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Columnea antennifera, a new species of Gesneriaceae from the Cordillera Central of the Columbian Andes

Authors:
  • Lawrenceville School
COLUMNEA ANTENNIFERA, A NEW SPECIES OF GESNERIACEAE
FROM THE CORDILLERA CENTRAL OF THE COLOMBIAN ANDES
John L. Clark Laura Clavijo
Department of Biological Sciences Department of Biological Sciences
Box 870345 Box 870345
The University of Alabama The University of Alabama
Tuscaloosa, Alabama 35487, U.S.A. Tuscaloosa, Alabama 35487, U.S.A.
jlc@ua.edu lauriclav@gmail.com
abstr act
A recent e xpedition t o the Cordil lera Centra l of the Colombia n Andes resu lted in the di scovery of a new s pecies of Columnea (Gesneriaceae,
tribe Episcie ae). The new species, Columnea antennifera J.L. Clark & Clavijo, is distinguished from other congeners by t he presence of
elongate c orolla append ages that a lternate w ith the corol la lobes, a nisophyllo us leaves, an d a scandent obl igate epiphyt ic habit. A di scussio n
and images are provided to different iate Columnea antennifera from C. dissimilis, C. fil amentosa, and C. rosea. The characterist ic of elongate
corolla a ppendages i s discuss ed and Columnea antennifera is compared w ith other ta xa that shar e this unusu al feature.
resum en
Una reciente expedición al norte de la Cordillera Central de los Andes Colombianos permitió el descubrimiento de un a nueva especie de
Columnea (Gesneriace ae, tribu Episicieae); Columnea antennifera J.L. Clark & C lavijo, que se di stingue de ot ras espe cies del género p or la
prese ncia de apéndic es alargado s alternos a los l óbulos de la corola , hojas anisóf ilas y hábit o epífito esc andente obliga do. Se presenta n foto-
graf ías y una discusión para dife renciar Columnea antennifera de C. dissimilis , C. filam entosa y C. rosea. Se discut e la presencia de apéndices
alar gados en la corola d e Columnea antennifera y se compara con ot ras espe cies que compar ten este car ácter inusua l.
key words: Columnea, Colombia, Episcieae, Gesner iaceae, Taxonomy
int roduc tion
The genus Columnea L. is prima rily epiphytic a nd belongs to the New World subfami ly Gesnerioideae a nd tribe
Episcieae. Columnea ranges from Mexico south to Bolivia and is most diverse in the northern Andes of Colom-
bia and Ecuador. With over 200 species, Columnea is the la rgest genus in the subfamily Gesnerioideae (Burtt &
Wiehler 1995; Weber 2004; Skog & B oggan 2006). The genus is disting uished from other clo sely related genera
by an indehiscent berry instead of a fleshy bivalved capsule.
Columnea antennifera J.L. Clark & Clavijo was discovered during a 2012 research expedition to the Co-
lombian department of Antioquia in the Cordillera Central of the northern Andes. A remarkable character of
Columnea antennifera is the presence of five elongate appendages near the corolla sinuses. The presence of co-
rolla appendages and where they appear has been discussed in numerous artificial classifications of groups
now recognized as Columnea, such as the section Ortholoma Benth. and the genus Trichantha Hook. (Morton
1963, 1971; Morley 1976; Smith 1994). Corolla appendages have not been thoroughly evaluated in a phyloge-
netic context and most likely this character is convergent within Columnea.
Columnea antennifera J.L. Clark & Clavijo, sp. nov. (Fig. 1) type: COLOMBIA. antioquia: Municip io Valdiv ia, Cordille ra
Central, road Ventanas to Br iceño, before the quebrada El Oro, 07°05'20"N, 75°29'20"W, 1802 m, 19 May 2012 (fl), J.L. Clark, J.
Ander son, L. Clavijo, M. Ma zo & D. Suescún 13036 (hoLotype: COL; isotypes: BRIT, HUA, MO, NY, UNA, US).
