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Phytotaxa 598 (4): 293–300
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: 18 May 2023; published: 29 May 2023
https://doi.org/10.11646/phytotaxa.598.4.3
293
Restrepia santanderensis (Orchidaceae: Pleurothallidinae), a new species from the
western slope of the eastern Andes in Colombia
NICOLÁS GUTIÉRREZ MORALES1,2,6*, JUAN SEBASTIÁN MORENO2,3,7, ADAM P. KARREMANS4,5,8 &
KAREN GIL-AMAYA2,4,9
1 Instituto de Biociências de Rio Claro, Departamento de Botânica, Universidade Estadual Paulista - UNESP, Av. 24 A 1515, Bela Vista,
13506-900, Caixa Postal 199, Rio Claro, São Paulo, Brazil.
2 Grupo de Investigación Schultes, Fundación Ecotonos, Valle del Cauca, Cali, Colombia
3 Departamento de Biología, Universidad del Valle, Calle 13 # 100-00, Cali, Colombia
4 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Costa Rica
5 Naturalis Biodiversity Center, Evolutionary Ecology Group, Sylviusweg 72, 2333 BE, Leiden, The Netherlands
6
�
nicolaiequal@gmail.com; https://orcid.org/0000-0002-6592-9326
7
�
semoreno113@gmail.com; https://orcid.org/0000-0002-0875-9498
8
�
adam.karremans@ucr.ac.cr; https://orcid.org/0000-0001-5987-7710
9
�
kgilecologa@gmail.com; https://orcid.org/0000-0002-2088-1541
* Author for correspondence
Abstract
A new species of the orchid genus Restrepia from the department of Santander, in the western slope of the eastern Andes of
Colombia, is described and illustrated. Restrepia santanderensis is compared with the morphologically similar R. flosculata
and R. mendozae, which share having short inflorescences and small to medium yellow flowers. The new species can be
recognized by its ramicauls prominently covered by brown-dotted sheaths, with the uppermost sheath elongate as long as
the peduncle, the sepals partially spreading, the synsepal with glandular trichomes along the veins of the adaxial surface, the
straight petals and the rectangular lip with truncate apex. Comparisons with similar species and comments on its ecology
are provided.
Keywords: Colombia, eastern Andes, endemism, Restrepia flosculata, Restrepia mendozae, taxonomy
Resumen
Se describe e ilustra una nueva especie de orquídea del género Restrepia oriunda del departamento de Santander,
en el flanco occidental de los Andes Orientales de Colombia. Restrepia santanderensis se compara con las especies
morfológicamente similares, R. flosculata y R. mendozae, semejantes por sus inflorescencias cortas y las flores amarillas
pequeñas a medianas. La nueva especie se distingue de estas por sus ramicaules cubiertos por brácteas prominentemente
punteadas de marrón, con la bráctea superior tan larga como el pedúnculo, los sepalos parcialmente abiertos, el sinsépalo
tiene tricomas glandulares a lo largo de las nervaduras de la superficie adaxial y el labelo es rectangular y truncado en
el ápice. Se compara con especies similares y se proporcionan comentarios sobre su ecología.
Introduction
With about 60 species occurring along Central America and the tropical Andes (Karremans 2016, Karremans & Vieira-
Uribe 2020, Miller et al. 2020), the genus Restrepia Kunth (1815: 293) represents a challenge for taxonomists. The
species are easily recognized as belonging to the genus (Luer 1996), in contrast, the floral structure of most species
is extremely uniform. Except for a few species that are easily recognized, most taxa are difficult to distinguish based
on their morphology when placed side by side (Luer 1996). Populations of common and widespread species, such as
Restrepia antennifera Kunth (1815: 367), R. brachypus Reichenbach, H.G. (1886: 554), R. contorta (Ruiz & Pavón
GUTIERREZ MORALES ET AL.
294 • Phytotaxa 598 (4) © 2023 Magnolia Press
1798: 325) Luer (1996: 50), R. guttulata Lindley (1836: 357), R. muscifera (Lindley 1842: 77) Reichenbach, H.G. ex
Lindl (1859: 2) and R. trichoglossa F. Lehmann. ex Sander (1901: 215), often exhibit wide morphological variation
in the size of the vegetative portion and in the color of the flowers, with many intermediate forms within the spectrum
(Miller et al. 2020, Luer 1996).
