TWO NEW SPECIES AND TWO NEW VARIETIES OF COLUMNEA ( GESNERIACEAE )

In this paper two new species of Gesneriaceae (genus Columnea) are described and illustrated. Columnea chocoensis is distributed in the Colombian departments of Chocó and Valle del Cauca, while its variety, C. chocoensis var. altaquerensis is restricted to the Department of Nariño, Colombia. Columnea stilesiana was found in La Serranía de Los Paraguas located in the Cordillera Occidental between the Chocó and Valle del Cauca Departments in Colombia. Further, Columnea archidonae is here considered a variety of C. ericae; Columnea ericae var. archidonae is distributed in Colombia and Ecuador, in forests at elevations higher than those in which the typical variety is found.


INTRODUCTION
Taxonomic novelties and nomenclatural changes have arisen during the process of studying material for the taxonomic revision of Columnea L. section Collandra Lem.(Benth.)(Gesneriaceae).Columnea is the most diverse genus of the neotropical Gesneriaceae with more than 205 species (Möller & Clark 2013), and more than 80 species in Colombia, with the Chocó biogeographical region contributing ca.44% of that diversity (Rangel-Churrio & Rivera-Díaz 2004); recent reports of new species for this region (see Amaya-Márquez et al. 2015) as well as the new species that we report here, support this geographic patterns of diversity.Recent molecular studies have shown that Columnea is a monophyletic group, while the subgeneric classification is not fully supported (Smith et al. 2013).In this paper two species of Columnea, C. stilesiana and C. chocoensis, as well as the variety C. chocoensis var.altaquerensis are reported new for science; the taxonomical placement of these species in section Collandra is discussed.An examination of Columnea ericae Mansf.indicates a wider intraspecific morphological variation, as well as a wider range of geographical distribution for this species.Therefore, a new variety of C. ericae Mansf. is recognized.
Etymology: The name refers to Altaquer where the Natural Reserve of the Ñambí River is located (Department of Nariño, Colombia), and acknowledges the efforts made by local people to preserve the natural patrimony and the high biological diversity harbored in the area.Columnea chocoensis var.chocoensis and var.altaquerensis both exhibit the floral mechanism of dynamic herkogamy associated with protandry, which is known to occur in other species of Columnea (Morley 1974; Amaya-Márquez ined.).At the end of the male phase, the filaments usually coil and pull the anthers inside at the base of the corolla, creating an efficient physical separation between the female and male parts.After anthesis in both varieties of C. chocoensis the stamens are no longer functional and they bend down remaining outside the corolla (Figure 3E).Columnea chocoensis var.altaquerensis has been more widely collected than the typical variety, however all the collections come from the Natural Reserve Río Ñambi, suggesting that this variety might be endemic to the region, or that more botanical exploration is required in the Nariño and Cauca departments.(5-10-celled trichomes) with sparse white, setulose trichomes, abaxially green, sparsely sericeous (5-7-celled trichomes) with sparse white setulose trichomes.Inflorescence fasciculate (cymose), with 1-3 flowers in the axil of the larger leaf; bracts 2-5, yellow, lanceolate, 0.8-2.3× 0.1-0.6 cm, sericeous on both faces.Flowers pedicellate, pedicels 0.6-3.8cm long, densely golden-sericeous (10-14-celled trichomes).Calyx pale green, yellow, or yellow with a red midline on each sepal; sepals free, unequal, lanceolate, 1.7-3 × 0.3-0.4cm, adaxially and abaxially sericeous, margin dentate with 3-4 tiny teeth per side.

Etymology:
The species name 'ericae' honors the collector of the holotype, Miss Erica Heinrichs, who collected plants in Ecuador during the 1930s.

Phenology:
The examined specimens show that this species produces flowers all year round; fruits were recorded on a specimen collected in June.

Distinctive characters
Columnea ericae var.ericae can be recognized by having the following traits: (1) a sericeous dorsiventral vegetative shoot, with short internodes; (2) the larger leaf in a pair oblanceolate, adaxially green, abaxially green less often reddish with contrasting red or deep purple in the apical 1/3 or 1/5 of the foliar area, 11 veins (rarely 7-14); (3) inflorescences with two to three pedicellate flowers usually adpressed to the ventral side of the vegetative shoot, bracts lanceolate, 1-3.5 × 0.2-0.8cm; and (4) pedicels densely sericeous, 0.6-3.8cm long, corolla erect in the calyx, limb bilabiate, yellow with two deep purple vertical stripes along both sides of the ventral lobe, and a nectary of one dorsal gland (Figures 6 and  7).This variety is similar to C. ericae var.archidonae, and to C. stilesiana, although it can be differentiated from them by the shorter pedicel, the corolla color, and additional differences presented in Table 1.8).

