Three new species of Columnea (Gesneriaceae) from the western Andean slopes of Ecuador and Colombia

Abstract

Three new species of Columnea (Gesneriaceae, tribe Gesnerieae) are described from the western Andean slopes of northern Ecuador and southern Colombia. Columnea angulata J.L. Clark & F. Tobar and Columnea floribunda F. Tobar & J.L. Clark are described from northern Ecuador. Columnea tecta J.L. Clark & Clavijo is described from southern Colombia and northern Ecuador. The three new species are facultative epiphytes with dorsiventral shoots and are readily recognized by bright red tips on the abaxial and adaxial leaf surfaces. The species described here are vegetatively similar to the sympatric species Columnea picta H. Karst. and are readily differentiated by floral features that are illustrated, described and featured with digital images.

Full text

Three new species of Columnea (Gesneriaceae) from
the western Andean slopes of Ecuador and Colombia
John L. Clark1, Francisco Tobar2,4, Laura Clavijo3,
Mathieu Perret5, Catherine Helen Graham6
1Science Department, e Lawrenceville School, Lawrenceville, NJ 08648, USA 2Área de Investigación
y Monitoreo de Avifauna, Aves y Conservación – BirdLife en Ecuador, Quito, Ecuador 3Instituto de Cien-
cias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia 4Instituto Nacional de Biodiversidad,
Herbario Nacional del Ecuador QCNE, Quito, Ecuador 5Conservatoire et Jardin botaniques de la Ville de
Genève, Ch. de l’Impératrice 1, CH-1292 Chambésy, Switzerland 6Biodiversity and Conservation Biology
Unit, Swiss Federal Research Institute WSL, Birmensdorf, Switzerland
Corresponding author: John L. Clark (jclark@lawrenceville.org)
Academic editor: Ricardo Kriebel|Received 22 May 2021|Accepted 3 August 2021|Published 20 September 2021
Citation: Clark JL, Tobar F, Clavijo L, Perret M, Graham CH (2021) ree new species of Columnea (Gesneriaceae)
from the western Andean slopes of Ecuador and Colombia. PhytoKeys 182: 67–82. https://doi.org/10.3897/
phytokeys.182.69016
Abstract
ree new species of Columnea (Gesneriaceae, tribe Gesnerieae) are described from the western Andean
slopes of northern Ecuador and southern Colombia. Columnea angulata J.L. Clark & F. Tobar and Colum-
nea oribunda F. Tobar & J.L. Clark are described from northern Ecuador. Columnea tecta J.L. Clark &
Clavijo is described from southern Colombia and northern Ecuador. e three new species are facultative
epiphytes with dorsiventral shoots and are readily recognized by bright red tips on the abaxial and adaxial
leaf surfaces. e species described here are vegetatively similar to the sympatric species Columnea picta
H. Karst. and are readily dierentiated by oral features that are illustrated, described and featured with
digital images.
Abstract
Se describen tres especies nuevas de Columnea (Gesneriaceae, tribu Gesnerieae) originarias de la vertiente
occidental de los Andes de Ecuador y Colombia. Columnea angulata J.L. Clark & F. Tobar y Columnea
oribunda F. Tobar & J.L. Clark se describen del norte de Ecuador; Columnea tecta J.L. Clark & Clavijo
se describe del sur de Colombia y norte de Ecuador. Las tres especies nuevas son epítas facultativas con
vástagos dorsiventrales, las cuales se reconocen fácilmente por las puntas de las hojas de color rojo brillante
en ambas supercies; son vegetativamente similares y simpátricas con Columnea picta H. Karst., pero se
pueden diferenciar por las características orales que se ilustran, describen y detallan con imágenes digi-
tales en este artículo.
Copyright John L. Clark et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY
4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
PhytoKeys 182: 67–82 (2021)
doi: 10.3897/phytokeys.182.69016
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John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
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Keywords
Colombia, Columnea, Ecuador, Gesneriaceae, taxonomy
Introduction
e owering plant family Gesneriaceae, with over 3400 species and 150+ genera (Weber
2004; Weber et al. 2013), is in the order Lamiales. e family is divided into three sub-
families and seven tribes (Weber et al. 2013, 2020), which represent monophyletic lineages
(Ogutcen et al. 2021). e majority of New World members are in the subfamily Gesneri-
oideae and are represented by 1200+ species and 77 genera (Clark et al. 2020). Columnea
L. is classied in the tribe Gesnerieae and subtribe Columneinae (Weber et al. 2013, 2020).
e genus Columnea is primarily epiphytic. It ranges from Mexico south to Bolivia,
and is most diverse in the northern Andes of Colombia and Ecuador. With over 210
species (Clark et al. 2020), Columnea is the largest genus in the subfamily Gesnerioideae
(Weber et al. 2013, 2020). e genus is distinguished by fruits that are indehiscent ber-
ries in contrast to eshy bivalved capsules in closely related genera. Columnea is strongly
supported as a monophyletic genus based on molecular phylogenetic studies (Smith et
al. 2013; Schulte et al. 2014). e species described here were discovered during explora-
tory research expeditions and ongoing taxonomic research of herbarium specimens. e
three newly described species are similar to many taxa recognized in the section Col-
landra (Lem.) Benth. or previously classied as members of the genus Dalbergaria Tus -
sac. We refrain from classifying the new species to a subgeneric rank because most are
articially dened and not supported by phylogenetic studies (Smith and Carroll 1997;
Smith 2000; Clark and Zimmer 2003; Clark et al. 2012; Smith et al. 2013; Schulte et al.
2014). e three species described here are distributed on the western Andean slopes of
northern Ecuador and southern Colombia (Fig. 1). Herbarium specimens representing
these three species are often annotated as “Columnea a. picta” or “Columnea cf. picta”
because they share a similar vegetative feature of apical red leaf apices on the upper and
lower leaf surfaces. In contrast, most species of Columnea have red leaf apices on the
lower leaf surface, but not on the upper leaf surface. Table 1 summarizes prominent char-
acters to dierentiate the three new species from each other and from Columnea picta.
Taxonomic treatment
Columnea angulata J.L. Clark & Tobar, sp. nov.
urn:lsid:ipni.org:names:77219739-1
Figs 2, 3
Diagnosis. Diers from Columnea picta by a shallow bilabiate corolla limb (vs. deeply
bilabiate corolla limb) and a corolla perpendicular to oblique relative to the calyx (vs.
corolla straight relative to the calyx).