This has led to historical confusion among several taxa, both in herbaria and in personal collections (Luer 1996,
Millner et al. 2020), and to notorious taxonomic inflation through the recognition of numerous color forms as distinct
species, which can have important negative consequences outside of taxonomy in the overestimation in speciation
rates, the underestimation of ecological preferences and distribution ranges, false interpretations of endemisms
and hotspots, skewed origin and diversification patterns, mistaken conservation priorities, and increased interest of
collectors (Karremans et al. 2020).
Not all Restrepia species are broadly distributed and variable. Certain taxa are restricted to a few locations in
humid forest of mid to high elevations along the Andes of Colombia, Venezuela and Ecuador, and have been recorded
only a few times or just once (Millner et al. 2020). Due to their restricted distribution and rarity in the wild, narrow
endemics may be threatened by human pressures such as land use transformation (Millner et al. 2020). If proven to be
truly distinct, these taxa should be considered a conservation priority among Restrepia species.
Currently, 30 species of Restrepia are known to occur in Colombia, 18 of which are reportedly endemic to the
country (Luer 1996, Bernal et al. 2016). Several of these species occur in the eastern Andes of Colombia, a very
fragmented and anthropized region, mainly in the western slope, towards the Magdalena valley (Armenteras et al.
2003). Recent botanical efforts in this region, specifically in the department of Santander, yielded several novelties
for various plant families (Díaz-Piedrahita & Rodríguez-Cabeza 2011, 2012, Gutiérrez Morales et al. 2018, 2021;
Jara-Muñoz & Zabala-Rivera 2018, Mendoza-Cifuentes & Aguilar-Cano 2018, Moreno et al. 2018, Vieira-Uribe &
Gutiérrez 2020). Here we describe and illustrate a previously undescribed, apparently rare Restrepia species, found in
this highly threatened region during a field exploration in June of 2016.
Materials and methods
Description and drawings were prepared from living specimens. Digital images were taken using a Canon EOS T3
fitted with a Canon 60 mm f/2.8 macro lens. Sketches from living and preserved specimens were digitized to create
a composite template in Adobe Photoshop® CS6. A digital composite line drawing was then made in the Procreate
illustration application for IPad 6th and IPad 8th generation tablet computers (following Bogarín et al. 2019). Specimens
from COL, JBB, JAUM, VALLE, FMB, JAUM, HUA, TOLI, ICESI, CAUP, MO (online), were consulted, but no
additional material of the new species was found.
Taxonomy
Restrepia santanderensis N. Gut. & K. Gil-Amaya, sp nov. (Figures. 1, 2, 3C, 3D)
Type:—COLOMBIA. Santander: La Belleza, Vereda Berlin, 1950 m, January 2016, flowering under cultivation in January 2020, M.
Calderón et al. 191 (holotype: JBB!).
Restrepia santanderensis is similar to R. mendozae Luer (1996: 157), but can be distinguished by the sheaths of the
ramicauls completely brown-dotted (vs. only lowermost sheath dotted), the sepals partially spreading, yellow with
vinaceous stripes in the synsepal (vs. sepals completely spreading, yellow with purple spots), the dorsal sepal prostrate
(vs. dorsal sepal erect), the synsepal with trichomes along the veins on the adaxial surface (vs. synsepal glabrous),
the straight petals (vs. decurved petals), and the lip yellow with intense rose at base (vs. entirely yellow with purple
spots).
