SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 79 

Revision of Kohleria (Gesneriaceae) 

Lars Peter Kvist 

and 

Laurence E. Skog 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1992 
ABSTRACT 

Kvist, Lars Peter and Laurence E. Skog. Revision of Kohleria (Gesneriaceae). Smithsonian 

Contributions to Botany, number 19, 83 pages, 37 figures, 4 tables, 1992.-The genus Kohleria 

(Gesneriaceae: Gloxinieae) ranges from Mexico to Peru and east to Surinam, and from sea level 

to about 2500 meters altitude, but most commonly grows between 800 and 2000 meters. About 

100 species have been described in Kohleria, but in the present work only 17 are recognized. 

Fourteen species occur in Colombia, with 9, including four newly described species, K. 

diastemoides, K. grandiflora, K. longicalyx, and K. neglecta, being endemic to this country. 

Eleven Kohleria species occur in exposed, disturbed habitats along rivers, roads, etc. These 

species have capsular fruits that split apically into two valves and have wind-dispersed seeds. In 

contrast, the remaining species are found in shaded, humid forest understory, and have capsules 

that split by a slit from the apex to the base to expose a sticky mass of seeds that apparently are 

animal-dispersed. These latter species also usually lack the scaly rhizomes that enable 

exposed-habitat species to survive dry spells. 

Some Kohleria species are stable in most features even over wide distribution ranges, e.g., K. 

spicata and K. tubifora. In contrast, a few species are surprisingly variable. To circumscribe this 

variation the following varieties are recognized: K. amubilis var. bogotensis, K. hirsuta var. 

longipes, K. villosa var. anisophylla, K. inaequalis var. ocellata, and var. lindenii (none of these 

are new taxa). Kohleria hirsuta and K. trianae hybridize in the Central Colombian Cordillera. 

In the Eastern Colombian Cordillera, var. Iongipes of K. hirsuta may be a formerly 

well-separated semi-species that has hybridized extensively with the typical variety. The most 

variable species K. inaequulis apparently is a complex of three extensively hybridizing 

semi-species, and pure breeding populations still are found in the periphery of its range. 

Generally, the main mode of speciation of Kohleria appears to be the geographical isolation of 

peripheral populations, while hybridization breaks down species limits rather than creating new 

ones. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution’s annual report, Smithsonian Year, SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging-in-Publication Data 

Kvist, Lars Peter 

Revision of Kohleria (Ge.sneriaceae)/Lars Peter Kvist and 

p. cm.-(Smithsonian conmibutions to botany ; v. 79) 

Includes bibliographical references and index. 

Supt of Docs. no.: SI 1.29: 

1. KohleriaAlassification. I. Skog, Laurence E. I1 

QK495.G4K85 1992 92-5842 

583’.81-d~20 CIP 

aurence E. Skog. 

Title. 111. Series. 

@ The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials Z39.4&1984. 
Contents 

Page 

Introduction ........................................ 1 

Acknowledgments ................................... 1 

Materials and Methods .................................. 1 

Taxonomic History .................................... 1 

Delimitation of Kohleria Regel .............................. 4 

Leaves ......................................... 4 

Stomata ......................................... 4 

Indumentum ...................................... 4 

Inflorescences ..................................... 4 

Seeds .......................................... 8 

Pollen .......................................... 8 

Anatomy ........................................ 11 

Cytology ....................................... 11 

Chemistry ....................................... 11 

Scaly Rhizomes and Stolons ............................. 11 

Phenology ....................................... 13 

Flowers and Pollination ................................ 13 

Fruits and Dispersal .................................. 19 

Phytogeography .................................... 20 

Popular Names and Medicinal Uses ......................... 20 

Interspecific Hybridization .............................. 30 

Intraspecific Hybridization .............................. 32 

Intergeneric Hybridization .............................. 36 

Related Genera .................................... 36 

Evolution ....................................... 38 

Kohleria Regel ...................................... 38 

Key to the Species and Varieties of Kohleria Regel ................. 39 

1 . Kohleria allenii Standley & L.O. Williams .................. 42 

2 . Kohleria mbilis (Planchon & Linden) Fritsch ................ 44 

2a . var . amubilis ................................ 45 

2b . var . bogotensis (Nicholson) Kvist & L . Skog, new status ......... 45 

3 . Kohleria diastemoides Kvist & L . Skog, new species ............. 47 

4 . Kohleria grandflora Kvist & L . Skog, new species ............. 47 

5 . Kohleria hirsuta (Kunth) Regel ........................ 49 

5a . var . hirsuta ................................. 50 

5b . var . Iongipes (Bentham) Kvist & L . Skog, new status .......... 51 

6 . Kohleria hondensis (Kunth) Hanstein ..................... 52 

7 . Kohleria inaequalis (Bentham) Wiehler .................... 53 

7a . var . inaequalis ............................... 53 

7b . var . lindenii (Hanstein) Kvist & L . Skog. new status ........... 54 

7c . var . ocellata (W.J. Hooker) Kvist & L . Skog, new status ......... 54 

8 . Kohleria longicalyx Kvist & L . Skog. new species .............. 55 

9 . Kohleria neglecta Kvist & L . Skog, new species ............... 57 

10 . Kohleria peruviana Fritsch .......................... 57 

11 . Kohleria rugata (Scheidweiler) Kvist & L . Skog. new combination ..... 59 

12 . Kohleria spicata (Kunth) Oersted ....................... 60 

13 . Kohleria stuebeliana Fritsch .......................... 63 

... 

111 
iv SMITHSONIAN CONTRIBUTIONS TO BOTAhT 

14 . Kohleria trianae (Regel) Hanstein ....................... 65 

15 . Kohleria tubifloru (Cavanilles) Hanstein ................... 66 

16 . Kohleria villosa (Fritsch) Wiehler ...................... 68 

16a . var . villosa ................................ 70 

16b . var . anisophylla (Fritsch) Kvist & L . Skog. new status ......... 70 

17 . Kohleria warszewiczii (Regel) Hanstein .................... 70 

Appendix 1: List of Excluded Names .......................... 73 

Appendix 2: Names Based on Hybrids ......................... 74 

Literature Cited ..................................... 76 

Index to Scientific Names of Genera and Species ................... 80 
Revision of Kohleria (Gesneriaceae) 

Lars Peter Kvist and Laurence E. Skog 

Introduction 

The genus Kohleria has for many years needed revision. 

Species of Kohleria are among the most well-known members 

of New World Gesneriaceae, partly because they are frequently 

cultivated in Latin America, Europe, and in the United States, 

and partly because the most common Kohleria species, 

especially K. spicata and K. tubflora, are somewhat weedy and 

among the most frequently collected New World Gesneriaceae. 

Since 1847, when Kohleria was established by Regel, about 

100 species names have been added to the genus, mostly by 

European workers who never visited the New World but based 

their descriptions to a great extent on garden material. Some of 

the common and widespread species were repeatedly newly 

described. In addition, many species names were added to 

several synonymous genera. By the early 20th century Kohleria 

was in a seemingly hopeless taxonomic and nomenclatural 

mess, and most workers avoided the group. Wiehler (1975, 

1978a) split off Moussonia Regel and Parakohleria Wiehler 

distributed from Panama to Mexico and along the eastern 

Andean slopes, respectively. In the present work the remaining 

approximately 70 species names remaining in Kohleria are 

reduced to 17. The taxa are distributed from southern Mexico to 

Peru and east to Surinam. The center of diversity is in the 

Colombian highlands, where 14 species occur (including nine 

endemics). 

ACKNOWLEDGMENTS.-We greatly appreciate the assistance 

of Ximena Londoiio during field work and her continued 

collections and observations of Kohleria in Colombia, the 

excellent drawings of Alice Tangerini, the help of Marie 

Uehling and Leslie Brothers in manuscript and plate preparation, the EPR-spectroscopic investigations undertaken by 

Jens Arne Pedersen, the careful and critical reading of the 

Lars Peter Kvist, Royal Veterinary and Agricultural University, 

Department of Botany, Dendrology, and Forest Genetics, Rolighedsvej 23, D.K.-1958 Frederiksberg c. Denmark, and Laurence E. 

Skog, Department of Botany, NHB-166, Smithsonian Institution, 

Washington, D.C. 20560. 

Review Committee: John J. Wordack, Department of Botany, 

Smithsonian Institution, Chair: Susanne S. Renner, Botanical Institute, University of Aarhus, Denmark; and three arwnymus reviewers. 

manuscript by Susanne S. Renner and three anonymous 

reviewers, and the herbaria listed below for the loan of their 

specimens for study. 

The first author also acknowledges the support of the Danish 

Research Council, the Research Foundation of the University 

of Aarhus, the Research Academy of Denmark, the Research 

Fund of the American Gloxinia and Gesneriad Society, the 

Susan Liebers Erickson Danish Exchange Fund of the 

Smithsonian Institution, as well as a pre-doctoral fellowship 

that allowed him to undertake research at the National Museum 

of Natural History, Smithsonian Institution. The second author 

acknowledges the support of the Research Opportunity Fund of 

the National Museum of Natural History that provided travel 

funds to visit herbaria. 

MATERIALS AND METHODS 

The revision is based on studies of herbarium material (1762 

collections from the following herbaria: A, AAU, BH, BM, C, 

CGE, CM, COL, E, EAP, ECON, F, FAUC, G, GB, GH, HBG, 

JAUM, K, L, LE, MA, MANCH, MEDEL, MEXU, MO, NY, 

P, PORT, QCA, QCNE, S, SEL, TULV, U, UC, US, USM, 

VEN, W, WU, WIS), as well as living plants in the field and in 

cultivation. A complete list of collections studied is available 

from the second author. During fieldwork in Ecuador and 

Colombia (L.P. Kvist) and Peru and Panama (L.E. Skog) 13 of 

the 17 recognized species were studied in their native habitats: 

Kohleria amabilis, K. hirsuta, K. hondensis, K. longicalyx, K. 

neglecta, K. trianae, K. warszewiczii in Colombia, K. inaequalis and K. spicata in Colombia and Ecuador, K. villosa in 

Ecuador, K. peruviana in Peru and K. spicata and K. rubiflora 

in Panama. During the time of the study the following species 

have been in cultivation in the greenhouses of the Smithsonkin 

Institution and Botanical Institute of the University of Aarhus: 

K. allenii, K. hirsuta, K. hondensis, K. inaequalis, K. 

peruviana, K. rugata, K. spicata, K. trianae, K. villosa, and K. 

warszewiczii. 

TAXONOMIC HISTORY 

Until Regel published the name Kohleria in 1847, the 

species now referred to Kohleria were included in Gesneria 
2 SMITHSONIAV CONTRIBUTIONS TO BOTANY 

Linnaeus or in Gesnera Martius. The first species described 

was Kohleria tubflora (as Gesneria tubflora Cavanilles, 

1801). In 1818 Kunth added six species of Gesneria that 

currently are placed in Kohleria: K. hirsuta (as Gesneria 

hirsuta, G. longflora, and G. mollis), K. hondensis (as G. 

hondensis), K. spicata (as G. spicata), and K. trianae (as G. 

elongata). Toward the middle of the 19th century interest in 

New World Gesneriaceae increased. Additional species were 

described, mainly in Gesneria, which are, for the most part, 

synonyms of the current species Kohleria hirsuta (as Gesneria 

vestita Bentham, 1845b, Gesneria eriantha Bentham, 1846, G. 

longipes Bentham, 1846, and Gesneria rubricaulis Kunth & 

Boucht, 1847), Kohleria spicata (as Gesneria longifolia 

Lindley, 1841, Gesneria schiedeana (DC.) W.J. Hooker, 1845, 

and Gesneria petiolaris Bentham, 1845a), and Kohleria 

tubflora (as Gesneria incurva Bentham, 1845a, G. rhynchocarpa Bentham, 1845a, and Gesneria hondensis W.J. 

Hooker, 1846). The earliest descriptions of the following 

accepted species also come from this period: Kohleria amabilis 

(as Achimenes picta W.J. Hooker, 1844, Kohleria inaequalis 

(as Gesneria inaequalis Bentham, 1846), Kohleria rugata (as 

Gesneria rugata Scheidweiler, 1847) and Kohleria villosa (as 

Nematanthus erianthus Bentham, 1846). Thus, even at this 

early point most of the taxa described were synonyms of a few 

widespread species. From the turn of the 20th century this trend 

accelerated dramatically with new species named by Fritsch 

and new species based on hybrids. In addition, the species that 

currently are placed in Kohleria were simultaneously assigned 

to several synonymous genera, Brachyloma Hanstein, Isoloma 

(Bentham) Decaisne, Tydaea Decaisne, etc. The main workers 

in New World Gesneriaceae during this period (E. Regel, first 

in Ziirich and later in St. Petersburg (formerly Leningrad), J. 

Decaisne in Paris, G. Bentham and W.J. Hooker at Kew, and 

especially J. Hanstein in Berlin) described the same few species 

again and again, mainly from plants cultivated in Europe. 

The name Kohleria, with K. hirsuta as the type species, 

appeared for the first time in 1847 with a short diagnosis in a 

seedlist from the botanical garden in Ziirich, where Regel was 

at that time head gardener. In April of the following year Regel 

described Kohleria more fully and added K. triflora (now K. 

hondensis). He noted that Gesneria hondensis Kunth (1818) 

would belong to Kohleria, but Regel never published a new 

combination in the genus for the species. Hanstein, however, 

crediting Regel (1848) for the combination K. hondensis, 

actually published it in 1859. In December 1848 another paper, 

by Decaisne, appeared that dealt with the splitting of Gesneria. 

In this paper Decaisne elevated the taxon Isoloma, earlier 

established in 1846 as a section of Gesneria by Bentham, to 

generic rank. Decaisne transferred most of the abovementioned species of Gesneria and Kohleria to Isoloma. 

However, he did not select a type species, and it is remarkable 

that one of the species he did not transfer was G. hirsuta (the 

type species of Kohleria). Regel (1851) later followed Decaisne 

in believing that Isoloma had priority over Kohleria. He stated 

that Bentham chose the same type species for Isoloma as he 

himself had chosen for Kohleria in 1847, viz. K. hirsuta. This 

is incorrect, as Bentham (1846) chose no type species and only 

mentioned K. hirsuta in the description of K. eriantha. Much 

later, Morton (1973) finally lectotypified Isoloma with Kohleria hirsuta. Isoloma was actually illegitimate when elevated 

by Decaisne, because the name had been used previously for a 

fern genus by Smith (1841), and, in addition, the later 

published generic name Kohleria would have priority over the 

sectional name Isoloma. Unfortunately this error confused the 

nomenclature of Kohleria for the remainder of the 19th century. 

To add to the confusion Regel (185 1, 185%) described another 

new genus that he again named Kohleria but this time based on 

K. ignorata (described by Kunth and Boucht in 1848 as 

Gesneria ignorata) as the sole species. This species, based on 

material from Central America, is another synonym of K. 

spicata. This second genus established by Regel is obviously a 

later homonym. 

In addition to Isoloma, Decaisne (1848b) also established the 

monotypic genus Tydaea to include 'I: picta, first described by 

W.J. Hooker (1844) as Achimenes picta (now Kohleria 

amabilis var. bogotensis). Shortly thereafter, Regel (1849) 

based Giesleria, another new monotypic genus, on the same 

species. But he soon realized (1851) that 7'ydaea had priority. 

In the same paper Regel added by transfer a second species, 'I: 

ocellata, originally described by WJ. Hooker (1848) as 

Achimenes ocellata (now Kohleria inaequalis var. ocellata). In 

1853 Regel added another genus, Sciadocalyx, with S. 

warszewiczii as the sole species. The latter genus was expanded 

with S. digitalflora by Linden and Andrt in 1870. Both species 

are currently synonyms of Kohleria warszewiczii. 

Hanstein (1854, 1856,1859,1865) studied the taxa currently 

placed in Kohleria and published a large number of new 

species names and combinations. In 1854 he realized that the 

name Isoloma (Bentham) Decaisne was illegitimate, replaced 

the generic name with Brachyloma (with B. hirsutum as type 

species), and included B. hondense as a second species. 

Unfortunately, Hanstein (1854) did not realize that the later 

(1851) Kohleria of Regel, based on K. ignorata, was also 

illegitimate as a later homonym. He took up this later Regel 

genus and added to it another synonym of K. spicata, K. 

seemannii. Oersted (1858) also accepted both the second 

Kohleria based on K. ignorata, as well as Brachyloma, and 

described a number of new species, especially in Brachyloma. 

These are all currently reduced to synonyms of K. spicata or K. 

tubiflora. In 1858 Oersted transferred Gesneria spicata to 

Kohleria, but because he accepted the genus Brachyloma with 

B. hirsutum as type species, he actually transferred Gesneria 

spicata to the illegitimate Kohleria genus that Regel (1851) 

based on K. ignorata. Hanstein (1856, 1859), well aware of the 

work of Oersted, included the latter's new species and 

established two more genera, Calycostemma and Cryptoloma. 

Calycostemma was monotypic and based on C. lindenii, treated 

herein as the variety lindenii of the variable Kohleria 

inaequalis. Hanstein split off Cryptoloma from Brachyloma 

and included 10 species in that genus, but only Cryptoloma 
NUMBER 79 

hondense was transferred from Brachyloma to Cryptoloma. He 

did not choose any type species, and Cryptoloma hondense is 

herein selected as the lectotype. Besides the 10 species in 

Cryptoloma, Hanstein (1859) included 11 species in Kohleria 

and 15 species in Brachyloma. In the current work all 11 

species Hanstein referred to Kohleria are synonyms of K. 

spicata, 8 of the 10 species referred to Cryptoloma are 

synonyms of Kohleria tubzflora (the remaining two are 

synonyms of K. hondensis), and 12 of the 15 species referred to 

Brachyloma are synonyms of the correct Kohleria type species, 

K. hirsuta (two of the remaining are synonyms of K. trianae 

and one of K. spicata). Thus, the genera accepted by Hanstein 

in 1859 basically represent the three most widespread species 

in the current concept of Kohleria. 

After a few years Hanstein apparently recognized that his 

generic concepts were extremely narrow, and in 1865 he fused 

Brachyloma, Cryptoloma, Calycostemma, and Sciadocalyx 

with Kohleria. The only genus currently synonymized with 

Kohleria that he preserved was Tydaea. Into the latter genus 

Hanstein (1859) had placed four species, treated herein as 

synonyms of Kohleria amubilis and K. inaequalis. 

At the time of the original establishment of Kohleria, Regel 

(1847, 1848) also established the genus Moussonia. There 

remains some doubt regarding the identity of the type species 

M. elongata selected by Regel, because no authority other than 

“Auct.” was given after the basionym. If this is the species 

described as Gesneria elongata by Kunth in 1818 from 

Colombia, the name Moussonia evidently becomes a synonym 

of Kohleria. According to Wiehler (1975) plants originally 

collected in Mexico by Deppe were distributed in Europe under 

the name Gesneria elongata, and Regel probably had these 

plants in mind as Moussonia elongata, since they match his 

diagnosis of Moussonia more closely. The Mexican plants are 

Moussonia deppeana (Schlechtendal & Chamisso) Hanstein, 

while the Colombian plants of Gesneria elongata Kunth 

represent Kohleria trianue. 

The most recently described genus synonymous with 

Kohleria is Synepileana established by Baillon (1888a). The 

genus only included one species, S. granatensis, and was based 

on the same Linden collection number upon which Hanstein 

(1859) based his monotypic genus Calycostemma. It is now 

included in the variety lindenii of Kohleria inaequalis. 

Although Hanstein (1854) pointed out that Isoloma (Bentham) Decaisne was an illegitimate name, Bentham (in contrast 

to Regel) continued to use the name. The use by Bentham of 

Isoloma, in Genera Plantarum (1876) for example, resulted in 

Isoloma virtually replacing Kohleria in the literature (see works 

of Hemsley, 1882; Baillon, 1888b; Voss, 1894; Rusby, 1896, 

1900; Bailey, 1900, 1915; and Smith, 1916). Finally, through 

the works of Fritsch (1894, 1898, 1913a) Kohleria became 

generally accepted as the correct name. Fritsch (1894) also 

included Tydaea and Moussonia in Kohleria and treated these 

groups as two of the seven sections into which he divided 

Kohleria (the other sections were Sciadocalyx, Calycostemma, 

Cryptoloma, Isoloma, and Eukohleria). Apparently, Fritsch 

3 

also did not realize that Regel (1 85 1) had established a second 

Kohleria, being an illegitimate genus name based on K. 

ignorata. He placed the correct type species K. hirsuta in the 

section Isoloma, and Kohleria spicata (of which K. ignorata is 

a synonym) in the section Eukohleria. In later papers Fritsch 

(1913a, 1916) described many additional new species and 

varieties of Kohleria, of which only K. peruviana and K. 

stuebeliana are currently accepted. Two species that he 

described in Diastema Bentham (Fritsch 1913a) were transferred by Wiehler (1978b) to Kohleria as K. villosa and K. 

anisophylla and are herein treated as K. villosa var. villosa and 

var. anisophylla. 

After the work of Fritsch (l894,1913a, 1916) half a century 

passed with little work being done on Kohleria until Wiehler 

(1975, 1978a) split off Moussonia and his new genus 

Parakohleria. The most important investigations during this 

extended period were those of Moore (1953,1954) and Morton 

(1967a), but the only currently accepted new species described 

between 1916 and the present is K. allenii named by Standley 

and L.O. Williams (L.O. Willams, 1968). 

Wiehler (1975) re-elevated Moussonia to generic rank. This 

taxon had been treated by Bentham in 1876 as a synonym of the 

series Brachyloma in the genus Isoloma, and Fritsch (1894) had 

treated it as a section of Kohleria. The group does not belong in 

the present concept of Kohleria; among other characters it has 

a different basal chromosome number (11 versus 13 in 

Kohleria). However, as already mentioned, the name MousSonia published by Regel (1847) may actually be a synonym of 

Kohleria. Therefore, in order to stabilize the usage, we choose 

to accept the genus Moussonia based on Gesneria elongata of 

cultivation, originally from Mexico, and now Moussonia 

deppeana. 

In his Moussonia paper Wiehler (1975) sorted out a species 

group denoted as “Kohleria group three” from the eastern 

Andean slopes. Later (1978a) he formally split off these plants 

from Kohleria as the new genus Parakohleria. His belief that 

this group was misplaced in Kohleria seems justified. During 

fieldwork we discovered that the fruits of Kohleria and 

Parakohleria (Kvist, 1991) are fundamentally different, and we 

consider this difference to be the primary distinguishing feature 

(see section below under “Related Genera”). The species 

transferred to Parakohleria by Wiehler (1978a) came from 

several genera: Kohleria (including several that had never been 

transferred from Isoloma), Diastema Bentham, Moussonia 

Regel, and Pearcea Regel. 

Kohleria belongs to the tribe Gloxinieae sensu lato, which 

Wiehler (1983) established by fusing five tribes of Fritsch 

(1894): Bellonieae, Gloxinieae sensu stricto, Kohlerieae, 

Sinningieae, and Solenophoreae. Gloxinieae includes all the 

Gesneriaceae with perigynous to epigynous flowers, except for 

the Caribbean Gesneriaceae that are, for the most part, easily 

recognized by having an alternate leaf arrangement (Skog, 

1976). Wiehler (1983) included 24 genera and more than 300 

species in the reformed Gloxinieae. Recent revisions, e.g., 

Achimenes Persoon (Ramirez-Roa, 1987), Heppiella Regel 
4 SMITHSONIAV CONTRIBUTIONS TO BOTANY 

(Kvist, 1990), and other so far unpublished studies indicate the 

number to be smaller and probably about 200. 

Delimitation of Kohleria Regel 

As is often the case in genera of neotropical Gesneriaceae, no 

single feature distinguishes the genus Kohleria from related 

and similar genera in the tribe Gloxinieae, but the genus is set 

apart by a suite of characters: species of Kohleria are herbs, 

subshrubs, shrubs, or rarely scandent shrubs, but never lianas or 

epiphytes: the subtending leaves of the axillary inflorescences 

often become gradually reduced to bracts toward the shoot 

apex, but there never is a sharp gap between subtending leaves 

and bracts (Figure 3c.g): the corollas are mostly tubular often 

with a conspicuous limb and the tube is red or lavender outside; 

the nectary is usually reduced to free glands (Figure 10b-f) but 

consists occasionally of a five-lobed ring (Figure 10a); the 

stamens are included to subincluded with coherent anthers: the 

style is also included or subincluded and is conspicuously 

bilobed: the haploid chromosome number is 13: scaly rhizomes 

are often present but tubers never occur: and the capsules split 

apically into two valves (in species found in exposed habitats) 

or less commonly with a longitudinal slit to the base (in species 

found in the understory) (Figure 13c-e). The affinities with 

other genera in the tribe Gloxinieae are discussed in the section 

below, “Related Genera.” 

LEAVES 

The leaf arrangement in Kohleria is opposite or less 

commonly whorled (three or four leaves at one node). The latter 

condition is always found in K. allenii (Figure 29a) and K. 

neglecta (Figure 32a) and usually in K. spicata. However, even 

these species may have opposite leaf pairs intermixed with 

whorls. In addition, several other species occasionally have 

leaves in whorls of three, e.g., K. stuebeliana (Figure 35a). The 

leaves, when in a pair, are normally equal or subequal, but 

unequal pairs characterize the rare K. villosa var. anisophylla 

(Figure 37a). The shape of the blades is usually ovate to elliptic 

and shows little variation. Thus, leaf characters have obviously 

little taxonomic value. 

STOMATA 

Stomata are restricted to the lower leaf surfaces and are often 

raised on mounds. This feature is known from many other New 

World Gesneriaceae, both in genera related to Kohleria, e.g., 

Heppiella (Kvist, 1990), and in more distantly related genera, 

e.g., Reldia Wiehler in the tribe Beslerieae (Kvist and Skog, 

1989). In Kohleria (Figure 1) stomatal mounds are taller in 

species with a dense indumentum; the mounds are conspicuous 

in K. spicata (Figure lc), but absent in K. amabilis (Figure la). 

This indicates that the mounds may improve the transpiration 

by raising the stomata above the calm air-layer created by the 

indumentum. There is no correlation between permanently 

humid habitats and the presence of stornatal mounds, in conflict 

with Wiehler’s (1983) statement that “stomatal mounds occur 

only in terrestrial species in humid, rainforest habitats.” 

Understory species, e.g., K. inaequalis and K. villosa, do 

possess stomatal mounds, but these are not as well developed 

as in the exposed-habitat K. spicata (Figure lb-d). 

INDUMENTUM 

Kohleria has four main types of uichomes, all exclusively 

uniseriate: (a) slender, eglandular trichomes of 3-15 cells: (b) 

bristle-like eglandular trichomes of 1-3 cells: (c) glandular 

trichomes with a slender stalk of 5-10 cells: and (d) tiny 

glandular trichomes with a stalk of 1-2 cells. 

The prevalent indumentum of stems, leaves and inflorescences consists of slender, eglandular trichomes (Figures 2a, 

30b). The up-to-10 mm long trichomes of 10-15 cells that 

cover the stems of Kohleria longicalyx are among the longest 

known in New World Gesneriaceae, and those of K. grandiflora (Figure 31a) and K. peruviana (Figure 33a) are only 

slightly shorter. 

The bristle-like trichomes mainly are found on upper leaf 

surfaces (Figure 2b). The species of Kohleria can be sorted out 

into two groups: one with a pilose to velutinous upper-leaf 

surface that is soft and pleasant to touch, and another with a 

hispid upper leaf surface that feels very scabrous, each 

exemplified by K. hirsuta and K. hondensis, respectively 

(Figure 2). This difference is more conspicuous in dried than in 

living material. 

Glandular trichomes with a 1-7 mm long slender stalk are 

rare, but occur consistently on Kohleria allenii, frequently on 

K. grandijlora, and on K. hirsuta collections coming from 

elevations below 500 meters in the Rio Magdalena valley of 

Colombia. The glandular spheres of these latter plants often 

bear curiously branched appendages. This same phenomenon 

was illustrated by Wiehler (1983) from Rhytidophyllum 

auriculatum W.J. Hooker (as Gesneria auriculata (W.J. 

Hooker) 0. Kuntze, 1891) and from Achimenes dulcis C. 

Morton. 

The throat of corollas of Kohleria and numerous other 

Gesneriaceae always possess at least a few approximately 0.2 

mm long glandular trichomes. In addition, these often occur on 

the style and stigma, and occasionally on vegetative parts such 

as the stems. Wiehler (1983) illustrated both uni- and 

multicellular glandular spheres from the throat of K. villosa. 

According to Steiner (1985), working with Drymonia serrulata 

(Jacquin) Martius (tribe Episcieae), the multicellular nature of 

the glandular spheres is not revealed in SEM-micrographs due 

to a thick, surrounding, cuticular layer. Steiner also reported 

that the glandular spheres of D. serrulata exude substances that 

cause the pollen to stick to flower visitors. 

INFLORESCENCES 

Inflorescence characters are important both to distinguish 

species of Kohleria and to set the genus apart from related 
NUMBER 79 

FIGW 1.Stomata of Kohleria (scale length = 20 pm): a, K. amabilis var. amabilir; b, K. inaequalis var. 

inaequalis; c, K. spicata; d, K. villosa var. villosa. Micrograph a of Killip et al. 38338 (US); b of Balslev 1939 

(US); c of Daviahe et al. 29560 (MO); d of Harling & Andersson 23222 (US). 

5 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 2.-Upper leaf surfaces of Kohlerio (scale length = 250 pm): a, K. hirsutu var. hirsuta; b, K. hondemir. 

Micrograph a of Plowman & Berry 13356 (US); b of Petme113578 (US). 

genera. Gesneriaceae have open (monopodial) flowering shoots 

with closed (sympodial) inflorescences in the axils of the 

individual subtending leaves or bracts (Figure 3). The 

terminology applied to describe the inflorescences of the 

Gesneriaceae has often been inconsistent. The term “inflorescence” is alternately used for the flowers located in the axil of 

a single subtending leaf, and for flowering shoots with few to 

many subtending leaves each with one to several flowers in 

their ails. Troll (1964) classified the inflorescences of 

Gesneriaceae as indeterminate, designated the closed inflorescences located in the ails of the single subtending leaves as 

“partial florescences” (Figure 3d-f), the open flowering shoots 

with several to many partial florescences as “florescences” 

(Figure 3b,c,g), and the complex systems made up of several 

florescences as “synflorescences” (Figure 3a). Furthermore he 

designated florescences with un-reduced subtending leaves and 

reduced subtending leaves (bracts) as “frondose” and 

“bracteose,” respectively (Figure 3b,g). Finally, he designated 

synflorescences with subtending leaves that gradually are 

reduced to bracts toward the apex as “frondo-bracteose” 

(Figure 3c). The partial florescences of Gesneriaceae were 

described by Weber (1973, 1978, 1982) as “pair-flowered 

cymes,” cymes in which each regular flower is accompanied by 

an additional bractless flower in frontal-median position 

(“front-flower”). 

The number of flowers in the partial florescences of Kohleria 

varies from 1 to 10. Only few species have a constant number 

of flowers, e.g., K. rugata (Figure 3b) with one flower and K. 

trianae (Figure 3d) with four flowers, including three regular 

flowers and the front-flowers of the central flower. In many 

species the partial florescences mostly produce 1-2 flowers, 

although they occasionally produce four to six flowers 

including up to three front-flowers, e.g., K. spicata (Figure 3c) 

and K. stuebeliana (Figure 35a). Only three species, K. 

neglecta, K. peruviana, and K. warszewiczii, may produce up to 

10 flowers, including three or four front-flowers, although in 

most cases their partial florescences only develop six or fewer 

flowers. The latter two of these species have pedunculate 

partial florescences (Figure 3e, f), while the up-to-10 flowers of 

K. neglecta (Figure 32a) are densely congested in the axils of 

the subtending leaves. Nine species always lack peduncles, and 

the remaining eight species either always have pedunculate 

partial florescences (4 species) or both states occur (4 species) 

(Figure 4). The bracts subtending the regular flowers of the 
NUMBER 79 7 

f a 

C 

d e f 

FWmE 3.-Inflorescences of Kohleria and Diastema (open circle = flowers and buds, solid circle = old capsules, 

thick crescents = bracts, thin crescents = leaves and leaves reduced to bracts): a, biennial flowering shoot of K. 

hirsufa; b, frondose florescence of K. rugata with partial florescences of single, axillary flowers; c, 

frondo-bracteose florescence of K. spicara with partial florescences of often several axillary flowers; d. partial 

florescence of K. trianae, pair-flowered cyme with three regular flowers and one front-flower. e, partial 

florescence of K. warszewicrii, pair-flowered cyme with 5 regular flowers and one front-flower; f, partial 

florescence of K. peruviano, pair-flowered cyme with seven regular and three additional bractless flowers; g, 

bracteose florescence of Diastema afine Fritsch. 

partial florescences often are reduced and caducous, and are 

even absent in a few species (Figure 30. The flowering shoots 

of Kohleria are frondose, apart from K. allenii (Figure 29c), K. 

spicata (Figure 3c), and K. stuebeliana (Figure 35a), which are 

frondo-bracteose. Diastema is set apart from Kohleria by 

having only bracteose florescences (Figure 3g). 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Species/ length (cm) 

1. K. allenii 

2. K. amabilis 

3. K. diastemoides 

4. K. grandiflora 

5. K. hirsuta 

6. K. hondensis 

7. K. inaequalis 

8. K. longicalyx 

9. K. neglecta 

10. K. peruviana 

11. K. rugata 

12. K. spicata 

13. K. stuebeliana 

14. K. trianae 

15. K. tubiflora 

16. K. villosa 

17. K. warszewiczii 

0 12 3 4 5 6 7 8 9 10 

- 

- - 

- 

. . . . . . . 

0 12 3 4 5 6 7 8 9 10 

FIGURE 4.-Peduncle length variation (in cm) in the 17 Kohlerio species. Nine species are always epedunculate. 

In biennial or perennial shoots, the apex continues to grow at 

the end of the flowering period (Figure 3a). However, in this 

study the flowering shoots or florescences of one flowering 

season are treated as the basic inflorescence-unit, as they are 

separated by periods of vegetative growth. 

SEEDS 

Seeds in the tribe Gloxinieae rarely exceed 0.6 mm in length. 

Ivanina (1965) illustrated seeds from Kohleria bogoremis 

(probably a hybrid). Beaufort-Murphy (1983) included SEMmicrographs of seeds from K. brachycalyx (= K. hirsuta) and K. 

digitaliflora (= K. warszewiczii). For the present study 

SEM-micrographs were taken of seeds from all Kohleria 

species except K. longicalyx (Figures 5, 6). Light-microscope 

studies show the seeds to be brown or tawny. 

The seeds of exposed-habitat and understory Kohleria 

species differ. The first group has narrowly ellipsoid to 

ellipsoid seeds with longitudinally, or to some extent, 

transverse striations (Figures 5a,e, 6a-c,e,f, and can also be 

seen in the seeds of K. allenii, K. hondensis, K. rugata, K. 

stuebeliana, and K. trianae, which not are illustrated). All of 

these seeds are typical of most wind-dispersed genera and 

species in the tribe Gloxinieae judged from the illustrations in 

the paper by Beaufort-Murphy (1983). 

The seeds of the understory species Kohleria diastemoides 

(Figure 5c), K. inaequalis (Figure 50, K. longicalyx (only 

studied with light-microscope), and K. villosa (Figure 6d) differ 

by being sub-globose, and by having irregularly rather than 

longitudinally striated surfaces. The seeds of K. amabilis 

(Figure 5b) are intermediate between these two seed types. The 

seeds of the last understory species, K. grandifora (Figure 5d), 

have the combination of the two seed types: ellipsoid shape (as 

in exposed-habitat species) and an irregular surface (as in 

understory species). The striations are narrow in this species. 

The understory Gesneriaceae have seeds similar to those 

found in the genera Diastema, Monopyle, and Parakohleria as 

illustrated by Beaufort-Murphy (1983). All these genera mostly 

occur in the calm understory, and have types of capsule 

dehiscence (and probably dispersal) different from the winddispersed Kohleria species (Figure 13). 

POLLEN 

Few studies of Gesneriaceae pollen exist, and the only one 

that included species of Kohleria was that of Williams (1978) 

with SEM-micrographs of pollen from K. allenii, K. digitaliflora (= K. warszewiczii), and K. spicata. During this study 

SEM-micrographs were taken of pollen from 12 Kohleria 

species, and apart from K. peruviana and K. trianae, these are 

all illustrated (Figure 7). Neither size, shape, nor surface of the 

pollen shows much variation among the species of Kohleria. 
NUMBER 79 9 

FIGURE 5.4eeds of Kohleria (scale length = 50 pm): a, K. neglecia; b, K. amabilis var. bogoiensis; c, K. 

diastemoides; d, K. grandiflora; e, K. hirsuia var. hirsuia; f. K. inaequalis var. inaequalis. Micrograph a of 

Garcia-Barriga 10737 (US); b of Pwdie s.n. 6); c of Lehmann 8892 (NY); d of Cuairecasas 11683 (US); e of 

Barkley 35221 (NY); f of Lehmann 1236 (NY). 

The size of the grains is 19-25 pm along the polar axis and 

15-18 pm along the equatorial axis. The sexine is reticulate 

and only K. rugata (Figure 71,m) is set apart by having a 

verrucose surface. Skog (1976) found the same feature in a few 

Gesneria species, and suspected that this was a morphological 

modification for bat pollination. Kohleria rugata, however, is 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 6.-Seeds of Kohleria (scale length = 50 p): a, K. perwiana; b, K. spicata; c, K. tubifora; d, K. villosa 

var. villosa; e,f, seed and seed surface of K. warszewiczii. Micrograph a of Schunke 1472 0: b of GwiCrrez 35 

(US); c of Davidse & Gonzalez 19038 (US); d of Harling & Andersson 23222 (US); e.f of Pirez-Arbeldez & 

Cuatrecasas 5865 (US). 
NUMBER 79 11 

likely not bat pollinated, although the identity of its pollinators 

is unknown. 