ree new species of Columnea (Gesneriaceae) 69
Type. E Imbabura: cantón Cotacachi, parroquia García Moreno, Cordil-
lera de Toisán, Cerro de la Plata, Bosque Protector Los Cedros, sendero Camino del
Oso, north of lodge, 0°18'N, 78°46'W, 1500–2600 m, 19 Mar 2003, J.L. Clark, F.
Figure 1. Distribution of Columnea angulata (circles), C. oribunda (triangles), C. tecta (squares), and
C. picta (asterisks). Note that C. picta ranges from southern Ecuador to northwestern Colombia (exceed-
ing the range of the currently described species) (Map generated by Marco Monteros).

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
70
Figure 2. Columnea angulata J.L. Clark & F. Tobar A mature ower B front view of ower C ventral
view of ower showing laterally compressed corolla tube D dorsiventral habit (A, D from J.L. Clark et al.
12198 B from J.L. Clark 10968 C from J.L. Clark et al. 7413). Photos by J.L. Clark.

ree new species of Columnea (Gesneriaceae) 71
Nicolalde & R. Hall 7413 (holotype: US [US3492386]; isotypes: AAU, COL, K, MO,
QCA, QCNE, SEL, UNA).
Description. Facultative epiphyte with dorsiventral shoots to 1.5 m long, sub-
woody, surutescent, glabrescent below, sparsely pilose above; internodes 7–10 cm
Figure 3. Columnea angulata J.L. Clark & F. Tobar A dorsiventral habit B lateral view of ower C dis-
sected corolla showing laments and ventral view of the anthers D dorsal view of the anthers E lateral
view of ovary and nectary gland F dorsal view of nectary gland G lateral view of calyx and style, showing
shallowly bid stigma H lateral view of ower. Illustrated by M.J. Gavilanes, based on F. Tobar et al 3409.

