Research Article |
Corresponding author: John L. Clark ( jclark@lawrenceville.org ) Academic editor: Laura Clavijo
© 2022 Francisco Tobar, James F. Smith, John L. Clark.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Tobar F, Smith JF, Clark JL (2022) Two new pendulous epiphytic Columnea L. (Gesneriaceae) species from the Chocó forests of the Northern Andes. PhytoKeys 196: 49-61. https://doi.org/10.3897/phytokeys.196.79673
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Exploratory field expeditions to the Chocó forests in the northwestern slopes of the Ecuadorian and Colombian Andes resulted in the discovery of two new species of Columnea (Gesneriaceae). Columnea fluidifolia J.L.Clark & F.Tobar, sp. nov., is described as a narrow endemic from Bosque Protector Mashpi and surrounding areas in the province of Pichincha in northern Ecuador. Columnea pendens F.Tobar, J.L.Clark & J.F.Sm., sp. nov., is described from recently discovered populations in the provinces of Carchi and Santo Domingo de los Tsáchilas (Ecuador) and the departments of Cauca and Nariño in southwestern Colombia. The two new species are pendent epiphytes with elongate shoots and shallowly bilabiate to nearly tubular corollas. Descriptions, complete specimen citations, and a distribution map are provided. Based on IUCN guidelines, a preliminary conservation status of Critically Endangered (CR) is provided for C. fluidifolia and Endangered (EN) is provided for C. pendens.
Las expediciones de campo a los bosques del Chocó en las laderas noroccidentales de los Andes de Ecuador y Colombia dieron como resultado el descubrimiento de dos nuevas especies de Columnea (Gesneriaceae). Columnea fluidifolia J.L.Clark & F.Tobar, sp. nov., descrita como endémica de una reducida área del Bosque Protector Mashpi y zonas circundantes en la provincia de Pichincha. Columnea pendens F.Tobar, J.L.Clark & J.F.Sm., sp. nov., se describe a partir de poblaciones recientemente descubiertas en las provincias de Carchi, Santo Domingo de los Tsáchilas (Ecuador) y los departamentos de Cauca y Nariño en el suroeste de Colombia. Las dos nuevas especies son epífitas pendulares con brotes alargados y corolas ligeramente bilabiadas a casi tubulares. Descripciones, citaciones completas de los especímenes y mapa de distribución son incluidas. Basados en los criterios de la IUCN, se asigna un estado de conservación preliminar de En Peligro Critico (CR) para C. fluidifolia y a C. pendens se le asigna la categoría de En Peligro (EN).
Chocó, Colombia, Columnea, Ecuador, Gesneriaceae, taxonomy
The flowering plant family Gesneriaceae, with more than 3400 species and 150 + genera (
Most Columnea are epiphytic with primary shoots that are characterized as erect, horizontal, dorsiventral (associated with facultative epiphytes), or pendent. The two species described here are characterized as epiphytes with elongate pendent shoots, a habit more typical of Columnea from Central America. In contrast, most Columnea in South America are facultative epiphytes with dorsiventral shoots. For example, Panama and Costa Rica have collectively more than 20 species of Columnea that are characterized as pendent epiphytes. Some examples of species in the northern Andes with elongate pendent shoots include Columnea billbergiana Beurl., C. kienastiana Regal., and C. minor Hanst. Columnea fluidifolia is endemic to the Bosque Protector Mashpi and surrounding areas in the province of Pichincha in northern Ecuador (Fig.
Columnea ranges from Mexico south to Bolivia, and is most diverse in the northern Andes of Colombia and Ecuador. With more than 210 species (
Plants were photographed in the field and subsequently pressed and dried. Specimens are currently deposited at the National Herbarium of Ecuador (QCNE) and the herbarium at the Pontificia Universidad Católica del Ecuador (
We assessed the extinction risk of the two new species following the
Ecuador. Pichincha: cantón Quito, parroquia Pacto, primary road between the town of Pacto and Mashpi Lodge, 0°9'49.3"N, 78°49'14.6"W, 1662 m, 15 Mar 2019, J.L. Clark & L. Jost 16286 (holotype: US; isotypes:
Similar to Columnea ceticeps, differing in calyx and corolla uniformly orange (vs. calyx green and corolla bright red in C. ceticeps) and corolla shallowly bilabiate (vs. deeply bilabiate in C. ceticeps).