Williams (1978) also illustrated pollen from other genera of 

the tribe Gloxinieae, viz., species of Bellonia Linnaeus, 

Gloxinia L’Heritier, Moussonia and Sinningia Nees, and Kvist 

(1990) illustrated pollen from Heppiella, another close relative 

of Kohleria. All these genera have pollen very similar to those 

of Kohleria, and pollen characters apparently have little 

taxonomic value in the Gloxinieae. 

ANATOMY 

Very little work has so far been published on the anatomy of 

New World Gesneriaceae, but see Weber (1971), Wilson 

(1974), Skog (1976), and Wiehler (1983). The first two 

publications include sections of the gynoecium morphology of 

Kohleria. 

CYTOLOGY 

The chromosome numbers of the comparably sized genera 

Kohleria, Diastema, and Parakohleria have been reported 37 

times, five times, and once, respectively (Skog, 1984). This 

difference reflects that Kohleria is much more commonly 

cultivated than the other two genera. The 37 counts of Kohleria 

represent 9 of the currently accepted species and some hybrids. 

Since all species have a haploid chromosome number of 13, 

tetra- or polyploidy is unimportant or even absent in the 

evolution of Kohleria. A majority of the genera in the tribe 

Gloxinieae also have 13 haploid chromosomes, and tetraploids 

are not common (Skog, 1984). 

CHEMISTRY 

The chemistry of the Gesneriaceae is little known. Morley 

included some chemical data in Steam (1969), and Harborne 

(1966, 1967) included some Kohleria species in his work on 

the flavonoids. As in most other New World Gesneriaceae the 

leaves and the petals of Kohleria contain an unusual class of 

flavonoids, 3-deoxyanthocyanins, responsible for the intensely 

red and orange colors of many members of the Gesnerioideae. 

Kvist and Pedersen (1986) included six species of Kohleria in 

their EPR-spectroscopic study of dihydric phenolic compounds 

in the Gesneriaceae. Additional EPR-studies were undertaken 

during this study (see Kvist and Pedersen for methods) and all 

the 17 recognized Kohleria species have now been investigated. 

The occurrence of 3-4 dihydroxy-phenylethoxy glycosides 

(probably Acteoside) have been demonstrated from all KohIeria species apart from K. tubifora. In this species another 

phenolic compound, Aesculetin, appears to replace Acteoside; 

the phenomenon was demonstrated by Kvist and Pedersen 

(1986) in a number of Gesneriaceae. In addition to Acteoside, 

a related compound Conandroside apparently was recorded 

from K. hondensis. Until now Conandroside had been isolated 

only from Conandron ramondioides in the Old World 

subfamily Cyrtandroideae (Nonaka and Nishioka, 1977), and 

Kvist and Pedersen (1986) recorded both compounds only from 

C. ramondioides. 

SCALY RHIZOMES AND STOLONS 

“Scaly rhizomes” is the phrase used for the subterranean, 

creeping stems with scale-like leaves densely congested into a 

somewhat cone-like structure (Figure 8b-d). In Kohleria their 

diameter typically is about one cm and their length up to five 

cm, but in native habitats the scaly rhizomes of K. hirsuta 

occasionally attain a length of 15 cm and a diameter up to two 

cm (pers. obs.). The scaly rhizomes allow the plants to survive 

unfavorable dry spells or seasons in dormancy and to re-sprout 

from the axils of the scale-like leaves when conditions improve. 

The majority of the Kohleria species produce scaly rhizomes in 

their native habitats, but this feature was not found in 

populations of K. inaequalis, K. longicalyx, and K. villosa. 

These latter species mostly occur in permanently humid 

understory and scaly rhizomes may not provide an advantage in 

this environment. Most exposed-habitat species, e.g., K. hirsuta 

(Figure 8b) and K. warszewiczii (Figure 8c), nearly always 

possess well-developed scaly rhizomes, but in K. neglecta and 

K. trianae (pers. obs.), and possibly K. allenii (Figure 29b), 

these are weakly developed. The possible explanation is that 

the roots of these species are the stoutest and the deepest found 

in the genus, probably supplying water to counteract the effects 

of dry periods. 

The dormant organs densely covered with scales are called 

rhizomes, whereas stolons are the actively growing organs 

often with scales widely separated. The latter may be 

subterranean (Figures 8a, 36a, 37a). They are typically 10-30 

cm long, and serve for vegetative propagation by giving rise to 

new shoots at the tips. They may also branch and thus give rise 

to several shoots. Stolons are nearly always present at least in 

species growing on soil substrate, and only epilithic plants, e.g., 

Kohleria hondensis (Figure 8d), growing in confining rockcrevices lack this feature (pers. obs.). 

The transition between scaly rhizomes and stolons actually is 

gradual as the two features display the extremes of a 

continuum. The tips of stolons may form scaly rhizomes from 

which new plants later arise or conversely, dormant scaly 

rhizomes often re-sprout with stolons from which shoots arise. 

As a consequence, populations of Kohleria often occur in 

clones with few to many shoots linked together by stolons and 

scaly rhizomes. In K. warszewiczii the flowering of the 

individual clones, often consisting of many shoots, was found 

to be synchronized (pers. obs.). 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 79 13 

PHENOLOGY 

Species or hybrids of Kohleria cultivated in temperate 

regions tend to flower during the northern summer, the shoots 

then die back and the plants survive the winter as dormant 

rhizomes. However, most scaly-rhizomed Kohleria species in 

their natural habitats are not strongly seasonal. The only 

exception is K. rugata (Figure 9) from Mexico, which flowers 

exclusively during the humid period from September to 

December; it apparently never has been collected during the dry 

season, indicating that during that time the species is dormant 

and probably existing only as rhizomes. In Ecuador and 

Colombia not only the shaded-habitat species but also more 

surprisingly the exposed-habitat ones flower throughout almost 

the entire year (Figure 9; pers. obs.). Exposed-habitat species, 

however, often only have a few flowers during the driest 

period. 

In native habitats the shoots of several Kohleria species may 

become biennial or even perennial (pers. obs.). The age of a 

shoot (the number of flowering seasons) can be easily 

determined. First-season shoots never branch apically, and 

shoots that continue growth after flowering and fruiting 

typically branch right above the old capsules, or at least possess 

a vegetative section of stem between the old capsules and the 

new buds or flowers (Figure 3a). Kohleria hirsuta is frequently 

biennial; also numerous shoots of K. warsrewicrii and K. 

neglecta are at least biennial and some may grow three to four 

seasons, but the most fruticose, much-branched, and long-lived 

species is K. trianae. The shoots of the understory species K. 

inaequalis, K. longicalyx, and K. villosa, do not show any 

obvious seasonality (pers. obs. and data from herbarium 

sheets). 

Observations in the Department of Cundinamarca, Colombia, demonstrated that the size, longevity, and branching of 

Kohleria hirsuta shoots (and the size of flowers) are correlated 

with the precipitation and the elevation. Two populations were 

studied at about 1000 meters altitude in a dry region with a 

sparse and degraded vegetation (Londonio & Kvist 81). and the 

second in a more humid area with a more luxuriant vegetation 

(Londorio & Kvist 72). At the first locality the shoots rarely 

exceeded 30 cm; these were all unbranched with no remnants of 

old capsules, but all had well-developed scaly rhizomes. 

According to the local inhabitants the plants flowered in the 

rainy season. At the second locality many plants exceeded a 

meter in height and were branched. Many plants had buds 

FIGURE 7 (left).-Pollen of Kohleria (scale length = 5 pn): a,b. K. allenii; c,d, 

K. amabilk var. bogotensk; e, K. hirsu!a var. hirsuta; f, K. hondensis; gb, K. 

inaequalir var. ocellata; i-k, K. neglecta; l,m, K. rugata; n-p, K. stuebeliana; 

q, K. villosa var. villosa; r, K. warszewiczii. Micrographs a,b, of cultivated 

material, US acc. 2469223; c,d, of Purdie s.n. (K); e, of Aymard C. 2191 (US); 

f, of Mason 13806 (US); g,h, of Toms 2447 (US); i-k of Garcia-Barriga 

10737 (US); l,m, of Breedlove 11885 (US); n-p of Cuatrecasas3511 (US); q, 

of Harling & Andersson 23222 (US); r, of Agredo 119 (US). 

toward the apex and numerous degraded capsules or at least 

pedicels left from previous flowers below (Figure 3a). 

FLOWERS AND POLLINATION 

The nectary morphology shows considerable variation in 

Kohleria. The dominant type consists of five free glands 

(Figure lob). Fritsch (1894) characterized the tribe Kohlerieae 

as having five nectary glands (Figure lob-d), in contrast to the 

annular nectaries in his tribe Gloxinieae. However, K. allenii 

(Figures 10a, 29g) apparently always has annular, but usually 

shallowly to deeply five-lobed nectaries; this feature has also 

been observed in populations of K. hirsuta, K. neglecta (Figure 

32e), K. peruviana (Figure 330, K. rugata, K. tubiflora, and K. 

warszewiczii. The arrangement and the shape of nectary glands 

also varies; K. amabilis and usually K. warszewiczii (Figure 

1Oc) have three free glands while the two dorsal are basally 

fused, K. inaequalis (Figure 10d) has five free glands, but often 

two of these are placed closely together, and K. villosa (Figures 

10e, f, 36f, 37e) and K. diastemoides (Figure 3Oe) have glands 

that are two to four times as long as wide at the base. These 

glands are similar to those found in Diastema, while those of 

other Kohleria species are at most 1.5 times as long as wide at 

the base. 

The size of the flowers of some Kohleria species is unusually 

variable (Figure 11). The shape of the corolla tube varies from 

narrow to broadly tubular (Figure 12) or less commonly 

campanulate (Figure 29c). The size of the limb varies from very 

small in K. hondensis and K. tubij7ora (Figure 12g,h) to large 

and conspicuous in K. Iongicalyx (Figure 120, and in K. 

inaequalis var. inaequalis (Figure 250. The tube has some 

shade of red outside and frequently yellow basally and 

ventrally, The limb usually is yellow or red with a pattern of 

darker red, violet, or purple dots or lines. The four anthers are 

coherent by their apices and sides (Figure 33d) and included or 

subincluded, and a dorsal staminode is always present (Figure 

Virtually all Gesneriaceae, including Kohleria, have protandrous flowers. A brief apparently bisexual stage has been 

recorded in K. hirsuta, K. hondensis, K. spicata, K. trianae, and 

K. warsrewiczii (pers. obs.). During this stage the process of 

style elongation causes the stigma to be temporarily located 

near the anthers, and these may shed pollen onto a stigma that 

may already be receptive. Thereupon the filaments coil and pull 

the anthers away from the stigma, and later in the female stage 

the anthers are found back into the flowers as a collapsed 

fungus-infested mass. The temporary bisexual stage indicates 

that self-pollination is a possibility. However, both personal 

experiences, mostly with K. hirsuta, and discussions with other 

growers support the idea that at least the commonly grown and 

therefore well-known Kohleria species require crosspollination. 

Weihler (1975, 1976) considered the genera Kohleria, 

Moussonia, and Parakohleria to be adapted to hummingbird 

3%). 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 8.Stolons and scaly rhizomes of Kohteria: a, stolons of K. amabilis var. bogofensir; b, scaly rhizome 

of K. hirsuta var. hirsuta; c, seedling of K. warszewiczii with scaly rhizome; d, scaly rhizomes and habit of K. 

hondensir, clone of plants from rock-crevice. a of Londono & Kvirl78 (AAU); b of Londono & Kvisf 75 (AAU); 

c of Londono & Kvirf 62 (AAU); d oLondonoio & Kvisf 69 (AAU). 
NUMBER 79 15 

9 

8 

7 

6 

5 

4 

3 

2 

1 

0 

9 

8 

7 

6 

5 

4 

3 

2 

1 

0 

9 

8 

7 

6 

5 

4 

3 

2 

1 

0 

K. rugata 

*$an Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 

K. villosa 

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 

K. warsze w iczii 

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 

FIGURE 9.-Flowering phenology of Kohleria species. The number given is that of specimens with flowers 

recorded from each month. 
16 SMITHSONIAV CONTRIBUTIONS TO BOTANY 

a 

d 

b 

e 

C 

f 

FIGURE 10 (left).-Nectary types of Kohleria: a, five-lobed ring as found in K. 

allenii; b, five free glands as found in K. spicafa; c, three free and two basally 

fused glands, the usual state found in K. warszewiczii; d, five free glands with 

two of these placed close together, the usual state found in K. inaequalis; e, 

three free and two basally fused narrow glands, a rare state found in K. villosa; 

f, five free narrow glands, the usual state found in K. villosa. 

FIGURE 11 (below).-CoroUa length variation (in mm) found in the 17 

Kohleria species. Two of the four most frequently collected species, K. spicata 

(762 collections) and K. tubiji'ora (260 collections), show little variation, in 

contrast with the two other commonly collected species, K. hirsuta (255 

collections) and K. inaequalis (153 collections). 

FIGURE 12 (right).-Flowers of Kohleria: a, K. ambilis var. bogotensis, lateral 

and front views; b, K. amabilis var. amobilis, lateral view; c, K. rugata, lateral 

view; d, K. rugata, lateral view; e, K. spicafa, lateral and front views; f, K. 

Iongicalyx, lateral and front views (note the scale is reduced compared with that 

of the other flowers); g, K. hondensk, lateral and front views; h, K. fubifora, 

lateral view; i, K. trianae, lateral and front views; j, K. warszewiczii, lateral and 

front views. a of Londono & Kvist 78 (AAU); b of Haught 2422 (US); c of Ton 

2934 (US); d of Nelson 2890 (US); e of Haughr 4634 (US); f of Londono & 

Kvist 153 (AAU); g of Londono & Kvist 69 (US); h of Nee 8598 (US); i of 

Londono & Kvist 51 (US); j of Londono & Kvkf 62 (US). 

Speciesflength (cm) 0 1 2 3 4 5 6 

1. K. allenii 

2. K. amabilis 

3. K. diastemoides . - 

4. K. grandiflora . 5. K. hirsuta 

6. K. hondensis . - 

7. K. inaequalis . 8. K. longicalyx . 9. K. neglecta - 

10. K. peruviana 

11. K. rugata 

12. K. spicata 

13. K. stuebeliana 

14. K. trianae 

15. K. tubiflora 

16. K. villosa 

17. K. warszewiczii 

0 1 2 3 4 5 6 
NUMBER 79 17 

NUMBER 79 19 

pollination, which promoted their superficial similarity through 

parallel evolution. Their red, often funnel-shaped flowers 

conform to the “bird-pollination syndrome.” However, many 

populations of Kohleria, as well as Parukohleria, were 

observed in Ecuador and Colombia, without seeing hummingbirds or other visitors to their flowers (pers. obs.). The few 

observations reported on herbarium labels do not identify a 

single group of visitors, as hummingbirds, bees, wasps, and 

butterflies have all been reported to visit Kohleria flowers 

(Table 1). 

Some Kohleria species, especially K. hirsuta, may be 

pollinated by a series of unspecialized pollinators. Both the 

flowers and the habitats are extremely variable, making it 

unlikely that a single agent or even a few will function as 

pollinators. The length of K. hirsuta corollas ranges from two 

to five cm (Figures 11,24), and in central Colombia the species 

occurs from 200 meters to 2700 meters elevation. The 

small-flowered populations occur at low elevations in hot, dry, 

open scrubland, and the large-flowered ones in chilly, humid, 

high-montane forest. The flower variation in K. inaequalis is 

even more extreme, as the corolla length ranges from 1.5 to 5.5 

cm (Figures 11, 25), but in this species there is no correlation 

between flower size, precipitation, habitats, and elevations. 

FRUITS AND DISPERSAL 

Kohleria has unilocular capsules with parietal placentation 

and loculicidal dehiscence. Exposed-habitat, wind-dispersed 

species and understory, probably animal-dispersed species 

have capsules that split by two apical valves and by a slit from 

the apex to the base, respectively. However, the capsules of 

some understory species have intermediate features. 

The 11 exposed-habitat Kohleria species all have ovoid 

capsules that dehisce into two apical valves and persist in this 

stage for weeks or months, during which several hundred 

minute seeds are dispersed by wind (Figures 13c, 33h). A row 

of stiff trichomes located at the edges of the valves further 

restricts the relatively narrow opening and probably extends the 

period during which dispersal occurs. Herbarium sheets often 

have numerous bivalved capsules. In contrast, open capsules 

FIGURE 13 (left).-Capsule dehiscence in species of Kohleria and in some 

other genera in the tribe Gloxiniese: a, Parakohleria abd Wiehler, two 

fleshy valves that split from apex to base and reflex completely; b. Diastema 

racemiferum Bentham, two apical valves (2 slits) with one of the slits 

continuing to the base (similar to the capsules of Kohleria villosa); c, Kohleria 

allenii, two apical valves; d, Kohleria longicalyx, a single longitudinal slit from 

apex to base; e, Kohleria inaequalis, a single longitudinal slit from apex to 

base, but in this case possibly first forming two apical valves (most K. 

inaequalis capsules usually are more oblong and similar to those of K. 

longicalyx); f, Monopyle macrocarpa Bentham, a single longitudinal slit from 

the middle of the capsule toward the apex and base. a of Kvist et al. 60450 

(AAU); b of Steyermark & Rube 97441 (US); c of Liesner 2106 (US); d of 

Londono & Kvist 153 (AAU); e of Londono & Kvist 190 (AAU); f of Chwchill 

et al. 4735 (US). 

TABLE 1.4bservations of potential pollinators visiting Kohleria flowers 

(l = two different species seen, 

species seen, = documented with photos). 

= Lumpornir calolaema, = three different 

Species 

K. spicata 

K. tubijlora 

K. warszewiczii 

Pollinator Country Reference 

Bees’ Colombia Folsom 10437 

Hummingbirds2 Guatemala Atwood 7737 

Butterflies3 Panama Davidse & Pohl1321 

Hummingbirds Panama Croat (1978) 

Wasps4 Colombia Londono (pen. comm.) 

Hummingbirds Colombia Londono (pers. comm.) 

are rare on sheets of understory species. The capsules mostly 

are oblong (Figure 300 rather than ovoid (Figure 13c). 

During fieldwork in Colombia it was discovered that the 

oblong capsules of the understory species Kohleria inaequalis 

(Figure 13e) and K. longicalyx (Figure 13d) dehisce by a single 

slit from the apex to the base, exposing the glutinous seed mass. 

The edges of the slit have no row of trichomes as seen in the 

exposed-habitat species. The rarity of open capsules both on 

herbarium sheets and in natural habitats indicates the capsules 

to be soon shed (and consequently the seeds quickly dispersed). 

The oblong capsules of K. diastemoides (Figure 30f) are 

similar, and their dehiscence is probably also unilateral. 

The capsules of another understory species, Kohleria villosa, 

were found to have features intermediate between the bivalved 

and the unilaterally dehiscing fruits. The ovoid (Figure 36g) 

capsules first split apically into two valves, but one of the two 

slits continues to the base (Figure 13b). The valves have a few 

trichomes at the edge. The capsules of K. amubilis apparently 

are similar to those of K. villosa, but open capsules have only 

been observed on herbarium sheets. The dehiscence of the last 

understory species, K. grandiflora, is unknown as no mature 

fruits were seen. 

The understory Kohleria species probably have switched 

from wind- to animal-dispersal. The mode of seed presentation 

presumably is an adaptation to attract animals that actively 

search the exposed glutinous seed-mass. Unfortunately no 

dispersing animals have so far been observed, but insects may 

prey on the seed-masses and become covered with the seeds, 

thus dispersing them. Burtt (1970, 1976) mentions that 

water-dispersal and the casual passive transportation by 

adherence of the tiny seeds to animals contribute to the 

dispersal of understory Gesneriaceae. However, these vectors 

are hardly the primary ones for species exposing a glutinous 

seed- m ass. 

Numerous seedlings are often found close to plants of 

exposed-habitat species, e.g., Kohleria hirsuta, K. hondensis, 

and K. warszewiczii (pers. obs.). These seedlings develop scaly 

rhizomes very early. Seedlings only a few cm tall of K. 

warszewiczii have tiny, scaly rhizomes (Figure &). The 

seedlings apparently must establish scaly rhizomes before they 

flower or form stolons. This may explain why K. hirsuta, K. 

spicata, and K. tubiflora, although they thrive in dsturbed 
20 SMITHSONIAV CONTRIBUTIONS TO BOTANY 

habitats, are never me annual weeds. 

Animal-dispersal is often considered more efficient over 

longer distances than wind-dispersal (see, for example, 

Stebbins, 1971), but in Kohleria, with its numerous dust-like 

seeds, the situation may be quite the opposite. The exposedhabitat species have a much larger distribution than understory 

species (Figure 14). and the occurrence of K. spicata and K. 

tubzflora on many coastal islands (Figures 15, 16), and the 

former even on Cocos Island, -500 km southwest of the nearest 

mainland in Costa Rica, demonstrates that long-distance wind 

dispersal must occur. 

PHYTWEOGRAPHY 

Kohleria ranges from Mexico to Peru and east to Surinam 

(Figure 14). The 11 exposed-habitat species occur widely 

throughout the range, while the ranges of the six understory 

species are restricted to near the diversity center of Kohleria in 

Colombia and northwestern Ecuador (Figure 14). The entire 

ranges of all 17 Kohleria species are first discussed below, 

followed by the distributions within Colombia (14 species) in 

more detail. 

Three exposed-habitat species are widely distributed, and to 

each of these a locally distributed, allopatric species appears to 

have close affinity. The most common species, Kohleria 

spicata (Figure 15), occurs from Mexico to Peru, and is 

replaced by K. stuebeliana on the eastern Andean slopes of 

Colombia. The second most common species, K. tubifora 

(Figure 16), is widespread in Central America and northern 

South America, and is replaced by K. hondensis in the Rio 

Magdalena valley of Colombia. The third widespread species, 

K. hirsuta (Figure 17), ranges from the Colombian highlands 

through Venezuela to Surinam. Further to the south, in Peru, 

the species is replaced by K. peruviana (Figure 17). Two 

species, K. allenii and K. rugata (Figure 18), are endemic to 

Costa Rica and adjacent Panama, and southern Mexico, 

respectively. The remaining three exposed-habitat species, K. 

trianae, K. neglecta, and K. warszewiczii (Figure 19), are 

endemic to Colombia. The only common understory species, 

K. inaequalis, is widespread in the Colombian highlands and 

northwestern Ecuador (Figure 28). The five other understory 

species, K. amabilis, K. diastemoides, K. grandflora, K. 

longicalyx, and K. villosa, are rare and local endemics in the 

Same region, although the latter species is locally common in 

Ecuador (Figure 20). 

The valleys of the south- to north-flowing Rios Cauca and 

Magdalena divide the Colombian highlands into three Cordilleras, delimiting and isolating the ranges of the 14 Colombian 

Kohleria species (Figure 21). On the eastern slopes of the 

Eastern Cordillera (the upper Amazon and Orinoco watersheds) 

six Kohleria species occur (Figure 21) including three 

endemics, K. grandflora, K. longicalyx (Figure 20), and K. 

stuebeliana (Figure 19). In the Rio Magdalena watershed eight 

species occur, making this the area of greatest species diversity 

(Figure 21). Among these are three endemics, K. amabilis 

(Figure 20), K. hondensis (Figure 16), and K. neglecta (Figure 

19). In the Rio Cauca watershed six species occur (Figure 21) 

but only the rare K. diastemoides (Figure 20) is endemic. 

Finally five species also occur in the area draining to the Pacific 

Ocean (Figure 21), but only K. villosa (Figure 20) is restricted 

to the western Andean slopes. Kohleria trianae is endemic to 

the Central Cordillera, including both the eastern and western 

slopes (draining to Rio Magdalena and Rio Cauca, respectively 

(Figures 21, 22)). The related K. hirsuta (Figures 21, 22) 

mostly is found in the Eastern Cordillera. K. inaequalis (Figure 

28), K. spicata (Figure 15), and K. warszewiczii (Figure 19) are 

widespread in the Magdalena and Cauca watersheds as well as 

in the Pacific area (Figure 21). Finally K. tubiflora (Figure 16) 

is widespread but absent from the Magdalena watershed (where 

K. hondensis occurs). 

Kohleria species often have a broad altitudinal distribution 

range, e.g., K. hirsuta and K. spicata (Figure 21). However, 

even the species concentrated within a specific altitudinal belt 

are often represented by a few collections from much higher, 

e.g., K. stuebeliana and K. tubifora, or much lower altitudes, 

e.g., K. inaequalis and K. warszewiczii. The latter two species 

only reach the lowland in the extremely humid Pacific 

Colombia (Figure 21). 

POPULAR NAh4ES AND MEDICINAL USES 

Popular names have been recorded for species of Kohleria in 

Venezuela, Colombia, and Central America (Table 2). In the 

Central and Western Cordilleras of Colombia the most frequent 

name for Kohleria species that possess scaly rhizomes is 

“caracola,” a name that is likewise used for Heppiella ulmifolia, 

which also has scaly rhizomes (Kvist, 1987). Perez-Arbeahez 

(1947) confirmed that the name “caracola” referred to the scaly 

rhizomes. In the Eastern Colombian Cordillera and Venezuela 

an often-used name for Kohleria hirsuta is “tusilla” (Table 2). 

Garcia-Barriga (1975) reported that both “caracola” and 

“tusilla” refer to a drug produced from the scaly rhizomes. 

In Colombia and Venezuela, the scaly rhizomes of Kohleria 

are boiled for a beverage drunk as a treatment for kidney 

afflictions. This use has been recorded for K. hirsuta in the 

Colombian Dept. Tolima (Londono & Kvist 72, 81), from 

Venezuela by Steyermark and Huber (1978), and for K. spicata 

in the Colombian Dept. Valle (Herrera 975). The latter species 

was also claimed to be useful for treatment of venereal disease. 

In Guatemala K. spicata is said to be an astringent (Ruana 393). 

This effect is confirmed by Garcia-Barriga (1975) in Colombia, 

and the above-mentioned applications may very well also be 

based on the astringent properties. PCrez-Arbeliez (1947) 

reported that Kohleria spp. were used to regulate liver and 

kidney functions in Colombia. In addition, label data of an 

apparently Colombian K. tubifora collection (collector unknown, at K) indicate the plant was used “in cases of dysentery 

with effect (as a decoction).” 
NUMBER 79 21 

FIOL?RE 14.-Entire range of Kohleria and the diversity center of the genus where 19 of the 22 accepted taxa 

occur. The latter area is also the apmximate range of the Kohleria species that have capsules splitting all the way 

to the base as an apparent adaptation to animal-dispersal. 

20 

10 

0 

I 

I 

ia 
0 

0 

NVINOSHLINS SNOLLflBRILN03 OL WLOB 

0 

DC 

zz 
NUMBER 79 23 

Kohleria hondensis 

0 K. tubiflora 

10 

0 

FZomE 16.--Distributions of Kohleria fubifora and K. hodensis. The barriers formed by the Central and 

Eastern Colombian Cordilleras appear to isolate the two related species. 
24 SMITHSONIAV CONTRIBUTIONS TO BOTANY 

FIGURE 17.-Distributions of Kohleria hirsufa and K. peruviunu. (Figure 22 shows the distribution of K. hirsua 

in Colombia, including var. iongipes endemic to that country.) 
NUMBER 79 25 

FIGURE lS.-Distributions of the two Mexico and Central American endemics, Kohleria allenii and K. rugata. 

20 

10 
26 SMITHSONAN CONTRIBUTIONS TO BOTAhY 

FIGURE 19.-Distributions of Kohleria neglecta, K. stuebeliana, and K. warszewiczii (note the single collection 

of K. stuebeliana from the western slopes of the Eastern Cordillera within the limited range of K. neglecta). 
NUMBER 79 27 

Kohleria amabilis 

o var. amabilis 

m var. bogotensis 

A K. diastemoides 

A K. grandiflora 

@ K. longicalyx 

K. villosa 

ROW 20.-Distribution of all Kohleria species and varieties (except K. inaequulir; see Figure 28) having 

capsules that split all the way to the base. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Western Cord. Central Cord. Eastern Cord. 

AMABlLlS -I---- 

DIASTEMOIDES---- 

GRANDIFLORA ...... 

Pacific Ocean 

A 

Magdalena Valley 

HIRSUTA ---__--___ 3000rn 

HONDENSIS........ 

INAEQUALIS --I_ 

LONGICALYX --- A 3000rn 

NEGLECTA ________ SPICATA ------ 

STUEBELIANA ...... 

TRIANAE ............ 3000rn 

TUBIFLORA ____ VILLOSA __________ WARSZEWICZllIFIGURE 21.--Distributions and altitudinal ranges of the 14 Kohleria species that occur in the Colombian 

highlands. The three Colombian Cordilleras have been transected east-west, and the ranges of the individual 

species within each of the four watersheds are shown. 
NUMBER 79 29 

Kohleria hirsuta 

var. hirsuta 

o var. longipes 

A K. trianae / 

FIGURE ZZ.-Estributions in Colombia and adjacent Venezuela of Kohleria trianae, K. hirsuiu var. longipes, and 

K. hirsuiu var. hirsufu (see Figure 17 for the entire distribution of the variety hirsufa). 
TABLE 2.-Vernacular names of Kohleria species given on herbarium labels and on a Venezuelan stamp. 

SueciesKountrv 

K. grandiflora 

Colombia 

K. hirsufa 

Colombia 

Venezuela 

K. hondensis 

Colombia 

K. spicata 

Colombia 

El Salvador 

Guatemala 

Honduras 

K. stuebeliana 

Colombia 

K. trianae 

Colombia 

K. tubifora 

Panama 

K. warszewiczii 

Colombia 

Colombia 

Cultivated hybrid 

Honduras 

Province Collector Vernacular name 

Putumayo 

Cundinamarca 

Cundinamarca 

Cundinamarca 

Dist. Federal 

Miranda 

unknown 

(Dist. Federal?) 

Tolima 

Antioquia 

Antioquia 

Antioquia 

Antioquia 

Antioquia 

Valle 

Valle 

Valle 

El Salvador 

San Vicente 

Guatemala 

Tegucigalpa 

Meta 

Antioquia 

unknown 

Canal area 

Antioquia 

Antioquia 

Caldas 

Cauca 

Morazan 

Lehmann 4018 

Cuatrecasas 13578 

Fermandez 1309 

Forero 139 

Elks 411 

FerMandez 4 

from a stamp 

Steyermark & Huber 

(1978) 

Andre 1949 

Daniel 809 

Daniel 1355 

Daniel 3 92 1 

Rodriguez 24 

Tomas 977 

Cuatrecasas 22568 

Figuieras 8198 

Herrera 975 

Calderon I74 

Standley 21462 

Ruana 393 

Hagen et al. 1211 

Fosberg I9468 

Daniel 232 7 

unknown 

Paul 161 

Daniel 391 7 

Tomas 612 

Tomas I91 9 

Cabrera R. 7869 

Molina R. 14581 

Flor de Bolivar 

Tusilla 

’hcilla 

El bum cuidadano 

Caljois del Diablo 

Tusilla 

Calmcito del diablo 

Tusilla 

Yerba de almonan 

Caracola 

Caracola 

Caracola roja 

San Juanita 

Caracola 

Caracola 

Caracola 

Caracola 

Digital Montes 

Diente de Perro 

Mazorco 

Sulfatillo 

Pana silvesto 

Caracola 

Sangremaria 

Pemto 

Caracola 

Caracola 

Caracola 

Chande, Yute 

Flor de Pena 

A hybrid, possibly between Kohleria amabilis and K. 

hirsuta, is cultivated in Ecuador and used by the Cayapa 

(Chachi) and Colorado (Tsatchela) Amerindian tribes for 

treating snake bite (Kvist, 1986). Other Gesneriaceae have the 

same utility (Holm-Nielsen et al., 1983; Kvist, 1989; Kvist and 

Holm-Nielsen, 1987). In Carchi province of Ecuador K. 

inaequalis (Boeke 838) was used to “provoke menstrual flow,” 

probably by the local Coaiquer (Awa) Indians who often rely 

on the “doctrine of signatures” in their use of plants 

(Holm-Nielsen and Barfod, 1984). In this case, the red flowers 

and brown-red pubescence are probably associated with blood. 

LNTERSPECIFIC HYBRIDIZATION 

Until now few interspecific hybrids have been reported from 

Gesneriaceae in native habitats. Morley (1971) described a 

Columnea hybrid-swarm, Kvist and Skog (in press) mention 

extensive hybridization between C. ericae Mansfeld and C. 

guttata Poeppig, and Wiehler (1983) mentions introgression 

between Kohleria spicata and K. villosa in northwestern 

Ecuador. On the other hand numerous Kohleria hybrids have 

been produced by hand-pollination in cultivation since the 

middle of the 19th century. All Kohleria species may 

potentially be able to interbreed, and the hybrids normally are 

partially fertile. The parentage of many early hybrids is 

unknown, but the following species were in cultivation in the 

last century and have probably been involved in artificial 

hybridization: K. amabilis, K. hirsuta, K. hondensis, K. 

inaequalis, K. rugata, K. spicata, K. trianae, K. tubijlora, and 
NUMBER 79 

75 

50 

25 

.o 

K. trianae K. hirsuta 

75 

50 

25 

-0 

31 

CALYXLOBESLENGTHS I 1 [I 0 

INFLORESCENCES I 1 p i] not exceeding 5 mm 

pedunculate 

Colombian Cordilleras Venezuela 

Central Central Eastern 

Number of collections 17 18 18 32 

FIOW 23.-Introgression between Kohferia hirsufo and K. trionoe. The frequencies of pedunculate 

inflorescences and calyx lobes being shorter than five mm in populations of K. trionae-a species restricted to the 

Central Colombian Cordillera-and in populations of K. hirsuto in the Central Colombian Cordillera, in the 

Eastem Colombian Cordillera, and in Venezuela. Both characters characterize K. triome but are only common in 

K. hirsuto populations sympatric with K. triome indicating introgression. 

K. warszewiczii. Some species, e.g., K. hondensis and K. 

trianue, died out in cultivation early and were reintroduced into 

cultivation only during this study. Kohleria hybrids still enjoy 

widespread horticultural popularity. The currently available 

hybrids and cultivars were discussed by Batcheller (1985). 

Interspecific hybridization is a fairly common phenomenon 

in native populations of Kohleria. Three cases are discussed 

below; the first describes first generation hybrids between two 

apparently distantly related Kohleria species; the second, local 

hybridization between two widely distributed sympatric species; and the third, hybridization and introgression between two 

closely related species whose ranges mostly are allopauic 

separated by a topographic barrier. Hybridization between 

subspecific taxa (varieties) and between Kohleria and related 

genera are discussed in the sections below, “Intraspecific 

Hybridization” and “Intergeneric Hybridization.” 

In northwestern Ecuador, where both the understory species 

Kohleria inaequalis and the exposed-habitat species K. spicata 

are common, a putative hybrid between them, growing in a 

large clonal population, was studied (Kvist et al. 48659). Its 

indumentum, inflorescences, and flowers (Figure 25b) were 

intermediate between the parent species (Figures 12e, 25d-f). 

A specimen from adjacent Colombia is similar (Soejarto 942), 

and both these specimens apparently are first generation 

hybrids, while another Ecuadorian collection may be a 

backcross with K. inaequalis (Jativa & Epling 843). 

The two exposed-habitat species, Kohleria hirsuta and K. 

tubflora, are sympatric throughout much of their ranges, but 

hybridization only has been recorded from Trinidad. Specimens of the putative hybrids (e.g., Adam 14358, Simmonds 

275) have tubular, ventricose corollas with a reduced limb 

resembling those of K. tubflora (Figure 12h), although the 

throat is wider, but the calyx lobes are lanceolate like those of 

K. hirsuta (Figure 24). The hybrid differs from both parents in 

having red or purple lower leaf surfaces. Hybrid material was 

introduced into cultivation and considered to be me K. hirsuta 

by Raymond (1968), Stone (1978, with photo), and Bornstein 

(1986, with photo). 

In Colombia, the main ranges of Kohleria hirsuta and the 

related K. trianue are separated by the low, hot, and dry Rio 

Magdalena valley (Figure 22). Kohleria trianae is endemic to 

the Central Cordillera between 900 and 2500 meters elevation 

but is only common between 1700 and 2200 meters (Figure 

21). Plants of Kohleria hirsuta mainly are found in the Eastern 

Cordillera, but also occur on the eastern slopes of the Central 

Cordillera at elevations between 400 and 1200 meters (Figure 

21). The latter populations frequently have features that 

characterize K. trianae and almost certainly indicate hybridization and introgression with this species; e.g., pedunculate 

partial florescences (but their lengths do not exceed the length 

of the pedicels as in K. trianae) and less than five mm long 

triangular calyx lobes rather than longer, lanceolate ones 

(Figures 23, 24). Kohleria hirsuta may only have been able to 

colonize the lower slopes of the Central Cordillera where K. 

trianae is rare or absent (Figure 21). At higher elevations 

extensive hybridization with K. trianae may “eliminate” 

occasional K. hirsuta plants. Kohleria hirsuta populations in 

the Eastern Cordillera have a smaller frequency of K. trianae 

attributes than those in the Central Cordillera but a higher 

frequency than in the more distant populations of Venezuela 

indicating introgression (Figure 23). Kohleria trianae may 

occasionally wind-disperse across the Rio Magdalena valley 
SMITHSONIAN CONTRIBUTIONS TO BOTANY

32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 24.-Rowers of Kohleriu hirsuta: a. var. hirsuta, lateral and front views; b, var. longipes, lateral view; 

c, var. hirsufu, lateral view: d, var. hirsuta, lateral view (example showing introgression with K. friunoe). a of 

Kvirl416 [cultivated material ofLondono & Kvirt 721 (AAU); b of Schultes 5692 (US); c of Kvirt 41 7 [cultivated 

material of Londono& Kvist 811 (AAU); d of Andre1967 (NY). 

and hybridize with K. hirsuta. A single old collection of K. 

trianae actually came from the Eastern Cordillera (Holton 

611A). 