Diffe rs from all othe r Columnea by the combin ation of the pre sence of broad ca lyx lobes, elong ate appendage s near corolla si nuses, oblig ate
scand ent epiphytic h abit, and strong ly anisophyl lous opposite le aves.
Obligate scandent epiphytic climber; stems elongate and horizontal, 2–3 m long, suffrutescent, glabrescent
below, sparsely pilose above. Leaves opposite, strongly anisophyllous in a pair; larger leaf with petioles terete,
4–10 mm long, blade coriaceous when dry, elliptic to oblong, 3–12 × 1.3–3.4 cm, base rounded to oblique,
J. Bot. Res. Inst. Texas 6(2): 385 – 390. 2012
386 Journal of the Botanical Research Institute of Texas 6(2)
Fig. 1. Columnea antennifera J.L. Clark & Clavijo. A. Lateral view of flower. B. Front view of flower. C. Immature flower showing corolla appendages. D.
Habit showing elongate stems and strongly anisophyllous leaf arrangement. (A–D from the holotype, J.L. Clark et al. 13036).
Clark and Clavijo, A new Columnea from Colombia 387
sometimes asymmetrical, apex acute, margin entire, adaxially shiny green, abaxially light green, sparsely pi-
lose on upper surface and densely pilose on lower surface; smaller leaf greatly reduced relative to larger leaf,
nearly sessile, orbicular to ovate, 1–2.7 × 0.5–1.5 cm, base rounded to cordate, apex acute, margin entire, sur-
faces and vestiture similar to larger leaf. Flowers solitary and erect; pedicels 1.3–2 cm long, red, pilose; calyx
2–3.5 cm long, uniformly bright red, inside and outside sparsely pilose, inside lanate at the base, lobes 5, erect
at anthesis, each lobe tightly appressed to adjacent lobe and folded lengthwise, ovate, apex broadly acuminate,
margin entire, 4 lobes nearly equal 0.7–1 × 1.2–1.5 cm, fused at the base for 1.5–2.6 cm, 5th lobe (dorsal) slight-
ly smaller, fused at the base for 1.1–1.5 cm; corolla tubular, posture erect in calyx, 3.0– 4.5 × 0.6–1.0 cm; outside
uniformly bright red and tomentose, internally glabrate, lobes bright yellow, appressed, 5–6 × 2–2.5 mm,
ovate; corolla appendages present in immature and mature flowers, located in each sinus alternate the corolla
lobes, 14–20 mm long, pilose, bright yellow with or without a dark spot at the base; stamens 4, didynamous,
included; filaments 12–15 mm long, coiled after anthesis, connate and adnate to the base of the corolla tube,
glabrous; anthers conn ate, longer than broad, 2–2.5 × 1–1.5 mm, dehiscing by longitudinal slits; staminode not
seen; nectary a dorsal gland, glabrous; ovary superior, lanate, ca. 5 × 4 mm, style ca. 30 mm long, glabrous,
stigma included and capitate. Fruits not seen.
Columnea antennifera is morphologically similar to C. dissimilis C.V. Morton (Fig. 2, A–D). These two spe-
cies are easily differentiated by the elongate corolla appendages in Columnea antennifera (Fig. 1) in contrast to
the relatively short corolla appendages in C. dissimilis (Fig. 2A). The corolla appendages are developed in im-
mature flowers of Columnea ante nnifera (Fig. 1C). In contrast, the corolla appendages in Columnea dissimilis are
either absent or significantly reduced when the flowers are immature (Fig. 2B). Another species that has co-
rolla appendages and is morphologically similar to C. antennifera is C. filamentosa (Figs. 2E, F). These two spe-
cies are readily differentiated by the uniformly red corolla in Columnea filamentosa (Fig. 2E) in contrast to the
red corolla with bright yellow lobes in C. antennifera (Fig. 1). Vegetatively these two species are differentiated
by the isophyllous leaf a rrangement in Columnea filamentosa in contrast to t he strongly anisophyllous C. ante n-
nifera. A third species that has corolla appendages is Columnea rosea (C.V. Morton) C.V. Morton (Fig. 3E). How-
ever, the calyx lobes of Columnea rosea are deeply serrate to fimbriate (Fig. 3E) in contrast to the entire calyx
margins of C. antennifera (Fig. 1).