Epiphytic herb with elongate pendent shoots, 1.0–1.5 m long, red-brown, with zigzag appearance, densely pilose with gold-colored multicellular trichomes; internodes 2.0–4.0 cm long. Petioles 0.2–0.4 cm long, pilose with multicellular trichomes; leaves opposite, pairs either strongly anisophyllous or isophyllous, sometimes anisophyllous and isophyllous on the same shoot, larger leaf 9.5–10.5 cm long, 3.9–4.5 cm wide, ovate-elliptic, apex long acuminate, base slightly oblique, lateral veins 5–8 per side, adaxially dark-green, with multicellular white-transparent trichomes, abaxially light-green, densely pilose with multicellular white transparent trichomes, more densely pubescent on veins, margin serrulate; smaller leaf 1.3–1.9 cm long, 0.2–0.3 cm wide, lateral veins 2–3 per side, petiole 0.1–0.2 cm long, otherwise similar to larger leaf. Inflorescence reduced to a single axillary flower; bracts not seen, presumably caducous. Pedicels 2.6–3.7 cm long, bright orange, densely pilose with multicellular rust-colored trichomes. Calyx loosely clasping corolla, uniformly bright orange, lobes 2.6–4.2 cm long, 0.4–0.6 cm wide, oblong to narrowly-elliptic, apex acuminate to obtusely acuminate, exterior pilose, with multicellular rust-colored trichomes, more pubescent on veins and margins, interior glabrous, margin serrulate. Corolla 4.5–4.9 cm long, 0.6–0.8 cm at widest point, tubular throughout, inflated near center, slightly gibbous at the base, 0.6–0.7 cm wide before the limb, 0.2–0.4 cm at narrowest point of the base, bright orange externally, densely pubescent with multicellular rust-colored trichomes, interior with short white trichomes; ventral and lateral lobes reflexed, ventral lobe narrowly triangular to oblong, 0.4–0.5 cm long 0.2–0.3 cm wide, dorsal lobes fused, 0.2–0.3 cm long, 0.4–0.5 cm wide, lateral lobes triangular to narrowly triangular, 0.2–0.3 cm long, 0.2–0.3 cm wide. Filaments ca. 4.5 cm long, connate at base for 0.3 cm and adnate to corolla, anthers ca. 1.0 mm long, 1.0 mm wide, included in the corolla throat, quadrangular. Ovary ca. 4.0 mm long, conical, densely pubescent, with multicellular transparent trichomes; style 3.5–4.0 cm long, white with yellow apex, glabrescent; stigma unlobed, papillate, included in corolla tube. Nectary a bilobed dorsal gland. Fruit an indehiscent succulent berry, globose, distally acuminate, uniformly white, pilose; seeds not seen.
Flowering during June, August, and November. Mature fruits observed in June.
The phyllotaxy in most neotropical Gesneriaceae is opposite with pairs that are either equal in size (isophyllous) or strongly unequal in size (anisophyllous). The leaves in C. fluidifolia are unusual for having anisophyllous leaf pairs (Fig.
Columnea fluidifolia J.L.Clark & F.Tobar A lateral view of mature flower B anisophyllous leaf pairs C isophyllous leaf pairs (base & apex) and anisophyllous leaf pairs (middle) D immature berry E mature berry (A, B, D from J.L. Clark 16286 C, E from A.J. Perez & F. Tobar sn). Photos A, B, D by J.L. Clark & C, E by F. Tobar.
Columnea fluidifolia is endemic to the northwestern Andean slopes of Ecuador. The only known populations are located in the Mashpi-Pachijal conservation area, in the northwestern province of Pichincha (Fig.
Columnea fluidifolia differs from most Columnea by the presence of pendent elongate shoots and paired leaves that are both anisophyllous (Fig.
Ecuador. Pichincha: Distrito Metropolitano de Quito, Bosque protector Mashpi, Trasecto Mashpi Laguna 0.169750°N, 78.872120°W, 1150 m, 27 Jun 2017, F. Tobar et al. 2908 (
Ecuador. Carchi, Espejo, road between Goaltal and Chical, near to río Gualpi, 0.890555°N, 78.2188889°W, 1786 m, 12 Sep 2021, F. Tobar 3638 (holotype:
Similar to Columnea ceticeps, differing in the bright orange corollas (vs. bright red corollas in C. ceticeps), calyx lobes broadly ovate (vs. narrowly elongate in C. ceticeps), corolla tube ventricose (vs. uniformly tubular in C. ceticeps), and corolla tube densely pubescent (vs. corolla tube sparsely pubescent in C. ceticeps).