INTRASPECIFIC HYBRIDIZATION 

Kohleria hirsuta and K. inaequalis (Figures 4, 11) are 

extremely variable when compared with the 15 other species in 

the genus. Both “species” apparently are complexes of formerly 

well-delimited semi-species that have hybridized extensively 

and broken down their limits. 

In Colombia and Venezuela Kohleria hirsuta is sympatric 

with 10 and 2 other Kohleria species, respectively. The species 

is only unusually variable in the former country. Inrrogression 

from K. trianae (see “Interspecific Hybridization” section 

above) obviously adds to this variation (Figure 23). In addition, 

two semi-species apparently are fusing in the Eastern Colombian Cordillera. In this region the populations found above 

2000 meters elevation usually are distinct by having larger 

flowers (Figure 24b) and scandent shoots. These plants are 
NUMBER 79 

Locality 

Colombia 

2000-2700 

1500-m 

1000-1500 

0-lo00 

Venezuela 

33 

Lobe width (mm) Limb width (mm) 

-2 2-3 3-4 4- -8 8-10 10-12 12-14 14-16 16-18 18-20 

14187 -227655 

64 41 1224211 

55 - 213-2- 

10 2 - 2244- 

26 6 - 14 17 7 - 

TABLE 3.-Variation in width of calyx lobes at base, and in width of corolla limb in Kohleria hirsuta populations 

at different elevations (m) in Eastern Colombian Cordillera and in Venezuela vachira, Aragua, Federal District). 

Colombian high-elevation populations referred to var. longipes have calyx lobes at least 3 mm wide, and mrolla 

limbs at least 12 mm wide. 

herein referred to K. hirsuta var. longipes, as there is a 

continuous variation from pure breeding large-flowered var. 

longipes at high elevation to typical smaller-flowered var. 

hirsuta below 1500 meters (Table 3). The variety longipes 

possibly has a common ancestor with the species K. peruviana 

from the Peruvian highlands, making the evolution truly 

reticulate (in contrast to a single species diverging into two 

species in isolation followed by hybridization after renewed 

contact). Both taxa have similar unusually large flowers 

(Figures 24b, 33c) and leaves. Annular nectaries (Figure 33g) 

also occur in both taxa (in contrast to the typical variety of K. 

hirsuta). The Colombian populations may have diverged from 

the putative ancestor, developing a climbing habit in contrast to 

K. peruviana. The rather weedy, smaller-flowered, typical form 

of K. hirsuta later may have become more common in the Rio 

Magdalena valley (possibly promoted by the disturbances of 

man) and have started to interbreed with the scandent 

populations from higher elevations-a process that has reduced 

the former semi-species to its current status as var. longipes of 

K. hirsuta. 

Kohleria inuequalis shows a variation unmatched in any 

known New World Gesneriaceae, e.g., in peduncle length, 

corolla length, limb size, indumentum, etc. (Figures 4, 11,25). 

The variation of the former two characters within Ecuador and 

several departments of Colombia (Figures 26,27) indicates that 

K. inaequalis is a complex of three extensively hybridizing 

semi-species. The pure breeding populations are still found at 

the periphery of the “species” range (Figure 28); viz., the 

large-flowered, pedunculate var. inaequalis (Figures 25f, 27) in 

Ecuador and adjacent Colombia, the small-flowered, pedunculate var. ocellata (Figures 25a, 27) in the Colombian Rio Cauca 

valley northward from Pereira, and the large-flowered, epedunculate var. lindenii (Figures 25c, 27) on the western slopes of 

the Western Cordillera in the Colombian Depts. Choco and 

Antioquia. If Dept. Antioquia alone were considered, the 

isolated populations of K. inuequalis probably would be treated 

as two well-segregated species, since both var. ocellata and var. 

Iindenii (but no intermediate forms) occur in the department 

(Figure 27). In Depts. Tolima, Risaralda, Valle, and Cauca 

(Figure 26), all three varieties blend together (Figure 27), but 

these plants mostly are referred to var. inuequalis (Figure 28). 

The indumentum, inflorescences, and corolla shapes of these 

populations mostly are intermediate between var. inaequalis 

and var. lindenii, but their flowers are considerably smaller 

than those of both of these varieties, indicating introgression 

from var. ocellata (Figure 25d,e). 

During glacial periods the three semi-species of Kohleria 

inaequdis may have speciated in isolation. The cooler and drier 

climate in those periods extended the size of the paramos 

(Simpson, 1975), with the montane forests occuring lower on 

the slopes. Consequently, from the perspective of montane 

forest species, the Andean Cordilleras were more formidable 

barriers, restricting the dispersal compared with the present 

situation, The relatively low Western Cordillera presumably 

isolated the small-flowered variety, ocellata, in the Cauca 

valley from the large-flowered varieties, inuequalis and 

Iindenii, to the southwest and the northwest respectively. 

Currently the Western Cordillera mostly is covered with 

montane forest and hardly restricts the dispersal of K. 

inaequalis significantly (Figure 21). The elevation is lowest 

west of Cali in Dept. Valle (Figures 26, 28). This is precisely 

the region where all three K. inaequalis semi-species interbreed 

extensively. 

The extreme variation in Kohleria inuequalis may also in 

part reflect a faster speciation in isolated populations of 

understory species compared to wind-dispersed, exposedhabitat populations, possibly due to the fact that the seeds of the 

former species only are dispersed locally by understory insects. 

This theory is supported by the fact that, in the tribe Gloxinieae, 

the species delimitation apparently is more problematic in the 

presumably animal-dispersed understory genera Diastema, 

Monopyle, and Parukohleria (pers. obs.) than in the winddispersed genera Achimenes (Ramirez-Roa, 1987), Heppiella 

(Kvist, 1990), and Sinningia (A. Chautems, pers. comm.). The 

species delimitation has also proven to be difficult in 

understory genera belonging to other New World Gesneriaceae 

tribes; e.g., Cremospermu (Kvist & Skog, 1988) and Reldia 

(Kvist & Skog, 1989). 
34 SMITHSONIAN CONTRIBUTIONS TO BOTAhY 

FIGURE 25.--Flowers of Kohleria inaequalis: a, var. ocellata; b, var. inaequalis x Kohleria spicata; c, var. 

lindenii; d, var. lindenii; e, var. inaequalis; f, var. inaequalis. a of Juncosa 1971 (US); b of Kvisr et al. 48659 

(AAU) and Soejarto 942 (GH, SEL); c of Forero etal. 2138 (US); d of Londono& Kvbt 68 (AAU); e of LLondono

& Kvist 190 (AAU); f of Balslev 1939 (AAU) and Kvirr et al. 48619 (AAU). 
NUMBER 79 35 

I I I 

n 

2 

Meta 

FIGURE 26.-Departments of Colombia in which species of Kohleria occur, and the number of species that occur 

in each department (and in Ecuador, Panama, and Venezuela). The approximate range of K. inaequdis is 

indicated by hatching. 
36 SMITHSON” CONTRIBUTIONS TO BOTANY 

Region 

Ecuador 

Nariiio 

Cauca 

Huila 

Valle 

Risar alda 

Quindio 

Caldas 

Antioquia 

Choc6 

Tolima 

Corolla length (em) 

12345 

Peduncle length (cm) 

0123456 

I 

I 

I 

I 

ILO 

ILO 

0 

0 

LO 

L 

L 

123 45 0123456 

FIGURE 27.-The variation in corolla length and peduncle length (in mm) of KohIeria imequalk in Ecuador and 

in the Colombian departments where the species occurs (epedunculate populations are illustrated as having very 

short peduncles). In addition, the varieties that occur in the single areas are indicated (r = var. inuequalk. 0 = var. 

ocellara, and L = var. lindenii-). 

INTERGENERIC HYBRIDIZATION 

Intergeneric hybrids have been produced between Kohleria 

and species of Koellikeria Regel (Wiehler, 1968), Parakohleria 

(Wiehler, 1976a), Diastema, Gloxinia, and Moussonia (Wiehler, 1976b). No intergeneric hybrids with Kohleria are known 

from natural habitats. The possibility that K. diastemoides 

(newly described below) is a hybrid between a species of 

Diastema and Kohferia inaequalis has been considered. The 

fact that the only known K. diastenwides collection shows a 

combination of features characterizing these two taxa rather 

than intermediate features (Table 4), and that the seed 

production of K. diastemoides apparently is abundant, argues 

against the plant being an intergeneric hybrid. Intergeneric 

hybrids usually are entirely sterile. However, Wiehler (1976b) 

reported that hybrids between K. spicata and both Diastema 

vexans H.E. Moore and Koellikeria erinoides (DC.) Mansfeld 

were weakly fertile, which may indicate a close affinity with 

these genera. Diastema is related to Kohleria (see “Related 

Genera” section below) but the affinities of Koellikeria are 

uncertain. The habit, inflorescences, and flowers of Koellikeria 

do not suggest an affinity with Kohferia. The unusual 

inflorescences with alternate subtending prophylls are similar 

to those found in the small Mexican genus Smithiantha 0. 

Kuntze, but the two genera differ in other respects. Koellikeria 

is monotypic and occurs in montane forests from Mexico to 

Argentina. 

RELATED GENERA 

The following genera are discussed: Moussonia and Parakohleria, both recently split off from Kohleria; Diastema and 

Capanea, genera that possibly have evolved from ancestors 

that might fit into the current concept of Kohleria; and 

Heppieffa and Vanhouttea as collections of these genera often 

look very similar to Kohleria. Achimenes, Eucodonia, and 

Gloxinia are discussed in the section on “Evolution’’ below 

because these genera may be close to the ancestors of Kohleria. 

The Central American genus Moussonia was until recently 

placed in Kohferia due to its similar tubular flowers and 

inflorescences. Moussonia (Wiehler, 1975) differs from Kohleria by having the basic chromosome number of 11 (rather 

than 13), a capitate stigma, and an annular nectary. Wiehler 

(1978b) split off the genus Parakohleria from Kohleria based 

on the following characters: capitate stigmas, absence of scaly 

rhizomes, basally bent corolla tubes, and congested inflorescences. Only the first of these characters separates the two 

genera consistently, but the two genera also are set apart by 

their very different capsule dehiscence. Parakohleria capsules 

differ from both the bivalved and the unilaterally dehiscing 

capsules of Kohleria (Figure 13c-e) by splitting from the apex 

to base into two fleshy valves that reflex completely to expose 

the glutinous seed mass (Figure 13a). Parakohferia (Kvist, 

1991) is an understory genus, ranging along the eastern Andean 

slopes from Colombia to Bolivia (and locally on the western 

slopes in Ecuador). 

Diastema, an understory genus of small herbs ranging from 

Mexico to Bolivia, shares many attributes with Kohleria, 

especially with the understory species K. villosa (Figures 36, 

37). This species shares the following features with Diastema: 

long narrow nect&y glands, capsules that dehisce by a slit 

down to the base, and sub-globose seeds with an irregular 

surface. However, Diastema is consistently set apart from 
hWMBER 79 

Character 

Inflorescence 

Inflorescence 

Corolla shape 

Corolla color 

Corolla inside 

Filament base 

Capsule 

Nectary glands 

Stigma 

37 

~~ 

K. diastemoides K. inaequalis Diastema 

frondose frondose bracteose 

pedunculate pedunculate epedunculate 

narrowly tubular broadly tubular narrowly tubular 

red or purple red to purple white 

villous villous glabrous 

villous villous glabrous 

oblong oblong ovoid/oblong 

narrow and high broad and short narrow and high 

bilobed bilobed bilabiate 

FIGURE 28.-Distributions of the varieties of Kohleria inaequalis: var. inaequalis, var. ocellata, and var. lindenii. 

The Pacific coast collection is from Gorgona Island in the Dept. Cauca of Colombia (Guano & Rangel 5280). 

Kohleria by having bracteose inflorescences (Figure 3g), and 

by mostly having small white or yellowish flowers often with 

blue or purple spots or lines on the limb. In addition, Diastema 

has scaly rhizomes (in contrast to the most similar Kohleria 

species), and the stigma is not conspicuously bilobed. 

Diastema possibly is a satellite genus derived from Kohleria, 

but additional studies are needed. The two Cupanea species 

mostly are epiphytic climbers with capitate stigmas, exserted 

stamens, and capsules that split with four apical valves. The 

tubular to campanulate corollas, as well as the pedunculate 

axillary inflorescences and the rotundate calyx lobes of 

Capanea affinis Fritsch and Kohleria warszewiczii, are 

TABLE 4.4omparison of Kohleria diastemoides with Kohleria innequalis 

and the genus Diastema. In size and habit Kohleria diastemoides resembles 

Diastema rather than Kohleria inaequalis. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTAhY 

smkingly similar. Capanea grandflora (Kunth) Decaisne ex 

Planchon, has campanulate corollas similar to those of 

Kohleria allenii, but the latter species has different frondosebracteose inflorescences. The similarity between Capanea and 

Kohleria probably reflects a parallel adaptation to pollinators, 

e.g., bats, but the possibility that Capanea has descended from 

Kohleria still exists. 

The four species of Heppiella from the Andean highlands 

and the approximately five species of Vanhouttea from Brazil 

all have tubular red flowers making them superficially similar 

to Kohleria. Heppiella differs from Kohleria by having free 

anthers, capitate stigmas, annular nectaries, and the plants are 

often climbing or even epiphytic. Heppiella has scaly rhizomes 

and is probably a close relative of Kohleria (Kvist, 1990). The 

resemblance of Vanhouttea and Kohleria is likely caused by 

parallel evolution, as Vanhouttea occasionally develops tubers, 

suggesting a close affinity with Sinningia (A. Chautems, pers. 

comm.). Vanhouttea also differs from Kohleria by having 

annular nectaries, capitate stigmas, and by being woody shrubs. 

EVOLUTION 

Originally it was the intention to present a cladistic analysis 

of Kohleria. This has been abandoned for two reasons: (1) The 

relationships among the 24 genera currently included in the 

tribe Gloxinieae are little understood, making the selection of 

an outgroup unreliable; and (2) Populations of Kohleria 

speciate in isolation and hybridize after renewed contacts 

making the evolution reticulate. The following aspects are 

discussed below: the hypothetical ancestor of Kohleria, as well 

as the diverging of the understory species from exposed-habitat 

ancestors and the reticulate evolution of Kohleria. 

The species with fewest advanced features among the current 

Kohleria species may be the wind-dispersed K. rugata from 

Mexico. The only conspicuous feature that separates K. rugata 

populations with annular nectaries from all species in the genus 

Gloxinia (from South America) and most species of Achimenes 

and Eucodonia (from Mexico and Central America) is the 

presence of bilobed rather than capitate stigmas (the latter two 

genera differ from Kohleria and Gloxinia by having 11 and 12 

rather than 13 basic chromosomes). Achimenes, Eucodonia, 

and Gloxinia (except some advanced species that may fit better 

into Monopyle) possibly are the most primitive members of the 

Gloxinieae, being seasonal herbs with scaly rhizomes, bivalved 

capsules, annular nectaries, usually capitate stigmas, epedunculate partial florescences with 1 or 2 flowers, and frondose 

florescences. Consequently, wind-dispersal and occurrence in 

exposed habitats are primitive states in Gloxinieae. The 

understory groups, e.g., Parakohleria, Diastema, Monopyle, 

and some Kohleria species have individually advanced from 

exposed-habitat to understory habitats. During this process the 

scaly rhizomes have been reduced and the plants have 

apparently switched to animal-dispersal, evolving different 

types of capsule dehiscence in the different genera (Figure 13). 

Kohleria amabilis apparently has some features intermediate 

between exposed-habitat and understory Kohleria species, e.g., 

the capsules (Figure 13b-d) and the seeds (Figure 5a-c). In 

addition, the species is at least superficially similar to both the 

exposed-habitat species K. rugata and the understory species 

K. villosa. 

The main mode of speciation in Kohleria Seems to be 

geographical isolation; this is evident in the diverging of K. 

hondensis from K. tubflora, and K. stuebeliana from K. 

spicata. None of the 17 species apparently has evolved by 

interspecific hybridization. This is surprising as hybridization 

seems to be a common phenomenon in Kohleria, and hybrids 

usually are fertile. One explanation may be that hybrids never 

become stabilized by chromosome doubling, as tetraploids are 

unknown in Kohleria. On the contrary, the effect of hybridization has been seen to weaken or even break down the species 

limits. The two most variable species, K. hirsuta and K. 

inaequalis, apparently consist of extensively interbreeding 

semi-species herein taxonomically treated as varieties. The 

variety longipes of K. hirsuta may share a common ancestor 

with a third semi-species, K. peruviana, making the evolution 

truly reticulate. Kohleria has evolved in the dynamic environment of northwestern South America. The climate changed, 

becoming cooler and drier during glacial periods, and volcanic 

eruptions and earthquakes occasionally have caused massive 

destruction of the vegetation cover. Populations of Kohleria 

must recurrently have been isolated during extended periods 

succeeded by new contacts. The current cases of speciation in 

isolation and hybridization after renewed contacts may be the 

most recent examples of an evolutionary pattern that has 

characterized the evolution of Kohleria, possibly during 

millions of years in the rising Andean highlands. 

Kohleria Regel 

Kohleria Regel, 1847:(4). [Type: K. hirsufa (Kunth) Regel (Gesneria hirsufa 

Kunth).] 

Gesneria sect. Isoloma Bentham, 1846:230. [Lectotype: Kohleria hirsuta 

(Kunth) Regel (selected by C. Morton, 1973:317) = K. hirsuta (Kunth) Regel 

var. hirsuta.] 

Isoloma (Bentham) Decaisne, 1848a:465 [not lsoloma J. Smith, 1841:414]. 

[Lectotype: Kohleria hirsufa (Kunth) Regel (selected by C. Morton, 

1973:317) = K. hirsufa (Kunth) Regel var. hirsufa.1 

Tydaea Decaisne, 1848b468. [Type: T. picfa (Bentham ex J.D. Hooker) 

Decaisne = Kohieria amabiiir (Planchon & Linden) Fritsch var. bogofensir 

(Nicholson) Kvist & L. Skog.] 

Giesleria Regel, 1849:181. [Type.: Achimenespicta Bentham ex J.D. Hodter = 

Kohleria amabilis (Planchon & Linden) Fritsch var. bogorensis (Nicholson) 

Kvist & L. Skog.] 

Kohleria Regel, 1851:893 [not Kohleria Regel, 1847:(4)]. [Type: K. ignorafa 

(Kunth) Regel = Kohleria spicufa (Kunth) Oersted.] 

Sciadocalyx Regel, 1853a:257. [Type: S. warszewiczii Regel = Kohleria 

warszewiczii (Regel) Hanstein.] 

Brachyloma Hanstein, 1854:303. [Type: B. hirsufwn (Kunth) Hanstein = 

Kohleria hirsufa (Kunth) Regel var. hirsufa.] 

Crypfoloma Hanstein. 1859:506, 578. [Lectotype: C. hondense (Kunth) 

Hanstein (selected herein) = Kohleria hondensb (Kunth) Hanstein.] 
NUMBER 79 39 

Calycostemma Hanstein, 1859506, 556. [Type: C. lindenii Hanstein = 

Kohleria imequalis (Bentham) Wiehler var. lindenii (Hanstein) Kvist & L. 

Skog.] 

Synepileam BaiUon, 1888a:725. [Type: S. gramtensis Baillon = Kohleria 

inuequulis (Bentham) Wiehler var. lindenii (Hanstein) Kvist & L. Skog.] 

Herbs, subshrubs, or shrubs, terrestrial or epilithic, most 

parts usually with conspicuous indumentum, scaly rhizomes 

and stolons often present, adventitious roots occasionally 

present; stems decumbent to erect, rarely pendant or scandent, 

terete (usually bisulcate or subquadrangular in dried material), 

(5-) 20-150 (-400) cm long, green to yellow or purplish, 

indumentum usually pilose to villous or sericeous towards the 

apex, basally often glabrescent, shoots annual and unbranched, 

less commonly biennial or perennial and branched. Leaves 

opposite, less commonly whorled (3 or 4 at each node), equal 

to subequal or rarely unequal in pairs or whorls, petiolate; 

blades usually ovate, elliptic or lanceolate, less commonly 

cordate or oblanceolate, base oblique, often acute or rounded, 

apex acute to acuminate, margin crenate to serrate, often hispid 

or pilose-hirsute above, green to dull green, often pilose or 

tomentose below, lighter green to purple or variegated, 

secondary veins (5-) 7-9 (- 11) per side, above obscure, below 

emersed, villous or sericeous, often yellow-green. Florescences 

frondose or frondose-bracteose, partial florescences with 1 to 6 

(-10) flowers, congested in leaf ails or less commonly 

pedunculate; peduncles and pedicels sericeous, villous or 

hispid, often purplish, bracts (0-) 2-3 (-6), sessile, lanceolate 

or scaly, often caducous. Flowers protandrous; floral tube often 

villous, green, yellow-green or purple; calyx lobes equal to 

occasionally somewhat unequal, usually erect, occasionally 

reflexed, often connate --1/3 of length, or less commonly free 

nearly to base, the 5 lobes with margin entire, often sericeous 

outside and appressed pilose inside, with 3-5 sometimes 

prominent, longitudinal veins; corolla usually oblique in calyx, 

zygomorphic, occasionally subactinomorphic, tubular to campanulate, often subventricose, 1.2-6.0 cm long, tube outside 

villous to sericeous, red, often ventrally yellow, rarely 

yellowish red, inside glabrous or sparsely glandular-hairy , less 

commonly basally pilose to villous, lighter red, cream, or 

yellow, often with purple or red dots, base occasionally saccate, 

often dorsally gibbous, never spurred, outside cream to yellow, 

glabrous, throat often constricted, inside sparsely glandularhairy, limb subregular to bilabiate, of 5 ovate to rotundate 

lobes, outside usually red, rarely orange, yellow, or yellowish 

green, villous or sericeous to glabrous along the margin, inside 

usually with red or purple dots and/or lines on a paler red, 

orange, or yellowish green background, rarely entirely red; 

stamens 4, didynamous, filaments never connate but adnate to 

the base of the corolla tube for 0.3-0.8 mm, usually glabrous, 

occasionally pilose to villous, yellowish, anthers included or 

subincluded, oblong, coherent at apices and sides, glabrous, 

cream or yellowish, thecae distinct, dehiscing by longitudinal 

slits, the filaments coiling in the later female flowering stage to 

pull the anthers deeper into the flower; one dorsal staminode 

present, 1-2 mm long; nectary fleshy, yellow and glabrous, 

usually of 5 free glands, less commonly with 3 free and 2 dorsal 

basally united glands or a 5-lobed ring; ovary half-inferior to 

almost completely inferior, sericeous to villous, style usually 

pilose and often glandular-hairy, rarely glabrous, cream or 

yellow-white, stigma included or subincluded, bilobed. Fruit a 

capsule, ovoid or less commonly oblong, apex conic or rostrate, 

dehiscing apically and loculicidally into 2 valves (by 2 apical 

slits), less commonly by 1 slit from apex to base (splitting the 

ovary and the floral tube) or by 2 apical slits with 1 of these 

continuing to the base; seeds numerous, 0.3-0.6 mm long, 

often narrowly ellipsoid to ellipsoid, less commonly subglobose, surface longitudinally to transversely reticulate striate, 

less commonly irregularly striate, tawny to brown. 

Key to the Species.and Varieties of Kohleria Regel 

Fertile material is essential for identification, and fully developed flowers are often 

important. Any key is of little use for cultivated material, even within the native range of 

Kohleria, as cultivars are mainly hybrids of partly or completely unknown parentage. 

Putative natural hybrids usually have either Kohleria hirsuta or K. inaequalis in their 

ancestry, and tend to sort out with those species in the following key. Within the key the 

terms “partial florescences” and “florescences” are used for the closed inflorescences in the 

axils of subtending leaves and bracts, and for the open flowering shoots with several to 

many partial florescences, respectively (see Figure 3 for illustration of inflorescence 

terminology). 

1. Corolla campanulate, outside yellow with a flush of red trichomes, 25-32 mm long, 

diameter at throat 15-20 mm; styles glabrous; subtending leaves of florescences 

(flowering shoots) gradually reduced to bracts toward the apex, partial 

florescences epedunculate, with 1 or 2 flowers; leaves 3 at a node, glandular-hairy 

(plants of Costa Rica and Panama) . . . . . . . . . . . . . . . . . . 1. K. allenii 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

2. 

Corolla tubular or tubular to campanulate, outside usually red, 12-60 mm long, 

diameter at throat 2-21 mm; styles pubescent; subtending leaves of florescences 

(flowering shoots) mostly not reduced to bracts toward the apex and only in 

combination with 12-20 mm long corollas, partial florescences epedunculate or 

pedunculate, with 1-10 flowers: leaves opposite or less commonly 3 at a node, 

rarely glandular-hairy (plants of Mexico, Central America, and South America) 

........................................... 2 

Subtending leaves of florescences (flowering shoots) gradually reduced to bracts 

toward the apex, partial florescences epedunculate; corolla 12-20 mm long, 

diameter of limb 5-10 mm; calyx lobes triangular to ovate, 1.5-3.5 mm long 

........................................... 3 

Subtending leaves of florescences (flowering shoots) not reduced to bracts toward 

the apex, partial florescences epedunculate or pedunculate: corolla 14-60 mm 

long, diameter of limb 3-35 mm; calyx lobes triangular, ovate, rotundate, 

lanceolate, or subulate, 2-40 mm long ..................... .4 

3. Limb orange-red with red to purple dotsflines, nearly glabrous: upper leaf surface 

usually hispid, leaf arrangement frequently 3 at a node: widespread and common 

12. K. spicata 

Limb yellow-green with brown to black dots, appressed pilose and glandular-hairy: 

upper leaf surface velutinous, leaf arrangement rarely 3 at a node (plants of 

Colombia) ........................... 13. K. stuebeliana 

4. Corolla 16-30 mm long, diameter in throat 1.5-4 mm, diameter of limb 3-7 mm, 

length of lobes 1-1.5 mm; calyx lobes triangular, ovate or lanceolate, 2-8 mm 

long; partial florescences epedunculate: upper leaf surface usually hispid to 

extremely hispid ................................. 5 

Corolla 14-60 mm long, diameter in throat 3-21 mm, diam. of limb 5-35 mm, 

length of lobes 2-12 mm; calyx lobes triangular, ovate, rotundate, lanceolate or 

subulate, 2-40 mm long: partial florescences epedunculate or pedunculate; upper 

leaf surface hispid or velutinous ......................... 6 

5. Calyx lobes ovate to lanceolate, 5-8 mm long; corolla 16-20 mm long: upper leaf 

Calyx lobes triangular to ovate, 2-4 mm long: corolla 20-30 mm long; upper leaf 

6. Limb red with white edge (diameter at throat 3-5 mm) or yellow-green with purple 

dots (diameter at throat 7-13 mm); corolla narrowly tubular or tubular to 

campanulate, 25-42 mm long; partial florescences with peduncles 2-3 times as 

long as pedicels: calyx lobes triangular or rotundate, 2-6 mm long; upper leaf 

surface velutinous. ............................... .7 

Limb yellow to red with darker red or purple dotsflines, occasionally uniformly red, 

diameter at throat 3-21 mm; corolla narrow to broadly tubular, 15-60 mm long; 

partial florescences epedunculate or pedunculate; calyx lobes rotundate, ovate, 

lanceolate or subulate, 3-40 mm long: upper leaf surface velutinous or hispid 

........................................... 8 

7. Limb red with white edge: corolla narrowly tubular, tube red outside: partial 

florescences with 4 flowers ..................... 14. K. trhnae 

Limb yellow-green with purple dots; corolla tubular to campanulate, tube lavender 

outside; partial florescences with 3-10 flowers ....... 17. K. warszewiczii 

8. Partial florescences always of single flowers, epedunculate; calyx lobes ovate, 3-8 

mm long: corolla 18-32 mm long, limb yellow with red dots: leaf pairs equal, 

upper surface velutinous, lower surface uniformly colored (plants of Mexico; only 

.................................... 

surface extremely hispid (plants of Colombia) ......... .6. K. hondensis 

surface usually hispid; widespread and common ......... 15. K. tubiflora 

sympatric with K. spicutu) ............ 11. K. rugata, new combination 
NUMBER 79 41 

Partial florescences with 1 to several flowers, epedunculate or pedunculate; calyx 

lobes ovate, rotundate, lanceolate or subulate, 3-40 mm long; corolla 14-60 mm 

long, limb yellow to red with darker red or purple dotsflines; leaf pairs equal or 

rarely unequal, upper surface velutinous or hispid, lower surface uniformly 

Calyx lobes subulate, 8-11 mm long; corolla 14-18 mm long, 3-4 mm wide at 

throat; partial florescences epedunculate, of 5-6 (-10) flowers; leaf arrangement 

colored or with a pattern of purple and green (plants of South America) .... 9 

3 at a node, upper surface hispid (plants of Colombia) .............. ............................ .9. K. neglecta, new species 

9. 

Calyx lobes ovate, rotundate or lanceolate, 3-40 mm long; corolla 15-60 mm long, 

3-21 mm wide at throat; partial florescences epedunculate or pedunculate, of 1-4 

(-10) flowers; leaf arrangement rarely 3 at a node, upper surfaces hispid or 

Corolla 15-20 mm long, limb uniformly red or red with purple lines; nectary glands 

often 2-4 times taller than broad at base; calyx lobes 3-5 mm long . . , . . 11 

Corolla 20-60 mm long, limb yellow to red with darker red or purple dotsflines; 

nectary glands less than 2 times taller than broad at base; calyx lobes 3-40 mm 

long .......................................14 

11. Partial florescences usually pedunculate; leaves equal or subequal in pairs, upper 

leaf surface often hispid, lower surface uniformly colored; inside base of corolla 

and base of filaments villous .......................... 12 

Partial florescences epedunculate; leaves equal, subequal or unequal in pairs, upper 

leaf surface never hispid, lower surface often with a pattern of purple and light 

green; inside base of corolla and filaments glabrous (but corolla toward the throat 

12. Partial florescences with 1 or 2 flowers; corolla narrowly tubular, 3-4 mm wide at 

throat; nectary glands 2-4 times taller than broad at base; upper leaf surface 

Partial florescences usually with 4 flowers; corolla broadly tubular, 5-9 mm wide 

at throat; nectary glands equally tall and broad at base; upper leaf surface hispid 

13. Leaves ovate with acute apex, equal or subequal in pairs; corolla tube not with 

long villous trichomes outside, limb 12-19 mm wide. .............. ............................. 16a. K. villosa var. villosa 

haves ovate to lanceolate with acute to acuminate apex, most leaf pairs 

conspicuously unequal in pairs; corolla tube with villous trichomes outside, limb 

9-13 mm wide ........... 16b. K. villosa var. anisophylla, new status 

14. Calyx lobes ovate, 3-5 mm long; corolla 24-32 mm long; upper leaf surface 

velutinous; partial florescences epedunculate (plants of Colombia) ...... 15 

Calyx lobes usually lanceolate, rarely ovate, occasionally rotundate to spathulate 

(but then in combination with 45-60 mm long corollas), 3-40 mm long; corolla 

20-60 mm long; upper leaf surface hispid or velutinous; partial florescences 

epedunculate or pedunculate (plants of South America) ............ 16 

15. Diameter of limb 20-27 mm, ventral lobe up to 10 mm long, all lobes inside with 

a pattern of red dots on a paler pink background; lower leaf surface with a 

conspicuous pattern of light green blotches on a purple background ....... .2a. K. amabilis var. amabilis 

Diameter of limb 13-18 mm, ventral lobe up to 6 mm long, lateral and ventral lobes 

with red or purple lines on a yellow background and dorsal lobes red; lower leaf 

surface without a conspicuous purple and green pattern .............. 

velutinous (plants of South America) ...................... 10 

10. 

and apical part of filaments usually villous) .................. 13 

velutinous .................... 3. K. diastemoides, new species 

........................... 7c. K. inaequalis var. ocellata 

........................... 

.................... 2b. K. amabilis var. bogotensis, new status 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

16. Calyx lobes rotundate to spathulate; corolla 45-60 mm long, tube 38-46 mm long, 

inside glabrous, limb red with purple lines; partial florescences with 1 flower 

............................ 4. K. grandiflora, new species 

Calyx lobes lanceolate, rarely ovate; corolla 20-60 mm long, tube 18-50 mm long, 

inside glabrous or especially basally pilose to villous, limb orange-red with purple 

dots or red with purple lines; partial florescences with 1-10 flowers ..... 17 

17. Calyx lobes 2-25 mm long, usually reaching half of corolla length; corolla 45-60 

mm long, tube 40-50 mm long, inside glabrous; upper leaf surface velutinous; 

partial florescences pedunculate .......... 8. K. longicalyx, new species 

Calyx lobes 3-25 mm long, rarely reaching half of the corolla length; corolla 20-52 

mm long, tube 18-45 mm long, inside glabrous or especially basally pilose to 

villous; partial florescences pedunculate or epedunculate; upper leaf surface 

velutinous or hispid ............................... 18 

18. Limb red with purple lines, 9-32 mm wide; mature capsules 2-3 times as long as 

wide; corolla basally villous to pilose inside; upper leaf surface usually hispid 

.......................................... 19 

Limb orange-red with purple dots, 6-22 mm wide; corolla glabrous inside; mature 

capsules 1-2 times as long as wide; upper leaf surface soft-velutinous .... 20 

19. Partial florescences of several flowers, usually pedunculate; corolla 22-52 mm 

long, tubular to broadly tubular, not subventricose; upper leaf surface usually 

hispid ...................... 7a. K. inaequalis var. inaequalis 

Partial florescences of single epedunculate flowers; Corolla (25-) 35-52 mm long, 

narrowly tubular to tubular, often subventricose; upper leaf surface occasionally 

hispid ................. 7b. K. inaequalis var. lindenii, new status 

20. Base of corolla saccate and sharply delimited, corolla 35-52 mm long; stems and 

lower leaf surfaces densely villous, whitish; length of leaves usually 8-18 cm 

(plants of Peru) ......................... 10. K. peruviana 

Base of corolla not saccate, corolla 22-52 mm long; stems and lower leaf surfaces 

pilose, greenish or purplish; length of leaves usually 4-12 cm (plants of northern 

South America) ................................. 21 

21. Width of calyx lobes at base 1-2 mm; length of corolla 20-50 mm, diameter 3-8 

mm at throat, limb 6-12 mm wide; rarely above 2000 meters .......... ............................. 5a. K. hirsuta var. hirsuta 

Width of calyx lobes at base 2-6 mm; length of corolla 32-50 mm, diameter 8-15 

mm at throat, limb 12-22 mm wide; rarely below 1800 meters. ......... ...................... 5b. K. hirsuta var. longipes, new status 

1. Kohleria allenii Standley & L.O. Williams 

Figures 7a.b (pollen), 13c (capsule dehiscence), 29 (habit and floral 

features); L.O. Williams, 1968:293 (photo). 

Kohleria allenii Standley & L.O. Williams in LO. Williams, 1968:291. [Type: 

Costa Kca, Puntarenas, near Palmar Sur, rocky railroad cuts, Allen 6300 (F, 

holotype: EAP, GH, US, isotypes).] 

Herbs or subshrubs, often epilithic, scaly rhizomes weakly 

developed or absent, stolons not seen; stems erect, rarely 

branching, glandular-hairy, purplish, 50-150 cm long, basal 

diameter 5-8 mm. Leaves temate or whorled (4 at each node), 

internodes 4-11 cm; the blades ovate to elliptic, less 

commonly lanceolate to oblanceolate, 6-15 x 3-7 cm, above 

pilose and usually hispid, often glandular-hairy, below pilose 

and usually glandular-hairy, veins below villous, yellow-green 

or purplish; petioles 1-5 cm. Florescences frondose-bracteose, 

partial florescences with 1 or 2 flowers, the subtending leaves 

toward the apex reduced to subsessile, ovate to obovate 

prophylls; pedicels 2-5 cm long, villous and glandular-hairy, 

bracts scaly, caducous. Floral tube villous and glandular-hairy; 

calyx 6-8 mm long, glandular-hairy, lobes erect, occasionally 

reflexed, ovate to triangular, 4-5 mm long and 3-4 mm wide 

at base, outside pilose to villous, inside sericeous, veins 

prominent; corolla 25-32 mm long, campanulate, tube 20-25 

mm long, diameter 6-8 mm at base and 15-20 mm at throat, 

outside sericeous and glandular-hairy, yellow with a flush of 

red trichomes, inside glabrous, yellow with red blotches, the 

base dorsally gibbous, limb 22-30 mm wide, lobes 7-9 mm 

long and 7-9 mm wide at base, inside sparsely glandular-hairy, 

ventral and lateral lobes green with purplish dots and lines, 

dorsal lobes yellow-red; filaments 18-22 mm, glabrous, 

anthers -3.2 x 3.0 mm, included; nectary a 5-lobed ring, up to 

1 mm tall; ovary half-inferior, 5-6 mm tall, diameter 4-5 mm, 

style 15-20 mm long, glabrous, stigma subincluded. Capsule 
NUMBER 79 43 

FIGURE 29.-Kohleria allenii: a, stem and leaves; b. sturdy root with small scaly rhizome; c, frondose-bracteose 

florescence; d, flower, e, ventrally opened corolla showing parts of androecium; f, calyx and gynoecium; g, 

capsule. a,b,g, from Davidse & Pohl 1401 (MO); c, from Allen 5339 (MO); d,e.f. from Allen 6300 WO). 
44 SMITHSOhTAN CONTRIBUTIONS TO BOTANY 

ovoid, 10-12 mm long, diameter 6-7 mm, bivalved; seeds 

ellipsoid, longitudinally striate, -0.40 x 0.15 mm. Chromosome number n = 13 (Wiehler, 1975). 