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
72
near base, then clustered at branch apex. Leaves opposite, strongly anisophyllous, pa-
pyraceous when dry; larger leaf nearly sessile, petioles 0.1–0.4 cm long, pilose; blade
asymmetric, oblanceolate to oblong, 7–28 × 2–9.5 cm, base oblique, apex acuminate,
margin serrate, adaxially uniformly dark green to red with dark red apex, glabrous,
abaxially light green, upper regions of margins, and apical third of leaf dark red, sparse-
ly pilose to densely pilose along the venation, lateral veins 7–15, primary and second-
ary veins occasionally red; smaller leaf sessile, sometimes appressed to the dorsal surface
of stem; blade asymmetric, lanceolate 0.5–2 × 0.4–0.6 cm, base oblique, apex acumi-
nate, margin serrate, green with red apex on both surfaces, glabrous adaxially, sparsely
pilose to densely pilose along the venation and margins abaxially. Inorescence reduced
to a single axillary ower (rarely 2–3); peduncles absent or highly reduced (< 0.2 cm);
bracts 1–2, light green, lanceolate, 0.7–1.2 × 0.2–0.4 cm, glabrous on both sides.
Flowers subtended by elongate pedicels, 1.5–2.5 cm long, sparsely to densely pilose,
tightly appressed to the abaxial leaf surface when immature, becoming pendent during
anthesis; calyx lobes 5, nearly free, mostly equal in size and shape, dorsal lobe slightly
smaller, lobes appressed to ower when immature and spreading during anthesis, from
uniformly yellow, to red with yellow margins, to yellow with a large reddish mid-
region, 1.5–3.5 × 0.5–1 cm, ovate to broadly oblong, apex acuminate to acute, margin
serrate, pilose on both surfaces; corolla tubular, appearing perpendicular to calyx via
a sigmoid-shaped corolla tube, 2.1–3.5 cm long, outer and inner surfaces pilose, base
appearing laterally compressed, limb shallowly bilabiate, white suused with yellow
on lower two thirds, more yellow toward apex, splotches of dark red on lower portion
of lateral and ventral lobes, light yellow patch below lobes, red streaks abaxially, lobes
0.3–0.4 × 0.3–0.5 cm. Androecium of 4 stamens, laments connate at the base and
forming a lament curtain for 0.2–0.4 cm, free portion of laments 3–3.5cm long,
Table 1. General geographic distribution (names in parentheses indicate Ecuadorian
provinces and Colombian departments) and comparison of morphological characters
between Columnea angulata, C. oribunda, C. tecta, and C. picta.
Columnea angulata J.L.
Clark & F. Tobar
Columnea oribunda F.
Tobar & J.L. Clark
Columnea picta H.
Karst
Columnea tecta J.L.
Clark & L. Clavijo
Calyx lobe margin serrate towards apex serrate entire serrate towards apex
Calyx lobe shape elongate to lanceolate ovate broadly ovate ovate
Corolla posture relative
to calyx
oblique to perpendicular straight straight straight
Corolla tube angulation angulate not angulate not angulate not angulate
Corolla deeply or
shallowly bilabiate
shallowly bilabiate (nearly
tubular)
shallowly bilabiate
(nearly tubular)
deeply bilabiate shallowly bilabiate
(nearly tubular)
Number of owers/axil single (rarely 2–3) 4–6 2–3 2–4
Relative length of
corolla tube vs. calyx
lobes
exceeds length of calyx lobes exceeds length of calyx
lobes
exceeds length of calyx
lobes
equal to or less than
length of calyx lobes
Distribution western Andean slopes
in northern Ecuador
(Cotopaxi, Imbabura,
Pichincha, and Santo
Domingo de los Tsáchilas)
western Andean slopes
of northern Ecuador
(Pichincha)
widespread on the
western Andean slopes
in Colombia & Ecuador
western Andean slopes
of northern Ecuador
(Esmeraldas) and
southern Colombia
(Nariño)

