New circumscriptions add two northern Andean species to Kohleria (Gesneriaceae) Author Clark, John L. https://orcid.org/0000-0002-1414-6380 Science Department, The Lawrenceville School, Lawrenceville, NJ 08648, USA jclark@lawrenceville.org Author Jost, Lou Lou Jost, Fundacion EcoMinga, Banos, Tungurahua, Ecuador text PhytoKeys 2021 2021-07-12 179 99 110 http://dx.doi.org/10.3897/phytokeys.179.65990 journal article http://dx.doi.org/10.3897/phytokeys.179.65990 1314-2003-179-99 F9007B81A3025C47AF6B5D7A9EF49B0E Kohleria andina (Fritsch) J.L. Clark & Jost comb. nov. Capanea andina Fritsch, Bot. Jahrb. Syst. 50: 431-432. 1913 ( "1914" ). Type: Ecuador. Andes Quitenses, Tunguragua, 1857, R. Spruce 5178 (lectotype K000395097, designated here; isolectotypes: BM000953512, E00062367, G00370826, G00370838, K000395097). Remarks. One of the key characters discussed by Kvist and Skog (1992) as the basis for the generic circumscription of Kohleria was a terrestrial (i.e., non-epiphytic) habit. In the generic delimitation of Kohleria ( Kvist and Skog 1992 ), the habit is described as herbs, subshrubs, shrubs, or rarely scandent shrubs. The terrestrial habit was considered a character by Kvist and Skog (1992) to differentiate Kohleria from closely related genera that are epiphytes or lianas. Phylogenetic studies by Roalson et al. (2005b) showed that Kohleria was paraphyletic with the exclusion of " Capanea ", a group of epiphytic subshrubs from the Andes. Thus, many of the features that differentiated " Capanea " from Kohleria , such as an epiphytic habit and four-valved capsules, are autapomorphic. The transfer of two species from " Capanea " to Kohleria is well-supported, and combinations were made by Roalson et al. (2005b) . Roalson et al. (2005b) did not make a combination for Kohleria andina because it was considered a heterotypic synonym of Kohleria affinis . Examination of material in the field and in herbaria allowed us to recognize K. affinis and K. andina as different species. Outlined here are characters to differentiate K. andina from K. affinis (see Table 1 for a summary of the characters that are discussed below). Table 1. Morphological differences and general distribution of Kohleria affinis and K. andina .

-	Kohleria affinis (Fritsch) Roalson & Boggan	Kohleria andina (Fritsch) J.L. Clark & Jost
Corolla tube shape	usually narrow, rarely broad (Colombia)	broad
Corolla tube color	dark red to bright purple	white
Corolla tube trichome color	transparent	yellow
Corolla tube length	3-6 cm	<3.5 cm
Peduncle and pedicel trichome color	transparent	purple
Distribution	widespread in Colombia, Ecuador, and northern Peru	endemic to the Ecuadorian province of Tungurahua)

Another feature that defines the clade previously recognized as " Capanea " is the presence of resupinate flowers via a twisted pedicel. The androecium and gynoecium are located in the lower region of the corolla tube (Fig. 1B, C and Fig. 2B, C ). In contrast, all other Kohleria and closely related genera have the androecium and gynoecium in the upper region of the corolla tube. Figure 1. Kohleria affinis (Fritsch) Roalson & Boggan A lateral view of flower B female phase of mature flower C male phase of mature flower D habit ( A Clark et al. 7698 B Clark s.n . C Clark et al. 12979 D Clark et al. 15845 ). Photos by J.L. Clark. The flowers of Kohleria affinis are often photographed because of their conspicuous clusters of brightly colored purple-red corolla tubes with contrasting green lobes (Fig. 1 ). It is common to see individuals with 50+ pendent flowers, especially in abandoned cow pastures or recently cleared forests. Herbarium specimens do not preserve floral colors and most corollas dry uniformly black. Thus, corolla colors are challenging to determine on dried herbarium specimens unless noted by collectors in the descriptions. Use of field-based images, review of taxonomic literature, and examination of type specimens provided information for re-assessing the circumscription of Kohleria affinis and K. andina . The corolla tube of Kohleria andina is white, but appears bright yellow from dense tomentose yellow trichomes (Fig. 2 ). In contrast, the corolla tube of Kohleria affinis is dark red to bright purple (Fig. 1 ). The corolla tube in most Kohleria affinis is narrow, but some populations from Colombia are broad. The corolla tube of Kohleria andina is consistently broad. Corolla length in Kohleria affinis is highly variable and ranges from 3 to 6 cm. In contrast, the corolla tubes of Kohleria andina are usually less than 3.5 cm long. Both species have bright green corolla lobes that contrast with dark purple spots on the inner surface (Figs 1 , 2 ). Figure 2. Kohleria andina (Fritsch) J.L. Clark & Jost A lateral view of flower B female phase of mature flower C male phase of mature flower D habit ( A-D Clark et al. 7750 ). Photos by J.L. Clark. An additional character useful for differentiating Kohleria andina from K. affinis is the presence of dark red-purple trichomes on the peduncles and pedicels (Fig. 2A, D ). The red-purple trichomes on the peduncles were noted by Fritsch (1913 : page 432) in the protologue, "Pedunculi axillares elongati purpureo-villosi." In contrast, the peduncles in Kohleria affinis are sparsely pilose and appear green due to transparent trichomes. Kohleria andina and K. affinis are geographically separated by elevation. K. andina is endemic to elevations above 2500 meters on the western Andean slopes (Cordillera Oriental) in the Tungurahua province of Ecuador. In contrast, Kohleria affinis is widespread in the northern Andes of Colombia (Antioquia, Boyaca , Caldas, Caqueta , Cauca, Choco , Cundimarca, Huila, Narino , Putumayo, Quindio , Risaralda, and Valle del Cauca), Ecuador (Azuay, Bolivar , Carchi, Chimborazo, Cotopaxi, Esmeraldas, Imbabura, Loja, Napo, Pichincha, Santo Doingo, Tungurahua, and Zamora-Chinchipe), and northern Peru (Amazonas and Cajamarca). The authors' field work from the upper slopes of Tungurahua, an active volcano in the western Andean slopes of the Cordillera Oriental, revealed little overlap. Kohleria andina is locally endemic to elevations above 2500 meters and K. affinis is widespread and located in forests below 2500 meters. Intermediate forms were not found here, indicating that these two forms are geographically separated by elevation and supported as different biological species. Lectotypification. Syntypes are from two distinct localities: F.C. Lehmann 4869 (F0060498) from Colombia and R. Spruce 5178 from K (K000395097) from Tungurahua, Ecuador. The specimen of F.C. Lehmann 4869 (F) is more similar to the widespread Kohleria affinis . The specimen of R. Spruce 5178 from (K) is similar to the locally endemic Kohleria andina , and is designated as the lectotype to stabilize this species concept. According to Fritsch (1913) , Richard Spruce cites Tunguarahua as a locality and the specimens have characters that are congruent with the Tungurahua populations featured in the images here (Fig. 2 ). The lectotype has a corolla that is wide and more ampliate (Fig. 2 ) relative to the narrower corolla tube of K. affinis (Fig. 1 ). An additional character that is congruent with material from the type locality and the lectotype ( R. Spruce 5178 ) is the presence of dark red trichomes on the peduncles and pedicels. In contrast, the peduncle and pedicel trichomes on F.C. Lehmann 4869 are transparent and more similar to K. affinis .