Description:—Plant epiphytic, caespitose, occasionally prolific, erect to sub-erect, up to 24 cm tall. Roots
slender, flexuous, ca. 1 mm in diameter. Ramicauls erect, 2–13 cm long, enclosed by 5–6, whitish, brown-spotted,
papery, loose, imbricating, oblique, laterally compressed, acute sheaths, the uppermost prominent, extended as long
as the pedicel, acuminate. Leaves erect, elliptical-ovate, coriaceous, sub-acute, minutely tri-denticulate, 4.5–11.2 ×
2.2–4.3 cm, the base cuneate contracted into a short, twisted petiole. Inflorescence up to 3.9 cm, emerging without an
annulus 5–9 mm below the apex of the ramicaul, with a solitary yellow flower, produced successively in a fascicle
A NEW SPECIES OF RESTREPIA FROM COLOMBIA Phytotaxa 598 (4) © 2023 Magnolia Press • 295
FIGURE 1. Illustration of Restrepia santanderensis, A. Habit. B. Flower. C. Dissected perianth with magnified trichomes of the synsepal.
D. Lip, column, ovary, floral bract in lateral view. E. Lip, lateral and adaxial view. F. Column in ventral view, without the anther G. Anther
cap and pollinaria. Illustration by Nicolás Gutiérrez Morales and Sebastián Moreno from the holotype.
GUTIERREZ MORALES ET AL.
296 • Phytotaxa 598 (4) © 2023 Magnolia Press
FIGURE 2. Lankester composite dissection plate (LCDP) of Restrepia santanderensis. A. Habit. B. Flower. C. Dissected perianth with
magnified trichomes of the synsepal. D. Lip and column, lateral view. E. Lip, adaxial view. F. Column in ventral view. Photographs by
Nicolás Gutiérrez Morales, from the holotype.
A NEW SPECIES OF RESTREPIA FROM COLOMBIA Phytotaxa 598 (4) © 2023 Magnolia Press • 297
FIGURE 3. Restrepia santanderensis N. Gutiérrez & K. Gil-Amaya, and similar morphological species. A. R. flosculata. B. R. mendozae.
C. R. santanderensis. D. Capsules of R. santanderensis. Photographs by Andreas Key (A), Joost Riksen (B) from cultivated specimens and
Nicolás Gutiérrez Morales (C, D) from the paratype.
GUTIERREZ MORALES ET AL.
298 • Phytotaxa 598 (4) © 2023 Magnolia Press
behind the leaf, in a slender, sub-erect, 2.0–2.7 cm long peduncle, subtended by a tubular, oblique spathe, 9.2–9.9 mm
long, floral bract translucent, oblique, slightly inflated, apiculate, to 6.9 mm long, enclosing the short, stout pedicel,
1.0–1.3 mm long, with a filament 2.8–3.5 mm long. Ovary slightly arcuate, lightly costate, 4.7–6.17 mm long. Flowers
resupinate, intermediate-sized, to 2.5 cm long. Sepals pale yellow, membranous, not completely spreading, the dorsal
sepal free, narrowly ovate and slightly concave below the lower third, 5-veined, contracted into a long tail above the
lower third, with the apex clavate-thickened, 1 mm thick at the apex, 17.0–22.6 mm long, 1.91–2.56 mm wide above
the base, the lateral sepals somewhat striped in vinaceous near the margins of the basal third, each 7-veined, connate
ca. 19.0–21.8 mm into a narrowly oblong, slightly concave at base, bifid, subacute lamina, with glandular trichomes in
the middle region and in the apex of the adaxial surface, along the veins, 22–25 mm long, 7.35–7.8 mm wide. Petals
translucent yellow, slender, narrowly linear-triangular, membranous, 3-veined, the margins minutely serrate near the
base, attenuated above the middle, the apex clavate-thickened, 11.0–11.6 mm long, 1.0–1.2 mm wide above the base.
Lip yellow, vinaceous along the veins, the base intense rose and white, rectangular, 3-veined, truncated, 3-veined, 7.6–
10.9 mm long, 2.3–3.6 mm wide, the base a slightly concave, with erect margins, the sides with a translucent, uncinate
process or cirrhi, ca. 1.4–1.8 mm long, each with a minute tooth at the base, the lip with a pair of low calli extending
forward from the base of near the middle, the base connected to the column-foot by a white, rigid, cylindrical neck, the
apical portion microscopically verrucose. Column greenish-white, the basal half slender, clavate, the margins of the
apical half irregular, marked with intense yellow, 4.6–6.1 mm long, the base pedestal-like with a pair of longitudinal,
minutely papillose calli, ending in a pair of light orange, obtuse points, the anther and stigma ventral, the stigma
subcuneiform. Anther cap incumbent, deciduous, whitish. Pollinia four, ovoid, in two pairs, united by a viscidium.