ree new species of Columnea (Gesneriaceae) 73
glabrous; anthers longer than broad, ca. 2 × 1.5 mm, dehiscing by longitudinal slits;
staminode absent; nectary a bilobed dorsal gland, glabrous; ovary superior, densely pi-
lose, 0.2–0.4 × 0.2 cm, style ca. 2.5 cm long, glabrous, stigma included and shallowly
bid. Fruit not observed.
Phenology. is species has been found with owers in two periods: February to
May and August to October.
Etymology. e specic epithet is in reference to the angulate or bent corolla tube.
e corolla is nearly perpendicular to the calyx lobes because of the sigmoid-shaped tube.
Distribution and preliminary assessment of conservation status. Columnea an-
gulata is locally abundant in forests along the western slopes of the Ecuadorian Andes
in the provinces of Cotopaxi, Imbabura, Pichincha, and Santo Domingo de los Tsáchi-
las (Fig. 1) where it grows in mature forests and the shaded understory of recently
cleared forests, from 1500 to 2600 m in elevation. It is especially common along the
old highway between Quito and Santo Domingo. It has been documented in two
protected areas: Reserva Florística-Ecológica Río Guajalito and Bosque Protector Los
Cedros. According to the IUCN Red List criteria (IUCN 2001) for limited geographic
range (B1, less than 20,000 km2) and considering the uncertain future of habitat con-
servation of western Andean forests (B2b, c), Columnea angulata should be listed in the
category Vulnerable (VU).
Comments. Columnea angulata is unique from other Columnea by the posture of
the pendent mature owers where the corolla tubes are oriented oblique to perpendicu-
lar relative to the calyx (Figs 2A, 3B). Another dening character is a constriction at the
base of the corolla tube that makes it appear laterally compressed (Fig. 2C). Columnea
picta and C. angulata are vegetatively similar and grow sympatrically. ese two species
are dierentiated by the presence of deeply bilabiate corolla tubes in Columnea picta
(Fig. 6A) in contrast to the shallowly bilabiate corolla tubes in C. angulata (Fig. 2B, C).
Columnea angulata diers from C. tecta by an elongate corolla tube (vs. corolla tube
that does not exceed the calyx lobes in C. tecta) and single axially owers (rarely 2–3)
in contrast to the abundant clusters of 3–5 axially owers in C. oribunda. Columnea
picta and C. angulata are the two most commonly collected species in this complex and
readily dierentiated by the entire calyx margin in C. picta and serrate calyx margin in
C. angulata.
Specimens examined. E Cotopaxi: cantón Sigchos, parroquia San Fran-
cisco de las Pampas, Bosque Integral Otonga, 0°25.17'S, 79°0.19'W, 1900 m, 26 Jan
2001, J.L. Clark and Muñoz 6125 (QCA, QCNE, SEL, UNA, US); Pichincha: cantón
Quito, parroquia Nono, El Pahuma Orchid Reserve, 17 km east of Nanegalito, 0°1'S,
78°37'W, 1700 m, 17 Apr 2003, J.L. Clark et al. 7648 (QCA, QCNE, SEL, UNA, US);
cantón San Miguel de los Bancos, Mindo Loma Cloud Forest Reserve, km 73.5 via
Calacali-La Independencia, 3 km past the entrance to the village of Mindo, 0°0'44"S,
78°44'29"W, 1800 m, 23 May 2011, J.L. Clark & C. Aulestia 12198 (QCNE, UNA,
US); cantón San Miguel de los Bancos, Las Gralarias Reserve, 1.2 km east of the lodge,
0°05'N, 78°43'W, 1900 m, 15 Aug 2017, F. Tobar, A. Nieto, A. Marcayata & S. Imba
2832 (QCA); cantón San Miguel de los Bancos, Las Gralarias Reserve, Puma trail,

