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Drymonia betancurii (Gesneriaceae), a new species from northwestern Colombia

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A new species of Gesneriaceae from the Pacific slopes of the Colombian Andes is described and illustrated. The new species, Drymonia betancurii, is differentiated from other congeners by the following combination of characters: upper leaf surface with papillose-hispid trichomes, dark green and often covered with white spots; lower surface pitted; and corolla lobes orange-red with white to yellow margins.
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Phytotaxa 221 (1): 077–082
www.mapress.com/phytotaxa/
Copyright © 2015 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Zhi-Qiang Zhang: 3 Jul. 2015; published: 28 Jul. 2015
http://dx.doi.org/10.11646/phytotaxa.221.1.8
77
Drymonia betancurii (Gesneriaceae), a new species from northwestern Colombia
LAURA CLAVIJO1,2 & JOHN L. CLARK1
1Department of Biological Sciences, The University of Alabama, Box 870345, Tuscaloosa, Alabama 35487 USA
2E-mail: lvclavijoromero@crimson.ua.edu
Abstract
A new species of Gesneriaceae from the Pacific slopes of the Colombian Andes is described and illustrated. The new species,
Drymonia betancurii, is differentiated from other congeners by the following combination of characters: upper leaf surface
with papillose-hispid trichomes, dark green and often covered with white spots; lower surface pitted; and corolla lobes or-
ange-red with white to yellow margins.
Resumen
Se describe e ilustra una nueva especie de Gesneriaceae de la vertiente pacífica de los Andes Colombianos. La nueva espe-
cie, Drymonia betancurii, se diferencia de las otras especies del género por la siguiente combinación de caracteres: haz con
indumento papiloso-híspido, verde oscuro y usualmente con manchas blancas; envés con pequeñas depresiones; y lóbulos
de la corola anaranjado-rojo con margen blanca a amarilla.
Introduction
The flowering plant family Gesneriaceae Richad & Jussieu in Candolle (1816: 182) is represented in the Neotropics by
more than 1200 species (Weber et al. 2013). The highest diversity is found in Colombia with 32 genera and over 400
species (Kvist et al. 1998), followed by Ecuador with 29 genera and 240 species (Skog & Kvist 1997), and Brazil with
28 genera and 207 species (Forzza et al. 2010). The third largest genus in the Neotropics is Drymonia Martius (1829:
57) with 75+ species (Möller & Clark 2013) where most of them are in northwestern South America, particularly along
the Pacific slopes of the Andes in Colombia and Ecuador (Clark et al. 2006, Clavijo & Clark 2009). In Colombia the
genus is distributed from sea level to 3000 m, and the highest species richness is in the Tropical rain forest (bp-T) and
the Premontane Rain forests (bp-PM) (Holdridge 1978) at low to mid-elevations (0–1400 m).
Drymonia is one of the most morphologically diverse genera among the members of the neotropical Gesneriaceae
(Clark et al. 2012; Clark et al. 2015), displaying a wide range of habits, such as herbs, subshrubs, shrubs, and lianas that
can be terrestrial, hemiepiphytic, or epiphytic (facultative or obligate). Corolla shapes can be campanulate, funnelform,
tubular, laterally compressed, urceolate, or hypocyrtoid (with a ventral pouch). Fruit types range from fleshy bivalved
capsules to indehiscent berries. The most distinctive characteristic of Drymonia is the presence of basal poricidal
anther dehiscence (Fig. 2G), which is lost in several lineages within the genus (Clark et al. 2006; Clark et al. 2015).
Recent transfers from Nautilocalyx Linden ex Hanstein (1854: 207) (Clark et al. 2011) and Alloplectus Martius (1829:
55) (Clark 2005) have been supported by molecular sequence data that strongly support a monophyletic Drymonia
represented by highly divergent morphologies in the above mentioned vegetative and reproductive characters.
High humidity, and a heterogeneous landscape associated with the Andean orogeny have promoted the
diversification of several plant lineages on the Pacific slopes of the Andes (Gentry 1989), which are considered among
the most biologically diverse regions on the planet with numerous endemic taxa (Gentry 1982, 1989, Mittermeier et al.
2004). Recent expeditions to poorly explored areas on the Pacific slopes of the Andes have resulted in the discovery
and description of several new species of Gesneriaceae (e.g., Amaya-Márquez 2010, Amaya-Márquez & Marín-Gómez
2012, Amaya-Márquez & Smith 2012, Smith et al. 2013), including three new species of Drymonia (Clavijo & Clark
2010, 2012, 2014). In this paper we describe a new species from the Pacific slopes of the Colombian Andes and discuss
its morphological similarities with other congeners.
