–45.
https://doi.org/10.3100/hpib.v23iss1.2018.n6
Luer, C.A. (2005) Icones Pleurothallidinarum XXVII: Dryadella and Acronia section Macrophyllae Fasciculatae. Monographs in
Systematic Botany from Missouri Botanical Garden 103: 1–311.
Luer, C.A & Thoerle, L. (2012) Miscellaneous new species in the Pleurothallidinae (Orchidaceae). Harvard Papers in Botany 17: 355.
https://doi.org/10.3100/025.017.0214
Marchal, E. (1880) Notice sur les Hédéracées récoltées by Élie Marchal. Bulletin de la Société Royale de Botanique de Belgique 19: 90.
Martius, C.F.P. (1824) 1. Iriartea exorhiza Tab. 33. 34. Historia Naturalis Palmarum 2: 36–37.
Mez, C.C. (1902) Myrsinaceae. In: Engler, H.G.A. Das Pflanzenreich IV. no. 236. pp. 378.
Monteros, M.F., Mogrovejo, A.L., Iturralde, G.A. & Baquero, L.E. (2022) Masdevallia purocafeana, a new species of Orchidaceae from
Cordillera del Toisán, northwestern Ecuador. Phytotaxa 547: 223–231.
https://doi.org/10.11646/phytotaxa.547.3.1
Monteros, M.F., Restrepo, E. & Baquero, L.E. (2022) Platystele finleyae (Pleurothallidinae), a new species from northwestern Ecuador.
Lankesteriana 22: 63–72.
https://doi.org/10.15517/lank.v22i1.50869
Pipoly, J.J. (1992) Estudios en el género Myrsine (Myrsinaceae) de Colombia. Caldasia 17: 7.
Planchon, J.E. (1846) Voisin du Cliftonia. London Journal of Botany 5: 251.
Poeppig, E.F. & Endlicher, S.F.L. (1835) Maxillaria. Nova Genera ac Species Plantarum 1: 38.
Pridgeon, A., Solano, R. & Chase, M.W. (2001) Phylogenetic relationships in Pleurothallidinae (Orchidaceae): combined evidence from
nuclear and plastid DNA sequences. American Journal of Botany 88: 2286–2308.
https://doi.org/10.2307/3558390
Pridgeon, A.M. (2005) Subtribe Pleurothallidinae. In: Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (Eds.) Genera
Orchidacearum. Volume 4 Epidendroideae (Part One). Oxford University, Oxford, pp. 319–422.
Pupulin, F., Aguilar, J., Belfort-Oconitrillo, N., Díaz-Morales, M. & Bogarín, D. (2021) Florae Costaricensis subtribui Pleurothallidinis
(Orchidaceae) Prodromus II. Systematics of the Pleurothallis cardiothallis and P. phyllocardia groups, and other related groups of
Pleurothallis with large vegetative habit. Harvard Papers in Botany 26: 203–295.
https://doi.org/10.3100/hpib.v26iss1.2021.n14
Raldkofer, L.A.T. (1882) Sitzungsberichte der Mathematisch-Physikalischen Classe (Klasse) der K. B. Akademie der Wissenschaften zu
München 12: 333.
Reichenbach, H.G. (1859) Nederlandsch Kruidkundig Archief. Verslangen en Mededelingen der Nederlandsche Botanische Vereeniging
4: 330.
Reichenbach, H.G. (1878) 36. Zygopetalum (Kefersteinia) bicallosum. Otia Botanica Hamburgensia 1: 9.
Ruiz, H. & Pavón, J. (1798) Systema vegetabilium Florae Peruvianae et Chilensis, characteres prodromi genericos differentiales,
specierum omnium differentias, durationem, loca natalia, tempus florendi, nomina vernacula, vires etusus nonnullis illustrationibus
interspersis complectens. Typis Gabrielis de Sancha. Madrid.
https://doi.org/10.5962/bhl.title.887
Ruiz, H.L. & Pavón, J.A. (1802) Flora Peruviana, et Chilensis, sive descripciones, et ícones plantarum peruvianum, et chilensium,
secundum systema Linnaeanum digesta, cum haracteribus luriumgenerum evulgatorum reformati 3. Madrid, España, 95 pp.
Schlechter, F.R.R. (1920) Die Orchideenfloren der südamerikanischen Kordillerenstaaten: II. Colombia. Repertorium Specierum Novarum
Regni Vegetabilis, Beihefte 7: 121.
Shaw, J.M.H. (2016) Registrar’s notes on names in the register. Orchid Review Supplement 124: 39.
Swartz, O.P. (1799a) Dianome epidendri generis. Linn. Nova Acta Regiae Societatis Scientiarum Upsaliensis 6: 61–88.
Swartz, O. (1799b) Dianome epidendri generis Linn. Journal für die Botanik 2: 201–244.
Vélez-Abarca, L., Jiménez, M.M., Gutiérrez del Pozo, D. & Baquero, L.E. (2022) Pleurothallis ariana-dayanae, a new species in
subsection Macrophyllae-Fasciculatae (Pleurothallidinae) from the Cordillera del Cóndor, Ecuador. Lankesteriana 22: 101–109.
https://doi.org/10.15517/lank.v22i2.51750
Wendland, H.A. (1860) Bemerkungen über einige Palmengattungen Amerika’s. Ueber Iriartea R. & P. und die ihr zunächst stehenden
Gattung mit Einschluss von Wettinia Poepp. & Endl. Bonplandia 8: 103.
Willdenow, C.L. (1806) Caroli a Linne: species plantarum. Editio quarta Vol. 4. Berlín: G. C. Nauk.
Wurdack, J.J. (1976) Certamen Melastomataceis XXV. Phytologia 35: 1–12.
FIVE NEW SPECIES OF PLEUROTHALLIS FROM ECUADOR Phytotaxa 607 (3) © 2023 Magnolia Press • 181
https://doi.org/10.5962/bhl.part.2606
Zambrano, B., Solano-Gómez, R. & Wilson, M. (2017) A new species of Pleurothallis (Orchidaceae: Pleurothallidinae) from Southwestern
Ecuador: Pleurothallis marioi. Phytotaxa 308: 80–88.
https://doi.org/10.11646/phytotaxa.308.1.6