DISTRIBUTION.-Kohhleria allenii is endemic to the Prov. 

Puntarenas in southwestern Costa Rica, and the Prov. Chiriqui 

of adjacent Panama (Figure 18). The species ranges from sea 

level up to 1000 meters altitude. 

OTHER COLLECTIONS.--€OSTA RIGA: PUNTARENAS: Galfit0 de Golfo Dulce, Allen 5339 (BM, COL, F(2), G, GH, 

M0(2), NY(2), US(2)); Palmar Sur, Allen 6300 GAP, F, GH, 

US); between Agua Buena and Villa Neily, Burger & Matta 

4595 (F, NY); Golfito, Burger & Matta 4753 (NY); Rio 

Catarata along Carretera Interamericana, canyon of Rio 

Terraba, Davidse & Pohll401 (F, MO, US); 8 km N of Villa 

Neily, Gillis & Plowman 10074 (GH); road from San Vito to 

Ciudad Neily, Grayum 3393 (MO); Rincon de Osa, Grayum 

4020 (BM, US); between Agua Buena and Villa Neily, Golfito, 

Jimenez M. 2445 (BM, F, MO); Rincon de Osa near Riyitos, 

Kennedy 1917 (MO); 5 km W of Rincon, Liesner 2106 (MO, 

NY, US(2)); Liesner 2115 (MO); between Rio Neily and 

Palmar, Rio Coto, McKee 11190 (US); Rincon de Osa, Utley & 

Utley 11.50 (F, SEL). LOCALITY UNKNOWN: Laskowski 1402 

PANAMA: CHIRIQLI~: Burica Peninsula, 19 km W of Puerto 

Armuelles, Busey 609 (MO, NY); Rio Escarrea, Buenavista, 

Butcher s.n. (SEL). Total of 16 collections seen. 

DISTINGUISHING FEATURES.-The combination of long 

spicate inflorescences (frondose-bracteose synflorescences) 

and large campanulate flowers makes Kohleria allenii a very 

distinctive species (Figure 29). Glandular trichomes are more 

prevalent in K. allenii than in any other Kohleria species. 

Kohleria allenii is the only species that has glabrous styles. 

HABITAT.-Kohleria allenii occurs in calcareous areas, 

possibly the original habitat, and on roadsides. Most plants 

apparently grow on steep slopes. 

PHENOL0GY.-Kohleria allenii has been collected with 

flowers from October to February and in July. 

NOTES.-The species may survive drought by having 

well-developed roots rather than scaly rhizomes. The latter 

feature has only been observed on one herbarium sheet 

(Davidse & Pohll401; Figure 29b). However, scaly rhizomes 

may be more common in natural habitats. Plant collectors could 

easily fail to collect rhizomes deeply buried in rock crevices. 

The corollas of Kohleria allenii are similar to those of other 

presumably bat-pollinated species of New World Gesneriaceae, 

e.g., Capanea grandiflora (Vogel, 1958), species of Paliavana 

and Lietzia (A. Chautems, pers. comm.), and some species of 

Gesneria (Skog, 1976). 

03. 

2. Kohleria amabilis (Planchon & Linden) Fritsch 

Kohleriu ombilk (Planchon & Linden) Fritsch, 1913a:413. 

Herbs, often epilithic, scaly rhizomes rarely observed, 

stolons abundant; stems decumbent to erect, branching not 

observed, pilose to villous near apex, 15-30 cm long, basal 

diameter 1.5-3.0 mm. Leaves opposite, internodes 0.5-8.0 

cm; the blades ovate to rotundate, 4-9 x 3-7 cm, above 

pubescent and sparsely pilose, below pubescent and sparsely 

pilose, green or purplish with a pattern of light green blotches, 

veins below pilose to villous; petioles 0.5-4.0 cm. Florescences short, frondose, with 1 to 6 (-numerous) flowers, partial 

florescences with 1 (-5) flower(s); pedicels 4-8 cm, pilose to 

villous, yellow to purple, bracts absent. Floral tube villous; 

calyx 5-7 mm long, lobes reflexed, ovate, 3-5 mm long and 

3-7 rnm wide at base, outside pilose, inside appressed 

pubescent; corolla 24-32 mm long, broadly tubular, sometimes 

subventricose, tube 17-25 mm long, diameter 3-4 mm at base 

and 6-12 mm at throat, outside pilose, red, ventrally and 

basally often yellow, inside glabrous, yellow, limb 13-27 mm 

wide, lobes 3-10 mm long and 3-9 mm wide at base, inside 

glabrous, yellow to pink with red dots; filaments 14-20 mm 

long, glabrous, anthers -1.8 x 1.3 mm, included; nectary of 3 

free and 2 dorsal basally united glands, each -1.2 mm high and 

1 mm wide at base; ovary half-inferior, 4-5 mm long, diameter 

2-3 mm, style 15-18 mm long, sparsely pubescent, stigma 

subincluded. Capsule ovoid, 6-8 mm long, diameter 4-6 mm, 

apically dehiscing into 2 valves (by 2 slits) and 1 of these slits 

appears to continue to the base; seeds sub-globose, longitudinally to transversely striate, -0.4 x 0.3 mm. 

DISTRIBUTION.-Kohleria amabilis is a rare endemic of the 

Rio Magdalena valley in central Colombia (Figure 20), where 

it occurs below 1100 m altitude (Figure 21). Apparently the 

species has become rarer, since nearly all collections are at least 

50 years old. However, in 1987 populations of the variety 

bogotensis were discovered on the banks of Rio Dulce between 

Villeta and Sasaima in the Dept. Cundinamarca (Londono & 

Kvist 78). Kohleria amabilis may disappear due to the 

advanced deforestation of the Rio Magdalena valley. An old 

locality near Tocaima (Killip et al. 38338) was visited, but here 

the species has disappeared. 

DISTINGUISHING FEATuREs.-Specimens of Kohleria amabilis from Colombia and K. rugata from Mexico are difficult to 

distinguish, e.g., the flowers are similar (Figure 12a-d). The 

former has an often conspicuous pattern of purple and light 

green blotches on the lower leaf surfaces, in contrast to the 

uniformly colored lower leaf surface of the latter. However, K. 

amabilis occurs in relatively humid and shaded habitats, 

whereas K. rugata is a hardy exposed-habitat species. Kohleria 

rugata has bivalved capsules, while those of K. amabilis 

apparently split to the base similar to Diastema racemiferum 

shown in Figure 13b. 

Another similar species is K. villosa (Figure 13b), which also 

has a lower leaf surface pattern of purple and light green 

(Figure 36a) and the same capsule dehiscence as K. amabile but 

differs in having smaller corollas that are villous inside, villous 

filaments (Figures 36e, 37d), a uniformly red limb and long 

narrow nectary glands (Figures 36f, 37e). Kohleria amabilis 
NUMBER 79 45 

and K. villosa both have an abundance of stolons not seen in 

other species (Figures 8a, 36a, 37a). 

HABITAT.-Kohleria amabilis occurs on calcare~us rocks, 

steep slopes along streams, etc. The populations along Rio 

Dulce 50w down to the highest water level on steep, fragile, 

slippery, and mostly shaded cliffs (Londono & Kvist 78). 

The label of the type collection of Achimenes picta (now K. 

amabilis var. bogotensis) relates that the plants occur in 

shade. 

PHEN0LOGY.-The few collections have flowers during the 

first half of the year. However, very few flowers but many 

immature capsules were present along Rio Dulce late in 

January 1987 (Londono & Kvist 78). 

NOTES.-NO open capsules were found in the field (Londono

& Kvist 78), but herbarium sheets show that the capsules split 

to the base (Karsten s.n., Pennell3552). Wiehler (1983) stated 

that Kohleria amabilis capsules split by a longitudinal slit, but 

his observation may have been based on a hybrid. 

Scaly rhizomes of Kohleria amabilis have not been observed 

on herbarium specimens or in the field. Their presence may 

have been overlooked in the populations along Rio Dulce 

because of the difficult collecting conditions (Kvist, 1987). 

However, the label of one collection relates that the plants 

possessed “peculiar scaly rhizomes” (Haught 2422), and Andre 

(1876) in describing Tydaea ceciliae also mentions the 

presence of scaly rhizomes. 

2a. Kohleria amabilis var. amabilis 

Figures la (stomata), 12b (flower); Planchon and Linden, 1856a. pl. 1070; 

Lescuyer, 1861, pl. 20; Andre, 1876, pl. 260. 

Kohleria amabilis (Planchon & Linden) Fritsch, 1913a:413; Tydaea amabilis 

Planchon & Linden, 1856a:237, 1856b:4; Achimenes amabilis (Planchon & 

Linden) Van Hwtte. 1856:253; Isoloma bogofenre Nicholson var. amobile 

(Planchon & Linden) Voss. 1894793; Isoloma amabile (Planchon & 

Linden) Hortorum ex Bellair & Saint-eger, 1899:951 [not seen]. [Type: 

Illustration in Planchon and Linden, 1856a, pl. 1070. drawn from cultivated 

material, originally collected by Triana in Colombia.] 

Ty&ea ceciliae And& 1876: 178; Isoloma ceciliae (Andre) Nicholson, 

1885:201; Isolomo ceciliae (Andre Hortorum ex Bellair & Saint-Uger, 

1899:952 [not seen]. [Type: Andre 1838, Colombia, Cundinamarca, close to 

Tocaima, Alto de Limb (K, holotype; NY, isotype).] 

Tydaea venosa Lescuyer, 1861:248. [Type: Illustration in Lescuyer, 1861, pl. 

20, drawn from cultivated material.] 

Stems and inflorescences villous. Lower leaf surface green to 

purplish with a conspicuous pattern of light green blotches. 

Limb 20-27 mm wide, inside pink with red or purple dots, 

corolla lobes unequal, ventral lobe up to 10 mm long. 

DISTFuBUTION.-The variety occurs only in Colombia both 

east and west of the Rio Magdalena in the Depts. Cundinamarca and Tolima, respectively (Figure 20). In addition, 

garden-collections from the Dept. Antioquia in Colombia 

(Sandeman 556I), from Cuba (Morton 10753) and from Puerto 

Rico (Britton & Britton s.n.) show that Kohleria amabilis var. 

amabilis is in cultivation in Latin America outside its native 

range, There are no indications that cultivated plants have 

naturalized. 

tween Tocaima and Pubenza, Quebrada Cabana, Hacienda el 

Cucharo, Killip et al. 38338 (US); Alto de Limba, Triana 2525 

(P); Triana 4164 -I (BM). TOLIMA: Guataquicito, Haught 

2422 (COL, US); Honda, Pennell3552 (GH, NY, US). Total of 

6 collections seen. 

NOMENCLATURE AND TYPIFICATION.-~dUeU amabilis: 

Planchon and Linden (1856a) wrote that Triana had introduced 

the living material, on which they based their description of 

Tydaea amabilis from the Colombian department of Cauca. 

However, no Triana collections have been seen from Cauca, but 

there do exist two from the Dept. Cundinamarca (Triana 2525, 

4164-1). Consequently, Triana may have collected the live 

material from which Tydaea amabilis was described in the 

Magdalena valley within the current range of the species. 

Unfortunately, the possibility also exists that Tydaea amubilis 

was described from a hybrid collected by Triana in the Dept. 

Cauca; a later collection from the Dept. Cauca is probably a 

hybrid (Garcia-Barriga 6492). The type drawing of Tydaea 

amabilis provides inconclusive evidence, but we accept it as the 

type to stabilize the name. 

Tydaea venosa: The species was possibly described from 

hybrid material, but there only exists an inconclusive illustration for evidence. 

OTHER COLLECTIONS.~OLOMBIA: CUNDINAMARCA: Be2b. Kohleria amabilis var. bogotensis (Nicholson) 

Kvist & L. Skog, new status 

Figures 5b (seed), 7c,d (pollen), 8a (stolons), 12a (flower); Bentham ex W.J. 

Hooker, 1844, pl. 4126; Linden, 1845, pl. 42; Hanstein, 1854. pl. 1: fig. 13; 

Bailey, 1900, pl. 1184; 1915, pl. 1997. 

Isoloma bogofense Nicholson, 1885:201; Kohleriu bogofensis (Nicholson) 

Fritsch, 1894:178 [based on Achimenes pictu Bentham ex W.J. Hooker, 

1844: pl. 4126, not Kohleria picfa (W.J. Hooker) Hanstein, 1865:442]; 

Tydaea picta (Bentham ex W.J. Hooker) Decaisne, 1848b3468; Diasfemo 

pictlun (Bentham ex W.J. Hooker) Walpers, 1849:472; Giesleria picta 

(Bentham ex W.J. Hooker) Regel, 1849:182; Isoloma tydaea L.H. Bailey, 

1900.836. [Type: Colombia, Cundinamarca, close to Guaduas, Hacienda de 

Palmar, Hurhveg 1263 (K. holotype; BM, CGE(2), E, G, K, NY, P, W(2), 

isotypes).] 

Stems and inflorescences pilose. Lower leaf surface green, 

not with a conspicuous pattern of light green blotches. Limb 

13-18 mm wide, corolla lobes subequal, inside yellow with 

purple lines, dorsal 2 lobes red, ventral lobe up to 6 mm long. 

DISTRIBUTION.-The variety is endemic to Colombia in a 

small area in the Dept. Cundinamarca east of the Rio 

Magdalena (Figure 20). Pure-breeding var. bogotensis is 

apparently not cultivated in Latin America as is var. amabilis. 

Close to Villeta, Goudot s.n. 0; Guaduas, Karsten sen. (LE, 

W); between Villeta and Sasaima, steep banks of Rio Dulce, 

Londono & Kvist 78 (AAU, COL, QCA, TULV, US). 

LOCALITY UNKNOWN: Purdie s.n. (GH, K(3)). Total of 5 

OTHER COLLECTIONS-COLOMBIA: CUNDINAMARCA: 
46 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

FIGURE 30.-Kohleria diasfemoides, new species: a, habit and inflorescence: b, indument of upper leaf surface; 

c, flower, d, laterally opened corolla showing androecium; e, calyx, nectary, and gynoecium; f, capsule; g, seeds. 

All from Lehmann 8892 (GH, NY). 

collections seen. 

NOMENCLATURE AND TYPIFICATION.-Achimenes picta: 

The two isotypes cited from CGE may be later clonal cultivated 

material of the type collection. 

NOTES.-The most popular Kohleria in cultivation is known 

as "K. bogotensis." This is apparently a hybrid between K. 

amubilis var. bogotensis and K. hirsuta. It may have originated 

as a natural hybrid brought into cultivation first in Latin 
NUMBER 79 47 

America and later introduced from there to Europe, or it may 

have been produced in Europe. This hybrid is cultivated in 

Latin America, and occasionally naturalizes from cultivation, 

especially in Jamaica (e.g., Lun&n & Nilssen 26; Stearn 265; 

Sucker 17418) and also in Brazil (Ball s.n., Moroes 2232). 

However, no collections are known from the small Colombian 

range of true K. amabilis. The features of the hybrid are 

variable, and the delimitation toward both parent-species are 

somewhat arbitrary. The hybrid produces scaly rhizomes, and 

its haploid chromosome number has been reported to be 13 

(Eberle, 1956; Fussell, 1958). Kohleria amabilis may also 

hybridize with K. warszewiczii (see K. warszewiczii). 

3. Kohletia diastemoides Kvist & L. Skog, new species 

Figures Sc (seed), 30 (habit and floral features). 

Differt a Kohleria inaequalis (Bentham) Wiehler et K. 

villosa (Fritsch) Wiehler corollis parvis anguste infundibuliformibus. 

Herbs, scaly rhizomes, or stolons not seen; stems erect, 

branching not observed, villous, 5-10 cm long, basal diameter 

1-3 mm. Leaves opposite, internodes 0.5-2.5 cm long; the 

blades ovate, 3-7 x 1.5-4.0 cm, above appressed pilose, below 

purplish, pilose, veins below villous: petioles 1.5-3.5 cm long. 

Florescences frondose, partial florescences with 1 or 2 flowers; 

peduncles 1.5-2.5 cm long, pedicels 1.5-2.5 cm long, villous, 

bracts absent. Floral tube villous; calyx 4-5 mm long, lobes 

erect to somewhat reflexed, ovate to lanceolate, 3-4 mm long 

and 1.5-2.0 mm wide at base, pilose; corolla 18-20 mm long, 

narrowly tubular, subventricose, tube 16-18 mm long, diameter 1.5-2.5 mm at base and 3-4 mm at throat, outside villous, 

deep purple (?), inside basally villous, glabrous toward the 4-6 

mm wide limb, lobes 1.5-2.0 mm long and 1.5-2.0 mm wide 

at base, inside glabrous, deep purple (?); filaments 12-14 mm 

long, basally villous, apically glabrous, anthers -1.2 x 1.0 mm, 

included; nectary of 5 separate glands, each -1 mm high and 

0.3 mm wide at base; ovary inferior, 2-3 mm long, diameter 

2.0-2.5 mm long, sericeous, style 10-12 mm long, pilose, 

stigma included. Capsule oblong, 10-14 mm long, diameter 

2.0-3.5 mm, probably dehiscing from apex to base by a single 

slit; seeds subglobose, irregularly striate, 4.3 x 0.3 mm. 

TYPE.-COLOMBIA: CAUCA: Western slopes of the WestAndes near Popaah, 900-1300 m, Dec 1899, Lehmann 8892, 

(NY, holotype; GH, K, isotypes). 

DISTRIBUTION.-Only known from the type collection from 

Dept. Cauca, Colombia Figure 20). 

HABITAT.-PlantS only found on slopes in dense, humid 

forest. 

PHENOL0GY.-The type collections show flowers in December. 

DISTINGUISHING FEATURES.-The oblong capsules, the 

pedunculate partial florescences, and the basally villous 

filaments and corollas are shared with Kohleria inaequalis, and 

the long, narrow, nectary glands, the velutinous leaves, and the 

habit and size of the shoots are shared with K. villosa. The 

small narrowly tubular corollas set K. diastemoides apart from 

both K. inaequalis and K. villosa. Kohleria diastemoides is 

sympatric only with the former species (Figures 20,21). 

NoTES.-The possibility that Kohleria diastemoides may be 

an intergeneric hybrid between K. inaequalis and a species of 

Diastema was discussed in the section above (see “Intergeneric 

Hybridization” and Table 4). 

No dehiscing capsules have been seen, but the shape of the 

capsules (Figure 3Of) is similar to those of K. inaequalis and K. 

longicalyx (Figure 13d,e), which dehisce by a slit from the apex 

to base. Scaly rhizomes may be absent (as in K. inaequalis and 

K. villosa), while stolons very likely are present. 

4. Kohleria grandjflora Kvist & L. Skog, new species 

Figures Sd (seed), 31 (habit and floral features). 

Differt a Kohleria longicalyx Kvist & L. Skog lobis calycum 

rotundis ad spathulatis, inflorescentiis partialibus l-floribus, a 

K. inaequalis (Bentham) Wiehler corollis glabris intus, 

filamentibus glabris basalibus, foliis paginis superioribus 

velutinis. 

Herbs to subshrubs, scaly rhizomes not observed, stolons 

presenc stems erect, rarely branching, villous with yellowbrown 3-5 mm long trichomes and smaller glandular 

trichomes, 50-150 cm long, basal diameter 5-8 mm. Leaves 

opposite, internodes 3-15 cm long; the blades ovate, 6-18 x 

4-12 cm, above puberulent and sparsely pubescent, below 

pilose, light green to yellow-green, veins below pilose to 

villous, yellow; petioles 2-8 (-15) cm long. Florescences 

frondose, flowers single in leaf axil, often pedunculate but 

appearing epedunculate due to reduced bracts and buds; 

peduncles 0-3 cm long, pedicels 4-8 cm long. Floral tube 

villous; calyx 8-14 mm long, lobes erect, free nearly to base, 

rotundate to spathulate, basally 3-7 mm wide, ampliate above 

to 4-15 mm, puberulent and pilose to villous, with 5 prominent 

veins; corolla 45-60 mm long, tubular, tube 38-46 mm long, 

diameter 5-6 mm at base and 10-16 mm at throat, outside 

sericeous, red, inside glabrous, limb 23-35 mm wide, lobes 

6-11 mm long and 8-13 mm wide at base, inside glabrous, red 

with purple-black dotted lines with white edges; filaments 

30-35 mm long, apical half with a few trichomes, anthers -2.5 

x 2.0 mm, included; nectary of 5 free glands, each -2.5 mm 

high and 1.5 mm wide at base; ovary inferior, 6-8 mm long, 

diameter 4-5 mm, villous, style 35-42 mm long, villous, 

stigma subincluded. Capsules ovoid to oblong, 16-20 mm 

long, diameter 8-12 mm, probably dehiscing apically into 2 

valves (2 slits) and one of these slits later continues to the base; 

seeds ellipsoid, irregularly striate, 4.35 x 0.15 mm. 

TYPE.-COLOMBIA: PUTUMAYO: Mocoa-Pasto road, 1320 

m, 6 Mar 1987, Londono & Quinrero 215 (COL, holotype; 

AAU (2). TULV, US, isotypes). 

DISTRIBUTION.-AII collections are from Colombia and near 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

RGURE 31.-Kohleria grandiflora, new species: a, habit and flowers; b, indurnent of upper leaf surface; c, 

indurnent of lower leaf surface; d. flower, e, laterally opened corolla showing parts of androecium; f, calyx and 

gynoeciurn. All from Fosberg 20369 (US). 
NUMBER 79 49 

the roads toward Mocoa on the eastern slopes of the Eastern 

Cordillera in Dept. Putumayo (Figure 20) at altitudes 450 to 

2200 meters (Figure 21). 

COLLECTIONS.-COLOMBIA: PUTUMAYO: Between Achipayaco and Mocoa, Cuutrecasas 11273 (COL, F(2), US); Valle 

de Sibundoy, Cuutrecasas 11683 (F(2), US); 17 km W of 

Mocoa, below Sachamates, Fosberg 20369 (US); by Putumayo, Lehmann 522a (BM, LE); Mocoa-Pasto road, 3 km from 

Rio Pepinito, Londono & Quintero 216A (AAU, COL); road to 

Mocoa, between El Pepino and Mirador, Idrobo & OspinaHernandez 2404 (COL); N of Mocoa, Schultes & Smith 2013 

(COL, GH, US); Mocoa, Vogel 316 (US). LOCALITY UNKNOWN. Lehmann 4018 (BM, LE). Total of 10 collections seen. 

DISTINGUISHING FEATURES.-Kohferia grandiflora differs 

from K. longicalyx by having rotundate to spathulate calyx 

lobes instead of lanceolate lobes (Figures 12f, 31d), partial 

florescences with one rather than two to three flowers, and 

ellipsoid rather than subglobose seeds (and possibly a different 

fruit dehiscence). Kohleria grandiflora and K. longicalyx both 

differ from K. inaequalis by having corolla tubes that are 

glabrous inside instead of basally villous, filaments that are 

basally glabrous instead of villous to pilose, considerably 

longer corolla tubes, and by having puberulous or velutinous 

instead of usually hispid upper leaf surfaces. 

HABITAT.-Kohleria grandiflora occurs in shaded, humid 

forest understory. 

PHEN0LOGY.-The species has been collected with flowers 

from January to April and in July. 

NOTES-The capsules of Kohleria grandiflora apparently 

are considerably shorter than those of K. longicalyx (or the 

capsules that have been seen are not mature). They may split 

into two valves apically with one of these slits continuing to the 

base (Figure 13b), rather than only having a single longitudinal 

slit as in K. longicalyx (Figure 13d). 

Scaly rhizomes are absent according to X. Londono (pers. 

comm .). 

5. Kohleria hirsuta (Kunth) Regel 

Kohleria hirsuta (Kunth) Regel, 1848:250. 

Herbs or shrubs, frequently epilithic, scaly rhizomes and 

stolons usually present: stems erect, occasionally decumbent, 

pendent or scandent, branching variable, villous, less commonly tomentose or lanate, rarely glandular-hairy, erect part 

30-150 (-200) cm long, scandent shoots occasionally reaching 

400 cm, basal diameter 3-8 mm; internodes (1-) 2-7 (-14) 

cm long. Leaves opposite, occasionally ternate; the blades 

elliptic, less commonly ovate to elliptic, rarely appearing 

cordate due to an oblique base, 4-12 (-18) x 2-6 (-10) cm, 

above pubescent to sericeous, occasionally purplish especially 

along the margin, below villous to tomentose, lighter green to 

red or purplish, veins below sericeous or villous, yellow-green 

to purplish; petioles 1-4 (-8) cm long. Florescences frondose, 

partial florescences of 1 to 4 (-6) flowers, epedunculate or less 

commonly pedunculate; peduncles 0-2 (-4) cm long, always 

shorter than the 2-6 (-10) cm long pedicels, villous, 

yellow-green to purple, bracts' usually scaly, occasionally 

lanceolate and up to 2 cm long, often caducous. Floral tube 

villous; calyx (4-) 7-12 (-20) mm long, lobes erect, 

lanceolate to subulate, rarely triangular, (3-) 6-10 (-15) mm 

long and 1-6 mm wide at base, outside villous or sericeous, 

green or purplish, inside appressed pubescent to sericeous; 

corolla (20-) 27-40 (-50) mm long, tubular, occasionally 

subventricose, tube (18-) 25-38 (-45) mm long, diameter 2-6 

(-10) mm at base and 3-15 mm at throat, outside villous, red, 

less commonly yellow-red or orange-red, inside glabrous, 

yellow, with purple dots, the base usually saccate but not 

sharply delimited, limb 6-22 mm wide, lobes 3-6 mm long 

and 4-7 mm wide at base, inside glabrous, rarely sparsely 

glandular-hairy, yellow with numerous purple dots; filaments 

18-35 mm, glabrous, anthers 2.5-4.0 x 1.5-2.5 mm, 

subincluded; nectary often of 5 free glands, each -1 mm tall 

and 0.8 mm wide at base, less commonly a 5-lobed 1-2 mm tall 

ring or irregularly lobed; ovary half-inferior, 5-7 mm long, 

diameter 3-4 mm, style (22-) 27-33 (-37) mm long, sparsely 

pilose and glandular-hairy, stigma subincluded. Capsule ovoid, 

10-14 mm long, diameter 5-9 mm, bivalved; seeds narrowly 

ellipsoid to ellipsoid, longitudinally striate, -0.50 x 0.15 mm. 

Chromosome number n = 13 (Davidse, 1971, 1981; Wiehler, 

1972). 

DISTRIBUTION.-Plants of Kohleria hirsuta occur in Venezuela, Guyana, Surinam, Trinidad, and in Colombia (Figure 17) 

in the Eastern Cordillera and Rio Magdalena valley (Figure 22). 

Collections from Venezuela and Guyana close to the Brazilian 

territory of Roraima make it likely that K. hirsuta also occurs in 

Brazil. The lowland along the Rio Orinoco apparently isolates 

the Guayana shield populations (the collections from southeastern Venezuela and western Guyana) from the main range of K. 

hirsuta (Figure 17). A few recent collections from south of its 

Colombian main range may represent plants escaped from 

cultivation; viz., from the Dept. Putumayo in southern 

Colombia (Londono 203, 212), and from the Prov. MoronaSantiago in southeastern Ecuador (Holm-Nielsen 20472). 

Collections from the state of Chiapas in Mexico (Schayffer s. 

n.), and from the island of Guadeloupe in the Lesser Antilles 

(Questel 3840) must have escaped from cultivation. Gardencollections from Puerto Rico (Britton & Britton 9048, 9819, 

9939) and the Dominican Republic (Jimenez 5483) show the 

species to be elsewhere cultivated in Latin America outside its 

native range. Kohleria hirsuta occurs from sea level to above 

2600 meters altitude (Figure 21). 

DISTINGUISHING FEATURES.-The to the species epithet, 

Kohleria hirsuta never has hispid nor scabrous leaves. A 

constant feature of this variable species is its pilose or hirsute 

leaves and stems that distinguishes K. hirsuta (Figure 2a) 

vegetatively from species with hispid or scabrous leaves 

(Figure 2b). Kohleria trianae and K. warszewiczii are 

vegetatively indistinguishable, but their long-pedunculate 

inflorescences set fertile plants apart. Kohleria peruviana 

(Figure 33) differs by having conspicuously saccate corolla 

bases, larger leaves, and a dense gray-white pubescence. 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

HABITAT.-The species is hardy and is often seen on 

exposed, dry, rocky slopes, The habitats of Kohleria hirsuta in 

Dept. Cundinamarca, Colombia, are described above (see 

“Phenology” section). 

F’HEN0LOGY.-Kohleria hirsuta definitely is seasonal at 

lower elevations in the Colombian department of Cundinamarca (pers. obs., Jan-Feb 1987). However, data from 

herbarium labels do not show this seasonality. The species has 

been collected with flowers throughout the year, possibly 

because there is some reduced flowering throughout the dry 

season. The conspicuous flowers of K. hirsuta and the frequent 

Occurrence on roadsides make it likely to be collected even 

during periods of reduced flowering. 

NOTES.-The extreme variation of Kohleria hirsuta in 

Colombia is discussed above (Figures 4, 11, 24) The species 

has adapted to very different habitats (see “Phenology” 

section), and is hybridizing with other species, especially K. 

trianae (Figure 23; see “Interspecific Hybridization” section). 

In addition, the variety longipes apparently is a previously 

isolated semi-species that has hybridized extensively with the 

typical variety (Table 3; see “Intraspecific Hybridization” 

scetion). 

The scaly rhizomes of Kohleria hirsuta have medicinal uses 

(see “Popular Names and Medicinal Uses” section above). 

5a. Kohleria hirsuta var. hirsula 

Figures 2a (indument of upper leaf surface), 5e (seed), 7e (pollen), 8b (scaly 

rhizome), 24a,c,d (flowers); Kunth, 1818, pls. 189, 191; W.J. Hooker, 1841. pl. 

3815; Regel. 1852c, pl. 7; Hemsley, 1903, pl. 7907; Hoehne, 1970, pls. 164, 

173, 176. 

Kohleria hirsuia (Kunth) Regel, 1848:ZO; Gesneria hirsufa Kunth, 1818: qto. 

ed. 394, fol. ed. 317; Isoloma hirsuiwn (Kunth) Regel, 1851:893; 

Brachyloma hirsutwn (Kunth) Hanstein, 1854:203. [?Lpe: Venezuela, 

Monagas (on border with Sucre), Caripe, Cumana, Bonpland 330 (P-BO, 

holotype; B-W, P, isotypes).] 

Gesneria mollk Kunth, 1818: qto. ed. 395, fol. ed. 317; Isoloma molle 

Decaisne, 1848a:465; Brachyloma mole (Kunth) Hanstein, 1859530, 571; 

Kohleria mollk (Kunth) Hanstein, 1865441; Isoloma molle (Kunth) 

Nicholson, 1885:201. [Type: Probably collected in the Rio Magdalena valley 

of Colmbia, Bonpland s.n. (P, holotype).] 

Gesneria longiflora Kunth, 1818: qto. ed. 396, fol. ed. 317; Isoloma 

longiforwn (Kunth) Regel, 1854c: 108; Brachyloma longiflorwn (Kunth) 

Hanstein, 1859530, 572; Kohleria longiflora (Kunth) Hanstein, 1865:442. 

[Type: Probably collected by Bonpland in the Rio Magdalena valley of 

Colombia but no specimen has been located; Colombia, Tolima, New 

Quindio trail, No Coello, 1000-1500 m, 7 Aug 1922, Hazen 96Sl (NY, 

neotype, selected herein).] 

Gesneria vestifa Bentham, 1845b: 19; lsoloma vestifwn (Bentham) Decaisne, 

1848a:465; Brachyloma vestifwn (Bentham) Hanstein, 1859530, 573; 

Kohleria vestifa (Bentham) Hanstein, 1865:442. [Type: Colombia, Cundinamarca, Tena, Hartweg 1259 (K, holotype; CGE, K, isotypes).] 

Gesneria eriantha Bentham, 1846:228; Isoloma erianfhwn (Bentham) Decaisne, 1848a:465; Brachyloma erianthwn (Bentham) Hanstein, 1859530, 

573; Kohleria eriantha (Bentham) Hanstein, 1865:442. [Type: Colombia, 

Cundinamarca, between Villeta and Guaduas, Harfweg 1256 (K, holotype; 

BM, CGE, G, K, isotypes).] 

Gesneria rubricaulir Kunth & Bouche, 1847:12; Isoloma rubricaule (Kunth & 

Bouche) Regel, 1851 $393; Brachyloma rubricaule (Kunth & Bouche) 

Hanstein, 1859526. 569; Kohleria rubricadis (Kunth & Bouche) Hasskarl, 

1860:97., [Type: Venezuela, Distrito Federal, “Caracas” Moritz 1127 (B, 

holotype not extant; BM, lectotype selected herein).] 

Brachyloma leuomallon Hanstein. 1859528, 570; Kohleria leucomallas 

(Hanstein) Hanstein, 1865:441. [Type: Colombia, Cundinamarca, “Bogotii,” 

Karsten s.n. (W. holotype).] 

Brachyloma moritzianwn Bouche & Hanstein in Hanstein, 1859524, 568; 

Kohleria moritziana (Bouch6 & Hanstein) Hanstein, 1865:441. [Type: 

Described from cultivated material collected by Moritz in Colombia (B, no 

longer extant); Colombia, Santander, close to Bucaramanga, 1500 m, 27 Aug 

1948, Barkley & Araque M. 316 (US, neotype selected herein).] 

Brachyloma rhodomallon Hanstein, 1859:526. 570; Kohleria rhcdomallos 

(Hanstein) Hanstein, 1865:441. [Type: Venezuela, Distrito Federal, “Caracas,”Morifz414 (LE, lectotype selected herein, BM, BR, CGE. G(3), HBG, 

L, W, P, isolectotypes).] 

Brachyloma ventricawn Hanstein, 1859:428, 572; Gesnera [sic] ventricosa 

Hortom ex Hanstein, 1859529 [nom. nud. pro syn.]; Kohleria venfricosa 

(Hanstein) Hanstein, 1865:441. [Type: Described from cultivated material, 

Regel s.n. (LE, neotype selected herein).] 

Kohleria longipedunculata Hanstein, 1865:441; Isoloma longipedunculatum 

Hortom ex Hanstein, 1865:442 [nom. nud. pro syn.]. [Type: Apparently 

described from cultivated material, no specimens located; Venezuela, 

Miranda, Parque Nacional de Guatopo, slopes between Rio San Lorenm and 

ridge leading to Aguaramal, 600-700 m, 25 Nov 1%1. Steyermark 90076 

(US, neotype selected herein).] 

Kohleria brachycalyx Fritsch, 1913a:424. [Type: Colombia, Tolima. Ibague, 

Triana 2519 (W, holotype; G, isotype).] 

Kohleria lanigera Fntsch, 1913a:423. [Type: Colombia, Anticquia, Aguadas, 

Lehmann 4620 (B, holotype not extant; K, lectotype selected herein).] 

Kohleria sfraussiana Fritsch, 1913a:34. [Type: Described from cultivated 

material (B, holotype not extant); Colombia, Cundinamarca, Guaduas to 

Pahnar and road to Guaduero, 1040-1 150 m, 6 Nov 1945, Garcia-Barriga 

11775 (US, neotype selected herein).] 

Isoloma pycnosuzygiwn J.D. Smith, 1916:382; Kohleria pycnosuzygia (J.D. 

Smith) Badillo in Pittier et al., 1947:397. [Type: Venezuela, Distrito Federal, 

Los Chorros, Eggers 13078 (US, holotype).] 

Herbs or subshrubs; stems not scandent, 30-150 cm long. 

Stems, pedicels, peduncles and calyces rarely purple, pedicels 

rarely exceeding 7 cm long. Diameter of floral tube 3-6 mm; 

calyx 4-20 mm long, lobes 3-15 mm long, width at base 1-2 

mm; corolla 22-50 mm long, diameter of throat 3-8 mm, 

diameter of limb 6-12 mm; nectary of 5 free glands. 

NOMENCLATURE AND TYPXFICATION.-Gesneria moilis: 

Kunth (1818) only gave the locality as “Regno Novogranatensi ?” and no additional information appears on the type 

specimen sheet. However, the type collection has features 

indicating introgression with K. trianae, a phenomenon 

restricted to the Rio Magdalena valley (see “Interspecific 

Hybridization” section above). Thus, the species apparently 

was collected during Humboldt and Bonpland’s sojourn 

through the Magdalena valley in 1801. When Decaisne 

published the name Isoloma molle in 1848, he probably 

intended to transfer the Kunth (1818) species, Gesneria molle, 

to Isoloma, but neglected to add the author of the basionym. 

However, the possibility also exists that Isoloma molle is an 

unidentified cultivar. 