CLAVIJO & CLARK
78 Phytotaxa 221 (1) © 2015 Magnolia Press
Taxonomy
Drymonia betancurii Clavijo & J.L. Clark, sp. nov. (Figs. 1 & 2)
Diagnosis: Differs from other species of Drymonia by the presence of papillose-hispid trichomes on upper leaf surface and pitted on the
lower surface; upper leaf surface dark green, often covered with white spots; calyx lobes densely pilose on both surfaces; corolla limb
orange-red with white to yellow lobe margins; style with glandular trichomes.
Type:—COLOMBIA. Antioquia: Municipio Frontino, Parque Nacional Natural Las Orquídeas, vereda Venados Abajo, sector de Venados,
sitio Arenales. 6º32’25.2”N, 76º18’38.7”W, 950–1000 m, 26 July 2011 (fl), J. Betancur, P. Pedraza-Peñalosa, M.F. González, R.
Arévalo, D. Sanín, A. Zuluaga, J. Serna & A. Duque 15434 (holotype COL!, isotypes HUA!, NY!).
Herb, subshrub, or liana; terrestrial, hemiepiphyte, or epiphyte. Stem prostrate, scandent or appressed to tree, herbaceous
to subwoody, with adventitious roots, branched, subquadrangular in cross-section, 2.5–3.9 mm in diameter, scarcely
pilose to pilose basally, pilose to lanate apically; trichomes whitish, 1–2 mm long, unbranched; internodes 4.4–10.2 cm
long. Leaves opposite, evenly spaced, decussate, subequal in a pair; petiole 0.5–2.5 cm long, terete in cross-section,
green, with a gland at the base, densely pilose to lanate, trichomes 0.8–1.6 mm long; blade ovate to oblong, 3.1–8.1
× 2.5–6.0 cm, cartaceous, upper surface dark green, usually with white spots along the veins and sometimes with the
venation light green, lower surface purple with light green venation, apex acuminate, base rounded to truncate, usually
oblique, margin crenate to serrulate, upper surface papillose-hispid, lower surface pitted; 5–6 (–7) pairs of secondary
veins, only evident abaxially, main vein sparsely pilose adaxially, densely pilose abaxially, secondary veins glabrate
adaxially, densely pilose abaxially, higher order of venation only evident abaxially, pilose. Inflorescence reduced to an
axillary solitary flower; bracts absent; flowers protandrous. Pedicel perpendicular or oblique relative to stem, 8–29 mm
long, green, densely pilose. Calyx green to green suffused with red, membranous, venation conspicuous; calyx lobes
5, 4 nearly equal, dorsal lobe slightly reduced, free to nearly free, when nearly free fused at base for 2–5 mm, apex
acute, base rounded to truncate, margin serrate, reflexed when in bud, pilose, dense at base on both surfaces; ventral
and lateral lobes 13–26 × 7–18 mm, rhombic, ovate or oblong, dorsal lobe 12–19 × 6–13 mm, ovate to oblong. Corolla
zygomorphic, 4.7–5 cm long, oblique to perpendicular relative to calyx, infundibuliform; tube constricted at base, 2.8–
3.2 cm long, 1.3–1.6 cm wide, outer surface white, sometimes pink ventrally, pilose, inner surface orange-red; base
gibbous, 6–5 mm in diameter, gibbosity 6–7 mm long; throat 17–19 mm in diameter, outer surface white and pilose,
inner surface orange-red with short glandular trichomes dorsally; corolla lobes subequal, orange-red, margin white to
yellow, 9–12 × 9–14 mm, orbicular, apex rounded, margin slightly erose, glabrous, ventral lobe slightly larger, lateral
and dorsal lobes spreading, sometimes reflexed. Androecium of 4 stamens, didynamous, filaments 26–32 mm long,
adnate to the corolla tube for 13–15 mm, white, glabrous, coiling after anthesis; staminode absent; anthers oblong,
coherent by the lateral walls, dehiscence by basal pores that develop into longitudinal slits, 6–7 × 2–3 mm. Gynoecium
with a single dorsal nectary gland, ovate, apex acute, 2–3 mm long, glabrous; ovary superior, 5–6 × 4–5 mm, ovate,
orange, sericeous; style 17–19 mm long with sparse glandular trichomes, stigma stomatomorphic. Fruit not observed.