Epiphytic herb with elongate pendent shoots, 1.2–1.5 m long, brown, with zigzag appearance, glabrous near base, pilose with gold-colored trichomes near apex; internodes 2.0–4.5 cm long. Petioles 0.1–0.2 cm long, pilose with multicellular gold-colored trichomes; leaves opposite, pairs usually, nearly equal to isophyllous, rarely anisophyllous, 0.6–6.0 cm long, 0.3–2.3 cm wide, ovate to elliptic, apex acuminate, base slightly oblique, lateral veins 3–8 per side, adaxially dark-green, with white trichomes, abaxially green to red-purple, densely pilose with multicellular gold-colored trichomes and single-celled white trichomes, margin crenulate to serrulate. Inflorescence reduced to 1 or 2 flowers per axil; bracts not seen, presumably caducous. Pedicels 0.3–1.3 cm long, green or red, appressed pilose with multicellular gold-colored trichomes. Calyx clasping, lobes 1.2–2.3 cm long, 0.25 cm wide, triangular-ovate to broadly triangular-ovate, apex long acuminate, exterior densely pilose, with multicellular gold-colored trichomes, interior glabrous, margin serrulate. Corolla 5.2–6.1 cm long, 0.6–1.1 cm at widest point (apex of corolla limb, near throat), tubular, medially ventricose, gibbous at base, 2.8–3.5 mm wide at narrowest point at the base, bright orange externally, interior yellow, exterior densely pubescent with multicellular red-colored trichomes, interior with sparse short trichomes and some stalked glandular trichomes; limb bilabiate, upper lip with two fused dorsal lobes and two lateral lobes, lower limb with an extended ventral lip; dorsal lobes connate, rounded to subquadrate, ca. 6.2 mm long, ca. 6.5 mm wide; lateral lobes triangular to narrowly triangular, ca. 1.5 mm long, ca. 2.0 mm wide; ventral lobe, narrowly oblong to linear ca. 1.9 cm long and ca. 2.8 mm wide, galea 1.9 cm long. Filaments ca. 5.1 cm long, connate for ca. 0.5 cm and adnate to base of corolla, anthers ca. 1.4 mm long, ca. 1.3 mm wide, included in the corolla throat, quadrangular. Ovary 3.4 mm long, conical, pubescent or glabrous, with multicellular transparent trichomes; style pale yellow to white, pilose with multicellular transparent and short stalked glandular trichomes; stigma unlobed, papillate, included in corolla tube. Nectary a bilobed dorsal gland. Mature fruit and seeds not seen.
Collected in flower during February-April, and September. Immature fruits observed in April.
The specific epithet refers to the pendent epiphytic habit.
Columnea pendens is endemic to the western Andean slopes of Ecuador and Colombia (Fig.
The Ecuadorian populations are known from the provinces of Santo Domingo de los Tsáchilas and Carchi (Fig.
The Colombia populations are known from Cerro Plateado, Cerro Pinche, and Munchique in the Cauca department, and La Planada in the Nariño department (Fig.
Based on the available information and According to the IUCN Red List criteria (
Columnea pendens is similar to C. ceticeps, which was recently described from Colombia and northern Ecuador (
Colombia. Cauca: toward south end of La Depresion, between Cerro Pinche and Cerro Plateado, 26 Sep 1994, E.L. Core 1358 (US); El Tambo, O. Haught 5200 (US); El Tambo, Munchique Jul 1948 S. Yepes Agredo 472 (US). Nariño: Reserva Natural La Planada, Ricaurte, on the road near Quebrada Dulce, 3 Mar 1989, J.F. Smith & P. Galeano 1522, (COL, WIS). Ecuador. Carchi: canton Mira, via Gualchan-El Carmen-Chical, 0.4166667°N, 78.216666°W, 1900–2000 m, 2 Nov 2014, AJ Perez et al. 7716 (
Support for F. Tobar was provided by the Swiss Federal Research Institute (WSL) – National Geographic Society (Grant N° 9952-16); Swiss National Science Foundation (SNF Grant N° 173342) – European Research Council Advanced Grants (ERC Grant N° 787638) Aves y Conservación. We thank the Ministry of the Environment of Ecuador for providing specimen collection and transportation permits (Research permit Aves y Conservación N° 007-2018-IC-FLO-FAU and mobilization permit Aves y Conservación 005-FLO-2019-DPAP-MA). Maria José Gavilanes is acknowledged for illustrating Columnea fluidifolia (Fig.