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
74
0°05'N, 78°43'W, 1900 m, 21 May 2018, F. Tobar, F. Richter 3280 (QCA); cantón
San Miguel de los Bancos, Puyucunapi Reserve, cultivada cerca de la casa de la reserva,
0°01'N, 78°41'W, 1800 m, 13 Oct 2019, F. Tobar & M. Gavilanes 3409 (HPUCESI,
QCNE); cantón San Miguel de los Bancos, a 2.6 km al este de San Tadeo en la vía a
Bellavista Lodge, 0°01'N, 78°44'W, 1893 m, 11 Mar 2020, F. Tobar & M. Gavilanes
3475 (QCNE); cantón San Miguel de los Bancos, Puyucunapi Reserve, a 800 m de la
entrada del transecto principal, 0°01'N, 78°41'W, 1995 m, 12 Mar 2020, F. Tobar &
M. Gavilanes 3479 (QCA); Santo Domingo de los Tsáchilas: cantón Santo Domingo
de los Colorados, Bosque Protector Rio Guajalito, located on the old Quito-Santo
Domingo road, between Chiriboga and La Palma, 0°18'50"S, 78°55'35"W, 1796 m,
30 May 2009, J.L. Clark et al. 10968 (NY, QCNE, SEL, US); Reserva Florística-
Ecológica Río Guajalito, km 59 de la carretera antigua Quito-Sto. Domingo de los
Colorados, a 3.5 km al NE de la carretera, 0°13'53"S, 78°48'10"W, 1800–2200 m, 3
Apr 2003, J.L. Clark, N. Muchhala & A. Hoyos 7618 (QCA, QCNE, SEL, UNA, US).
Columnea oribunda Tobar & J.L. Clark, sp. nov.
urn:lsid:ipni.org:names:77219740-1
Figs 4, 5
Diagnosis. Diers from Columnea picta by a nearly tubular corolla (vs. deeply bilabi-
ate corolla). Diers from Columnea angulata by the straight corolla relative to the calyx
(vs. oblique to perpendicular corolla relative to the calyx). Diers from Columnea tecta
by corollas that exceed the length of the calyx lobes (vs. corollas that are equal to or less
than the length of the calyx lobes).
Type. E Pichincha: cantón Pichincha, parroquia Pacto, Bosque Protec-
tor Mashpi, sendero Mashpi Capuchin, 5 km al norte de Lodge, 0°09'N, 78°52'W,
900–1200 m, 18 Jan 2020, F. Tobar, C.H. Graham, T. Santander & E. Guevara 3527
(holotype: QCA; isotypes: QCNE, US).
Description. Facultative epiphyte with dorsiventral shoots to 2–3 m long, sub-
woody, surutescent, glabrescent below, sparsely pilose above; internodes 3–16 cm
near base, then clustered at branch apex. Leaves opposite, strongly anisophyllous, pa-
pyraceous when dry; larger leaf nearly sessile, petioles 0.1–0.2 cm long, pilose; blade
asymmetric, oblanceolate to oblong, 1–28 × 4.8–6.2 cm, base oblique, apex acuminate,
margin serrate, adaxially uniformly dark green with bright red, glabrous, abaxially light
green with bright red apex, sparsely pilose, lateral veins 7–12; smaller leaf sessile, blade
asymmetric, lanceolate 1.4–2.5 × 0.3–0.5 cm, base oblique, apex acuminate, margin
serrate, green with red apex on both surfaces, adaxially glabrous, abaxially sparsely
pilose. Inorescence reduced to axillary clusters of 3–5 owers; peduncles absent or
highly reduced (< 0.2 cm long); bracts 1–2, light green, oblong, 0.5–1.2 × 0.2–0.3cm,
glabrous on both sides. Flowers subtended by elongate pedicels, 2.2–3.3cm long,
sparsely pilose; calyx lobes 5, nearly free, mostly equal in size and shape, dorsal lobe
elongate and slender, 1.7–2.1 × 0.7–1.3 cm, ovate, apex acute, margin serrate, mostly

ree new species of Columnea (Gesneriaceae) 75
Figure 4. Columnea oribunda F. Tobar & J.L. Clark A mature ower B front view of ower during
anthesis C mature fruit D dorsiventral habit A–D from F. Tobar et al. 3527 (A–D from F. Tobar et al.
3527, holotype). Photos by F. Tobar.

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
76
yellow with red splotches in the center, inner and outer surfaces pilose; corolla tubular,
0.6–2.1 cm long, mostly yellow with whitish base, outer and inner surfaces pilose, limb
shallowly bilabiate, corolla lobes 0.3–0.5 × 0.2–0.4 cm, lateral and lower lobes red, up-
per lobes yellow. Androecium of 4 stamens, laments connate at the base and forming
a lament curtain for 0.2–0.3 cm, free portion of laments 1.5–1.9 cm long, minutely
Figure 5. Columnea oribunda F. Tobar & J.L. Clark A mature ower B Lateral view of corolla
C dorsiventral habit D lateral view of calyx and style, showing shallowly bid stigma E corolla dissected
showing laments and ventral view of anthers F lateral view of ovary and nectary gland G dorsal view of
nectary gland. Illustrated by M.J. Gavilanes, based on F. Tobar et al. 3527, holotype.

ree new species of Columnea (Gesneriaceae) 77
Figure 6. Columnea picta H. Karst A lateral view of ower featuring deeply bilobed corolla B ven-
tral view of ower during anthesis C mature ower featuring curled lower lobe D dorsiventral habit
(A, B from J.L. Clark & L. Jost 16301 C from J.L. Clark et al. 15393 D from J.L. Clark, M. Mailloux &
S. Seger 7942). Photos by J.L. Clark.