Distribution and habitat.—Columnea antennifera is known from the northern Cordillera Central of the
Colombian Andes in the department of Antioquia from montane forests (1800 m). Herbarium collections of
Columnea antennifera were not seen during recent visits to the National University of Colombia (COL) or the
University of Antioquia (HUA), but an additional population was observed and photographed near the type
locality between Yarumal and Ventanas during a 1996 field expedition by Günter Gerlach from the Munich
Botanical Garden (Botanischer Garten München-Nymphenburg).
Etymology.—The specific epithet, antennifera, refers to the resemblance of the elongate appendages at the
apex of the corolla tube to insect antennae.
Classification.— Columnea antennifera appears to belong to section Ortholoma Benth. because of the pres-
ence of corolla appendages and an obligate epiphytic habit. However, the traditional sectional classification of
Columnea is artificial and arbitrary. As an example, the section Ortholoma has been recognized at the generic
level as Trichantha Hook. by previous authors (Morton 1963; Wiehler 1973, 1975). A monographic revision of
Trichantha by Morton (1963) was followed by another paper by the same author (Morton 1971) with a reduc-
tion of all species recognized as Trichantha to Columnea. The type species for Trichantha is Columnea minor
(Hook.) Hanst. and is characterized by the presence of appendages at the sinuses of the corolla (Fig. 3C, D). It
is important to note that corolla appendages are not a unifying character for section Ortholoma or genus
Trichantha. The type species for section Ortholoma is Columnea anisophylla DC., which lacks corolla append-
ages as do many other species that have been assigned to t his section. The traditional sectional classification of
Columnea has been shown to be artificial because many sections do not represent monophyletic lineages
(Smith 1994; Smith & Sytsma 1994; Clark et al. 2006). A revised sectional classification system based on mo-
lecular sequence data is currently a collaborative research focus by numerous authors (e.g., James Smith, John
L. Clark, Lacie Schulte and others).
388 Journal of the Botanical Research Institute of Texas 6(2)
Fig. 2. Columnea dissimilis C.V. Morton (A–D) and C. filamentosa L.E. Skog (E–F). A. Front view of flower showing corolla appendages. B. Lateral view of
immature flower. C. Habit showing anisophyllous leaf arrangement. D. Lateral view of flower. E. Lateral view of flower. F. Mature calyx showing dorsal
nectary gland. (Photos A & D from J.L. Clark 8629; B from J.L. Clark & J. de Gracia 12451; C from J.L. Clark & A. Zapata 12495; D & F from H. Wiehler et al.
1631 field collection that was cultivated and then vouchered for the holotype).
Clark and Clavijo, A new Columnea from Colombia 389
Fig. 3. Variation in corolla appendages present in Columnea. A & B. Columnea coronata Amaya, L.E. Skog & L.P. Kvist. C & D. Columnea minor (Hook.)
Hanst. E. Columnea rosea (C.V. Morton) C.V. Morton. F. Columnea filifera (Wiehler) L.E. Skog & L.P. Kvist. (Photos A & B from J.L. Clark et al. 12990; C from
J.L. Clark et al. 10870; D from J.L. Clark et al. 9647; E from J. Betancur 12394; F from J.L. Clark et al. 7140).
390 Journal of the Botanical Research Institute of Texas 6(2)
Corolla appendages vs. corolla lobes.—The presence of corolla appendages is widespread in Columnea and
this character is often not accurately distinguished from corolla lobes. Various taxa from different sections
have corolla appendages a nd this char acter is probably convergent w ithin Columnea. For example, Columnea fi-
lifera (Fig. 3F) has been assigned to section Collandra Lem. (Kvist & Skog 1993) and genus Dalbergaria Tu ss ac.