Capsule dehiscent, ca. 1.5 cm long
Distribution and Ecology:—Restrepia santanderensis occurs in a small area the eastern Andes of Colombia. The
specimen that served as type was found in 2016 in the municipality of La Belleza, Santander, near to the top of the
waterfall called “El chorro de la Humareda” (Figure 4). The plant was found growing epiphytically, partially exposed
to direct sunlight, at around four meters above the ground, on an isolated tree among pastures, not so far from the
edge of the forest. Other orchid species in the area were Pleurothallis furcifera Luer (1976: 108), Maxillaria carrilloi
Christenson (2013: 174), Epidendrum cupreum F. Lehmann. & Kraenzl (1899: 476) and Epidendrum fusagasugaense
E. Parra, Hágsater & L. Sánchez (2013: 69). A short expedition in May of 2022 failed to recover more individuals of
the new species, plantations of Solanum quitoense Lam (1794: 16), “lulo”, were introduced in recent years, and the
tree where the original specimen was collected was not found. However, the steep walls of the canyon that receives
the “Peña Bonita” river, which falls for more than 220 meters into a waterfall, has a well-preserved forest remnant in
a gradient of elevation of 1,700 to 1,900 m, and may host a population of Restrepia santanderensis.
Etymology:—Named for the Department of Santander, Colombia, where the species was discovered.
Additional material examined (paratype):—COLOMBIA. Santander: La Belleza, Vereda Berlin, 1950 m,
January 2016, flowered under cultivation in January 2023, N. Gutiérrez & J. de Jesus 241 (JBB!).
Taxonomic Discussion:—Restrepia santanderensis (Figures. 1, 2, 3C, 3D) can be distinguished from other species
in the genus by the ramicauls covered by densely brown-spotted sheaths, the elongate uppermost sheath extending as
long as the pedicel, the short inflorescences with intermediate-sized, partially spreading, yellow flowers, the elongate,
prostrate dorsal sepal, the synsepal with trichomes along the veins on the adaxial surface, and the rectangular lip,
truncate at the apex, with intense rose at base and microscopically verrucose.
The most similar species morphologically are R. flosculata Luer (1982: 127) (Figure 3A), from Valle del Cauca
department in Colombia and northwestern Ecuador, a species that present various color forms, dotted or stripped in
purple to orange (Luer 1996), and R. mendozae (Figure 3B) known from southeastern Ecuador (Luer 1996), with pale
yellow flowers dotted in dark purple. They all share the overall similar size, shape of the leaves, the short-pedunculate
inflorescence, and the pale yellow, intermediate to small-sized flowers.
The flowers of R. flosculata may have a similar color pattern to that of R. santanderensis. However, in R.
flosculata, only the basal sheaths of the ramicauls are brown-dotted (vs. all sheaths brown-dotted), the uppermost
sheath is oblique, acute (vs. elongate, acuminate), the sepals are shorter, to 14 mm long (vs. sepals to 25 mm long),
the dorsal sepal is narrowly triangular below the middle, attenuate above (vs. dorsal sepal narrowly ovate below the
lower third, contracted into a tail above), with a notoriously glabrous, obovate synsepal (vs. synsepal with evident
trichomes, oblong), decurved petals (vs. straight petals), and a broadly oblong lip with retuse apex (vs. rectangular lip
with truncate apex).
In R. mendozae the shapes and size of sepals and petals are very similar to that of R. santanderensis, but only
the lowermost sheath is brown-dotted. In addition, the sepals are spreading, yellow with purple dots (vs. partially
spreading sepals, yellow with vinaceous strips in the synsepal), the dorsal sepal is erect (vs. prostrate), the synsepal is
A NEW SPECIES OF RESTREPIA FROM COLOMBIA Phytotaxa 598 (4) © 2023 Magnolia Press • 299
glabrous (vs. synsepal with evident trichomes), the petals are decurved (vs. straight), and the lip is entirely yellow, with
dark purple spots (vs. yellow with intense rose at the base and vinaceous, longitudinal stripes).