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
78
pubescent; anthers longer than broad, ca. 0.3 × 0.2 mm, dehiscing by longitudinal slits;
staminode absent; nectary a trilobed dorsal gland, glabrous; ovary superior, densely
pilose, 0.2–0.4 × 0.2 cm, style 1.5–1.8 cm long, minutely pubescent, stigma included
and shallowly bid. Fruit an indehiscent globose white berry. Seeds not observed.
Phenology. Collections of Columnea oribunda are documented with owers be-
tween January and April and between June and October. Fruits have been recorded
during March.
Etymology. e specic epithet refers to axillary clusters of several owers (3–5).
Distribution and preliminary assessment of conservation status. Columnea o-
ribunda is locally abundant in the Mashpi Rainforest Biodiversity Reserve (900–1340
m) and the surrounding roads, a Chocó biogeographic forest relict in northern Ec-
uador. It is likely that additional populations are located in the adjacent provinces
of Imbabura and Esmeraldas. Future research expeditions to unexplored areas of the
Cotacachi Cayapas Ecological Reserve will hopefully result in additional documented
populations of C. oribunda. According to the IUCN Red List criteria (IUCN 2001)
for limited geographic range (B2a, less than ve locations) and considering the un-
certain future of habitat conservation, Columnea oribunda should be listed in the
category Endangered (EN).
Comments. Columnea oribunda is readily distinguished from all other congeners
by the elongate corolla tubes that exceed the length of the calyx lobes (Fig. 4A), in
contrast to the corolla tubes of C. tecta that are equal to or less than the length of the
calyx lobes (Fig. 7C); the corolla posture relative to the calyx that is straight (Fig. 4A),
in contrast to the oblique to perpendicular corolla relative to the calyx of C. angulata
(Fig. 2A); and the axillary clusters of three or more owers (Fig. 4D).
Specimens examined. E Pichincha: cantón Pacto, Mashpi Lodge, transec-
to Mashpi Laguna, a 500 m de la entrada del transecto, 0°09'N, 78°52'W, 880 m, 21
Sep 2017, F. Tobar & A. Nieto 2903 (QCA); cantón Pacto, transecto Mashpi Capuchin,
entrada del transecto, 0°10'N, 78°52'W, 800 m, 20 Mar 2018, F. Tobar, A. Marcayata
& K. Cortez 3161 (QCA); cantón Pacto, km 20, carretero entre La Delicia y el pueblo
de Mashpi, 0°09'N, 78°51'W, 1200 m, 18 Dec 2019, F. Tobar & M. Gavilanes 3509
(QCA); cantón Pichincha, Amagusa Reserve, 1200 m dentro del sendero principal ha-
cia el rio, 0°09'N, 78°51'W, 1213 m, 18 Mar 2018, F. Tobar, C. Poveda, S. Basantes &
M. Gavilanes 3465 (HPUCESI, QCNE); cantón Pacto, Mashpi reserve, road to lodge,
0°09'38"N, 78°50'58"W, 1338 m, 7 Feb 2019, M. Perret & F. Tobar 258 (QCNE).
Columnea tecta J.L. Clark & Clavijo, sp. nov.
urn:lsid:ipni.org:names:77219741-1
Fig. 7
Diagnosis. Diers from Columnea picta by a nearly tubular corolla (vs. deeply bila-
biate corolla) that is equal to or shorter than the calyx lobes (vs. corolla that extends
beyond the calyx lobes).

ree new species of Columnea (Gesneriaceae) 79
Figure 7. Columnea tecta J.L. Clark & Clavijo A mature ower with lateral calyx lobe pulled back
B Oblong white berry C mature ower D dorsiventral habit (A–D from J.L. Clark et al. 13433).
Photos by J.L. Clark.

