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 anisophylla (Fritsch) Wiehler Kohleria anisophylla (Fritsch) Wiehler. Kohleria anisophylla (Fritsch) Wiehler, Selbyana 5: 62. 1978. Type: Based on Diastema anisophyllum Fritsch. Kohleria villosa var. anisophylla (Fritsch) Kvist & Skog, Smithsonian Contr. Bot. 79: 70. 1992. Type: Based on Diastema anisophyllum Fritsch. Basionym. Diastema anisophyllum Fritsch, Bot. Jahrb. Syst. 50: 408. 1913 ( "1914" ). Type: Colombia. [ Narino ] Piedra Ancha, West of Andes of Tuquerres, F.C. Lehmann 5843 (B, holotype not extant, lectotype K000509983, designated by Wiehler (1978 : 62), isolectotype K000509984). Nematanthus erianthus Bentham, Pl. Hartw: 231. 1846. Type: Ecuador. Pichincha: Quito towards Nanegal, Hartweg s.n. (holotype K000509985). Columnea eriantha (Bentham) Hanstein, Linnaea 34: 391. 1865. Type: Based on Nematanthus erianthus Fritsch. Diastema anisophyllum Fritsch var. quitense Fritsch. Bot. Jahrb. Syst. 50(4): 408. 1913 ( "1914" ). Type: Ecuador. [Pichincha] Quito, W. Jameson s.n. (holotype W). Remarks. Kohleria anisophylla (Fig. 3 ) was previously recognized by Kvist and Skog (1992) as a variety of Kohleria villosa (Fig. 4 ). The strongly anisophyllous leaves and dorsiventral shoots (Fig. 3D ) are more similar to Kohleria hypertrichosa (Fig. 5D ) than K. villosa (Fig. 4D ). All three species are found on the northwestern Andean slopes of Ecuador. Only Kohleria anisophylla is documented from Colombia ( Narino department). Wiehler (1978) made the combination Kohleria anisophylla and recognized it at the rank of species. Kvist and Skog (1992) recognized this taxon as Kohleria villosa var. anisophylla . Based on limited material, Wiehler (1978) cited the type ( F.C. Lehmann 5843 ) and a recently collected specimen from Ecuador ( C. Luer & A. Hirtz 2672 ). Kvist and Skog (1992) cited the same Ecuadorian collection and mentioned the study of eleven additional specimens. This species is common along the northwestern slopes of the Ecuadorian Andes, especially along the old road between Quito and Santo Domingo where many of the images were taken for Figure 3 . Outlined here are characters to differentiate K. anisophylla , K. villosa , and K. hypertrichosa (see Table 2 for a comparison of characters that are discussed below). Table 2. Morphological differences and general distribution of Kohleria anisophylla , K. villosa , and K. hypertrichosa .

-	Kohleria anisophylla (Fritsch) Wiehler	Kohleria villosa (Fritsch) Wiehler	Kohleria hypertrichosa J.L. Clark & L.E. Skog
Habit	facultative epiphyte	terrestrial	facultative epiphyte
Shoots	dorsiventral	erect	dorsiventral
Relative leaf size	anisophyllous	isophyllous	anisophyllous
Corolla vestiture	villous	villous	tomentose
Distribution	Ecuador ( Bolivar , Carchi, Pichincha) and Colombia ( Narino )	Ecuador ( Bolivar , Carchi, Cotopaxi, Esmeraldas, Imbabura, Pichincha)	northern Ecuador (Carchi and Esmeraldas)

Figure 3. Kohleria anisophylla (Fritsch) Wiehler A lateral view of flower B front view of corolla C mature fruit D dorsiventral habit with anisophyllous leaves ( A, B Clark et al. 10981 C Clark 10948 D Clark et al. 14295 ). Photos by J.L. Clark. The recent transfer ( Roalson et al. 2005b ) of Kohleria affinis and K. tigridia (Ohlend.) Roalson and Boggan represented an autapomorphic synapomorphy of epiphytism in traditionally recognized Kohleria . What is noteworthy about Kohleria anisophylla and K. hypertrichosa is their previously unreported epiphytic habits. Thus, the presence of an epiphytic habit in K. anisophylla and K. hypertrichosa could represent an additional independent origin of epiphytism in Kohleria . Several populations of Kohleria anisophylla were observed and documented with dorsiventral shoots, a feature that is common in facultative epiphytes in other Gesneriaceae genera. Many members of Columnea have strongly anisophyllous leaves - especially species that are facultative epiphytes with dorsiventral shoots. Other species of Gesneriaceae that are facultative epiphytes with dorsiventral shoots include Cremosperma anisophylla J.L. Clark & L.E. Skog, Drymonia anisophylla L.E. Skog & L.P. Kvist, and the majority of species in Monopyle Moritz ex Benth. and Trichodrymonia Oerst. Likewise, Kohleria anisophylla and K. hypertrichosa are facultative epiphytes with dorsiventral shoots and anisophyllous leaves. In contrast, Kohleria villosa is a terrestrial herb with isophyllous leaves (Fig. 4D ). Figure 4. Kohleria villosa (Fritsch) Wiehler A lateral view of flower B front view of corolla C mature fruit D erect herbaceous habit with isophyllous leaves ( A Clark et al. 14295 B Clark 7331 C Clark et al. 7400 D Clark et al. 14295 ). Photos by J.L. Clark. The corollas of Kohleria villosa and K. anisophylla are villous (Figs 3 , 4 ). The corollas of Kohleria hypertrichosa are densely tomentose (Fig. 5 ). The specific epiphyte, " Kohleria hypertrichosa " refers to the abundance of trichomes, which is why it is commonly known in the horticultural community as "Chewbacca," a reference to the Wookie (fictional character) in the movie Star Wars. Figure 5. Kohleria hypertrichosa J.L. Clark & L.E. Skog A lateral view of flower B front view of corolla C mature fruit D dorsiventral habit with anisophyllous leaves ( A Clark et al. 15900 B Clark 6539 C Clark et al. 10310 D Clark et al. 14942 ). Photos by J.L. Clark. Kohleria villosa and K. anisophylla are easily recognized when sterile. The opposite leaves of Kohleria anisophylla are consistently unequal in size or anisophyllous (Fig. 3D ). In contrast, the opposite leaves of Kohleria villosa are consistently equal in size or isophyllous (Fig. 4D ). In addition, the dorsiventral shoots distinguishes K. anisophylla from the erect shoots of K. villosus .