Gesneria longflora: Kunth (1818) said that Gesneria 

longijlora came from the same locality as Gesneria mollis and 

that these two species were similar. Consequently, a Rio 
NUMBER 79 51 

Magdalena valley collection showing introgression with Kohleria trianae is selected as neotype (see Gesneria mollis). 

Gesneria eriantha: This species has features intermediate 

between Kohleria hirsuta var. hirsuta and var. longipes, but is 

here referred to the former, as the latter usually occurs above 

1800 meters. Hartweg collected the type of Gesneria eriantha 

at about 1200 meters altitude. The large-flowered, highelevation plants currently referred to the variety longipes, have 

often been called Kohleria eriantha. 

Brachyloma moritzianum: No specimens annotated with this 

name have been located, but Bouche and Hanstein in Hanstein 

(1859) wrote that the species was similar to Brachyloma 

rhodomallon. According to the description there also are 

similarities with Gesneria vestita. A neotype similar to these 

species has been selected. 

Brachyloma rhodomallon: Hanstein (1859) cited two specimens but selected no type (B, LE). The former is no longer 

extant. 

Brachyloma ventricosum: The only specimen seen annotated 

with this name was cultivated by Regel in Saint Petersburg 

(formerly Leningrad). This specimen was selected as neotype, 

because Hanstein received the material on which he described 

Brachyloma wentricosum (1859) from Zurich, where Regel 

worked before he went to Saint Petersburg. Thus, it is likely 

that the name Brachyloma ventricosum and the material from 

which Hanstein described Brachyloma ventricosum came from 

Regel. 

Kohleria longipedunculata: The species epithet apparently 

refers to the Occurrence of long pedicels, because Hanstein’s 

(1865) description demonstrates the partial florescences consist 

of single, axillary flowers. Kohleria longipedunculata was 

described from cultivated material of apparently unknown 

origin. Hanstein suspected K. longipedunculata was identical 

with K. lanata (now K. rugata), but the species was described 

as having lanceolate-lineate calyx lobes, while those of K. 

rugata are always ovate. Hanstein (1865) referred K. longipedunculata to the subgenus Brachyloma together with 13 

other current synonyms of K. hirsuta. In addition, the 

description of K. longipedunculata supports the idea that the 

species is a synonym of K. hirsuta var. hirsuta. 

Kohleriu brachycalyx: Fritsch (1913a) described K. 

brachycalyx from K. hirsuta plants showing introgression with 

K. trianae. Consequently, a Rio Magdalena valley collection 

has been selected as neotype (see Gesneria mollis). 

Kohleria lanigera: The single specimen of this species has 

pedunculate inflorescences and ternate calyx lobes, but is rather 

poor in having no fully developed flowers. It seems to have 

some features of K. warszewiczii (or possibly K. trianae), 

which may be due to introgression with one of these species. 

Kohleria straussiana: Fritsch (1913a) described Kohleria 

straussiana from cultivated material possibly of hybrid origin 

(he reported the pollen quality to be low). He mentioned four 

species he considered to be the closest relatives of K. 

straussiana, all currently synonyms of K. hirsuta. Kohleria 

straussiana apparently was described on plants similar to the 

material on which Brachyloma rhodomallon was based. 

D1sTmunoN.-The variety occurs throughout the range of 

Kohleria hirsuta (Figures 17, 22), and is common between 

1000 and 2000 meters elevation. 

Below San Luis, Rio Clara, Folsom & Renteria 10342 (US). 

BOYACA: Region of Mt. Chapon, El Umbo, Lawrance 469 

(BM, F, GH(2), MO, NY, S). CUNDINAMARCA: Honda-Guadua 

road, Londono & Kwist 72 (AAU, COL, QCA, TULV, US). 

HUILA: La Plata, Lehmann 478 (GH(2), NY). SANTANDER: 

Ocana Schlim 93 (BM, CGE, G(3), K). TOLIMA: Ibague,Andre

1967 (F, K, NY). 

GUYANA: UPPER TAKUTU REGION: Kanuku mountains, 

Goodland & Maycock 456A (NY, US). MAZARUNI REGION: 

Upper Mazaruni river basin, Kamarang, Tillett & Tillett 45809 

(Ny, US). 

SURINAM Wilhelmina Mts., Julianatop, Schulz-LBB 

10315 0. 

TRINIDAD: Between La Vache and Maracas Bay, Dawidse 

2565, (MO, NY). 

VENEZUELA: ARAGUA: Colonia Tovar, Fendler 793 (G(2), 

GH(2), K, MO, NY). BOLIVAR: SE Bolivar, Cerro Venamo, 

Steyermark 92776 (F, G, NY, US). DISTRITO FEDERAL: Old 

Caracas-La Guaira road, Rio Grande, Pittier 9864 (GH, NY, 

US). MERIDA: Merida-Carbonera road, km 35, Breteler 3128 

(G, NY, S, US). MIRANDA: Los Rques, Pittier 11251 (GH, NY, 

US). TACHIRA: San Juan de Colon, Bruijn 1341 (MO, NY, S, 

US). TERR. FED. AMAZONAS: Along the Brazilian border, Sierra 

Parima, Steyermark 106137 (SEL). Total of 211 collections 

seen. 

SELECTED COLLECTIONS.-COLOMBIA: ANTIOQUIA: 

5b. Kohleria hirsuta var. longipes (Bentham) 

Kvist & L. Skog, new status 

Figure 24b (flower); Kanten. 1860, pl. 50 

Gesneria longipes Bentham, 1846229; lsoloma longipes Decaisne, 1848a:465; 

Brachyloma longipes (Bentham) Hanstein, 1859530, 572; Kohleria longipes (Bentham) Hanstein, 1865:442. [Type: Colombia, Cundinamarca, 

Tequendama Falls, Hartweg s.n. (K, holotype; K, isotype).] 

Brachyloma karstenianwn Hanstein, 1859:532, 513; Kohleria karsteniana 

(Hanstein) Hanstein, 1865:442. [Type: Colombia, Cundinamarca, Tequendama Falls, Karsten s.n. (W. lectotype selected herein; F. LE. isoleaotypes).] 

Subshrubs or climbing shrubs; stems often scandent, 

100-250 (-400) cm long. Stems, petioles, peduncles, pedicels 

and calyces usually deep purple, length of pedicels often 

exceeding 7 cm. Diameter of floral tube 5-9 mm; calyx 12-20 

mm long, lobes 6-12 mm long, width at base 2-6 mm; corolla 

32-50 mm long, diameter of throat 8-15 mm, width of limb 

12-22 mm; nectary usually a 5-lobed ring. 

DIsTRIBun0N.-The variety is almost completely restricted 

to the Dept. Cundinamarca in the Eastern Colombian Cordillera 

(Figure 22) and to elevations above 1800 meters (Table 3). 

SELECTED COLLECTIONS.-COLOMBIA: BOYACA: Sierra 
52 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

del Cocuy, Bachira, Grubb 616 (K, US). CUNDINAMARCA: 

Salto de Tequendama, Cuatrecasas 71 (F(3), US); between El 

Salto and El Colegio, Cuatrecasas 8202 (F, US); Mosquera-La 

Mesa road, km 18, Gentry 17047 (MO, US); Salto de 

Tequendama, Killip 33996 (BM, S, US); Boyaca-Siveria road, 

2580 m, Londono & Kvist 80 (AAU, COL); Canasuca, Trianu 

2520 (G, K, US(2)). Total of 44 collections seen. 

NOMENCLATURE AND TYPFICATION.-Gesneria bngipes: 

When Decaisne (1848a) established the genus Zsoloma and 

included the species I. longipes, he apparently forgot to add its 

basionym. If he truly did not intend to transfer the Bentham 

species, then the taxonomic status of I. longipes Decaisne is 

currently unknown. 

Brachyloma kurstenianum: Hanstein (1859) cited two 

collections but selected neither as type. The Karsten collection 

was selected as lectotype because Karsten (1860) based his 

illustration of B. karstenianum on his own material. The 

syntype is Linden 394 at W. 

6. Kohleria hondensis (Kunth) Hanstein 

Figures 2b (indument of upper leaf surface), 8d (scaly rhizomes and habit). 

12g (flower); Kunth, 1818. pl. 190; W.J. Hooker. 1847, pl. 4342; Hanstein, 

1854. pl. 1: fig. 17; Fritsch, 1894, pl. 77: fig. L; Kvist, 1987:19 (photo from 

type locality). 

Kohleria hondensis (Kunth) Hanstein, 1865442; Gesneria hondensk Kunth, 

1818: qto. ed. 395, fol. ed. 317; Isoloma hondense (Kunth) Decaisne, 

1848a:465; Brachyloma hondense (Kunth) Hanstein, 1854:203; Isoloma 

hondense var. chloradhwn Regel, 1854a:2; Cryptoloma hondense (Kunth) 

Hanstein, 1859534, 579. [Kohleria hondensis Hanstein, 1859535 (cf. 

Regel, 1848:250) (nom. nud. pro syn.)]. [Type: Colombia, Tolima, Honda, 

Bonpland 1707 (P-BO, holotype, F, fragment; P, isotype).] 

Gesneria triflora W.J. Hooker, 1847, pl. 4342; Kohleria irifora (W.J. Hooker) 

Regel, 1848:250; Isoloma trijlorwn (W.J. Hooker) Decaisne, 1848a:465; 

Cryptoloma triforwn (W.J. Hooker) Hanstein, 1859534, 581. [Type: 

Illustration by Hooker, 1847, pl. 4342, originally described and drawn from 

cultivated material, collected by Purdie in Colombia.] 

Herbs or subshrubs, usually epilithic, with scaly rhizomes, 

stolons not seen; stems decumbent to erect, occasionally 

pendent, rarely branching, appressed pilose to sericeous near 

apex, erect shoots 5-120 cm long, basal diameter 3-5 mm. 

Leaves opposite, internodes 0.1- 1.5 cm long; the blades 

elliptic, 3-9 (-12) x 2-4 (-6) cm, above hispid, below 

tomentose, grayish white, veins below sericeous, yellow-green; 

petioles 0.5-2.0 cm long. Florescences frondose, partial 

florescences congested with 1 to 4 flowers; pedicels 1.5-3.0 

cm long, sericeous, light green, bracts scaly, caducous. Floral 

tube sericeous; calyx 7-12 mm long, lobes erect, ovate to 

lanceolate, 5-8 mm long and 3-5 mm wide at base, outside 

sericeous, inside appressed pubescent; corolla 16-22 mm long, 

tubular, subventricose, tube 15-20 mm long, diameter 3-4 mm 

at base, above widened to 4-8 mm, diameter 1.5-2.5 mm at 

throat, outside sericeous, orange-red, inside glabrous, yellow, 

basally saccate, yellow, limb 5-7 mm wide, lobes -1 mm long 

and 1.5 mm wide at base (the indument exceeding the length of 

the lobes), inside glabrous, yellow with 2 or 3 dots at each lobe; 

filaments 15-17 mm long, glabrous, anthers -2.5 x 2.0 mm, 

included; nectary of 5 free glands, each -1 mm high and 1 mm 

wide at base; ovary half-inferior, 5-6 mm long, diameter 2-4 

mm, style 11-13 mm long, glandular-hairy, stigma included. 

Capsule ovoid, 7-9 mm long, diameter 5-6 mm, bivalved; 

seeds narrowly ellipsoid to ellipsoid, longitudinally striate, 

-0.40 x 0.15 mm. 

DISTRIBUTION.-Kohleria hondemis is endemic to the Rio 

Magdalena valley in Colombia. The distance from the 

northernmost to the southernmost locality is about 350 km. 

Kohleria hondensis is restricted to elevations below 700 meters 

(Figure 21). The distributions of Kohleria hondensis and the 

related K. tubiflora are allopatric, with the latter occurring east, 

west, and north of K. hondensis (Figure 16). 

coli, Andre 509 (K, NY). CALDAS: La Dorada, Haught 2109 

(BH, COL, NY, UC, US). HUILA: 5 km N of Villavieja, Mason 

13806 (COL, GH, UC, US); Villavieja-Baraya road, km 11, 

Cabrera Lajas, Smith 1161 (COL, UC, US). TOLIMA: Piedras, 

Andre 1949 (F, GH, K, NY, US); between Honda and 

Maraquita, Garcia-Barriga 81 68 (US); Chicoral, Haught 6268 

(COL, US); Honda, Karsten s.n. (LE); Honda, Linden 1162 

(BM, CGE, G(3), K, LE, P); 2 km N of Honda toward La 

Dorado, Londono & Kvist 69 (AAU(3), COL, NY, QCA, 

TULV, US, USM, VEN); Honda, Pennell3578 (GH, MO, NY, 

US); Cuamo to Rio Saldana, Pennell & Rusby 211 (NY); 

Pennell & Ruby 212 (GH, NY, US); Maraquita, Schneider 479 

(COL, S); Piedras to Ibague, Triana 2516 (G); Palmilla to 

Piedras, Triana 41 65 (COL); Mariquita, Rio Guali, Uribe 2628 

(COL). LOCALITY UNKNOWN. Holton s.n. (NY). Total of 19 

collections seen. 

DISTINGUISHING FEATURES.-The very small limb distinguishes Kohleria hondensis (Figure 12g) from all species 

except K. tldiflora (Figure 12h). The latter species has longer 

corolla tubes, and the calyx lobes are triangular instead of 

lanceolate. Kohleria hondensis is also characterized by the very 

hispid upper leaf surface. 

HABITAT.-Kohleria hondensis is hardy, and most collections come from dry, rocky, overgrazed areas or from steep, 

exposed slopes, often along roads. 

PHEN0LOGY.-The species has been found with flowers 

from November to February and in July. 

NOMENCLATURE ANDTYPIFICATION.-Kohleria hondensis: 

Regel (1848) transferred Gesneria trifora to Kohleria and 

added that Gesneria hondensis also belonged here. However, 

Regel never made the new combination, and Hanstein (1859) 

listed Kohleria hondensis as a synonym of Cryptoloma 

hondense. By 1865 Hanstein fused Cryptoloma with Kohleria 

and finally made the valid combination. 

Isoloma hondense: Decaisne (1848a) cited W.J. Hooker, 

1846, pl. 4217, which actually shows a plant of Kohleria 

tubiflora. However, the Hooker description cites the Kunth 

publication of Gesneria hondensis. Thus, Isoloma hondense is 

OTHER COLLECI'IONS.-COLOMBIA: ANTIOQUIA: car& 
NUMBER 79 53 

included here as a synonym by indirect reference to the type. 

NOTES.-Kohleria hondensis shoots show a striking variation in size. In a population near the type locality at Honda the 

height of flowering shoots ranged from 5 to 120 cm. Small 

plants were epilithic, and larger ones were growing in porous 

soil. All plants had an abundance of scaly rhizomes often 

deeply buried in rock crevices, but no stolons were observed 

(Figure 8d; Londoiio & Kvist 69). 

The anthers of K. hondensis shed a considerable pollen 

quantity in the throat as a possible reward to visitors (Londono

& Kvist 69) 

7. Kohleria inaequalis (Bentham) Wiehler 

Kohlerio inaequa1i.s (Bentham) Wiehler, 1978a:62. 

Herbs or subshrubs, scaly rhizomes not observed, stolons 

often present; stems erect or decumbent, rarely branching, 

villous to hispid and puberulent or less commonly only 

puberulent, (15-) 30-60 (-100) cm long, basal diameter 3-9 

mm. Leaves opposite, internodes 2-6 (-10) cm long; the 

blades ovate to elliptic, less commonly lanceolate, (3-) 6-10 

(-20) x (15) 3-6 (-9) cm, above hispid, less commonly 

sparsely pilose, rarely pilose, hispid or puberulous, below 

puberulous or glabrous, veins villous and puberulous, rarely 

sericeous, yellow-green; petioles (0.5) 2-5 cm long. Florescences frondose, partial florescences with 1 to 4 flowers; 

peduncles (0-) 2-5 (-8) cm long, single flowers may appear 

epedunculate due to reduced additional flowers and bracts, 

pedicels 2-5 cm long, often villous to hispid with tawny, 

purple or blackish trichomes, less commonly with shorter 

appressed pubescence, bracts lanceolate, 4-13 x 1-3 mm, 

caducous. Floral tube villous to hispid, trichomes purple-black 

or yellow-green; calyx (3-) 6-1 1 (-25) mm long, lobes erect, 

free nearly to base, lanceolate, rarely ovate, base (1 -) 2-5 mm 

wide, upwards occasionally widened to 5-10 mm, outside 

scabrous and hispid, less commonly puberulous, inside 

appressed pubescent, veins occasionally prominent; corolla 

16-38 (-55) mm long, broadly tubular to narrowly tubular, 

occasionally subventricose, tube (12-) 20-35 (-50) mm long, 

diameter 3-5 mm at base and 5-21 mm at throat, outside 

villous, red to purple, inside basally villous, upwards pilose to 

glabrous, lighter yellow-red, the base dorsally gibbous, the 

limb 9-32 mm wide, lobes (3-) 5-7 (-15) mm long and (3-) 

5-7 (-12) mm wide at base, inside glabrous, red with purplish 

or black dotted lines, these often surrounded by a white edge: 

filaments (12-) 20-25 mm long, basally pilose to villous, 

apically pilose to glabrous, anthers 2-3 x 1.5-2.0 mm, 

included, rarely subincluded; nectary of 5 free glands, the two 

dorsal often located appressed to each other, each -1 mm high 

and 1 mm wide at base: ovary inferior, 4-7 mm long, diameter 

2-4 mm, style 12-20 mm long, pilose, occasionally glandularhairy, stigma subincluded. Capsule oblong, 9-15 (-20) mm 

long, diameter 4-6 (-8) mm, dehiscing by 1 slit from apex to 

base; seeds subglobose, irregularly striate, -0.4 x 0.3 mm. 

Chromosome number n = 13 (Wiehler, 1975, as K. magn6ca). 

DISTRIBUTION.-Kohleria inaequalis occurs on the WeStern 

Andean slopes of northern Ecuador, and in Colombia mainly in 

the Western and Central Cordilleras (Figures 21, 28). The 

species is frequent between 1000 and 2200 meters altitude, but 

occurs occasionally down to sea-level and up to 2600 meters. 

The collections from below 1000 meters come from Pacific 

coastal Ecuador and Colombia, the most humid part of the 

range (Figure 21). 

DISTINGUISHING FEATURES.-Kohleria inaequaks is extremely variable (Figures 4, 11,25,27), but most specimens are 

recognized by the combination of hispid upper leaf surfaces, 

pedunculate inflorescences, oblong capsules, and a brown-red 

villous to hispid indument that especially covers the inflorescences and the floral tube. In addition, more subtle but stable 

features set this species apart. The corolla and the filaments are 

always basally villous to pilose, a feature shared with K. 

diastemoides (Figure 30d), but this latter species has very long, 

narrow nectary glands and small, narrowly tubular corollas 

(Figure 30c,e). 

HABITAT.-Kohleria inaequalis mainly occurs in forest 

understory, and often along ravines with small streams. The 

species sometimes occurs along roads, but here usually under 

the cover of shrubs and protected from direct sun (Londono& 

Kvist 190). 

PHENOL0GY.-The species produces flowers throughout the 

year with little seasonality. 

NOTES.-The extremes in the “Kohleria inaequalis complex’’ are so diverse that it seems justifiable to give the 

variations taxonomic recognition, although the majority of the 

collections have intermediate features. These plants mainly 

come from the central part of its range, and are here mostly 

referred to var. inaequalis (Figures 25d,e, 28). The pure 

breeding varieties occur in the southern and the northern parts 

of the range (Figures 25a,c,f, 28). The extreme variation of K. 

inaequalis (Figures 4, 11, 25, 27) is discussed above (see 

“Intraspecific Variation” section). A hybrid between K. 

inaequalis and K. spicata (Figure 25b) also is discusssed above 

(see “Interspecific Hybridization” section). 

Scaly rhizomes are apparently never formed in native 

habitats but may occasionally develop in cultivation. Hooker 

(1848) wrote that the roots of var. ocellata (as Achimenes 

ocellata) were “tuberous, by which it is easily increased,” and 

Lindley (1856) noted that var. inaequalis (as Achimenes 

magn6ca) had scaly rhizomes. Thus, scaly rhizomes may arise, 

for example, under stress in cultivation. 

7a. Kohleria inaequalis var. inaequalis 

Figures lb (stomata), 5f (seed), 7g,h (pollen), 13e (capsule dehisoence), 

Ed-f (flowers); Regel, 1854b, pl. 79 (as 76); Regel. 1855b. pl. 120; Planchon 

and Linden,l855, pl. 1013. 

Kohleria inaequa1i.s (Bentham) Wiehler, 1978a62; Gesnerio inaequalis 
54 SMITHSONAN CONTRIBUTIONS TO BOTANY 

Bentham, 1846:230; Isoioma inaequaie (Bentham) Bentham ex Jackson, 

1895:1238 (cf. Bentham, 1876:lW). [Type: Colombia, Cauca, slopes of the 

Andes of Popayan toward the Pacific, Hartweg s.n. (K, holotype).] 

7jdaea warscewiczii [sic] Regel, 1854b73 [not Kohleria warszewiczii (Regel) 

Hanstein. 18654421 [Capanea warscewiczii [sic] Warscewicz [sic] ex 

Regel, 1854b73 (nom. nud. pro syn.)]; Isoloma bogotense Nicholson var. 

warscewiczii [sic] (Regel) Voss, 1894:794: Kohieria ferruginea Fritsch, 

1913a:413. [Type: Illustration by Regel, 1854b, pl. 79 [as pl. 761. described 

from cultivated material, originally collected by Warszewicz (?) in 

Colombia.] 

Tydaea oceliafa (W.J. Hooker) Regel var. formosa Regel, 1855b3181. [Type: 

Illustration by Regel, 1855, pl. 120, described from Cultivated material, 

originally collected by Warszewicz at St. Martha, Colombia.] 

Locheria magnifca Planchon & Linden, 1855:117: Achimenes magni3ca 

(Planchon & Linden) Lindley, 1856:116; 7jdaea magnifca (Planchon & 

Linden) Hanstein, 1865:440; Kohleria magn$ca (Planchon & Linden) H.E. 

Moore, 1954:383. [Type: Illustration by Planchon and Linden, 1855, pl. 

1013, described from cultivated material, originally collected by Triana at 

Cauca, “Popayan,” Colombia.] 

Kohleria lehmannii Fritsch, 1913a:419. [Type: Colombia, Valle, WestCordillera near Cali, Lehmann 3762 (B, holotype no longer extant K, 

lectotype selected herein; BM, LE, US, isotypes).] 

Stems usually hirsute to villous, the length of the trichomes 

often exceeding 2 mm. Upper leaf surface hispid, rarely nearly 

velutinous. Partial florescences pedunculate, of 2 to 4 flowers, 

rarely epedunculate. Calyx lobes 6-25 mm long, corolla 

broadly tubular to tubular, tube 20-35 mm long, throat 9-21 

mm wide, not constricted, limb 12-32 mm wide, lobes 5-15 

mm long and 5-12 mm wide at base. 

DISTRIBUTION.-The variety occurs on the western slopes 

of the Andes in northern Ecuador and in Colombia in the 

Central and Western Cordilleras northward to Dept. Risaralda 

(Figure 28). Var. inaequafis is only common between 1000 and 

2000 meters, but ranges from the lowland to 2500 meters (Figure 21). 

Agredo 130 (COL, F, US); Rio Pulace, Lehmann 1236 (F, GH, 

K, NY); Gorgona Island, road to Playa Gorgonilla, Guzano & 

Rangel 5280 (COL). HULA: Popaah-La Plata road, Santa 

Leticia, Garciu-Barriga & Hawkes 12870 (COL(2), US); 

Guadalupe-Florencia road, pass by Resinas, Vogef 205 (US). 

NARINO: Near Ricaurte, Afston 8425 (BM, F, S, US); road from 

Chucunes to Reserva La Planada, km 6, Londono 239 (AAU, 

COL, NY, TULV, US). RISARALDA: La Selva, Sneidern 5263 

(AAU, COL, F, MO, US(2)). VALLE: Valley of Rio Digua, La 

Elsa, Cuatrecusas 15301 (AAU, F(3). US); Cali-Buenaventura 

road, km 51, Londono& Kvist 190 (AAU(3), COL, QCA, NY, 

TLJLV, US, VEN, USM). 

ECUADOR: CARCHI: Maldonado, Balsfev 1939 (AAU, NY, 

QCA, US); Chical W of Maldonado, Kvist et al., 48619 (AAU, 

QCA, QCNE, US). ESMERALDAS: Lita, Jativa & Epling 844 

(AAU, BM, MO, NY, UC(2), US). IMBABURA: El Achiotal, 

Acosta-Sofis 12345 0. Total of 114 collections seen. 

NOMENCLATURE AND TYPIFICATION.-Gesneria inuequafis: 

Bentham (1876) noted that Gesneria inaequafis belonged to 

Isofomu, but he never made a new combination. 

SELECTED COLLECTIONS.--COLOMBIA: CAUCA: Popayan, 

7b. Kohleria inaequalis var. lindenii (Hanstein) 

Kvist & L. Skog, new status 

Figure 25c.d (flowers) 

Calycosfemma lindenii Hanstein, 1859516.557; Kohieria iindenii (Hanstein) 

Hanstein. 1865:441. [Type: Colombia, Tolima. Mariquita, Linden 955 (w, 

holotype; BM, GH, K, LE, P(2), W, isotypes).] 

Synepileana granafensk Baillon, 1888a:725. [Type: Colombia, Tolima, 

Mariquita Linden 955 (p, holotype: BM. GH, K, LE, P, W(2). isotypes).] 

Stems villous, trichomes up to 1.5 mm long. Upper leaf 

surface usually puberulous, rarely hispid. Partial florescences 

of single axillary, epedunculate flowers. Calyx lobes (5-) 7-12 

mm long; corolla narrowly tubular to tubular, often subconstricted, tube (25-) 30-50 mm long, throat 6-12 mm wide, 

limb 10-25 mm wide, lobes 4-8 mm long and 3-8 mm wide 

at base. 

DISTRIBUTION.-The variety has mainly been collected on 

the western slopes of the Western Cordillera in Dept. Choco 

and Antioquia (the most pure-breeding populations), and on the 

eastern slopes of the Central Cordillera in Dept. Tolima and 

Antioquia (Figure 28). The few collections from the Cauca 

valley are difficult to set apart from vat. inaequafis. The 

altitude ranges from 400 to 2100 meters. 

OTHER COLLECI’IONS.-COLOMBIA: ANTIOQUIA: w Of 

Argelia, Core 803 (US); trail from Encamacion to Parque 

Nacional de 10s Orchideas, Gentry & Renterfa 24579 (COL, 

MO, US); Santa Rita de Ituango, White s.n. (K). CHocO: Rio 

Ingar& between La Italia and Coruundo, Forero et af. 2138 

(MO, US); San Jose de Palmar, valley of Rio Torito (tributary 

of Rio Hiibita), Forero et af. 6807 (COL, US); Forero et al. 

7204 (COL); Forero et af. 7556 (COL); Ansermanuevo-San 

Josd de Palmar road, Luzano C. 3226 (COL). RISARALDA: 

Pueblo Rico, Sneidern 5470 0. TOLIMA: Libano,Londono & 

Kvist 68 (AAU, COL, QCA, TULV, US, USM, VEN). VALLE: 

NW of Darien, Rio Bravo, Robinson 84 (K, US). DEPARTMENT 

UNKNOWN: Road toward Quibdo, White s.n. (K). LOCALITY 

UNKNOWN: White 13 (K). Total of 14 collections seen. 

NOTES.-The collections by White are, unfortunately, 

rather poor, but have considerably larger limbs, and a pattern of 

purple and green on the lower leaf surfaces. 

7c. Kohleria inaequalis var. ocellata (WJ. Hooker) 

Kvist & L. Skog, new status 

Figures 7g,h (pollen), 25a (flower); W.J. Hooker, 1848, pl. 4359 (as 4357); 

Linden and Andre 1873, pl. 152; Hoehne, 1970, pl. 179. 

Achimenes oceilata W.J. Hooker, 1848. pl. 4359 [as 43571; Qdaea ocellafa 

(W.J. Hooker) Regel. 1851:894: Qdaea ocellafa (W.J. Hooker) Regel var. 

conjruens Regel, 1854b:74; Isoioma oceflatwn (W.J. Hooker) Bentham ex 

Jackson, 1895:1238 (cf. Bentham, 1876:1002); Kohleria ocellafa (W.J. 

Hooker) Fritsch, 1894: 178; Isoloma bogofense Nicholson var. oceifafum 

(W.J. Hooker) Voss, 1894:793. [Type: Illustration by W.J. Hooker, 1848, pl. 

4359 (as 4357), described and drawn from cultivated material, originally 

collected by F’urdie at El Roble, Dept. Valle, Colombia.] 
NUMBER 79 55 

Tydoea ocellata (W.J. Hooker) Regel var. picta Regel, 1854b:74 [Achimenes 

Wotzschii Wancewicz [sic] ex Regel, 1854b:74 (nom. nud. pro syn.)]. [Type: 

Described from material from cultivation, collected by Wanzewicz in 

Colombia; Colombia, Dep. Quindio, Salento, Rio Boquia, 1600-1900 m, 27 

Jul 1922 Kiffip & Hazen 8836 (US, neotype selected herein; GH, W, 

isonmtypes).] 

Tydoea pardina Linden & Andre 1873:212. [Type: Illustration by Linden & 

Andre 1873, pl. 152, described from cultivated material, originally collected 

by Red in Colombia.] 

Kohferia scabrida Fritsch. 1913a:420. [?Lpe: Colombia, Dept. Valle. El Roble, 

Triana 2531 (W, lectotype selected herein; G, K, MANCH, P, isolectoKohleria kalbreyeri Fritsch, 1916:33. [Type: Colombia, Dept. Antioquia, San 

Jose, Kalbreyer 1506 (B, holotype not longer extant; K, lectotype selected 

herein).] 

types).] 

Stems sericeous to villous, length of trichomes not exceeding 1 mm. The upper leaf surface hispid. Partial florescences 

pedunculate, usually of 4 flowers. Calyx lobes 3-6 mm long; 

corolla broadly tubular, tube 12-15 mm long, throat 5-9 mm 

wide, not constricted, limb 9-12 mm wide, lobes 2-4 mm long 

and 3-5 mm wide at base. 

DISTRIBUTION.-The variety occurs in Colombia in the Rio 

Cauca valley from Armenia in the south to Medellin in the 

north (Figure 28). The collections come from the eastern slopes 

of the valley, except for the southernmost collection from Dept. 

Valle. The altitude ranges from 1600 to 2300 meters. 

SELECTED COLLECTIONS.-COLOMBIA: ANTIOQUIA: Road 

between Medellin and El Retiro, Las Palmas, Barkley et al. 64 

(AAU, US); Rio Medellin, below Alto de Morrogil, Juncosa 

1971 (US). CALDAS: Manizales, road to Neira, Ceron 159 

(FAUC, US), Manizales, Monteleon, Fraume et al. 201 

(FAUC, US). QUINDIO: Boquia, Holton 608 (K, US). RISARALDA: Pereira, Tomus 2447 (US). VALLE: El Roble, Purdie 

s.n. (K(4)); close to El Roble, Triana 2523 (G); Triana 2532 

(NY); Triana 4166 (BM(2)). Total of 25 collections seen. 

NOMENCLATURE AND TYPIFICATION.-Achimenes ocellata: 

Bentham (1876) treated Achimenes ocellata in Isoloma but he 

never made a new combination. Hooker (1848) reported 

incorrectly that Seemann collected this species in Panama. 

Kohleria inuequalis does not occur in Panama, and var. 

ocellata does not occur along the Pacific coast of Colombia, but 

only in the Rio Cauca valley from where Purdie sent seeds to 

Hooker. 

Tydaea ocellata var. picta: The brief description of the small 

corollas by Regel (1854b) demonstrates that the species can 

only belong to Kohleria inaequalis var. ocellata. The neotype 

was selected according to this criterion. 

Kohleria scabrida: Fritsch (1913a) did not select a type, but 

cited both Triana (2531), herein selected to represent the name, 

and Triana (2523), a syntype currently represented at G and W 

(and by a fragment at F). 

8. Kohleria longicalyx Kvist 8i L. Skog, new species 

Figures 12f (flower), 13d (capsule dehiscence); Hoehne, 1970, pl. 171 (as 

Kohferia ferruginea). 

Differt a Kohleria inaequalis (Bentham) Wiehler corollis 

glabris intus, foliorum paginis superioribus velutinis, a K. 

grandiflora Kvist & L. Skog lobis calycum lanceolatis, 

inflorescentiis partialibus 2-3-floribus. 

Herbs or subshrubs, scaly rhizomes not observed, stolons 

present; stems erect, rarely branched, villous, with yellowbrown, 6-10 mm long trichomes, 50-150 cm long, basal 

diameter 5-8 mm. Leaves opposite, internodes 4-8 cm long; 

the blades ovate, 8-20 x 5-12 cm, above appressed pubescent, 

below pilose, light green to purplish, veins below villous, 

yellow; petioles 6-12 cm long. Florescences frondose, partial 

florescences with 2-3 flowers; peduncles 3-8 cm long, 

pedicels 2-6 cm long, villous, bracts lanceolate to subulate, up 

to 20 mm long. Floral tube villous; calyx 24-40 mm long, 

lobes erect, free nearly to the base, lanceolate, basally 2-3 mm 

wide, ampliate above to 6-15 mm, sericeous to villous on both 

sides, with 5 prominent veins; corolla 45-60 mm long, tubular, 

tube 40-50 mm long, diameter 5-6 mm at base and 12-17 mm 

at throat, outside villous, red, ventrally yellow-red, inside 

glabrous, yellow with purple dots, the base saccate, yellow, 

limb 23-35 mm wide, lobes 8-12 mm long and 10-14 mm 

wide at base, inside glabrous, yellow with purple-dotted lines 

and red margin; filaments 30-35 mm long, apically with a few 

trichomes, anthers -2.5 x 2.0 mm, subincluded; nectary of 5 

free glands, each -2.5 mm tall and 1.5 mm wide at base; ovary 

inferior, villous, 6-8 mm long, diameter 4-5 mm, style 40-50 

mm long, pilose and glandular-hairy, stigma subincluded. 

Capsule oblong, 28-32 mm long, diameter 8-12 mm, 

dehiscing by a single slit from apex to base: seeds subglobose, 

irregularly striate, -0.4 x 0.3 mm. 

TYPE.--COLOMBIA: CAQUETA: Florencia-Guadalupe road, 

km 26, 750 m, 2 Feb 1987, Londono & Kvist 153 (COL, 

holotype; AAU, QCA, TULV, US, isotypes). 

DISTRIBUTION.-Plants of Kohleria longicalyx are found on 

the eastern slopes of the Andes in the Colombian Dept. Caqueta

(Figure 20). The few collections come from near the 

Guadalupe-Florencia road, and from altitudes between 500 and 

1200 meters (Figure 21). 

atrecasas 9100 (CM, COL, F, US); Florencia-Guadalupe road, 

km 23, Davidse et al. 5763 (COL, MO, US); FlorenciaGuadalupe road, km 21, Luteyn 4943 (COL). Total of 4 

collections seen. 

DISTINGUISHING FEATURES.-Kohleria Iongicalyx differs 

from K. grandijlora mainly by having lanceolate instead of 

rotundate to spathulate calyx lobes (Figures 12f,13d), and from 

K. inaequalis mainly by having corollas that are glabrous inside 

(see "Discussion" under K. grandijlora and K. inaequalis). 

HABITAT.-Kohleria longicalyx occurs in humid forest 

understory, often in ravines along streams. Occasionally some 

plants were seen in direct sun on roadsides, but only shaded 

plants seem to flower (Londonio & Kvist 153). 

PH ENoLOGY.-The few collections were seen flowering 

during the first four months of the year. 

OTHER COLLECTIONS.--COLOMBIA: CAQUETA: sucre, cu- 
56 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 32.4ohlerk neglecta, new species: a, habit and inflorescences; b, calyx and calyx lobe indument of the 

inner side; c, flower, d, laterally opened corolla showing the androecium; e, calyx and gynoecium; f, capsule. All 

from Uribe 3155 (US). 
NUMBER 79 57 

9. Kohleria neglecta Kvist & L. Skog, new species 

Figures 5a (seed), 7i-k (pollen), 32 (habit and floral features). 

Differt a Kohferia spicata (Kunth) Oersted et K. tubij7ora 

(Cavanilles) Hanstein lobis calycum subulatis multo longioribus. 

Herbs or shrubs, often epilithic, scaly rhizomes absent or 

weakly developed, stolons present; stems erect, often branching, sericeous to villous, 50-150 cm long, basal diameter 5-15 

mm. Leaves 3 at a node, internodes 2-8 cm long; the blades 

lanceolate, 5-15 x 1.0-3.5 cm, above hispid, below tomentose, 

veins below sericeous; petioles 1-3 cm long. Florescences 

frondose, partial florescences of 5-6 (-10) flowers; pedicels 

0.8-3.0 cm, villous to sericeous, yellow-green, bracts subulate, 

5-20 x 1-3 mm. Floral tube sericeous; calyx 9-14 mm long, 

lobes erect, subulate, 8-11 mm long and 1-2 mm wide at base, 

outside sericeous, inside hispid; corolla 14-18 mm long, 

narrowly tubular to tubular, subventricose, tube 13-17 mm 

long, diameter 2.5-3.5 mm at base, above widened to 3-5 mm, 

diameter 3-4 mm at throat, outside sericeous, orange-red, 

inside glabrous, yellow, the base dorsally weakly gibbous, limb 

5-7 mm wide, lobes 2-3 mm long and 2-3 mm wide at base, 

inside glabrous, dorsal lobes reddish, lateral and ventral lobes 

yellow with red dots; filaments 13-16 mm long, glabrous, 

anthers -2.5 x 1.8 mm, included; nectary of 5 free glands, each 

-0.8 mm high and 1.2 mm wide at base, rarely a 5-lobed ring; 

ovary half-inferior, 6-7 mm long, diameter 4-5 mm, style 

9-11 mm long, pilose and glandular-hairy, stigma subincluded. Capsule ovoid, 8-9 mm long, diameter 4-5 mm, 

bivalved; seeds narrowly ellipsoid, longitudinally striate, -0.5 

x 0.15 mm. 