Columnea ericae
Columnea ericae var.archidonae differs from the typical variety by having smaller leaves, bracts, and corollas, while the internodes and the pedicels are longer.The corolla is oblique in the calyx, homogenously yellow (rarely with purple spots), remarkably curved dorsally, the red marks below the larger leaf do not extend to the apex, the spots do not have a defined shape, and are distributed along the arcs subtended by the two apical secondary veins; the flowers often present a staminode; the nectary consists of 2 (rarely 4 glands).In general, the flowers of this variety are more exposed, i.e., they are not adpressed to the vegetative shoot (Figure 8).

Etymology:
The epithet used by José    Columnea ericae var.archidonae was initially described as Columnea archidonae (Cuatrecasas 1935), later it was reduced to the synonymy of C. ericae Mansf.(Kvist & Skog 1993)  and it has a galea.Therefore, the taxonomic placement of the species described here might be temporal, waiting for molecular phylogenies that include these or similar species in their analysis.

Figure 1 .
Figure 1.Columnea chocoensis M. Amaya & L.E.Skog var.chocoensis A. Habit.B. Abaxial side of the larger leaf at a pair showing the veins.Note the base is slightly oblique and cordate, and the indumentum is denser toward one of the sides along the main vein.C. Detail of the abaxial indumenta.D. Bracts, pedicel, and calyx.Note the style is oblique to the ovary.E. Corolla of a flower in the female phase with the stamens pulled along the corolla tube.Note the limb is bilabiate and oblique.F. Frontal view of the corolla limb showing the zygomorphic symmetry, the two dorsal and the two lateral lobes markedly revolute form the upper lip, and the ventral lobe forms the lower lip with a tuft of trichomes located in the sinuses between the two dorsal lobes.G. Opened corolla showing the 2-3 celled-trichomes located toward the base inside.H. Pistil and nectary.I. Enlargement of the ovary and nectary.J. Young berry fruit.K. Seed.A-I based on Marín-Gómez & Gómez-Hoyos 77 (COL), J-K based on Ramos et al. 1199 (CUVC).

Figure 2 .
Figure 2. Columnea chocoensis M. Amaya & L.E.Skog var.chocoensis. A. Bilabiate corolla limb.B. Ventral view of the vegetative and reproductive shoots.C. Lateral view of the flower.D. Pedicel with bracteole, with the style oblique to the calyx, and corolla tube dorsally curved, with the base dorsally gibbous.E. Indument of pedicel and calyx, and the stomatomorphic stigma.

Figure 3 .
Figure 3. Columnea chocoensis M. Amaya & L.E.Skog var.altaquerensis L.E.Skog. A. Ventral view of the vegetative shoot, the red apex of each leaf appears like a "margin" for the whole vegetative shoot.B. Dorsal view of the vegetative shoot.C. Flowers hidden under the leaves with the pedicel perpendicular to the corolla.D. Flowers in the bud stage with pedicels erect in the calyx.Note the bracteoles with a medial red line.E. Flower at the female phase.F. Flower at the male phase.G. Bifid stigma.H. Young fruit.

Figure 5 .
Figure 5. Columnea stilesiana M. Amaya & L.P. Kvist. A. Branched dorsiventral vegetative shoot.B. Ventral view of the vegetative shoot.C. Bud and lateral view of an open flower.D. Corolla limb bilabiate and with galea.

Figure 6 .
Figure 6.Columnea ericae Mansf.var.ericae.A. Vegetative shoot.B. Axillary inflorescences located underneath the vegetative shoot.C. Bracts pale yellow and corolla bilabiate with two purple lines delimiting the entrance of the corolla.D. Lateral view of the flower.

Figure 8 .
Figure 8. Columnea ericae Mansf.var.archidonae (Cuatrec.)L.P. Kvist.A.Ventral view of the vegetative shoot with flowers exposed from the vegetative shoot.B. Pedicel.C. Corolla oblique in the calyx.D. Lateral view of the flower, showing the bilabiate limb.

:
Flowers have been recorded on the specimens collected in March, July, and August.