The arrangement of the trichomes along the veins of the synsepal in R. santanderensis, matches Millner &
Baldwin’s (2016) description of the papillae in R. brachypus, following the stripes of the synsepal. This as ‘false nectar
guides’, part of the non-rewarding pollination strategy that is supposed to occur in the genus Restrepia.
FIGURE 4. View of the Restrepia santanderensis habitat in the western slope of the eastern Andes in Colombia, near to the top of “La
Humareda” waterfall, in La Belleza (Santander) municipality.
Acknowledgements
We would like to thank the José Celestino Mutis Botanical Garden in Bogotá and all the people who helped with the
preparation of voucher specimens. Belisario Sutaterna and his family in La Belleza, Santander, whose friendship and
generosity were crucial for this discovery. Thanks to Melisa Alegría-Valencia from Universidad del Valle for assisting
with the flower dissection of the new species. We also thank Joost Riksen for the photograph of R. mendozae and the
anonymous reviewers for their comments and suggestions on the manuscript.
References
Armenteras, D., Gast, F. & Villareal, H. (2003) Andean forest fragmentation and the representativeness of protected natural areas in the
eastern Andes, Colombia. Biological conservation 113: 245–256.
https://doi.org/10.1016/S0006-3207(02)00359-2
Bernal, R., Gradstein, S.R. & Celis, M. (2016) Catálogo de plantas y líquenes de Colombia. Volumen II. Magnoliaceae a Zygophyllaceae
– especies introducidas y cultivadas. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá. pp. 1509–3058.
Bogarín, D., Eberhard, K. & Díaz-Morales, M. (2019) Lepanthes elusiva – a New Species of Lepanthes (Orchidaceae: Pleurothallidinae)
from Tapantí Area in Cartago, Costa Rica and a Glance to Other Species of the Genus in a Small Area Around the Río Grande de
Orosi in Costa Rica. Die Orchidee 5: 17–28.
Díaz-Piedrahita, S. & Rodríguez-Cabeza, B.V. (2011) Novedades en Asteraceas colombianas I. Revista de la Academia Colombiana de
Ciencias Exactas, Físicas y Naturales 35: 411–424.
Díaz-Piedrahita, S. & Rodríguez-Cabeza, B.V. (2012) Novedades en Asteraceas colombianas II. Revista de la Academia Colombiana de
Ciencias Exactas, Físicas y Naturales 36: 501–515.
GUTIERREZ MORALES ET AL.
300 • Phytotaxa 598 (4) © 2023 Magnolia Press
Christenson, E.A. (2013) Maxillaria carrilloi. In: Christenson, E.A. Maxillaria, an unfinished monograph, Volume 1. published for Robert
Christenson by Patricia Ann Harding, Lebanon, pp. 174–175.
Gutierrez Morales, N., Harding, P., Dalström, S. & Moreno Silva, J.S. (2018) A new species of Cyrtochilum (Orchidaceae: Oncidiinae)
from the Eastern Cordillera of Colombia. Lankesteriana 18: 63–68.
Gutierrez Morales, N., Vieira-Uribe, S. & Moreno, J.S. (2021) Lepanthes marielana (Orchidaceae, Pleurothallidinae), a new species from
the eastern Andes in Colombia. Phytotaxa 484: 113–120.
https://doi.org/10.11646/phytotaxa.484.1.5
Humboldt, F.W.H.A., Bonpland, A.J.A. & Kunth, C.S. (1815) Nova Genera et Species Plantarum (folio ed.). Librariae Graeco-Latino-
Germanicae, ex typographia d’Hautel. Paris. 302 pp.