Distribution and habitat:—Drymonia betancurii is endemic to Colombia and is only known from the Pacific
slopes of the Cordillera Occidental in the departments of Antioquia and Chocó, between 480 and 1000 m. Drymonia
betancurii grows in scattered populations in open areas that are often near forest edges or in the shade of intact
forests.
Phenology:—Flowers recorded from March to July; fruits not seen.
Etymology:—This species is named in honor of the Colombian botanist Julio Betancur, expert on Bromeliaceae
and Heliconiaceae of Colombia, who has made monumental contributions to the knowledge of Colombian flora and
has mentored several generations of Colombian botanists.
Additional specimens examined (paratypes):COLOMBIA. Antioquia: Municipio Frontino, Corregimiento
de Encarnación, Parque Nacional Natural Las Orquídeas, sector Venados, bosques cercanos a la cabaña de Parques
Nacionales, 11 April 2011, J. Betancur, P. Pedraza-Peñalosa, J.M. Vélez-Puerta, A. Orjuela & A. Duque 15165 (COL!,
HUA!, NY!); Parque Nacional Natural Las Orquídeas, sector Venados, vereda Venados Abajo, sitio La Esperanza,
cuenca de la quebrada Arenales, 6º42’6.8’’N, 76º18’46.03’’W, 880–920 m, 29 July 2011, P. Pedraza-Peñalosa, J.
Betancur, M.F. González, R. Arévalo, D. Sanín, A. Zuluaga, A. Duque & J. Serna 2443 (COL!, NY!); Parque Nacional
Natural Las Orquídeas, sector Venados, vereda Venados Abajo, sitio La Esperanza, cuenca de la quebrada Arenales,
6º42’6.8’’N, 76º18’46.03’’W, 880–920 m, 20 July 2011, P. Pedraza-Peñalosa, J. Betancur, M.F. González, R. Arévalo, D.
Sanín, A. Zuluaga, A. Duque & J. Serna 2486 (COL!, NY!, UNA!); vereda Cruces, sitio Piñares, camino a Perdidas,
DRYMONIA BETANCURII (GESNERIACEAE) Phytotaxa 221 (1) © 2015 Magnolia Press 79
FIGURE 1. Drymonia betancurii. A. Lower leaf surface. B. Upper leaf surface with inset showing the papillose-hispid indument. C. Lateral
view of flower. D. Lateral view of flower showing androecium and gynoecium. E. Gynoecium showing single dorsal nectary gland, glandular
trichomes on style, and stomatomorphic stigma. F. Habit. (from the holotype, J. Betancur 15434, and P. Pedraza-Peñalosa 2443).
CLAVIJO & CLARK
80 Phytotaxa 221 (1) © 2015 Magnolia Press
FIGURE 2. Drymonia betancurii. A–B. Habit. C. Lateral view of flower. D. Upper leaf surface. E. Front view of flower. F. Front view of
flower showing anthers and glandular hairs. G. Anther (from P. Pedraza-Peñalosa 2443).
DRYMONIA BETANCURII (GESNERIACEAE) Phytotaxa 221 (1) © 2015 Magnolia Press 81
poco después de la escuela La Esperanza, orilla izquierda del río Calles, Parque Nacional Natural Las Orquídeas,
bosque poco perturbado, 6º28’35.5’’N, 76º19’39.5’’W, 980 m, 3 May 2013, S.E. Hoyos-Gómez, J. Betancur, R. Arévalo,
M.F. González, M.S. Jaimes, F. Gómez, A. Duque, C. Rivera, W. Quinceno & F. Pino 2289 (COL!, NY!); Municipio
Urrao, vereda Cruces, camino al río Penderisco desde la escuela La Esperanza, alrededores de la quebrada La Balsora,
márgen izquierda del río Calles, Parque Nacional Natural Las Orquídeas, bosque al lado de una quebrada y potrero al
lado del camino, 6º27’58’’N, 76º19’19.11’’W, 880–900 m, 7 May 2013, J. Betancur, R. Arévalo, S.E. Hoyos-Gómez,
M.S. Jaimes, A. Duque, W. Quinceno & F. Pino (COL!, NY!). Chocó: Along road from Quibdó to Tutunendo, 1 hour
past Tutunendo, vereda 21, at Alto de Veinte, hacienda of Ruben Jaramillo, 1/2 hour walk down hill from roadside; rain
forest 18 km NE of Tutunendo, 480 m, 15 March 1987, H. Wiehler 8730B (SEL!).
Drymonia betancurii is similar to D. variegata Uribe (1952: 1), but they are readily differentiated when fertile.