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
80
Type. E Esmeraldas: cantón San Lorenzo, remnant patch of forest along
highway Ibarra-San Lorenzo, between the towns of Durango and Alto Tambo,
0°57'21"N, 78°33'38"W, 664 m, 3 Jun 2009, J.L. Clark & 2009 Gesneriad Research Ex-
pedition Participants 11104 (holotype: US [3693986]; isotypes: MO, NY, QCNE, SEL).
Description. Facultative epiphyte with dorsiventral shoots to 1.5 m long, sub-
woody, surutescent, glabrescent below, sparsely pilose above; internodes 5–10 cm
near base, then clustered at branch apex. Leaves opposite, strongly anisophyllous, pa-
pyraceous when dry; larger leaf nearly sessile, petioles succulent, 0.3–0.8 cm long, gla-
brous; blade asymmetric, broadly oblanceolate, 7–30 × 3–6.6 cm, base oblique, apex
acuminate, margin serrate, adaxially uniformly green with bright red apex, glabrous,
abaxially light green with bright red apex, sparsely pilose along the venation, lateral
veins 7–14, primary vein bright red, secondary veins red at base and green adaxially;
smaller leaf sessile and often clasping the base of the stem; blade asymmetric, lanceolate
0.5–1.5 × 0.4–0.5 cm, base oblique, apex acuminate, margin serrate, green with red
apex on both sides, adaxially glabrous, abaxially sparsely pilose. Inorescence reduced,
appearing in clusters of 1–4 axially owers; peduncles absent or highly reduced (< 0.2
cm long); bracts 1–2, light green, oblanceolate, 0.7–1 × 0.2–0.3 cm, glabrous on both
sides. Flowers subtended by elongate pedicels, 1.5–2.4 cm long, sparsely pilose, with
enations near the apex; calyx lobes 5, nearly free, mostly equal in size and shape, dorsal
lobe slightly smaller, 1.5–2.3 × 1–2 cm, ovate, apex acute, margin serrate, yellow with
red splotches in the center, inner and outer surfaces sparsely pilose; corolla tubular and
erect, 1–1.9 cm long, outer and inner surfaces pilose, limb shallowly bilabiate, mostly
yellow with red striations on lateral and ventral lobes, corolla lobes 0.2–0.5×0.2–0.4
cm. Androecium of 4 stamens, laments connate at the base for 0.1–0.3 cm and form-
ing a lament curtain, free portion of laments ca. 1.5 cm long, glabrous; anthers
longer than broad, ca. 2 × 1.5 mm, dehiscing by longitudinal slits; staminode absent;
nectary a bilobed dorsal gland, glabrous; ovary superior, densely pilose, 0.2–0.4 × 0.2
cm, style ca. 1.4 cm long, glabrous, stigma included and shallowly bid. Fruit an inde-
hiscent oblong white berry, 1.2 × 0.72 cm.
Phenology. is species was documented with owers in June and May. Fruits
have been recorded in June.
Etymology. e specic epithet tecta refers to the “hidden” or relatively short
corolla tube that does not exceed the length of the calyx lobes, an unusual character in
Columnea.
Distribution and preliminary assessment of conservation status. is species
has not been found in any formally protected areas. According to the IUCN Red List
criteria (IUCN 2001) for limited geographic range (B2a, less than ve locations) and
considering the uncertain future of habitat conservation of western Andean forests,
Columnea tecta should be listed in the category Endangered (EN).
Comments. Columnea tecta is readily distinguished from all other congeners by
relatively short corollas that barely exceed the length of the calyx lobes (Fig. 7). e
corollas of Columnea tecta have limbs that are shallowly bilabiate (Fig. 7A) in contrast
to the deeply bilabiate corollas of Columnea picta (Fig. 6A). Columnea tecta and C. picta