Jara-Muñoz, O.A. & Zabala-Rivera, J.C. (2018) Dos nuevas especies de Begonia (Begoniaceae) de la Serranía de los Yariguíes (Santander,
Colombia). Novon: A Journal for Botanical Nomenclature 26: 355–363.
https://doi.org/10.3417/2018226
Karremans, A.P. (2016) Genera Pleurothallidinarum: an updated phylogenetic overview of Pleurothallidinae. Lankesteriana 16: 219–
241.
https://doi.org/10.15517/lank.v16i2.26008
Karremans, A.P & Vieira-Uribe, S. (2020) Pleurothallids Neotropical Jewels - Volume I. Imprenta Mariscal, Quito. 320 pp.
Karremans, A.P., Chinchilla, I.F., Rojas-Alvarado, G., Cedeño-Fonseca, M. Damián, A. & Léotard, G. (2020) A reappraisal of Neotropical
Vanilla. With a note on taxonomic inflation and the importance of alpha taxonomy in biological studies. Lankesteriana 20: 395–
497.
https://doi.org/10.15517/lank.v20i3.45203
Lehmann, F. & Kraenzlin, F.W.L. (1899) Epidendrum cupreum. Botanische Jahrbücher für Systematik, Pflanzengeschichte und
Pflanzengeographie 26: 476.
Lindley, J. (1836) Notes of some genera and species of the American Orchidaceae. Companion to the Botanical Magazine 2: 353–357.
Lindley, J. (1842) Pleurothallis muscifera. Edwards’s Botanical Register 28: 79.
Luer, C.A. (1982) Miscellaneous new species and combinations in the Pleurothallidinae (Orchidaceae). Selbyana 7: 127.
Luer, C.A. (1976) Miscellaneous species of Pleurothallis. Selbyana 3: 38–201.
Luer, C.A. (1996) Icones pleurothallidinarum XIII. Systematics of the genus Restrepia (Orchidaceae). Monographs in Systematic Botany
from the Missouri Botanical Garden 59: 1–168.
Mendoza-Cifuentes, H. & Aguilar-Cano, J. (2018) A new species of Elaeagia (Rubiaceae) from the Cordillera Oriental of Colombia. Biota
Colombiana 19: 3–9.
https://doi.org/10.21068/c2018.v19s1a01
Millner, H.J. & Baldwin, T.C. (2016) Floral micromorphology of the genus Restrepia (Orchidaceae) and the potential consequences for
pollination. Flora-Morphology, Distribution, Functional Ecology of Plants 225: 10–19.
https://doi.org/10.1016/j.flora.2016.09.007
Millner, H.J., Bachman, S.P. & Baldwin, T.C. (2020) An assessment of the conservation status of Restrepia (Orchidaceae) reveals the
threatened status of the genus. Plant Ecology & Diversity 13: 115–131.
https://doi.org/10.1080/17550874.2020.1735553
Moreno, J.S., Gutiérrez, N. & Vieira-Uribe, S. (2018) Lepanthes agatarum (Orchidaceae: Pleurothallidinae), a new species honoring an
ancient tribe from the eastern Andes in Colombia. Phytotaxa 376: 89–96.
https://doi.org/10.11646/phytotaxa.376.2.2
Parra, E., Hágsater, E. & Sánchez, L. (2013) Epidendrum fusagasugaense. In: Hágsater, E. & Santiago, E. (eds.) The Genus Epidendrum,
Part 10. Icones Orchidacearum 14. t. 1430.
Reichenbach, H.G. (1886) Orchideae describuntur. Flora, oder allgemeine botanische Zeitung 69: 547–562.
Reichenbach, H.G. (1859) Restrepia muscifera (Lindl.) Rchb. f. ex Lindl. In: Lindley, J. Folia orchidacea: An enumeration of the known
species of orchids, Restrepia. Bradbury and Evans Printers, London. pp. 1–3.
Ruiz, H. & Pavón, J. (1798) Humboltia contorta. In: Ruiz, H & Pavón, J. Systema Vegetabilium Florae Peruvianae et Chilensis. Typis
Gabrielis de Sancha, Madrid. pp. 456.
Sander, H.F.C. & Lehmann, F.C. (1901) Restrepia trichoglossa. In: Sander, H.F.C. Sander’s Orchid Guide, F. Sander & Co., St. Albans.
pp. 215.