(Wiehler 1992; Kvist & Skog 2004) because of subsessile leaves, dorsiventral shoots, and a facultative epiphyt-
ic habit. A recently described species, Columnea coronata Amaya, L.E. Skog & L. P. Kvist, was assigned to sec-
tion Collandra (Amaya et al. 2004), where the specific epithet refers to a “corona” at the apex of the corolla tube.
More accurately, the “corona” in Columnea coronata is homologous to reduced corolla lobes like those found
in Columnea antennifera. Thus, what Amaya et al. (2004) referred to as “petals” in the description of Columnea
coronata are actually appendages that appear petaloid (Fig. 3A) and what was described in Amaya et al. (2004)
as the “corona” is homologous to reduced corolla lob es. Another species that has reduced corolla lobes and pet-
aloid appendages is Columnea filifera (Fig. 3F). The petaloid appendages in Columnea filifera (Fig. 3F) are visible
in the field and in photographs, but the corolla lobes are only visible with a hand lens or microscope. Phyloge-
netic studies on the evolution of corolla appendages and their presence in numerous lineages of Columnea will
play an important role in understanding their function, homology, and role in plant-pollinator interactions.
acknowl edg men ts
This study was supported by funds from the National Science Foundation (DEB-841958 and DEB-0949169).
We thank Christian Feuillet (US) and William R. Anderson (MICH) for help in selecting the specific epithet;
Alain Chautems (G), Laurence E. Skog (US), and Jeremy Keene (BHO) for providing helpful reviews of the
manuscript. Our 2012 research expedition to Colombia was a tremendous success because of logistical sup-
port from Álvaro Idárraga (HUA), Felipe Cardona (HUA), Julio Betancur (COL), Álvaro Cogollo (JAUM), and
Diego Suescún (JAUM). We thank Günter Gerlach from the Munich Botanical Garden (Botanischer Garten
München-Nymphenburg) for sharing his observations and images of Columnea antennifera. We gratefully ac-
knowledge Norris H. Williams (University of Florida) for his carefully curated slide collection that included
field images of voucher specimens (e.g., Columnea filamentosa – Fig. 2 E, F) from a 1972 research expedition to
Colombia wit h Hans Wiehler. We thank Julio Betancur from the Universidad Nacional de Colombia (COL) for
providing images of Columnea rosea (Fig. 3E).
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... años el conocimiento de las gesneriáceas en Colombia ha incrementado debido a las exploraciones recientes y la descripción de alrededor de 20 nuevas especies (Clavijo y Clark, 2010; Amaya-Márquez y Marín-Gómez, 2012; Clark y Clavijo, 2012; Mora y Clark, 2012; Amaya-Márquez y Smith, 2013; Amaya-Márquez et al., 2013; Smith, et al., 2013; Clavijo y Clark, 2014; Rodas y Clark, 2014), aunque aún se desconocen muchos aspectos de su ecología y distribución. Las Gesneriáceas se distinguen de otros miembros del orden Lamiales por las siguientes sinapomorfias: corola con cinco lóbulos, placentación parietal, ovarios uniloculares bicarpelados, inflorescencia en cimas y semillas pequeñas con presencia de un endospermo (Smith y Carroll, 1997). ...
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A key, typification and synonymy of the sections in the genus Columnea L. (Gesneriaceae) Contr
  • B Morley
morley, B. 1976. A key, typification and synonymy of the sections in the genus Columnea L. (Gesneriaceae) Contr. Natl. Bot. Gard. Glasnevin 1:1–11.
A new classification of the Western Hemisphere Gesneriaceae
  • L E Skog
  • J K And
  • Boggan
Skog, L.E. And J.K. BoggAn. 2006. A new classification of the Western Hemisphere Gesneriaceae. Gesneriads 56:12-17.