ree new species of Columnea (Gesneriaceae) 81
are vegetatively similar by the presence of a dorsiventral habit with red apices on both
leaf surfaces. e corolla tubes of C. tecta are short (less than the length of the calyx
lobes and shallowly bilabiate) relative to the longer corolla tubes of C. picta (exceed-
ing the length of the calyx lobes and deeply bilabiate). Columnea tecta diers from C.
angulata by a straight orientation of the corolla relative to the calyx (vs. oblique to
perpendicular in C. angulata).
Specimens examined. C Nariño: municipio Barbacoas, corregimiento
El Diviso, western slopes of the Cordillera Occidental, trail from El Diviso towards
Río Güiza, 1°21'21"N, 78°11'45"W, 404 m, 13 May 2013, J.L. Clark, L. Clavijo, O.
Marín & M. Flores 13433 (COL, CUVC); Altaquer to Junín, near Altaquer, 10 May
1972, H. Wiehler, R.L. Dressler, N.H. Williams & N.F. Williams 72222 (SEL).
Acknowledgements
Support for J.L. Clark was provided by the National Science Foundation (DEB-0841958
& DEB-0949169). Support for F. Tobar, M. Perret and C.H. Graham 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 permits for specimen
collection and transportation (Research permit Aves y Conservación N° 007-2018-IC-
FLO-FAU and mobilization permit Aves y Conservación 005-FLO-2019-DPAP-MA).
Maria Jose Gavilanes is acknowledged for the illustrations (Figs 3, 5). Marco Monteros
is acknowledged for producing the distribution map (Fig. 1). We are grateful to Chris-
tian Feuillet (US) for helping us select specic epithets. Oscar Marín, Mauricio Flórez
and the Fundación Ecológica Los Colibríes are acknowledged for facilitating the re-
search expedition that resulted in the discovery of Columnea tecta. We are grateful to
the Mashpi & Amagusa Reserve, Mashpi Lodge, and the Mindo Cloud Forest for sup-
porting our research by providing access to their reserves. We thank Fred R. Barrie and
Alain Chautems for providing valuable feedback on an earlier version of the manuscript.
References
Clark JL, Zimmer EA (2003) A preliminary phylogeny of Alloplectus (Gesneriaceae): Implica-
tions for the evolution of ower resupination. Systematic Botany 28: 365–375.
Clark JL, Funke MM, Duy AM, Smith JF (2012) Phylogeny of a Neotropical clade in the
Gesneriaceae: More tales of convergent evolution. International Journal of Plant Sciences
173(8): 894–916. https://doi.org/10.1086/667229
Clark JL, Skog LE, Boggan JK, Ginzbarg S (2020) Index to names of New World members of
the Gesneriaceae (Subfamilies Sanangoideae and Gesnerioideae). Rheedea 30: 190–256.
https://doi.org/10.22244/rheedea.2020.30.01.14

John L. Clark et al. / PhytoKeys 182: 67–82 (2021)
82
IUCN [Species Survival Commission] (2001) IUCN Red List Categories and Criteria: Version
3.1. IUCN, Gland and Cambridge.
Ogutcen E, Christe D, Nishii K, Salamin N, Möller M, Perret M (2021) Phylogenomics of
Gesneriaceae using targeted capture of nuclear genes. Molecular Phylogenetics and Evolu-
tion 157: e107068. https://doi.org/10.1016/j.ympev.2021.107068
Schulte LJ, Clark JL, Novak SJ, Ooi MT, Smith JF (2014) Paraphyly of Section Stygnanthe
(Columnea, Gesneriaceae) and a revision of the species of section Angustiorae, a new sec-
tion inferred from ITS and chloroplast DNA Data. Systematic Botany 39(2): 613–636.
https://doi.org/10.1600/036364414X680861
Smith JF (2000) Phylogenetic resolution within the tribe Episcieae (Gesneriaceae): Congruence
of ITS and ndhF sequences from parsimony and maximum-likelihood analyses. American
Journal of Botany 87(6): 883–897. https://doi.org/10.2307/2656896
Smith JF, Carroll CL (1997) A cladistic analysis of the tribe Episcieae (Gesneriaceae) based on
ndhF sequences: Origin of morphological characters. Systematic Botany 22(4): 713–724.
https://doi.org/10.2307/2419437
Smith JF, Ooi MT, Schulte LJ, Amaya-Márquez M, Pritchard R, Clark JL (2013) Searching for
monophyly in the subgeneric classication systems of Columnea (Gesneriaceae). Selbyana
31: 126–142.
Weber A (2004) Gesneriaceae. In: Kadereit J (Ed.) e Families and Genera of Vascular Plants.
Vol. 7. Flowering Plants. Dicotyledons. Lamiales (Except Acanthaceae Including Avicen-
niaceae). Springer, Berlin, 63–158. https://doi.org/10.1007/978-3-642-18617-2_8
Weber A, Clark JL, Möller M (2013) A New Formal Classication of Gesneriaceae. Selbyana
31(2): 68–94.
Weber A, Middelton DJ, Clark JL, Möller M (2020) Keys to the infrafamilial taxa and genera
of Gesneriaceae. Rheedea 30: 5–47. https://doi.org/10.22244/rheedea.2020.30.01.02