TYPE.--COLOMBIA: CUNDINAMARCA: Sasaima, towards 

Villeta, 900 m, 16 May 1958, Uribe 3155 (COL, holotype; US, 

isotype). 

DISTRIBUTION.-The few collections come from a small area 

in Dept. Cundinamarca west and northwest of Bogota (Figure 

19); the altitude ranges from 900 to 1600 meters (Figure 21). 

saima-Villeta road, km 4, banks of Rio Dulce, Barcfay 3675 

(COL, US); Sasaima-Villetaroad, Santa Cruz bridge across Rio 

Dulce, Dumont & Idrobo 36 (G); Pacho-Rio Negro road, 

Garcia-Barriga 10737 (COL, US); La Palma, road to Pacho, 

Rio Murca, Garcia-Barriga 12421 (COL, US); VilletaGuaduas road, Humbert 27121 (COL); Villeta-Guaduas road, 

close to Alto de Trigo, Londono & Kvist 79 (AAU, COL); 

Villeta, between town and Rio Dulce, Lozano C. & Torres C. 

211 (AAU, COL); Villeta, Alto del Trigo, canyons close to the 

road, Uribe 3706 (F, NY), Uribe 3707 (COL). Total of 10 

collections seen. 

DISTINGUISHING FEATURES.-Kohferia neglecta is vegetatively similar to, although often more sturdy and fruticose than 

K. spicata and K. tubiflora; it differs from both by having 

longer subulate calyx lobes instead of triangular to ovate lobes. 

The corollas are rather similar to those of K. spicata, but differ 

by having a smaller limb and by being subventricose. In 

OTHER COLLECTIONS.-COLOMBIA CUNDINAMARCA: Saaddition, the average pedicel length is considerably longer. The 

partial florescences of K. neglecta produce a higher average 

number of flowers (-6) than any other Kohferia species. 

HABITAT.-Kohleria neglecta has been collected from 

slopes of canyons, river banks, and roadsides. 

PHEN0LOGY.-The species has been collected with flowers 

during much of the year, although no plants were seen 

flowering in late January, 1987 (Londono & Kvist 79). 

NoTEs.-Scaly rhizomes are usually absent, but sometimes 

weakly developed, while the roots are stout and deeply rooted 

(Londono & Kvist 79). 

10. Kohleria peruviana Fritsch 

Figures 6a (seed), 33 (habit and floral features). 

KoNeria peruviam Fritsch, 1913a:421. [Type: Peru, Junin, Prov. Tama. La 

Merced, Weberbauer I869 (B, no longer extant); Peru, Junin. near La 

Merced, E of Quimiri Bridge, 800-1300 m, Jun 1929. Kiffip & Smith 23936 

(US, neotype selected herein; W, isoneotype).] 

KoNeriaperuviana Fritsch var. pall& Fritsch, 1913a:422. [Type: Peru, Junin, 

Prov. Tama, Huacapistana, 1600-1800 m, Weberbauer 2328 (B, no longer 

extant); Peru, Junin, Prov. Tarma, Utcayacu, 1800 m, 22 Feb 1948, 

Woytkowski 35366 (US, neotype selected herein; F, MO. UC, isoneotypes).] 

Herbs to subshrubs, often with scaly rhizomes, stolons not 

observed; stems erect, rarely branching, lanate, (40-) 60-120 

(-200) cm long, basal diameter 5-12 mm; internodes 1-9 cm 

long. Leaves opposite; the blades elliptic to obovate, (5-) 8-18 

(-25) x (25) 4-8 (-12) cm, above pubescent, below 

tomentose, yellow-green to gray or whitish, veins below lanate, 

yellow-green; petioles (05) 1.5-3.0 (-4.5) cm long. Florescences frondose, partial florescences in most cases pedunculate 

with 1-4 (-10) flowers; peduncles (0-) 2-5 cm long, pedicels 

(0.5-) 2.5-4.0 (-7.0) cm long, villous to lanate, yellow-green, 

bracts lanceolate, 2-10 x 1-3 mm. Floral tube lanate; calyx 

10-15 mm long, lobes erect, lanceolate, rarely ovate, 5-12 mm 

long and 1-4 mm wide at base, outside villous to sericeous, 

inside appressed pubescent; corolla 35-52 mm long, tubular, 

tube 28-45 mm long, diameter 4-5 mm at base and 8-17 mm 

at throat, outside sericeous, orange-red or yellow, inside 

glabrous, cream to yellow, the saccate base sharply delimited, 

yellow, limb 15-25 mm wide, dorsal lobes 4-5 mm long and 

5-6 mm wide at base, lateral and ventral lobes 7-8 mm long 

and 6-8 mm wide at base, inside sparsely appressed 

glandular-hairy, cream to yellow, with red dots; filaments 

28-35 mm long, glabrous, anthers -3.5 x 2.0 mm, subincluded; nectary often of 5 free glands, each -2 mm high and 1.5 

mm wide at base, less commonly a 5-lobed ring; ovary 

half-inferior, 5-7 mm long, diameter 2-4 mm, style 25-32 

mm, pilose and glandular-hairy, stigma subincluded. Capsule 

ovoid, 12-18 mm long, diameter 6-12 mm, bivalved; seeds 

ellipsoid, longitudinally striate, -0.5 x 0.2 mm. Chromosome 

number n = 13 (Wiehler, 1972). 

DISTRIBUTION.-Kohferia peruviana is endemic to Peru 

(Figure 17). Most collections come from Dept. San Martin in 

northern Peru and from Dept. Junin in central Peru, while the 

species is absent from the intervening Dept. Huanuco. In 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 33.40hIeria peruvia~: a, habic b, stem base with mots and scaly rhizome; c, flower, d, laterally 

opened corolla showing androecium; e, calyx and gynoecium; f, nectary; g, ovary and nectary; h, capsule; i, seeds. 

AU from Skog et al. 5003 (US). 
NUMBER 79 59 

addition, two apparently isolated collections from unusually 

low altitudes come from Depts. Loreto (Williams 7768) and 

Madre de Dios (Foster 5759). Kohleria peruviana occurs from 

(300-) 800 to 1700 (-2300) meters altitude. 

SELECTED COLLECTIONS.-PERU: AYACUCHO: Between 

Huanta and Rio Apurimac, Carrapa, Killip & Smith 22454 (NY, 

US). m“: Between Tarma and San Ramon, Huacapistana, 

Ferreyra 410 (US, USM); 8 km S of Vim, Rio Tulumayo, 

Chilpes, Gentry et al. 40166 (MO, US); La Merced, Killip & 

Smith 23544 (NY, US): San Ramon, Petersen & Hjerting 

1325A (C): Utcuyacu, Woytkowski 35366 (F, MO, UC, US). 

LORETO: San Roque, Williams 7768 0. MADRE DE DIOS: 

Parque Nacional de Manu, Rio Manu, Cocha Cashu Station, 

Foster 5759 0. PASCO: N of hente Paucartambo, Gentry et 

al. 39848 (MO, US). SAN MART~N: Tarapoto-Yurimaguas road, 

km 12, Rimuchi I: 3882 (F, MO, NY(2), US); road from 

Caserio de Santa Rosa to Mishollo, 4 km from Puerto Pizana, 

Schunke V. 4915 (COL, F(2), G, GH, K, MO, NY, P, US). Total 

of 45 collections seen. 

DISTINGUISHING FEATURES-Kohleria peruviana may easily be confused with K. hirsuta, especially with the variety 

longipes. In contrast to K. peruviana, shoots of K. hirsuta var. 

longipes are usually branched and often scandent. In addition, 

the sharply delimited Saccate base of the corolla, the larger 

leaves, and the whitish tomentose lower leaf surface and stems 

distinguish K. peruviana (Figure 33). 

HABITAT.-Kohleria peruviana occurs on rocky slopes 

along rivers, roads, etc. 

PHEN0LOGY.-The species flowers during the first eight 

months of the year. 

NOMENCLATURE AND TYPIFICATION.-Kohleria peruviana 

var. pallida: Fritsch (1913a) distinguished the variety pallidu 

from the typical variety solely on its paler yellowish instead of 

red corollas. This difference also exists between the neotypes 

selected herein. Both the type and the neotype of var. peruviana 

come from about 1000 meters altitude, while the type and the 

neotype of var. pallida come from about 1800 meters. 

NOTES.-About two-thirds of the collections have pedunculate partial florescences, and the epedunculate plants come 

mainly from Dept. San Martin. Otherwise the Junin and San 

Martin populations are similar. 

Kohleria peruviana is closely related to K. hirsuta, and could 

possibly be reduced to a subspecies of this variable species. The 

possibility that K. peruviana and K. hirsuta var. longipes have 

diverged from a common ancestor was discussed above (see 

“Intraspecific Hybridization” section). 

11. Kohleria ruguta (Scheidweiler) Kvist & L. Skog, 

new combination 

Figures 7l,m (pollen), 12c,d (flower); Lemaire, 1861, pl. 287. 

Gesneria rugafa Scheidweiler, 1847:226 (“Gesnera”). [Type: Described from 

cultivated material, originally collected by Ghiesbreght in Guerrero (?), 

Mexico; Mexico, Guerrero, cutover hillside with limestone outcrops at km 

338 beyond Acahuizotla on highway to Acapulco, lo00 m, 20 Aug 1948, 

Moore & Wood 4685 (US, neotype selected herein; BH, MEXU, GH, 

isoneotypes).] 

Gesneria maculata Mocino & Sess6eex A.P. de Candolle. 1839532 [not 

KoNeria maculata C. Morton, 1938:1180]. [Type: Icon. fl. Mex. ined., pl. 

721 (G); an original duplicate in the Tomer Coll., Acc. No. 6331.242 (Hunt 

Institute for Botanical Documentation).] 

Kohleria lanufa Lemaire, 1861, pl. 287. [Type: Illustration in Lemaire, 1861, 

pl. 287, described from cultivated material, originally collected by 

Ghiesbreght in Guerrero (?), Mexico.] 

Gesneria maculata Sesse & Mocino, 1889:97 [not Kohleria macdala C. 

Morton, 1938:1180, illegitimate name]. [Type: Mexico, Guerrero (?), Royal 

Botanical Eqedition (Sesse & Mocino) 2435 (MA, lectotype).] 

Herbs, often epilithic, with scaly rhizomes and occasionally 

with stolons; stems decumbent, rarely branching, villous to 

basally pilose, (5-) 10-30 (-50) cm long, basal diameter 

2.0-3.5 mm. Leaves opposite, rarely ternate, internodes 1-9 

cm long; the blades ovate, 2-8 (-22) x 1-5 (-8) cm, above 

appressed pilose, less commonly villous or tomentose, below 

pilose to villous or tomentose, veins below pilose to villous, 

less commonly sericeous; petioles 1-3 cm long. Florescences 

frondose with 1-6 (-12) flowers, partial florescences with 

single flowers; pedicels 2-20 cm long, villous, yellow-green, 

bracts absent. Floral tube villous; calyx 4-8 mm long, lobes 

often reflexed, ovate with acute apex, 4-6 mm long and 2-4 

mm wide at base, both sides sericeous: corolla 18-32 mm long, 

broadly tubular, tube 16-28 mm long, diameter 3-4 mm at 

base and 7-15 mm at throat, outside sericeous, orange-red or 

pink, inside glabrous, yellow, mottled with rose dots or 

blotches, limb 11-20 mm wide, lobes 3-8 mm long and 4-8 

mm wide at base, inside sparsely glandular-hairy, yellow with 

red dots; filaments 15-20 mm long, glabrous, anthers -2.2 x 

1.5 mm, included: nectary of 5 free glands, each -1.2 mm long 

and 0.8 mm wide at base, less commonly a 5-lobed ring; ovary 

half-inferior, 4-6 mm long, diameter 2-3 mm, style 12-15 

mm long, pilose and glandular-hairy, stigma included. Capsule 

ovoid, 8-11 mm long, diameter 5-7 mm, bivalved: seeds 

narrowly ellipsoid to ellipsoid, longitudinally to transversely 

striate, -0.6 x 0.3 mm. Chromosome number n = 13 (Wiehler, 

1975, 1976a). 

DISTRIBUTION.-Yohleria rugata is only known from the 

states of Chiapas and Guerrero of southern Mexico, with most 

collections from the former state. The species is absent from the 

intervening state of Oaxaca and thus occurs in two disjunct 

populations (Figure 18). Kohleria rugata ranges from 600 to 

1500 meters altitude. 

SELECTED COLLECTIONS.-MEXICO: CHIAPAS: Highway 

190, El Chorreadoro 8 km E of Chiapas de Cono, Breedlove 

11885 (F, US); 22 km N of Tuxtla Gutierrez, El Sumidora, 

Breedlove 14687 (F, US): 15 km NW of Suchiapa along road to 

Villa Flores, Breedlove 28259 (MEXU (2), MO): Tonala, 

Nelson 2890 (GH, US); N of Arriaga on highway 195, Roe et 

al. 851 (WIS(2)); Chorreadero, Ton 2934 (F, US). GUERRERO: 

2 km from Petaquillas on road to Colotlipa, Moore & Bunting 

8827 (BH, GH, MEXU, US); S of Chilpancingo, Pmon et al. 

17M785 (BH, BM, MEXU, NY, US). Total of 27 collections 

seen. 

DISTINGUISHING FEATURES.-Kohleria rugata is at least 

superficially similar to K. amabilis (Figure 12a-d: see the latter 

species). 
60 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

HABITAT.-KohIeria rugata occurs mainly in calcareous 

areas, possibly the original habitat, but also on rocky roadsides. 

PHEN0LOGY.-The flowering of Kohleria rugata seems to 

be restricted to a shorter period than that of any other Kohleria 

species, viz., August to November (Figure 9). During the dry 

months of the year Kohleria rugata probably survives as 

dormant rhizomes. The species is difficult to cultivate, because 

it demands a longer dormancy period than most Kohleria 

species (Batcheller, 1978, and pers. obs.). 

NOMENCLATURE AM> TYPIFICA’TION.-Gesneria rugata: 

Scheidweiler (1847) described this species on material collected by Ghiesbreght in Mexico in 1844. Later Lemaire (1861) 

described Kohleria lanata from cultivated plants probably 

derived from the Same Ghiesbreght collection. 

Gesneria maculata: A specimen at MA of Kohleria rugata 

and collected by the Sesse and Mocino expedition is herein 

selected as the lectotype because it closely resembles the 

description of the species and the illustrations (Fl. Mex. Ic. 330 

[IconesJorae mexicanae ineditae, pl. 721 in G and Tomer coll. 

acc. no. 6331.242 in Hunt Institute for Botanical Documentation both labeled “Gesneria maculata Sp. N.”]) cited by Sesse 

and Mocino, 1889. 

Kohleria lanata: See Gesneria rugata. 

NOTES.-Kohleria rugara shows a considerable variation in 

the length of the corolla tube and the size of the limb. The 

large-flowered specimens have longer florescences, while the 

small-flowered specimens have only a few flowers in apical 

leaf mils. The specimens from the state of Guerrero are all 

large-flowered, while both forms and intergradations occur in 

the state of Chiapas. For this reason, no varieties are 

established. Apparently both Ghiesbreght and the Sesse and 

Mocino expedition collected large-flowered plants in Guerrero. 

Scheidweiler (1847) described and Lemaire (1861) illustrated the nectary of K. rugata as a five-lobed ring (Figure lOa), 

but the flowers we have examined have a nectary of five free 

glands (Figure lob). 

12. Kohleria spicata (Kunth) Oersted 

Figures 6b (seed), 12e (flower); Kunth, 1818, pl. 188; Lindley, 1842, pl. 40; 

W.J. Hooker, 1845, pl. 4152; W.J. Hooker, 1850. pl. 4504; Regel, 1852a, pl. 1; 

Hanstein, 1854, pl. 1: fig. 18; Regel, 1854e. pl. 103; Oersted, 1858, pl. 4: figs. 

25-29; Fritsch, 1894, pl. 77: fig. K Hoehne, 1970, pl. 181; Skog, 1979, fig. 22. 

Kohleria spicata (Kunth) Oersted, 1858:27; Gesneria spicata Kunth, 181 8: qto. 

ed. 393, fol. ed. 316; lsoloma spicatwn (Kunth) Decaisne, 1848a:465. [Type: 

Colombia, Tolima, close to Ibague, Bonpland 2825 (P, holotype; PO), 

isotypes).] 

Gesneria spicata Kunth var. schiedeana A.P. de Candolle, 1839531; Gesneria 

schiedeana (A.P. de Candolle) W.J. Hooker, 1845. pl. 4152; Kohleria 

schiedeana (A.P. de Candolle) Hanstein, 1859518. 561; Isoloma schiedeanwn (A.P. de Candolle) Hemsley, 1882:479. [Type: Mexico, Veracruz, 

Hacienda de la Laguna, Schiede 185 @resumed to be at B, no longer extant); 

Mexico, Chiapas, near Tunbala, 1300-1800 m, 20 Oct 1895, Nelson 3367 

(US, neotype selected herein; GH. isoneotype).] 

Gesneria longifoh Lindley, 1841 : misc. 92 (“Gesnera”); Isoloma longifoliwn 

(Lindley) Decaisne. 1848a:465; Brachyloma longifoliwn (Lindley) Oersted, 

185830; Kohleria Iongifolia (Lindley) Hanstein, 1859524, 565. [Type: 

Described from cultivated material, originally collected by Hanweg in 

Guatemala; later clonal material of the Hanweg collection (K, neotype 

selected herein).] 

Gesneria petiolarir Bentham, 1845a:13 1; Isoloma petiolare (Bentham) 

Decaisne, 1848a:465; Brachyloma petiolare (Bentham) Oersted, 1858:30; 

Kohleria petiolarir (Bentham) Hanstein, 1865:442; Kohleria longifolia 

(Lindley) Hanstein var. petiolarir (Bentham) C. Morton, 1938:1180. [Type: 

Panama or Costa Rica, “Veragua, island along the coast,” Sinclair s.n.(K, 

holotype).] 

Gesneria ignorata Kunth in Kunth & Bouche, 1848:13 [Gesneria mollir 

Hortorum ex Kunth in Kunth & Bouche, 1848:14 (nm. nud. pro syn.)]; 

Kohleria ignorata (Kunth) Regel, 1851:893; Isoloma ignoratwn (Kunth) 

Hemsley, 1882:478. [Type: Illustration in Regel, 1852a, pl. 1. described from 

cultivated material, originally probably collected by Warszewicz in Central 

America.] 

Gesneria linkiana Kunth & Bouche, 1848:13; Kohleria linkiana (Kunth & 

Bouche) Hanstein, 1854:160, 213; Isoloma linkianum (Kunth & Bouche) 

Hemsley, 1882:478. [’&pe: Described from cultivated material, originally 

collected by Warszewicz in Guatemala; Guatemala, Izabal, 2000 m, 25 Feb 

1908, Kellerman 7358 (US, neotype selected herein).] 

Gesneria brevifora Lindley, 1848: 165 (“Gesnera”); Isoloma breviflorum 

(Lndley) Lindley, 1850:63. [Type: Described from cultivated material, 

originally collected by Seemann in Panama (CGE, holotype).] 

Gesneria seemannii W.J. Hooker, 1850, pl. 4504; Kohleria seemannii (W.J. 

Hooker) Hanstein, 1854:203; lsoloma seemannii (W.J. Hooker) Bentham, 

1876 1002. [Type: Described fran cultivated material, originally collected 

by Seemann in Panama (K, holotype).] 

Kohleria wageneri Regel, 1853b [not seen]; 1854~347; Kohleria spicata 

(Kunth) Oersted var. wageneri (Regel) Fritsch, 1913a: 417. [Type: 

Illustration in Regel, 1854e, pl. 103, described and drawn from cultivated 

material, originally collected by Wagener in Merida, Venezuela.] 

Isoloma kramerianwn Lehmann, 1854:458 [Gesneria mollir Hortorum ex 

Lehmann, 1854:459 (nom. nud. pro syn.)]; Kohferia krameriana (lehmann) 

Hanstein, 1859:524, 566. [Type: Described from cultivated material, 

originally collected by Warszewicz in Central America; Guatemala, Alta 

Verapaz, Cacao, Finca Trece Aguas, 300-400 m, 30 Sep 1905. Go11 25 (US, 

neotype selected herein).] 

Kohleria tetragona Oersted, 1858:27; Isoloma tetragonun (Oersted) Hemsley. 

1882:480. [Type: Costa Ria, Aguascaliente, Oersted 9309 (C, holotype).] 

Kohleria incam Klotzsch & Hanstein in Hanstein, 1859522, 564; Isoloma 

incanwn (Klotzsch & Hanstein) Hemsley, 1882:478. [Type: Costa Rica or 

Panama (“Veragua”), Warszewicz 29 (B, no longer extant); Costa Ria, Prov. 

Cartago, Tumalba, 600 m, 10 Oct 1949. Leon 2871 (US, neotype selected 

herein).] 

Kohleria tomentosa Hanstein, 1859:522, 561. [Type: Colombia (?), Warszewicz (B, no longer extant); Colombia, Antioquia, Medellin, 1800 m, Aug 

1945, Medina s.n. (US, neotype selected herein).] 

Kohleria rupestris Seemann, 1871:611; Isoloma rupestre (Seemann) Hemsley. 

1882479. [Type: Nicaragua, Chontales, Chontales Mountains, Seemann s.n. 

(BM, holotype).] 

Kohleria spicata Kunth var. hirpida Fritsch, 1913a:417. [Type: Ecuador, 

Sodiro 129/22 (B, no longer extant); Ecuador, Chimborazo, between Huigra 

and Naranjapata, 600-1200 m, 17 Jul 1923, Hitchcock20648 (US, neotype 

selected herein; GH, NY. isoneotypes).] 

Kohleria chiapensir T.S. Brandegee, 191466. [Type: Mexico, Chiapas, Finca 

Mexiquito, Pwpw 7049 (UC, holotype; BM, F, GH, MO, NY. US, 

isotypes).] 

Herbs or subshrubs, often epilithic and with scaly rhizomes 

and stolons; stems erect, rarely branched, basally pilose to 

sericeous or villous toward the apex, (20-) 50-100 (-200) cm 

long, basal diameter 4-7 (-10) mm. Leaves temate, less 

commonly opposite or whorled (4 leaves at nodes), internodes 
NUMBER 79 61 

2-8 cm long; the blades elliptic to lanceolate, (3-) 5-10 (-15) 

x 2.0-4.5 (-6.5) cm, above hispid, rarely velutinous or nearly 

glabrous, below tomentose, less commonly pubescent, veins 

below sericeous to villous, yellow-green; petioles 0.5-3.0 cm 

long. Florescences frondo-bracteose, partial florescences with 

1-3 (-6) flowers, epedunculate but occasionally with a 1-3 

mm long peduncle, subtending leaves toward the apex reduced 

to sessile ovate to lanceolate prophylls; pedicels 3-30 mm 

long, sericeous to villous, yellow-green to purple, bracts scaly, 

caducous. Floral tube sericeous to villous; calyx 4-8 mm long, 

lobes erect, 1.5-3.0 mm long and 2-3 mm wide at base, ovate 

to triangular, outside sericeous, inside appressed pubescent; 

corolla 12-20 mm long, tubular, tube 10-18 mm long, 

diameter 2-3 mm at base and 5-7 mm at throat, outside 

sericeous, red, inside glabrous, yellow-red, limb 5-8 mm wide, 

lobes 2-3 mm long and 2-3 mm wide at base, inside glabrous, 

yellow-red, with darker red to purplish dots, partly arranged in 

lines; filaments 10-15 mm long, glabrous, anthers 1.5-2.5 x 

1.2-1.7 mm, included; nectary of 5 free glands, each -1.2 mm 

high and 0.8 mm wide at base; ovary half-inferior, 5-6 mm 

long, diameter 3-4 mm, style 7-12 mm long, pubescent and 

usually glandular-hairy, stigma included. Capsule ovoid, 7- 10 

mm long, diameter 4-6 mm, bivalved; seeds ellipsoid to 

narrowly ellipsoid, longitudinally striate, -0.50 x 0.15 mm. 

Chromosome number n = 13 (Davidse, 1970; Rogers, 1954; 

Wiehler, 1975). 

DISTRIBUTION.-Kohleria spicata ranges from southern 

Mexico to northern Peru (only one collection) and east to 

northwestern Venezuela (Figure 15). The species is common 

throughout its range, but is especially abundant in Guatemala, 

Costa Rica, and western Colombia. In Colombia the species is 

absent from the eastern slopes of the Eastern Cordillera where 

K. stuebeliana occurs (Figure 21), in conerast to Ecuador where 

K. spicata occurs along the eastern foothills of the Andes. 

Kohleria spicata is the most frequently collected species of 

New World Gesneriaceae. 

Kohleria spicata is found from sea-level up to 2500 meters 

altitude, but is most common between 1000 and 2000 meters. 

In Colombia, the species mainly occurs above 1000 meters, and 

lower altitude collections come from the Pacific coast (Figure 

21). In Central America and Ecuador K. spicata often occurs 

down to sea-level. 

An ecotype of Kohleria spicata has adapted to saltinfluenced coastal rocks, occurring on the Pacific coasts of 

Costa Rica and Panama and the Cocos Island in the Pacific 

Ocean about 500 km southwest of Costa Rica (Figure 34). The 

label from one of the Cocos Island collections relates that 

“these plants are extremely rare, and are found on cliff faces on 

the northwestern shores” (Gomez 3296). 

SELECTED COLLECTIONS.-MEXICO: CHIAPAS: Near Tumbala, Nelson 3367 (GH, US). OAXACA: Between Puerto Eligio 

and Comaltepec, Calderon 369 (F, GH, US). VERACRUZ: 

Mirador, Liebman 9300 (AAU, C(2), GH, US). 

BELIZE: TOLEDO: Lower part of Maye mountains, Richardson Creek, Davidse & Brunt 31928 (US). 

GUATEMALA: ALTO VERAPAZ: Coban, von Tuerckheim 

1083 (ECON, F, G(2), GH, LE, MO, NY, US(2)). PETEN: Road 

to Poptun, km 67,Ortizfz 1662 (BM, F, MO, S). QUEZALTENANGO: Finca Helvetia, Skutch 1374 (GH, US). 

EL SALVADOR: SAN SALVADOR: Vicinity of San Salvador, 

Standley 19166 (GH, NY, US). 

HONDURAS: EL PARAfSO: Near Yuscaah, Williams & 

Molina R. 11201 (F, GH, MO, UC). SANTA BARBARA: Rio 

Sisicapa, Molina R. 13456 (BM, F, GH, MO, US). 

NICARAGUA: CHONTALES: Santo Domingo, Narvaez S. 

3374 (BM, GH, MO, Uc, Us). GRANADA: w slope of 

Mumbacho Volah, Stevens & Krukof4342 (BM, MO, US). 

MATAGALPA: 10 km E of Matagalpa, Williams & Molina 42558 

(BM, F, US). 

COSTA RICA: ALAJLJELA: 7 km S of Villa Quesada, Molina 

R. el al. 17481 (F, GH(2), NY, US). CARTAGO: 3 km SE of 

Turrialba, Holm & 11th 67 (BM, G, MO, US). SAN JOSE: 

Vicinity of El General, Skutch 2899 (GH, MO, NY, S, US). 

PANAMA: CHIRIQLI~: 4 km NW of Boquete, Skog 4013 

(AAU, MO, US). coc~8: 3km from El Valle on road to La 

Mesa, Croat 13315 (F, MO, NY). 

COLOMBIA: ANTIOQUIA: Dabeiba, Barkley & Gutierrez V. 

1794 (BM, NY, S, UC(2)). HUILA: Guadalupe-Florencia road, 

km 14, Londono & Kvist 98 (AAU, COL, QCA, TULV, US). 

MAGDALENA: Santa Marta, Manganares, Smith 2604 (F, GH, K, 

NY). TOLIMA: W of Santa Isabel, Londono & Kvist 60 (AAU, 

COL, TULV, US). VALLE: Old Cali-Buenaventura road, km 51, 

Londono & Kvist 179 (AAU, COL, QCA, TULV, US). 

ECUADOR: MORONA-SANTIAGO: Macas, Kvist 60449 

(AAU, QCA, QCNE, US). PASTAZA: Puyo-Tena road, Santa 

Clara, Kvist et al. 60350 (AAU, NY, QCA, QCNE, US). 

PICHINCHA: Santo Domingo-Quito road, km 45, Holm -Nielsen 

et al. 7185 (AAU, F, NY, MO, S, US). TUNGURAHUA: E of 

Banos, Kvist 60311 (AAU, NY, QCA, QCNE, US). 

PERU: CAJAMARCA: Rio Tabaconas valley, Weberbauer 

6263 (F). 

VENEZUELA: MEIDA: Tabay, Gehriger 356 (F, MO, NY, 

US). Total of 762 collections seen. 

DISTINGUISHING FEATURES-Kohleria spicata 1s easily 

distinguished from all other species except for the local K. 

stuebeliana (Figure 35; see discussion under K. stuebeliana for 

differences) by its frondose-bracteose florescences (Figure 3c), 

the small tubular flowers (Figure 12e), the ovate to triangular 

calyx lobes (Figure 12e), and the usually temate and hispid 

leaves. Vegetative plants are nearly impossible to distinguish 

from those of K. tubiflora or the rare and local K. neglecta 

(Figures 19,32). 

HABITAT.-Kohleria spicata occurs in exposed places of 

canyons, river banks, coastal cliffs, and roadsides. The 

disturbances caused by man promote the continued dispersal of 

this rather weedy species; e.g., in the Ecuadorian Amazon 

region, K. spicata colonizes along new roads into deforested 

areas (Kvist et al. 60350). Plants on coastal rocks occur in an 
62 SMITHSONIAN COSTRIBUTIONS TO BOTANY 

A Kohleria spicata 

0 K. tubiflora 

----7: c 

c 

20 

10 

0 

FIGURE 34.-Distributions of Kohleria spicufu and K. tubifloru salt-influenced ecotypes from coastal cliffs (the 

entire distributions of the two species appear in Figures 13 and 14, respectively). Note the Occurrence of K. spicatu 

on Cocos Island. 

especially harsh environment. 

PHEN0LOGY.-Based on data from herbarium sheets Kohleria spicata flowers with almost constant intensity throughout 

the year. However, field observations show that the intensity 

actually is reduced during dry periods, but there were always 

some flowers found. 

NOMENCLATURE AND TYPIFIcA??ON.-Gesneria spicata: 

An unusual nomenclatural problem results from the Oersted 

transfer of Gesneria spicata to Kohleria in 1858. In the section 

above, “Taxonomic History,” we discussed that Regel in 1851 

established a second illegitimate Kohleria with K. ignorata 

(currently a synonym of K. spicata) as type species. Although 

Oersted did not specifically state that he was transferring K. 

spicata to the second Kohleria genus, he made this transfer in 

practice. In the same work, Oersted accepted the genus Isoloma 

and included the type species of the legitimate Kohleria, K. 

hirsuta. However, according to the discussion by Nicolson 

(1975) and the response by Price (1976) on pseudo-isonymy, 

the transfer of K. spicata made by Oersted is available. 

Gesneria spicata var. schiedeana: A.P. de Candolle (1839) 

only gave the Mexican type locality but no collector’s name. 

However, the short description shows the variety to be a 
NUMBER 79 63 

synonym of K. spicata rather than the other Mexican species K. 

rugata. According to Hanstein (1859) there was a Schiede 

collection from this locality (later destroyed in Berlin). No 

duplicates of this collection have been traced. 

Gesneria petiolaris: The typification of the species by 

Bentham (1845a) is problematic due to fact that he gave neither 

collectors’ names nor collection numbers but only type 

localities. In the case of G. petiolaris the description reads 

“island off the coast of Veragua,” which is identical to data 

given on the label of a Sinclair collection assumed to be the 

type. The label of a Barclay collection at MANCH reads 

Atacamas (Veragua), but the specimen is not considered to be 

type material. The species was described from plants belonging 

to the coastal cliffs ecotype. 

Gesneria longifolia: Lindley originally described the species 

in 1841, illustrated the species in 1842, but the type specimen 

came from cultivation in 1843, and is consequently at most a 

neotype of possibly clonal material. 

Gesneria ignorata: This species, like Gesneria linkiana and 

Isoloma kramerianum, was apparently described from live 

material introduced by Warszewicz from Central America, 

while Kohleria incana and K. tomentosa were based on original 

herbarium specimens of the same Warszewicz collections. 

Kunth and Bouche (1848) did not credit Warszewicz as the 

collector of Gesneria ignorata. However, the material on which 

G. ignorata was based had previously been cultivated under the 

name G. mollis. According to Lehmann (1854), the material 

cultivated under this latter name was collected by Warszewicz 

in Central America. Regel (1852a) apparently illustrated 

Kohleria ignorata from clonal material of the Warszewicz 

collections. 

Gesneria linkiana: Kunth & Bouche (1848) credited Warszewicz for having collected the species in Guatemala. 

Hanstein (1854: 160) wrote that Gesneria linkiana belonged to 

Kohleria, but the combination K. linkiana only appears in the 

legend to his plate 1 (p. 213). 

Gesneria brevifora: According to Lindley (1850) both this 

species and G. seemannii were based on material from the same 

Seemann collection “Veragua” (Panama or Costa Rica). 

Gesneria seemannii: See Gesneria breviflora. 

Isoloma kramerianum: See Gesneria ignorata. 

Kohleria rupestris: According to Hemsley (1882) the type of 

this species was located at K, but only a specimen from BM has 

been located. Apparently Hemsley erred, as the first set of 

Seemann’s collections was deposited at BM. 

Kohleria spicata var. hispida: Judged from the description 

no major differences exist between the variety hispida and the 

typical variety; most K. hirsuta specimens have hispid or 

scabrous leaves. 

NOTES.-Kohleria spicata is surprisingly uniform throughout its range, but lowland collections often have less hispid or 

even velutinous upper leaf surfaces. The plants from saltinfluenced rocky cliffs have more succulent leaves with nearly 

glabrous upper surfaces. The coastal ecotype of K. tubifora has 

similar leaf characteristics. 

Hybrids with Kohleria inaequalis are discussed above (see 

“Interspecific Hybridization” section; Figure 25b). Kohleria 

spicata is sympatric with many other Kohleria species (Figure 

21), but the limited variation in K. spicata indicates that hybrids 

with other species are rare. 

The scaly rhizomes of Kohleria spicata have medicinal uses 

(see “Popular Names and Medicinal Uses” section above). 

13. Kohleria stuebeliana Fritsch 

Figures 7n-p (pollen). 35 (habit and floral features). 

Kohleria sfuebeliam Fritsch, 1913a:417. [Type: Colombia, Meta, Los Llanos 

de San Martin, Toquisa, Sfuibel 178f (B, holotype no longer extant); 

Colombia, Meta, Sabanas de San Juan de Arama, left bank of Rio Guejar,

“10s Micos,” 500 m, Dec 1950, Idrobo & Schulies 562 (GH, neotype selected 

herein; COL(2), US, isoneotypes).] 

Herbs, often epilithic, often with scaly rhizomes and stolons; 

stems decumbent to erect, rarely branching, sericeous to 

villous, (3-) 20-50 (-70) cm long, basal diameter 2-4 mm. 

Leaves opposite, less commonly ternate, internodes (1 .O-) 

1.5-5.0 (-10) cm long: the blades elliptic to lanceolate, 5-8 x 

1.0-2.5 cm, above velutinous, below pilose, veins below 

sericeous, green to reddish; petioles 0.3-1.5 (-4.0) cm long. 

Florescences frondo-bracteose, partial florescences with 1-2 

(-4) flowers, subtending leaves toward the apex reduced to 

subsessile lanceolate to ovate prophylls; pedicels 7-12 mm 

long, sericeous to villous, green to purple, bracts scale-like, 

caducous. Floral tube villous; calyx 3-6 mm long, lobes erect, 

ovate, 2.0-3.5 mm long and 1.5-2.0 mm wide at base, outside 

sericeous, inside pubescent; corolla 15-18 mm long, tubular, 

subventricose, tube 12-15 mm long, diameter 2-3 mm at base, 

above widened to 4-8 mm, diameter 4-6 mm at throat, outside 

villous, red, inside glabrous, sparsely glandular-hairy, yellowred, the base dorsally gibbous, limb 6-10 mm wide, lobes 

1.5-2.5 mm long and 2-3 mm wide at base, inside appressed 

pilose and glandular-hairy, yellow-green with numerous 

brown-black dots; filaments 12-15 mm long, glabrous, anthers 

-1.4 x 1.0 mm, included; nectary of 5 free glands, each -1 mm 

high and 0.8 mm wide at base; ovary half-inferior, 3-4 mm 

long, diameter 2-3 mm, style 7-10 mm, pilose and glandularhairy, stigma included. Capsule ovoid, 6-8 mm long, diameter 

4-5 mm, bivalved; seeds narrowly elliptic, longitudinally 

striate, -0.50 x 0.15 mm. 