The two species are similar in their diverse habits and foliage (e.g., dark green, papillose-hispid trichomes on upper
blade surface, and pitted and purple lower blade surface). Drymonia betancurii is differentiated from D. variegata by
the presence of white spots on the upper surface, although, at least one population of D. betancurii have light green
venation and lack white spots. When sterile, the non-spotted populations of D. betancurii are similar to D. variegata.
The two species are differentiated by the following characters in D. betancurii: petiole 0.5–2.5 cm long (vs. ca. 5 cm
long); blades ovate to oblong to 8.1 cm long (vs. lanceolate to elliptic to 15.2 cm long); calyx lobes rhombic, ovate or
oblong, 7–18 mm wide (vs. lanceolate, 1–6 mm wide); and corolla lobes orange with white to yellow margin (vs. white
with maroon or purple lines). Drymonia variegata is widely distributed from Panama to Ecuador, while D. betancurii
is restricted to the Pacific slopes of the Colombian Andes in the departments of Antioquia and Chocó, where these two
species are sympatric. Hans Wiehler and the Gesneriad Research Foundation team made the first known collection of
D. betancurii (H. Wiehler 8730B) during their expedition to Colombia in 1987, and although they noticed the unusual
white spots on the leaves they identified the collection as D. variegata (Milewiski 1987).
Drymonia betancurii and Drymonia droseroides J.L. Clark & Clavijo (2010: 190) are similar because of their
dark green and bullate foliage. The two species are differentiated by the following characters in D. betancurii: habit
of prostrate herb, liana, epiphyte or hemiepiphytic shrub (vs. terrestrial shrub); petioles 0.5–2.5 cm long (vs. 6–7.7 cm
long); leaves 3.1–8.1 × 2.5–6.0 cm (vs. 16.5–21.1 × 11.6–13 cm); calyx lobes rhombic, ovate or oblong and pilose (vs.
spatulate with glandular hairs); and an infundibuliform corolla with an orange-red limb (vs. campanulate with limb
yellow to pink).
The stem indument and the habit of D. betancurii are similar to D. alloplectoides Hanstein (1865: 358). However,
D. betancurii is differentiated by dark green, papillose-hispid leaves (vs. green, villous), calyx serrate (vs. entire),
corolla lobes orange-red with the margin white to yellow (vs. white), and ventral corolla lobe slightly erose (vs.
fimbriate).
Acknowledgments
We thank Mateo Jaimes (COL), Julio Betancur (COL) and Paola Pedraza-Peñalosa (NY) for providing access to
collections, and to Sue Blackshear for the illustration. We acknowledge the following botanists for providing images:
Paola Pedraza-Peñalosa (Figs. 2AE), David Sanín (Fig. 2B), María Fernanda González (Fig. 2C), and Mateo Jaimes
(Figs. 2DF). Collections of this species were made during expeditions for the project “Flora of Las Orquídeas National
Park” funded by the National Science Foundation (DEB 1020623) to Paola Pedraza-Peñalosa and Julio Betancur.
Steve Ginzbarg (UNA) and two anonymous reviewers are gratefully acknowledged for providing helpful feedback.
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... The tropical Andes represents one of the most biodiverse regions on the planet with 30,000-35,000 species of vascular plants ( Rodríguez-Maecha et al., 2004), of which, more than on the Pacific slopes, along with the ongoing taxonomic revision of the genus Drymonia, have resulted in the discovery and description of several new Drymonia species (Clavijo & Clark, 2010, 2012, 2014, 2015a, 2015b, including the species that is described and illustrated here. Terrestrial herb, up to 80 cm tall. ...
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Sparked by the publication of large phylogenetic studies and major generic redefinitions in the Gesneriaceae, we review this growing body of molecular studies on the family. Different aspects of molecular data and their use in Gesneriaceae systematics are considered including conceptual challenges on the phylogenetic work undertaken to date as well as an overview of taxon sampling in the family. Molecular data are currently available for 70 of 72 recognized New World genera and 64 of 68 Old World genera. Many of the smaller genera and some of the larger genera are completely sampled. Current knowledge of tribal and generic delineations and relationships among the New World genera is relatively advanced. In contrast, intergeneric relationships and tribal arrangements are mostly unresolved for the Old World genera. In this paper we illustrate and summarize the published phylogenetic work in composite phylogenies with an emphasis on the most pertinent and accurate molecular systematic studies. This paper provides the molecular-based background for a new formal classification of the family Gesneriaceae.
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