DISTRIBUTION.-Kohleria stuebeliana is locally common in 

the region around Villavicencio, Dept. Meta, Colombia, and the 

adjacent areas of Depts. Cundinamarca and Boyaca (Figure 

19). In addition, a single collection came from the western 

slopes of the Eastern Cordillera in Dept. Cundinamarca 

(Garcia-Barriga 10706). Kohleria stuebeliana evidently is 

very rare on the western slopes as this region is botanically well 

explored; the species may have gotten there casually by 
64 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

FIGURE 35.-Kohleria stuebelianu: a, habit and frondo-bracteose florescence; b, partial florescence with bud and 

flower. c. ventrally opened corolla showing androecium; d, calyx, nectary. and gynoecium; e, capsule. a,b,d,e, 

from ldrobo & Schultes 562 (US); c, from Fosberg 19468 (US). 
NUMBER 79 65 

wind-dispersal across the barrier of the mountains (the 

collection has not been added to Figure 21). Kohferia 

stuebefiana occurs on the Andean slopes and on the flat 

lowlands of Los Llanos, and has been collected from 300-1500 

(-2500) meters altitude (Figure 21). 

SELECTED COLLECTIONS.-COLOMBIA: BOYACA: Guateque-Santa Maria road, Piedra Campana, Garci-Barriga 17231 

(AAU, COL, NY, US); on border with the Dept. Casanare, 

Pajarito, Uribe 4516 (COL, F, NY). CUNDINAMARCA: Villavicencio-Bogota road, km 20, Davidse & Llanos 5524 (COL, 

NY, US); between Pacho and Rio Negro, Garcia-Barriga 

10766 (COL, US); SW of Quetame, “Sucumuco,” Penneff 

1357 (NY); Penneff 1728 (GH, NY, US). META: Villavicencio, 

Cuatrecasas 3511 (COL, F(2), US); Villavicencio, Haught 

2456 (AAU, BH, COL, NY, US). Total of 25 collections seen. 

DISTINGUISHING FEATURES.-Kohleria stuebefiana is the 

closest relative of K. spicata, but the yellow-green, dark-dotted, 

appressed pilose and glandular-hairy limbs, the delicate, 

soft-velutinous leaves, and the slender shoots distinguish K. 

stuebefiana (Figure 35). Occasionally, tiny K. stuebefiana 

plants flower when only 3-5 cm tall (e.g., Penneff 1357), a 

condition never found in K. spicata. 

HABITAT.-Kohferia stuebefiana grows in exposed places 

along rivers, roads, etc. 

PHENOLOGY-Kohferia stuebefiana flowers relatively constantly throughout the year, but less in the months April-July. 

NOTES.-One specimen label states that the dark glands on 

the corolla lobes are sticky and trap small insects (Uribe 4516). 

Kohferia stuebefiana apparently speciated from K. spicata 

when geographically isolated by the eastern Andean Cordillera. 

Kohferia stuebefiana could possibly be reduced to a subspecies 

of K. spicata, but is distinctive as a species. 

14. Kohleria trianae (Regel) Hanstein 

Figure 12i (flower); Kunth, 1818. pl. 192; Graham, 1840, pl. 3725; Regel, 

1854c, pl. 82; Lanaire, 1855, pl. 57; Planchon. 1855b,pl. 1057; Herincq, 1863, 

Kohleria frianae (Regel) Hanstein, 1865:442; Isoloma trianaei [sic] Regel, 

1854c:lCV; Brachyloma trianae (Regel) Hanstein 1859532, 577. [Type: 

Illustration in Regel, 1854c, pl. 82, described and drawn from cultivated 

material, originally collected by Warszewicz in Colombia at St. Martha.] 

Gesneria elongafa Kunth, 1818: qto. ed. 396, fol. ed. 318 [not Kohleria 

elongafa Regel, 1855a:4]; Gesneria elongala Humboldt in Maund, 1837, no. 

27 [Gesneria veldna Willdenow ex A.P. de Candolle in A.P. de Candolle, 

1839528 (nom. nud. pro syn.)]; Brachyloma elongatrun (Kunth) Hanstein, 

1859532; Kohleria elongafa (Kunth) Hanstein, 1865:442. [Type: “Ecuador” 

(locality probably incorrect as plants only known from Colombia), Bonpland 

s.n. (P. holotype; B-W, P(2), isotypes).] 

Gesneria elongala Kunth var.fruticosa Graham, 1840, pl. 3725 [not Kohleria 

fruficosa T.S. Brandegee, 1914:67; Gesneria oblongafa Honorurn ex 

Graham, 1840. pl. 3725 (nom. nud. pro syn.)]. [Type: Illustration in Graham, 

1840, pl. 3725. originally described from cultivated material.] 

Gesneria trianaei Herincq, 1863:30. [Type: Illustration in Herincq, 1863, pl. 4, 

described from cultivated material, originally collected by Triana in 

Colombia.] 

pl. 4. 

Herbs or shrubs, with stolons, scaly rhizomes occasionally 

present but weakly developed; stems erect, branching, sericeous, 50-150 (-250) cm long, basal diameter 4-10 (-15) 

mm. Leaves opposite; internodes 3-8 cm long; the blades 

ovate to lanceolate, 6-10 x 2-4 cm, above pilose, below 

tomentose, yellow-green, veins sericeous, often purplish; 

petioles 1.5-4.0 cm long. Florescences frondose, partial 

florescences of 4 flowers; peduncles 4-10 cm long, always 

longer than the 4-6 cm long pedicels, sericeous, purplish, 

bracts 2, lanceolate to ovate, 2-5 (-8) x 1-2 mm. Floral tube 

sericeous; calyx 3-5 mm long, lobes erect, triangular, 2.0-3.5 

mm long and 2.0-3.5 mm wide at base, both sides sericeous; 

corolla 26-40 mm long, narrowly tubular, subventricose, tube 

25-37 mm long, diameter 3-4 mm at base, above widened to 

6-9 mm, diameter 3-5 mm at throat, outside villous, red, 

inside glabrous, yellow, limb 6-10 mm wide, lobes 3-4 mm 

long and 2-3 mm wide at base, inside sparsely pilose and 

glandular-hairy, appearing red, but actually yellow with 

numerous red dots; filaments 25-35 mm long, glabrous, 

anthers -2.5 x 2.0 mm, subincluded; nectary of 5 free glands, 

each -1 mm high and 1 mm wide at base; ovary half-inferior, 

3-4 mm long, diameter 1.5-2.5 mm, style 20-33 mm long, 

glandular-hairy, stigma subincluded. Capsule ovoid, 9- 14 mm 

long, diameter 5-8 mm, bivalved; seeds narrowly elliptic, 

longitudinally striate, -0.5 x 0.2 mm. 

DISTRIBUTION.-Kohferia frianae is endemic to the Central 

Cordillera in Colombia (Figures 21, 22). The species is known 

from the eastern slopes in Depts. Quindio, Risaralda, and 

Caldas and from the western slopes in Depts. Tolima and 

Antioquia, but is only common in Quindio and Tolima. In 

addition, a single old K. trianae collection came from 

Tequendama in Dept. Cundinamarca (Hofton 611A (not shown 

in Figures 21, 22)). Numerous K. hirsuta var. fongipes 

collections also come from Tequendama, and K. trianae 

possibly got there by wind-dispersal, and later disappeared due 

to hybridization with K. hirsuta (see “Interspecific Hybridization” section above). 

Kohferia trianae has been collected from 900 to 2500 meters 

altitude (Figure 21), and is locally common between 1700 and 

2100 meters (pers. obs.). 

Felix, Daniel 2327 (US). CALDAS: Manizales, Yepes-Agredo 

827 (COL). CUNDINAMARCA: Cataracts of Tequendama, Hofton 61 1A (K). QUINDIO: Armenia-Cajamarca road, secondary 

road on the right toward Salento, Londono & Kvist 51 (AAU, 

COL, QCA, TULV, US, USM); Salento, Penneff et af. 8750 

(GH, K, NY, S, US). RISARALDA: Road to Herraduro, between 

El Cedral and Otun, &zfamurca 208 (COL). TOLIMA: LibanoMurillo, km 16, Alto de Pones, Garcia-Barriga 12272 

(COL(2), US); Trail Ibague-Salento, La Palmilla, Hawkes 474 

(BM, COL, K, US), Hawkes 479 (COL, K); CajamarcaArmenia road, before summit of divide, KifIip & Varefa 34635 

(BM, COL, GH, MO, US); Santa Isabel, Londono & Kvist 63 

(AAU, COL, NY, QCA, TULV, US, USM, VEN). Total of 35 

collections seen. 

DISTINGUISHING FEATURES.-Kohferia trianae is easily 

recognized by its four-flowered, long-pedunculate florescences 

(Figure 3d) with entirely red, narrowly tubular, sub-ventricose 

flowers (Figure 129. The species is vegetatively similar to, 

SELECTED COLLEC~ONS.-COLOMBIA:NTIOQUIA: San 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

although often more fruticose than, K. hirsura and K. 

warsrewiczii. 

HABITAT-AS the tallest and most fruticose species, 

Kohleria trianae thrives in denser and taller vegetation than 

any other species of Kohleria. In Dept. Quindio, the species 

was observed on steep slopes covered with low montane forest 

(Londono & Kvisr 51). Kohleria rrianae also is common in 

hedges along roads and between fields (Londono & Kvist 51, 

63). 

PHENOLOGY-The species flowers relatively constantly 

throughout the year, but the flowering intensity seems reduced 

from mid-March until mid-June. 

NOMENCLATURE AND TYpIFIcATI0N.-Gesneria trianaei: 

Herincq (1 863) based Gesneria trianaei on plants introduced in 

cultivation by Triana, while Regel (1854~) based Isoloma 

trianaei on material brought into cultivation by Warszewicz. 

According to Regel, Warszewicz asked him to name the species 

after his friend Triana. 

No~~s.--Many shoots of Kohleria trianae are strongly 

branched and certainly are several years old. However, even 

this fruticose species has shoots that die back after a single 

flowering. Kohleria trianae rarely possesses well-developed 

scaly rhizomes, but the roots are sturdy and deep-reaching 

(pers. obs.). 

15. Kohleria tubflora (Cavanilles) Hanstein 

Figures 6c (seed), 12h (flower); Cavanilles. 1801, pl. 584; W.J. Hooker, 

1846, pl. 4217; 1849, pl. 4431; Planchon, 1850. pl. 586; Raunkiaer, 1905418; 

Regel, 18548, pl. 74A,B; Oersted, 1858, pl: 4, figs. 9-24; Hoehne. 1970, pls. 

163, 175, 180. 

Kohlerio tubiflorn (Cavanilles) Hanstein. 1865:442; Gesnerio tubijroro 

Cavanilles, 1801:61; Isolomo tubiflorwn (Cavanilles) Decaisne, 1848a:465; 

Brochylomo tubiforwn (Cavanilles) Oersted, 1858:31. [Type: Panama,Nee

s.n. (MA, holotype, F (fragment).] 

Gesnerio incurvo Bentham, 1845a:131: Brochylomo incurvwn (Bentham) 

Oersted, 1858:31: Isolomo incurvwn (Bentham) Hemsley, 1882:478. [Type: 

Colombia, Cauca, GQrgona Island, Sinclair s.n. (K. holotype; K. isotype).] 

Cesnerio rhynchocorpo Bentham, 1845a:131; Isoloma rhynchocarpwn (Bentham) Decaisne, 1848a:465; Brochyloma rhynchocorpwn (Bentham) Oersted, 1858:29; dryptolomo rhynchocarpwn (Bentham) Hanstein, 1859538, 

586. [Type: Panama, Darien. Isthmus of Darien, Borcloy s.n. (K, holotype; 

MANCH, isc~ype).] 

Cesnerio hondensis sensu W.J. Hooker, 1846, pl. 4217 [not Gesnerio 

hondensis Kunth; Isoloma hondense sensu Decaisne, 1848a:465 (excluding 

the type of Gesnerio hondensis Kunth)]; Isolomo hondense var. hookeri 

Regel. 1854a:2; Cryptolomo hookerionwn (Regel) Hanstein, 1859536,581; 

Kohferio hookeri (Regel) Hanstein, 1865:442. [Type: Illustration in WJ. 

Hooker, 1846, pl. 4217. described and drawn from cultivated material, 

originally collected by Purdie in Colombia).] 

Gesnerio picta W.J. Hooker, 1849, pl. 4431; Isoloma pictwn (W.J. Hooker) 

Planchon, 1850:165; Isolomo pictwn (W.J. Hooker) Regel, 1854d:llO; 

Brochyloma pictwn (W.J. Hooker) Oersted, 1858:29; Cryptolomo pictwn 

(W.J. Hooker) Hanstein, 1859536, 582; Kohferio picfo (W.J. Hooker) 

Hanstein, 1865442. [Type: Described from cultivated material, originally 

collected by Seemann in Colombia (or possibly in Panama) (K. holotype).] 

Gesnerio picto W.J. Hooker var. minus-hirsufa W.J. Hooker 1849, sub pl. 

4431; Isolomo pictwn (W.J. Hooker) Planchon var. minus-hirsutwn (W.J. 

Hooker) Planchon, 1850:165. [Type: Described from cultivated material, 

originally collected by Lobb in Colombia; Colombia, Lobb s.n., (K. neotype 

selectedherein).] 

Cryptolomo colorotwn Hanstein, 1859536, 583 [Gesnerio picta Hortorum ex 

Hanstein, 1859537,583 (nan. nud. pro syn.)]; Kohlerio colorafo (Hanstein) 

Hanstein, 1865:442. [Type: Described from cultivated material, origin of 

original collection unknown; Panama, Canal Area, Anco Hill, 100-200 m, 

17 Oct 1922, Killip 12065 (US, neotype selected herein).] 

Isdomo hondense var. unforwn Regel, 1852b:326; Isolomo hondense var. 

wogeneri Regel, 1854a:2. [Type: Described from cultivated material, 

originally collected by Wagener in Colombia (or possibly by Warszewicz in 

Panama) (LE, neotype selected herein).] 

Isolomo hondense var. worsrewiczii Regel, 1854a:2. [Type: Described from 

cultivated material, originally collected by Warszewicz at Chagras. Panama; 

Panama, Canal Area, Balboa, 15 Sep 1936, Melt 13 (US, neotype selected 

herein).] 

Brachylomo piloswn Oersted, 1858:29; Cryptolomo pilarum (Oersted) Hanstein, 1859:538.586; Kohleria pilaro (Oersted) Hanstein, 1865:442; Isolomo 

piloswn (Oersted) Hemsley, 1882:479. [Type: Costa Rica, Aguacate, 

Oersted 9304 (C, holotype; C(2). US, isotypes).] 

Brochylomo strictwn Oersted, 1858:29; Cryptolomo strictwn (Oersted) 

Hanstein. 1859538, 585; Kohlerio stricta (Oersted) Hanstein. 1865:442; 

Isolomo sfrictwn (Oersted) Hemsley, 1882:479. [Type: Costa Rica, 

Aguacate, Oersted 9308 (C, holotype; US, isotype).] 

Cryptolomo ononymum Hanstein, 1859536, 584; Kohlerio anonym (Hanstein) Hanstein, 1865:442. [Type: Described from cultivated material (LE, 

holotype).] 

Cryptolomo cordifoliwn Hanstkn, 1859538; Kohlerio cordifolia (Hanstein) 

Hanstein, 1865:442; Isoloma cordifoliwn (Hanstein) Hemsley, 1882:478. 

[Type: Costa Rica, Hoffmn 672 (B, holotype no longer extant); Costa Rica, 

San lose, Basin of El General, 700-950 m, Jul-Aug 1943, Skurch & 

Barrontes 5152 (US, neotype selected herein; F, isoneotype).] 

Herbs or subshrubs, occasionally epilithic, often with scaly 

rhizomes and stolons; stems decumbent to erect, occasionally 

pendent, branching rare, pilose, 20-200 cm long, basal 

diameter 2-7 mm. Leaves opposite, less commonly 3 at a node, 

internodes 2-7 cm long; the blades ovate to lanceolate or 

elliptic, (1.S) 3-10 (-22) x (0.5-) 1.5-7 (-12) cm, above 

hispid to strigose, less commonly pilose, below pilose, 

occasionally purplish, veins below sericeous, yellow-green or 

purplish; petioles 1-6 cm long. Florescences frondose, partial 

florescences with 1-2 (-6) flowers; pedicels 1-3 cm long, 

pilose to villous, purplish, bracts usually present, scaly, 

caducous. Floral tube sericeous to villous; calyx 3-5 mm long, 

lobes erect, triangular to ovate, 2-4 mm long and 1.5-3.5 mm 

wide at base, outside sericeous to villous, inside appressed 

pubescent to sericeous; corolla 22-30 mm long, cylindric, 

subventricose, tube 20-28 mm long, diameter 3-5 mm at base, 

above widened to 6-10 mm, diameter 3-4 mm at throat, 

outside sericeous, inside glabrous, yellow to red, limb 3-5 mm 

wide, lobes 1.0-1.5 mm long and 1.5-2.5 mm wide at base, 

inside glabrous, yellow, with red to purplish dots; filaments 

20-25 mm long, glabrous, anthers -2.0 x 1.8 mm, included; 

nectary usually of 5 free glands, each -1.2 mm tall and 1 .O mm 

wide at base, less commonly a 5-lobed ring; ovary half-inferior, 

5-7 mm long, diameter 3-5 mm, style 18-25 mm, pilose and 

glandular-hairy, stigma subincluded. Capsule ovoid, 10-12 

mm long, diameter 6-7 mm, bivalved; seeds narrowly elliptic, 

longitudinally striate, -0.50 x 0.15 mm. Chromosome number 
NUMBER 79 67 

n = 13 (Davidse, 1970,1981; Lee, 1962; Wiehler, 1975). 

DISTRIBUTION.-P~~~~S of Kohleria tubflora are found from 

Nicaragua to Colombia and east along the Caribbean coast 

through Venezuela to Trinidad (Figure 16). Isolated populations occur in the Colombian Amazon region in Serrania de la 

Macarena, an isolated mountain range in Dept. Meta close to 

the Eastern Andean Cordillera but geologically the extreme 

western part of the Roraima sandstone. 

In Pacific Colombia, Kohleria tubifora is apparently only 

represented by the ecotype restricted to coastal cliffs (Figures 

16, 34). These plants are known from the Pacific coast of 

Panama south to Gorgona Island in the Dept. Cauca of 

Colombia. In addition, two collections come from the 

Caribbean coast from Prov. San Blas of Panama (Garwood 

II90), and the Prov. Limones of Costa Rica (Quiros 513) 

(Figure 34). In western Costa Rica, adjacent Panama, and on 

Cocos Island the coastal ecotype of K. spicata apparently 

replaces that of K. tubifora (Figure 34; see also “Discussion” 

under K. spicata). 

Most Kohleria tubiflora collections come from near sea level 

(Figure 21), and the species is rarely collected above 1000 

meters altitude. In Panama K. tubflora is the most frequently 

collected of all Gesneriaceae (Skog, 1979). 

SELECTED COLLECTIONS.-COLOMBIA: ANTIOQUIA: 

Cauca valley, 10 km N of Valdivia, Gentry & Fallen I7946 

(COL, MO, US). CAUCA: Gorgona Island, Fernander 418 

(COL, US). CHOCO: W of Mutaa4 Pavarano6 Grande, Gentry & 

Le6n 20245 (AAU, COL, MO, US). CORDOBA: Rio Ure, 

Espinu 660 (COL). MAGDALENA: Santa Marta, near Jiracasaea 

and OAaca, Smith 380 (BM, F, G(2), GH, MO, NY, UC, US, 

WIS). META: Rio Mico, S of El Mico airstrip, Philipson et al. 

1348 (BM, COL, F, NY, US). NARINO: Isla de Gallos, 

Dryander 2593 (US). 

COSTA RICA ALAJLJELA: Close to Rio Poas Brenes I7275 

(F, NY). GUANACASTE: 33 km N of Caiias, Frankie 266A (F, 

MO). PUNTARENAS: 8 km NW of junction of Highway 17 (to 

Puntarenas) and Carretera Interamericana, Davidse & Pohl 

1321 (F, MO, US). SAN JOSE: 3 km from Rivas and 7 km from 

San Isidro del General, Whitmore 80 (F, NY, MO). 

NICARAGUA: mAS: 3 km NE of San Juan del Sur, La 

Cuesta, Davidse & Pohl2284 (M0(2), NY, US). 

PANAMA: CANAL AREA: Barro Colorado Island, Croat 

41 74 (F, MO, NY), Croat 12441 (F, GH, MO(2)). CHIRIQUI: 

Cerro San Cristobal-David, Caballero 4 (US). COCEB: Vicinity 

of El Valle, Allen 743 (GH, MO, NY). COLO Portobelo, 

D’Arcy & D’Arcy 6687 (F, GH, MO, NY). DARIEN: Vicinity of 

Santa Fe, Rio Sabana, Duke 4134 (GH, MO, US). Los SANTOS: 

Headwaters of Rio Pedregal, fewis 2909 (MO, UC). PANAMA: 

3 km S of Accalda Dim, Cerro Penon, Nee 8598 (AAU, MO, 

US). SAN BLAS: Ustupo, D’Arcy 9487 (MO, US). VERAGUAS: 3 

km S of Canlitas, D’Arcy 15153 (MO, US). 

VENEZUELA: BARINAS: Baranitas-Apartaderos road, km 2, 

Breteler 4198 (F, G, MO, NY, US). CARABOBO: Between 

Valencia and Yuma, Alston 5667 (BM). LARA: NE of 

Barquisimeto, Guaremal River, Meijer & Smith 65 (MO, SEL). 

PORTUGUESA: 30 km NNE of Guanare, NE of Boca de Monte, 

Steyermark 127050 (US), Steyermark 127073 (US). SUCRE: 

Sucre district, dam site on the Rio Neveri, Davidse & Gonzailes, 

19038 (MO, NY, US). ZULIA: Valley of Rio Guasara, Bunting 

et al. 11970 (NY, US). 

TRINIDAD: ST. ANN’S, Jades Amandes, Broadway 7900 

(BM, K, MO). Total of 260 collections seen. 

DISTINGUISHING FEATURES.-The tubular ventricose flowers with a small limb and a narrow throat distinguish Kohleria 

tubij7ora (Figure 12h) from all species except K. hondensis 

(Figure 12g). The latter species has lanceolate calyx lobes (see 

discussion under K. hondensis). Kohleria tubflora and K. 

spicata are vegetatively almost indistinguishable. 

HABITAT-Kohleria tubflora is a rather weedy species of 

disturbed, open habitats, e.g., along rivers, on coastal cliffs, and 

roadsides and other man-created disturbances. Johnston (1949) 

wrote about the coastal cliff ecotype on San Jose Island of the 

Pearl Archipelago in the Gulf of Panama: “Here it grows on the 

barren islets and exposed tops of large rocks along the shore.” 

PHEN0LOGY.-According to Croat (1978) and observations 

on herbarium sheets, the species is strongly seasonal in 

Panama, flowering from September to January in the rainy 

season. In contrast, in Panama Kohleria spicata continues to 

flower during the dry season. 

NOMENCLATURE AND TYPIFIcAnoN.~esneria incurva: 

Bentham (1845a) described Gesneria incurva and G. rhynchocarpa from plants adapted to salt influence on coastal rocks. 

Gesneria rhynchocarpa: Barclay, Hinds, and Sinclair all 

collected plants on the voyage of the H.M.S. Sulphur. The 

typification is problematic because Bentham (1845a) gave 

localities but neither collectors’ names nor collection numbers. 

In the case of Gesneria rhynchocarpa the type locality was 

given as “Isthmus of Darien,” and there exists one Barclay 

collection labeled with this locality at K. In addition, a 

specimen at MANCH apparently is a duplicate (here the label 

reads “Panama @hien]”). The specimen at K is annotated as 

Gesneria rhynchocarpa var. oppositifolia, but no varieties have 

been published in G. rhynchocarpa. Two Sinclair specimens at 

K also annotated as G. rhynchocarpa come from the Pacific 

coast of Colombia. They have a ternate leaf arrangement in 

accordance with the description of G. rhynchocarpa. Thus, it 

appears that Bentham (1845a) originally intended to describe 

two G. rhynchocarpa varieties with opposite and ternate leaves, 

respectively, but ended with the specimen from Darien cited as 

the type, even though this collection has opposite leaves in 

conflict with the description. 

Gesneria hondensis: W.J. Hooker (1846) assumed he was 

undertaking a more detailed description based on live material 

of Gesneria hondensis of Kunth (1818) (now Kohleria 

hondensis), but W.J. Hooker was actually describing material 

of Kohleria tubifora. 

Gesneria picta var. minus- hirsuta: W.J. Hooker (1849) 

described the variety from live material introduced by Lobb 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

from Colombia, but no material called G. picta var. minushirsuta has been located. Consequently, the only Lobb 

collection of K. tubiflora seen from Colombia has been selected 

as neotype. 

Cryptoloma coloratum: Hanstein (1859) described C. coloratum from cultivated material received from England. He 

suspected the species to be identical with W.J. Hooker’s (1849) 

Gesneria picta var. minus- hirsuta. However, the description of 

the variety minus- hirsuta was brief and poor, and did not give 

sufficient evidence to conclude the two taxa to be identical. The 

description of Cryptoloma coloratum demonstrates that the 

species is a synonym of Kohleria tubifora, and that the species 

apparently was based on material not particularly distinct from 

typical K. tubiflora plants. 

Isoloma hondense var. uniforum: Regel (1852b) mainly 

distinguished the variety by its single flowers in the leaf axils, 

in contrast to several axillary flowers in Gesneria hondensis 

sensu W.J. Hooker (1846). However, he soon realized that his 

variety occasionally would develop several axillary flowers, 

and for the same material he created the name var. wageneri 

(1854a). Regel named the plant after Wagener despite his 

mentioning both Wagener in Colombia and Warszewicz in 

Panama as possible collectors of the material on which he based 

var. unzj7ora (1852b). The neotype selected herein was 

annotated several years later with the name Isoloma hondense 

var. wageneri by Regel in St. Petersburg (formerly now 

Len ingrad). 

Isoloma hondense var. warszewiczii: The description by 

Regel (1854a) is brief. However, no major differences 

apparently exist between the variety warszewiczii and typical 

K. tubiji’ora specimens. 

Cryptoloma cordijolium: The description indicates no major 

differences between Cryptoloma cordifohm and typical 

Kohleria tubzj7ora specimens. A similar Costa Rican neotype 

has been selected. 

NOTES.-Kohleria tubiflora is stable in most characters 

throughout its range. The ecotype of salt-influenced rocky 

coasts differs from typical K. tubiflora plants by having more 

succulent and less hispid leaves. 

Kohleria tubiflora hybridizes with K. hirsuta on Trinidad 

(see “Interspecific Hybridization” in section above). 

16. Kohleria villosa (Fritsch) Wiehler 

Kohleria villoso (Fritsch) Wiehler, 1978b:63. 

Herbs, rarely subshrubs, scaly rhizomes not observed, 

stolons abundant; stems decumbent to erect, pilose to villous, 

erect part 8-35 (-100) cm long, basal diameter 1.5-4.5 mm; 

lateral branches rare. Leaves opposite, internodes 2-7 cm long: 

the blades ovate, less commonly lanceolate, 2-8 (-13) x 1-5 

(-9) cm, above hispid, individual uichomes often located on 

small bullae, below pilose, usually purplish with a pattern of 

light green blotches, veins villous, yellow-green: petioles 

(0.3-) 1-3 (-5) cm long. Florescences frondose, partial 

florescences with 1 (-6) flower(s): pedicels 2-4 cm, villous, 

yellow-green to purple, bracts scale-like, caducous. Floral tube 

villous; calyx 4-6 mm long, lobes erect, ovate to lanceolate or 

spathulate, 3-4 mm long and 1-2 mm wide at base, outside 

pilose, inside basally glabrous to apically pilose; corolla 15-18 

mm long, broadly tubular, less commonly merely tubular, 

medially often sharply arcuate downward l/3 of length from 

base, tube 12-15 mm long, diameter 2-3 mm at base and 7-10 

mm at throat, outside sericeous or villous, dorsally deep red, 

ventrally yellow, inside pilose to villous but basally glabrous, 

glandular-hairy, yellow, ventrally with red spots, the base 

dorsally gibbous, limb 9-19 mm wide, inside glabrous, red, 

dorsal lobes 4-5 mm long and 4-5 mm wide at base, lateral 

and ventral lobes 3-5 mm long and 5-7 mm wide at base; 

filaments 8-11 mm long, basally glabrous, apically pilose to 

villous, anthers -1.8 x 1.5 mm, included; nectary of 5 separate 

glands, each 2.0-2.5 mm high and 0.5-1.2 mm wide at base, 

rarely 3 free and 2 basally fused glands; ovary half-inferior, 

3-4 mm long, diameter 1.5-2.5 mm, style 7-10 mm long, 

pilose and glandular-hairy, stigma subincluded. Capsule ovoid, 

6-8 mm long, diameter 3-5 mm, dehiscing apically into 2 

valves (by 2 slits) and later opening by one of the slits 

continuing to the base; seeds sub-globose, irregularly striate, 

-0.25 x 0.25 mm. Chromosome number n = 13 (Wiehler, 1975, 

as K. platylomata). 

DISTRIBUTION.-KoWeria villosa is restricted to the western 

slopes of the Andes in Ecuador and Colombia (Figure ZO), 

where it ranges from (500-) 900-2100 meters altitude (Figure 

21). The variety anisophylla occurs at higher elevations than 

does var. villosa. When Fritsch (1913a) described Diastema 

villosum he cited a Sodiro collection (11917) from Nap0 in 

eastern Ecuador. Kohleria villosa is otherwise unknown from 

the Eastern slopes of the Andes, and the specimen was probably 

misidentified. The collection was apparently destroyed at B. 

DISTINGUISHING FEATURES.-The purple and green mottled 

lower leaf surfaces (not conspicuous in var. anisophylla), the 

epedunculate inflorescences, the uniformly red corolla lobes, 

the weakly bullate upper leaf surfaces (Figures 36b, 37b), the 

long and narrow nectary glands (Figures 36f, 37e), the globose 

seeds (Figure 6d) and the ovoid capsules (Figure 36g) that 

dehisce into 2 apical valves with one of the slits continuing to 

the base (Figure 13b), make K. villosa distinctive. The latter 

four characters are shared with the genus Diastema, to which K. 

villosa thus shows an affinity (see “Related Genera” section 

above). 

HABITAT-The species may require less light than any other 

Kohleria species. Flowering plants were seen in deeply shaded, 

very humid forest understory, e.g., along small streams, and 

below steep rocks (Kvist et al. 60308). However, the species 

also thrives and flowers in low vegetation exposed to the sun, 

at least in permanently humid montane forest regions (Kvist et 

al. 60307). Plants growing in shade are more robust and have 

darker leaves than sun-exposed plants. 

PHEN0LOGY.-The flowering intensity is relatively stable 

throughout the year (Figure 9). 

NOTES.-Some specimens, especially from the area around 
NUMBER 79 69 

FIGURE 36.4ohleria villosa var. villosa; a, habit and inflorescences; b, indument of upper leaf surface; c, 

flower, d, limb and throat; e, laterally opened corolla showing androecium; f, calyx, nectary, and gynoecium; g, 

capsule. a-f, from Harling & Andersson 23222 (US); g, from Holmgren 820 (S). 
70 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

Nanegal in the Prov. Pichincha of Ecuador (e.g., Hurling & 

Andersson 11600) have features intermediate between the two 

varieties. However, because var. villosa and var. anisophylla 

are usually easily distinguished (Figures 36, 37) and differ in 

geographical (Figure 20) and altitudinal ranges, we find 

taxonomic recognition justifiable. 

16a. Kohleria villosa var. villosa 

Figures 6d (seed), 7q @ollen), 36 (habit and floral features). 

Kohleria villosa (Fritsch) Wiehler, 1978b:63; Diastema villoswn Fritsch, 

1913a:407. [Type: Ecuador, Jameson s.n. (W, lectotype selected herein; W, 

isolectotype (?)).I 

Diastema platylomatwn J.D. Smith, 1916:381; Kohleria platylomata (J.D. 

Smith) Wiehler, 1973:327. [Type: Ecuador, Pichincha, Rio Pilaoh, 

Lehmann 409a (US, lectotype selected herein; BM, LE, isolectotypes).] 

Herbs up to 40 cm tall; leaves equal to subequal in pairs, 

blades ovate with acute apex, below usually conspicuously 

purplish with a pattern of light green blotches. Corolla broadly 

tubular, outside sericeous, sharply arcuate downward l/3 

of length from base, limb 12-19 mm wide, lobes 3-5 mm 

long. 

DISTRIBUTION.-Kohleria villosa var. villosa is restricted to 

the western slopes of the Andes in Ecuador (Figure 20), and 

ranges from 900-1600 meters altitude. The variety is locally 

common in Prov. Pichincha. A single collection from Esmeraldas province (Balslev & Steere 1312) apparently comes from 

much lower elevations; according to the label it came from 500 

meters elevation, but the locality (located by coordinates given 

on the label) must be below 200 meters. 

SELECTED COLLECTIONS.-ECUADOR: BOLIVAR: Tablas 

valley, Balsapamba, Acosta-Solis 6157 0. COTOPAXI: 2 km S 

of El Corazon, Hurling & Andersson 19193 WS). ESMERALDAS: 20 km SE of San Mateo, Zapallo, Balslev & Steere 1312 

(AAU, NY). PICHINCHA: Old Quito-Santo Domingo road, 3 km 

from bridge over Rio Pilaon, Hurling & Andersson 23058 

(GB, US); 1 km above Tandapi on Quito-Santo Domingo road, 

Hurling & Andersson 23222 (GB, US); Tandapi, Holmgren 

820 (S(2)); new Quito-Santo Domingo road, La Union del 

Toachi, Kvist et al. 60307 (AAU, COL, NY, QCA, QCNE, 

US); Kvist et al. 60308 (AAU, COL, NY, QCA, QCNE, US); 

SW of Chiriboga, Ljtnant & Molau 15662 (AAU); 3 km from 

La Armenia, llgaard et al. 37851 (AAU). Total of 38 

collections seen. 

Fritsch (1913a) cited several collections but selected no type. 

Three Sodiro collections (119/2, 119/3, 119/7) were destroyed 

in Berlin, although (1 19/3) apparently is represented at BP and 

thus is a syntype. Another syntype at W is a Karsten collection. 

Diastemu platylomutum: J.D. Smith (1916) cited two 

Lehmann collections but selected no type. The syntype 

(Lehmann 481a) is represented in the same three herbaria as the 

lectotype (Lehmunn 409a). 

NOMENCLATURE AND TYPIFICATION.-Diustemu villosum: 

16b. Kohleria villosa var. anisophylla (Fritsch) 

Kvist & L. Skog, new status 

Figure 37 (habit and floral features). 

Diastema anisophyllwn Fritsch, 1913a:408; Kohleria anisophylla (Fritsch) 

Wiehler, 1978b:62. [Type: Colombia, Narino, W of Tuquerres, Piedra 

Ancha, Lehmann 5843 (B, holotype not extant, K, lectotype selected herein; 

K, isolectotype).] 

Nematanfhus erianthus Bentham, 1846:231 [not Kohleria eriantha (Bentham) 

Hanstein, 1865:442]; Colwnnea eriantha (Bentham) Hanstein, 1865:391. 

[Type: Ecuador, Pichincha, Nanegal, Hartweg s.n. (K, holotype).] 

Diastema anisophyllwn Fritsch var. quitense Fritsch, 1913a:408. [Type: 

Ecuador, Pichincha, “Quito,” Jameson s.n. (w, holotype).] 

Herbs to subshrubs, up to 1 m tall; leaves unequal in pairs, 

blades ovate to lanceolate with acuminate apex, below not with 

a conspicuous purplish and green pattern. Corolla tubular, 

outside densely villous, not arcuate downward ventrally, limb 

9-13 mm wide, lobes 2-3 mm long. 

DISTRIBUTION.-Kohleria villosa var. anisophylla comes 

from the western slopes of the Andes in Provinces Pichincha 

and Carchi of Ecuador and Dept. Narifio of Colombia (Figure 

20). The few collections come from altitudes between 1800 and 

2100 meters. 

OTHER COLLECTIONS.-COLOMBIA: NARINO: Near 

Chucunes, Finca La Planada, Libenson et al. 30538 (COL, US); 

Libenson et al. 30554 (COL, US); Libenson et al. 30555 

(COL). 

ECUADOR: CARCHI: Toward Maldonado from Tulain, 

1800 m, Luer et al. 2672 (SEL). PICHINCHA: Nanegal, Andre 

K39 (K); Andes near Quito, Couthouy sen. (GH); above Mindo, 

Luer et al. 4744 (SEL); old Quito-Santo Domingo road, km 59, 

Zuk & Jaramillo 578 (MO, US). LOCAL^ UNKNOWN: 

Jameson 111 (BM, K). Total of 12 collections seen. 

17. Kohleria warszewiczii (Regel) Hanstein 

Figures 6e,f (seed and seed surface), 7r (pollen), 8c (seedling with scaly 

rhizome), 12j (flower); Regel, 1853a, pl. 61; Planchon, 1853, pl. 941; Linden 

and And& 1870, pl. 17; Fritsch, 1894, pl. 77: fig. I; Kvist, 1987:18 (photo). 

Kohleria warszewiczii (Regel) Hanstein, 1865:441; Sciadocalyx warszewiczii 

Regel, 1853a:257; [Gesneria regeliana Warscewicz [sic] ex Planchon, 

1853:213 (nom. nud. pro syn.)]; Isoloma regelianwn Voss in Siebert C% Voss, 

1894:793 (nom. illeg., included Sciadocalyx warscewiczii [sic] Regel in 

synonymy); Isoloma warszewiczii (Regel) Bailey, 1917:3115. [Type: 

Illustration in Regel, 1853a, pl. 61, described and drawn from cultivated 

material, originally collected by Warszewicz in Colombia at St. Martha.] 

Sciadocalyx digitaliflora Linden & Andre 187095; Isoloma digitaliflorwn 

(Lmden & Andre) Voss in Siebert & Voss, 1894:793; Kohleria digitalijlora 

(Linden & Andre) Fritsch, 1894:178. [Type: Illustration in Linden and 

Andre 1870,pl. 17, described and drawn from cultivated material, originally 

collected by Wallis in Antioquia, Colombia.] 

FIGURE 37 (right).-Kohleria villosa var. anisophylla; a, habit and flowers; b, 

indument of upper leaf surface; c, flower. d, dorsally opened corolla showing 

androecium; e, calyx, nectary, and gynoecium. a,b, from Libenson et al. 30554 

(US); c,d.e, from Libenson et al. 30538 (US). 
NUMBER 79 71 
72 SMITHSONIAN CONTRIBUTIONS TO BOTAhY 

Kohleria violacea Fritsch, 1913a:414. [Type: Colombia, Cauca, Popayan, 

1700-2500 m, Lehmann 4495 (B, holotype not extant, K, lectotype selected 

herein; F. GH, US, isolectotypes).] 

Herbs or shrubs, often epipetric, with scaly rhizomes and 

stolons; stems erect, occasionally decumbent, usually branching, villous, covered with violet trichomes, 30-200 cm long, 

basal diameter 4-15 mm. Leaves opposite, less commonly 

ternate or whorled (4 at each node), internodes (1.5-) 3-9 cm 

long; the blades elliptic to ovate, occasionally lanceolate, 7-20 

x 2-9 cm, above pubescent to tomentose, below tomentose, 

less commonly villous or pilose, veins below villous, yellowgreen, occasionally violet; petioles 0.5-3.0 cm. Florescences 

frondose, partial florescences with 3-6 (-10) flowers; peduncles (3-) 5-9 (-13) cm, pedicels 1-3 (-6) cm, yellow-green, 

villous, mchomes violet, bracts lanceolate to ovate, 5-10 x 

2-3 mm, caducous. Floral tube tomentose; calyx 5-8 mm 

long, lobes often reflexed, triangular to rotundate, 5-6 mm 

long and 5-8 mm wide at base, outside villous, green suffused 

with red trichomes, inside sericeous; corolla (25-) 30-42 mm 

long, campanulate, subventricose, tube 22-40 mm long, 

diameter 4-6 mm at base, above widened to (6-) 9-12 (-16) 

mm, diameter 7-13 mm at throat, outside pilose to villous, 

lavender, ventrally lavender to cream, inside glabrous, lavender 

to cream, ventrally with purple dots, basally white, the limb 

12-24 mm wide, lobes 5-7 mm long and 4-6 mm wide at 

base, inside sparsely glandular-hairy, yellow-green with numerous purple dots; filaments 25-40 mm, white, sparsely 

pubescent, anthers -2.5 x 1.5 mm, subincluded; nectary often 

of 3 free and 2 dorsal basally united glands, each -1 mm high 

and 1.5 mm wide at base, less commonly a 5-lobed ring; ovary 

5-6 mm long, diameter 3-4 mm, style 24-32 mm, pilose and 

glandular-hairy, stigma subincluded. Capsule ovoid, 10- 15 

mm long, diameter 6-10 mm, bivalved; seeds elliptic, 

longitudinally to transversely striate, -0.6 x 0.3 mm. Chromosome number n = 13 (Wiehler, 1975, as K. digitaliflora). 

DISTRIBUTION.-Kohleria warszewiczii is widespread in the 

Colombian highlands (Figure 19). The species occurs in all 

three Cordilleras, but is apparently absent from the eastern 

slopes of the Eastern Cordillera (Figure 21). Kohleria 

warszewiczii ranges from 1300 to 2500 meters altitude, with a 

few collections from the Pacific lowlands of Dept. Cauca down 

to 300 meters altitude (Figure 21). The species is locally 

common between 1500 and 2100 meters altitude (pers. obs.). 

Medellin-Antioquia road, km 20, Metcalfe 30033 (GH, MO, 

uc, Us). CALDAS: Salamin, Tomus 1919 (US). CAUCA: 

Timbio, Hatoviejo, Arbelaez & Cuatrecasas 6079 (COL, F, 

US); Popaah, Lehmunn 9092 (G(2), GH(2), K, NY). cuh?)~ 

NAMARCA: Caqueza to Rio Sananie, Pennell 1337 (GH, MO, 

NY, US). HUILA: La Plata-Popaah road, km 25, Londono & 

Kvist 166 (AAU, COL, QCA, TULV, US, USM, VEN); km 42, 

Londono & Kvist 167 (AAU, COL, QCA, TULV, US). 

QUINDIO: Above Calarca, Alston 7784 (BM, S, US). RISARALDA: Apia-Pueblo Rico road, km 32, Luteyn et al. 7336 (COL, 

NY, SEL). TOLIMA: Below Santa Isabel, Londono & Kvist 62 

(AAU(2), COL, NY, QCA, TULV, US, USM, VEN). VALLE: 

Rio Cali, Pichinde, Cuatrecasas 18231 (F(3), US). Total of 72 

collections seen. 

DISTINGUISHING FEATURES.-Kohleria warszewiczii is set 

apart by the long-pedunculate inflorescences of 3-6 (-10) 

flowers (Figure 3e, f), the tubular to campanulate corolla with a 

usually lavender tube and purple-dotted, yellow-green limbs 

and the rotundate, often reflexed calyx lobes (Figure 12j). 

Kohleria trianue and K. hirsuta are vegetatively similar. 

Capanea affinis has flowers similar to those of Kohleria 

warszewiczii but its habit is scandent. However, the latter two 

species frequently are confused in herbaria (see “Related 

Genera” in Introduction). 

HABITAT.-Kohleria warszewiczii is found on exposed, 

often dry slopes along rivers, roads, etc. The species is light 

demanding, but can survive in fairly tall and dense vegetation 

due to its often fruticose habit. 

PHEN0LOGY.-The flowering intensity is greatest from 

March to September, but it is usually possible to find at least a 

few plants with flowers (Figure 9). 

NOTES.-The coloration and the size of the corolla vary 

considerably. Small-flowered, epipemc Kohleria warszewiczii 

plants may look rather similar to K. amubilis. This is especially 

true in Dept. Cundinamarca where the two co-occur (Figures 

19-21) and possibly hybridize. However, K. amabilis always 

differs from K. warszewiczii by having epedunculate inflorescences. 

SELECTED COLLECTIONS.-COLOMBIA: ANTIOQUIA: 
Appendix 1 

List of Excluded Names 

Isoloma bulbosum Baillon, 1888b, pls. 50, 51. [= Sinningia sp.] 

Isoloma costaricense (Klotzsch ex Oersted) Hemsley, 1882:478. 

[= Moussonia elegans Decaisne, 1849:489] 

Isoloma deppeanwn (Schlechtendal & Chamisso) Hemsley, 1882:478. 

[= Moussonia deppeana (Schlechtendal & Chamisso) Hanstein 

18652841 

Isoloma elatior Decaisne, 1848a:465. [= Simingia incarwta (Aublet) 

D. Denham, 1974:126] 

Isoloma elegans (Decaisne) Hemsley, 1882:478. [= Moussonia 

elegam Decaisne, 1849:489] 

Isoloma frexuosum Rusby, 1900:28. [= Parakohleria parviflora 

(Rusby ) Wiehler, 197 8 a: 81 

Isoloma hahnii Hortorum ex Baillon, 1888a:719. [= Moussonia sp.] 

Isoloma hypocyrtiflorum (W.J. Hooker) Bentham ex Jackson, 

18951238 (cf. Bentham, 1876:1002). [= Pearcea hypocyrtiJIora 

(W.J. Hooker) Regel, 1867:388] 

Isoloma jaliscanum S. Watson, 1890159. [= Moussonia elegans 

Decaisne, 1849:489] 

Isoloma lasianthum (Zuccarini) Decaisne, 1848a:465. [= Moussonia 

deppeana (Schlechtendal & Chamisso) Hanstein, 18652841 

Isoloma lindenianum (Regel) Nicholson, 1885:201. [= Gloxinia 

lindeniam (Regel) Fritsch, 191313366] 

Isoloma oblanceolatwn J.D. Smith, 1916:383. [= Capanea affinis 

(“Campanea”) Fritsch, 191 3a:4341 

Isoloma parviflorum Rusby, 1896:97. [= Parakohleria parviflora 

(Rusby) Wiehler, 1978a:8] 

Isoloma patentipilosum 0. Kuntze, 1898:242. [= Parakohleria 

parviflora (Rusby) Wiehler, 1978a:8] 

Isoloma purpureurn (Poeppig) Bentham, 1876: 1002. [= Parakohleria 

purpurea (Poeppig) Wiehler, 1978a:8] 

Isoloma sprucei Britton in Rusby, 1896:97. [= Parakohleria sprucei 

(Britton) Wiehler, 1978a:9] 

Isoloma urticifoliwn Rusby, 1900:28. [= Parakohleria parviflora 

(Rusby) Wiehler, 1978a:8] 

Isoloma vulcanicolwn J.D. Smith, 1916:382. [= Capanea affinis 

(“Campanea”) Fritsch, 19 13a:434] 

Kohleria avilana Cuatrecasas, 1935:256 (reprint page 53). 

[= Parakohleria avilana (Cuatrecasas) Wiehler, 1978a:5] 

Kohleria baezana Cuatrecasas, 1935:257 (reprint page 54). 

[= Parakohleria baezaw (Cuatrecasas) Wiehler, 1978a:6] 

Kohleria collina T.S. Brandegee, 1914:66. [= Moussonia elegans 

Decaisne, 1849:489] 

Kohleria deppeanu (Schlechtendal & Chamisso) Fritsch, 1894 179. 

[= Moussonia deppeana (Schlechtendal & Chamisso) Hanstein, 

1865:284] 

Kohleria deppeana var. lasiantha (Zuccarini) Fritsch, 1913a:425. [= 

Moussonia deppeam (Schlechtendal & Chamisso) Hanstein, 

1865:284] 

Kohleria elegans (Decaisne) Loesener, 1899574. [= Moussonia 

elegans Decaisne, 1849:489] 

Kohleria elegans var. pedunculata (T.S. Brandegee) C. Morton, 

1967a:75. [= Moussonia elegans Decaisne, 1849:489] 

Kohleria elongata Regel, 1855a:4. [= Moussonia deppeana 

(Schlechtendal & Chamisso) Hanstein, 1865:284] 

Kohleria jilisepala Standley, 1940: 107. [= Omiltemia filisepala 

(Standley) C. Morton, 1968:98] [Rubiaceae]) 

Kohleria fruticosa T.S. Brandegee, 1914:67. [= Moussonia fruticosa 

(T. S. Brandegee) Wiehler, 1975:25] 

Kohleria hirsutissima C. Morton, 1967a:65. [= Moussonia hirsutissima (C. Morton) Wiehler, 1975251 

Kohleria jmsoniana Fritsch, 1913a:429. [= Parakohleria jamesoniam (Fritsch) Wiehler, 1978a:6] 

Kohleria Iindeniam (Regel) H.E. Moore, 1954:380. [= Gloxinia 

lindeniana (Regel) Fritsch, 1913b:66] 

Kohleria martensii Fritsch, 1913a:428. [= Moussonia triflora (Martens & Galeotti) Hanstein, 18652861 

Kohleria papillosa (Oersted ex Hanstein) Fritsch, 1913a:427. 

[= Moussonia elegans Decaisne, 1849:489] 

Kohleria papillosa var. pendula C. Morton, 1967a:72. [= Moussonia 

eleganir Decaisne, 1849:489] 

Kohleria papillosa var. sericea Fritsch, 1913a:428. [= Moussonia 

elegans Decaisne, 1849:489] 

Kohleria papillosa var. solitaria C. Morton, 1967a:73. [= Moussonia 

elegans Decaisne, 1849:489] 

Kohleria patedpilosa (0. Kuntze) Schumann, 1898:386. [= Parakohleria parvifrora (Rusby) Wiehler, 1978a:8] 

Kohleriapedunculata T.S. Brandegee, 1914:67. [= Moussonia elegans 

Decaisne, 1849:489] 

Kohleria regelii Hanstein, 1865442 (nom. nud.). [= Moussonia 

deppeam (Schlechtendal & Chamisso) Hanstein, 1865:284] 

Kohleria reticulata Fritsch, 1913a:428. [= Parakohleria sprucei 

(Fritsch) Wiehler, 1978a:9] 

Kohleria rupicola Standley & L.O. Williams, 1952:62 [= Moussonia 

rupicola (Standley & L.O. Williams) Wiehler, 1975:25] 

Kohleria sancti-josephi Cuatrecasas, 1935:258 (reprint page 55). 

[= Parakohleria sancti-josephi (Cuatrecasas) Wiehler, 1978a:8] 

Kohleria saxicola T.S. Brandegee, 1914:66. [= Achirnenes jlma C. 

Morton, 1936:13] 

Kohleria septentrionalis D. Denham in C. Morton, 1967a:70. 

[= Moussonia septentrionalis (D. Denham) Wiehler, 1975:25] 

Kohleria serrulata C. Morton, 1939:309. [= Moussonia serrulata (C. 

Morton) Wiehler, 1975:25] 

Kohleria skutchii C. Morton & D. Gibson in D. Gibson, 1972:336. 

[= Moussonia skutchii (C. Morton & D. Gibson) Wiehler, 1975:25] 

Kohleria sprucei (Britton) Fritsch, 1913a:429. [= Parakohleria 

sprucei (Britton) Wiehler, 1978a:9] 

73 
74 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

Kohleria strigosa C. Morton, 1938:1181. [= Moussonia strigosa (C. 

Kohleria urticifolia (Rusby) Fritsch, 1913a:430. [= Parakohleria 

Kohleria viminalis TS. Brandegee, 1915:194. [= Moussonia viminalis 

Kohleria weberbaueri Fritsch, 1913a:430. [= Parakohleria weberbauTydaea elegans (Decaisne) Planchon & Linden in Linden, 18554 (not 

Morton) Wiehler, 1975251 

parvifora (Rusby) Wiehler, 1978a:8] 

(T.S. Brandegee) Wiehler, 1975261 

eri (Fritsch) Wiehler, 1978a:lOl 

seen). [= Moussonia elegam Decaisne, 1849:489] 

Tydaea lindeniana Regel, 1868:257. [= Gloxinia lindeniana (Regel) 

Fritsch, 19 13b:66] 

Tydaea lindenii Regel ex Andre, 1873:183, pl. 147. [= Gloxinia 

lindeniana (Regel) Fritsch, 1913b:66] 

NOTES ON NOMENCLATURE.-ISO~OVU elatior: Decaisne 

(1848a) probably intended to transfer Gesneria elatior Kunth 

(1818) to Isoloma, but neglected to include the basionym or the 

author. The possibility that Isolomu elatior is a currently 

unknown cultivar also exists. 

Appendix 2 

Names Based on Hybrids 

Only those taxa described as hybrids are marked as such (x). Types have not been 

selected herein and only those mentioned in the original descriptions are included. Moore 

(1954, 1957) gives the parentage of the majority of the hybrids included here. In many 

cases other hybrids constitute at least one of the parents. 

Gesneria oblongata Paxton, 1839:103. 

Isoloma bogoteme Nicholson var. gigdeum Voss, 1894:793. 

[Tydaea gigantea Van Houtte ex Voss, 1894:793 (nom. nud. pro 

Kohleria bella C. Morton, 1967b:79. [Type: H.E. Moore 7556 bis, 

Described from cultivation, originally collected by Horich in San 

Jose, Costa Rica, (US, holotype).] 

Kohleria behnickii Fritsch, 1916:32. [Tydaea decaisneana Hortorum 

ex Fritsch. 1916:32 (nom. nud. pro syn.); Tydaea grandiflora 

Hortomm Behnick ex Fritsch, 1916:33 (nom. nud. pro syn.).] 

[Type: Described from cultivation (B, no longer extant).] 

Kohleria dubia Fritsch, 1916:3 1; Isoloma giganteum Hortorum 

Bentham ex Fritsch, 1916:32. [Type: Described from cultivation 

(B, no longer extant).] 

Kohleria maculata C. Morton, 1938:1180. [Type: Costa Rica, San 

Jose, Morana 40 (F, holotype).] 

Sciadocalyx x lucianii Andre, 1874: 15 1; Kohleria x lucianii (Andre) 

Fritsch, 1894:178; Isoloma lucianii (Andre) Voss, 1894:793. 

Tydaea x decaisniana Regel, 1858:374. 

Tydaea x gigantea Planchon, 1855a:21; Kohleria x gigantea (Planchon) Fritsch, 1894178; Tydaea x hybrida Hortonun ex H.E. 

Moore, 1954:384. 

Tydaea x hansteinii Ortgies. 1858:65; Isoloma bogotense Nicholson 

var. hanrteinii (Ortgies) Voss, 1894:794; Kohleria x hansteinii 

(Ortgies) H.E. Moore, 1954:385. 

Tydaea hillii Regel, 1854b373. [Achimenes hillii Hortorum ex Regel, 

1854b73 (nom. nud. pro syn.); Achimenes kmemis Hortorum ex 

Regel, 1854b:73 (nom. nud. pro syn.)]; Isoloma bogotense 

Nicholson var. hillii (Regel) Voss, 1894:794; Kohleria x hillii 

(Regel) H.E. Moore, 1954:386. 

Tydaea x lenneana Ortgies, 1857:2; Isoloma bogotense Nicholson var. 

lewanum (Ortgies) Voss, 1894:794; Kohleria x lewana 

(Ortgies) H.E. Moore, 1954:386. 

SynJ.1 

Tydaea x lenneana var. viridis Ortgies, 1857:2. 

Tydaea x lenneana var. rutilans Ortgies, 1857:3; Isoloma bogotense 

Nicholson var. rutilans (Ortgies) Voss, 1894:794. 

Tydaea x meyendor-i Regel. 1858:374. 

Tydaea x meyendor-i var. rubra Regel, 1858:375. 

Tydaea x ortgiesii Hortorum Van Houtte ex Planchon, 1856:213; 

Isoloma x ortgiesii (Hortomm ex Planchon) Voss, 1894:793; 

Kohleria x ortgiesii (Hortorum ex Planchon) H.E. Moore, 

1954:387. 

Tydaea picta (Bentham ex W.J. Hooker) Decaisne var. karsteniana 

Hortomm ex Hanstein, 1865:439. [Tydueu karsteniana Hortorum 

ex Hanstein (nom. nud. pro syn.).] 

Tydaea x pulchra Herincq, 1856:98; Isoloma bogotense Nicholson 

var. pulchrum (Herincq) Voss, 1894:794; Kohleria x pulchra 

(Herincq) H.E. Moore, 1954:387. 

Tydaea x pulchra var. argyroneura Herincq, 1856:98. 

Tydaea x pulchra var. chloroneura Herincq, 1856:98. 

Tydaea xpulchra var. concolor Herincq, 1856:98. 

Tydaea x pyramidalis Regel va~. muhiflora Regel, 1861:193. 

Tydaea x regelii Herincq, 1856:97; Kohleria x heeriana H.E. Moore, 

Tydaea x regelii var. argentea Herincq, 1856:97. 

Tydaea x regelii var. chlorophylla Herincq 185698. 

Tydaea x regelii var. erythrichina Herincq, 1856:98. 

Tydaea x robert-le-diable Van Houtte, 1869:151. 

Tydaea x rossiana Ortgies, 1857: 1; Kohleria x rossiana (Ortgies) H.E. 

Tydaea x rossiana var. concolor Ortgies, 1857:2. 

Tydaea x rossiana var. lucida Ortgies, 1857:2. 

1954:386. 

Moore, 1954:387. 

NOT= ON NOMENCLATURE.-Gesneria oblongata: Paxton 

(1839) probably described this species from hybrid material, 

with Kohleria trianae as part of the parentage of this hybrid. 

Gesneria elongata Kunth var. fruticosa Graham (1840) was 

stated to be a synonym of Gesneria oblongata. The illustration 

of this variety, however, shows a different plant similar to 
NUMBER 79 75 

typical Kohleria trianae, and the variety fruticosa is herein 

treated as a synonym of K. trianae. 

Isoloma bogotense var. giganteum: This hybrid variety is 

probably identical to Tydaea x gigantea Planchon. Apparently 

Voss (1894) incorrectly credited Van Houtte as author of 

Tydaea x gigantea instead of Planchon (1855a); Van Houtte 

never published any Tydaea gigantea. 

Kohleria behnickii: Fritsch (19 16) described K. behnickii 

from cultivation, and added that the plant was known as Tydaea 

decaisniana, but had apparently never been described under 

this name. However, Regel (1858) published this name for a 

hybrid between Achimenes picta (now Kohleria amabilis var. 

bogotensis) and Sciadocalyx warszewiczii (Kohleria warszewiczii). The plant described by Fritsch as Kohleria behnickii 

has according to the description features intermediate between 

these two taxa, and K. behnickii is herein considered to be of 

hybrid origin. This condition is based on the Fritsch description, as no type material has been located. 

Kohleria bella: see Kohleria maculata. 

Kohleria dubia: Fritsch (1916) suggested that K. dubia 

might be a hybrid. Only minor differences distinguish K. dubia 

from Tydaea x gigantea Planchon (1855a), a hybrid between 

Kohleria amabilis var. bogotensis and K. warszewiczii. 

Kohleria maculata: Morton (1938) described K. maculata 

on material collected close to San Jose, Costa Rica, and later 

(1967b) he added the similar K. bella (mainly set apart by 

having larger flowers); the species was based on cultivated 

material originally collected in San Jose. A few additional 

collections from Costa Rica (Davidse & Pohl1310), Honduras 

(Molina R. 14581), Ecuador (Valverde 23) and Colombia 

(Sandeman 639:39) are similar; these latter collections are all 

from gardens. In addition, a hybrid cultivated at Cornell 

University in the 1950s under the name “Sunrise” has 

characteristics similar to K. maculata and especially the 

larger-flowered K. bella (Moore 7689). According to 

Batcheller (1985) “Sunrise” is unknown in origin and now 

believed to be extinct. Consequently, both K. maculata and K. 

bella apparently were described from a neotropical cultivated 

hybrid of unknown origin and parentage. Kohleria warszewiczii is probably part of the parentage. 
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1885. Isoloma. In The IllusfrafedDicfionary of Gardening, 2:201. London: 

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Isonyms and Pseudo-Isonyms: Identical Combinations with the 

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Bitter Phenyl Propanoid Glycosides from Conandron ramondioides. 

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Nicolson, D.H. 

1975. 

Nonaka, G., and I. Nishioka 

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Oersted, AS. 

1858. Centralamericas Gesneraceer, ef Systemafisk, Planfegeografisk 

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Tydaea (hybr.) Rossiana Ortgies; Tydaea (hybr.) Lenneana Ortgies. 

Garfenflora, 6:l-3, plates 181, 182. 

Tydaea (hybr.) Hansfeini Ortgies. Gartenflora, 7:65, plate 218. 

Gesneria oblongata (Oblong-Flowered Gesneria). Paxfon’s MagaOrtgies, E. 

1857. 

1858. 

Paxton, J. 

1839. 

zine ofBotany, 6:103-104. 

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Perez-Arbelaez, E. 

1947. 

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1850. Isoloma picfum: Isoloma a teintes vives. Flore des Serres et des 

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1853. Sciadocalyx warscewiczii [sic]. Flore &s Serres el des Jardins de 

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1855a. Tydaea (hybrid) Giganfea Planchon. Flore &s Serres ef &s Jardins 

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1855b. Isoloma frianaei, Regel. Flore &s Serres ef des Jardins de I’Europe, 

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1856. Tydaea (hybrid) Orfgiesii Hort. Van Houtte. Flore des Serres et des 

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1847. Index seminum in horfo bofanico Turicensi, anno 1847 collecforum, 

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1854c. Isoloma frianaei. Garfenflora, I, 3:107-108, plate 82. 

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1861. Tydaea pyramidalir multifora. Garfenflora, I, 10:193-194. 

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Raymond, M. 

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[not seen]. 

Gartenflora, I, 3:llO-111. 
NUMBER 79 79 

1867. 

1868. 

1954. 

1896. 

1900. 

Neue oder empfehlenswerthe Zierpflanzen. Gartenflora, 11,16383- 

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Tydaea lindeniana Rgl. Gartenflora, II,17:257, plate 589. 

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1898. 

Gesnera rugata Nob. Allgemeine Gartenzeitung, 15:226. 

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373 -3 87. 

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Stebbins, G.L. 

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1856. 

1869. 

Vogel, S. 

1958. 

1O4:49 1-530, 

Voss, A. 

1894. Gattung 858: Isoloma (Kohleria). In A. Sieben and A. Voss, 

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Verlagsbuchhandlung Paul Parey. 

Walpers, G.G. 

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Isoloma jolkcanm. Proceedings of the American Academy of Arts, 

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1973. 

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Die Struktur der paarbliitigen Partialinfloreszenzen der Gesneriaceen 

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1978. Transitions from Pair-flowered to Normal Cymes in Gesneriaceae. 

Notesfrom the Royal Botanic Garden, Edinburgh, 36355-368. 

1982. Evolution and Radiation of the Pair-flowered Cyme in Gesneriaceae. 

Australian Systematic Botany Society Newsletter, 30:23-47. 

Wiehler, H. 

1968. x Koellikohleria, a New Hybrid Genus (Gesneriaceae). Baileya, 

1972 [“1971”]. Chromosome Numbers in Some American Gesneriaceae. 

Baileya, 18: 118-120. 

1973. One Hundred Transfers from Alloplectus and Colmea (Gesneriaceae). Phytologia, 27:309-328. 

1975. The Re-establishment of Moussonia Regel (Gesneriaceae). Selby1976a. A Report on the Classification of Achimenes, Eucodonia, Gloxinia, 

Goyazia, and Anetanthus (Gesneriaceae). Selbyana, 1 :374-404. 

1976b. New Hybrid Genera in the Gesneriaceae. Selbyana, 1:405-409. 

1978a. Parakohleria, a New South American Genus in the Gesneriaceae. 

1978b. Miscellaneous Transfers and New Species of Neotropical Gesneri1983. 

1968. 

16:29-34. 

OM, 1~22-31. 

Selbyana, 5:4-10. 

aceae. Selbyana, 5:61-93. 

A Synopsis of the Neotropical Gesneriaceae. Selbyana, 6:l-219. 

A Beauuful Costa Rican Kohleria. Notes from the Royal Botanic 

Garden, Edinburgh, 28:291-293. 

Pollen Structure and Systematics of the Neotropical Gesneriaceae. 

Selbyana, 2:310-322. 

Williams, L.O. 

Williams, N.H. 

1978. 

Wilson, C.L. 

1974. Floral Anatomy in Gesneriaceae, 11: Gesnerioideae. Bolanical 

Gazette (Crawfordrville), 135:256-268. 
Index to Scientific Names of Genera and Species 

(Currently accepted names, apart from hybrids, are in italics.) 

Achimenes Persoon, 3,33,36,38 

amabilis (Planchon & Linden) Van Houtte, 45 

dulcis C. Morton, 4 

Java C. Morton, 73 

hillii Hortorum ex Regel, 74 

kewensis Hortorum ex Regel, 74 

klotzschii Warscewicz [sic] ex Regel, 55 

magnifica (Planchon & Linden) Lindley, 54 

ocellata W.J. Hooker, 2,53-55 

picta Bentham ex W.J. Hooker, 2,38,45,46,75 

Bellonia Linnaeus, 11 

Bellonieae Fritsch, 3 

Beslerieae Bartling & Wendland, 4 

Brachyloma Hanstein, 2,3, 38 

elongatum (Kunth) Hanstein, 65 

erianthum (Bentham) Hanstein, 50 

hirsutum (Kunth) Hanstein, 2, 38, 50 

hondense (Kunth) Hanstein, 2,52 

incurvum (Bentham) Oersted, 66 

karstenianum Hanstein, 51, 52 

leucomallon Hanstein, 50 

longiflorum (Kunth) Hanstein, 50 

longifolium (Lindley) Oersted, 60 

longipes (Bentham) Hanstein, 51 

moue (Kunth) Hanstein, 50 

moritzianum Bouche & Hanstein, 50.51 

petiolare (Bentham) Oersted, 60 

pictum (W.J. Hooker) Oersted, 66 

pilosum Oersted, 66 

rhodomallon Hanstein, 50, 51 

rhynchocarpum (Bentham) Oersted, 66 

rubricaule (Kunth & Bouche) Hanstein, 50 

strictum Oersted, 66 

trianae (Regel) Hanstein. 65 

tubiflorum (Cavanilles) Oersted, 66 

ventricosum Hanstein, 50.51 

vestitum (Bentham) Hanstein, 50 

Calycostemma Hanstein, 2, 3, 39 

lindenii Hanstein, 2,39,54 

Capanea Decaisne ex Planchon, 36, 37.38 

aftinis Fritsch. 37.72, 73 

grandgora (Kunth) Decaisne ex Planchon. 38.44 

warszewiczii Warszewicz ex Regel, 54 

Colmea Linnaeus. 30 

eriantha (Bentham) Hanstein, 70 

ericae Mansfeld, 30 

guttata Poeppig. 30 

Conandron ramondioides Sieber & Zuccarini, 11 

Cremosperma Bentham, 33 

Crypoloma Hanstein, 2, 3, 38, 52 

anonymum Hanstein, 55 

coloratum Hanstein, 66, 68 

cordifolium Hanstein, 66, 68 

hondense (Kunth) Hanstein, 2, 38.52 

hookerianum (Regel) Hanstein, 66 

pictum (W.J. Hooker) Hanstein, 66 

pilosum (Oersted) Hanstein, 66 

rhynchocarpum (Bentham) Hanstein, 66 

strictum (Oersted) Hanstein, 66 

triflorum (W.J. Hooker) Hanstein, 52 

Cyrtandroideae Jack, 11 

Diastema Bentham, 3,7.8, 11, 13, 33, 36. 37.38. 47,68 

afine Fritsch, 7 

anisophyllum Fritsch, 70 

var. quitense Fritsch, 70 

pictum (Bentham ex W.J. Hooker) Walpers, 45 

platylomatum J.D. Smith, 70 

racemiferum Bentham, 19,44 

vexam H.E. Moore, 36 

villosum Fritsch, 68,70 

Drymnia serrulata (Jacquin) Manius, 4 

Episcieae Endlicher, 4 

Eucodonia Hanstein, 36.38 

Gesneria Linnaeus [Gesnera], 1,2,9,44 

section Isoloma Bentham, 2,38 

auriculata (W.J. Hooker) 0. Kuntze, 4 

breviflora Lindley, 60.63 

elatior Kunth, 74 

elongata Humboldt. 65 

elongata Kunth, 2,3,65 

eriantha Bentham, 2.50.51 

hirsuta Kunth, 2, 50 

hmdensis W.J. Hooker, 2, 66,67,68 

hmdensis Kunth, 2, 52, 66, 67 

ignorata Kunth, 60,63 

inaequalis Bentham, 2, 53,54 

incurva Bentham, 2.66.67 

linkiana Kunth & Bouche,60,63 

longiflora Kunth, 2,50 

longifolia Lindley. 2, 60, 63 

longipes Bentham, 2,5 1.52 

maculata Sesse & Mocino, 59 

maculata Mocino & Sesse ex A.P. de Candolle, 59, 60 

mollis Hortorum ex Kunth, 60.63 

mollis Hortorum ex Lehmann, 60 

mollis Kunth, 2.50, 5 1 

oblongata Paxton. 74 

oblongata Hortorum ex Graham, 65 

petiolaris Bentham, 2, 60, 63 

picta W.J. Hooker. 66 

picta Hortorum ex Hanstein, 66 

regeliana Warszewicz ex Planchon, 70 

rhqnchocarpa Bentham, 2 66,67 

rubricaulis Kunth & Bouche, 2,50 

rugata Scheidweiler, 2,59,60 

schiedeana (A.P. de Candolle) W.J. Hooker, 2 

seemannii W.J. Hooker, 60.63 

spicata Kunth, 2, 60, 62 

var. fruticosa Graham, 65.74.75 

var. minus-hirsuta W.J. Hooker, 66, 67, 68 

80 
NUMBER 79 81 

var. schiedeana A.P. de Candolle, 60,62 

trianaei Herincq, 65.66 

triflora W.J. Hooker, 52 

tubiflora Cavanilles, 1, 66 

velutina Willdenow ex A.P. de Candolle, 65 

ventricosa Hortorum ex Hanstein, 50 

vestita Bentham, 2, 50, 51 

Gesneriaceae Dumortier, 1,2, 3, 4, 6, 8, 11, 44,76 

Giesleria Regel, 2, 38 

picta (Bentham ex W.J. Hooker) Regel, 45 

Gloxinia L’Heritier, 11, 36, 38 

lindenianu (Regel) Fritsch, 73,74 

Gloxinieae Fritsch, 3, 4, 8, 11, 13, 19, 33, 38 

Heppiella Regel, 3.4, 11, 33, 36,38 

Isoloma (Bentham) Decaisne, 2,3,38,50,54, 55, 62,74 

ulmifolia (Kunth) Hanstein, 20 

series Brachyloma (Hanstein) Bentham, 3 

amabile (Planchon & Linden) Hortorum ex Bellair & Saint-eger, 45 

bogotense Nicholson, 45 

var. amabile (Planchon & Linden) Voss, 45 

var. giganteum Voss, 74.75 

var. hansteinii (Ortgies) Voss, 74 

var. hillii (Regel) Voss, 74 

var. lenneanum (Ortgies) Voss, 74 

var. ocellatum (W.J. Hooker) Voss, 54 

var. pulchrum (Herincq) Voss, 74 

var. rutilans (Ongies) Voss, 74 

var. warszewiczii (Regel) Voss, 54 

breviflorum (Lmdley) Lindley, 60 

bulbosum Baillon, 73 

ceciliae (Andre) Nicholson, 45 

ceciliae (Andre) Hortorum ex Bellair & Saint-Leger. 45 

cordifolium (Hanstein) Hemsley, 66 

costaricense (Klotzsch ex Oersted) Hemsley, 73 

deppeanum (Schlechtendal & Chamisso) Hemsley, 73 

digitaliflorum (Lnden & Andre) Voss, 70 

elatior Decaisne, 73, 74 

elegans (Decaisne) Hemsley, 73 

erianthum (Bentham) Decaisne, 50 

flexuosum Rusby. 73 

giganteum Hortorum Bentham ex Fritsch, 74 

hahnii Hortorum ex Baillon. 73 

hirsutum (Kunth) Regel, 50 

hondense (W.J. Hooker) Decaisne, 66 

var. chloranthum Regel, 52 

var. hookeri Regel, 66 

var. uniflorum Regel, 66,68 

var. wageneri Regel, 66.68 

var. warszewiczii Regel, 66.68 

hondense (Kunth) Decaisne, 52 

hypocyrtiflorum (W.J. Hooker) Bentham ex Jackson, 73 

ignoratum (Kunth) Hemsley, 60 

inaequale (Bentham) Bentham ex Jackson, 53 

incanum (Klotzsch & Hanstein) Hemsley, 60 

incurvum (Bentham) Hemsley, 66 

jaliscanum S. Watson, 73 

kramerianum Lehmann, 60,63 

lasianthum (Zuccarini) Decaisne, 73 

lindenianum (Regel) Nicholson, 73 

linkianum (Kunth & Bouche) Hemsley, 60 

longiflorum (Kunth) Regel, 50 

longifolium (Lindley) Decaisne, 60 

longipenduculatum Hortorum ex Hanstein, 50 

longipes Decaisne, 51,52 

x luciani (Ande) Voss, 74 

molle Decaisne, 50 

molle (Kunth) Nicholson, 50 

oblanceolatum J.D. Smith, 73 

ocellatum (W.J. Hooker) Bentham ex Jackson, 54 

x ortgiesii (Honorurn ex Planchon) Voss, 74 

parviflorum Rusby, 73 

patentipilosum 0. Kuntze, 73 

petiolare (Bentham) Decaisne, 60 

pictum (W.J. Hooker) Regel, 66 

pictum (W.J. Hooker) Decaisne ex Planchon, 66 

pilosum (Oersted) Hemsley, 66 

purpureum (Poeppig) Bentham, 73 

pycnosuzygium J.D. Smith, 50 

regelianum (Warscewicz [sic] ex Planchon) Voss, 70 

rhynchocarpum (Bentham) Decaisne, 66 

rubricaule (Kunth & Bouche) Regel, 50 

rupestre (Seemann) Hemsley, 60 

schiedeanum (A.P. de Candolle) Hemsley, 60 

seemannii (W.J. Hooker) Bentham, 60 

spicatum (Kunth) Decaisne. 60 

sprucei Britton, 73 

strictum (Oersted) Hemsley, 66 

tetragonum (Oersted) Hemsley, 60 

trianaei Regel, 65, 66 

triflorum (W.J. Hooker) Decaisne, 52 

tubiflorum (Cavanilles) Decaisne, 66 

tydaea L.H. Bailey, 45 

urticifolium Rusby, 73 

vestitum (Bentham) Decaisne, 50 

vulcanicolum J.D. Smith, 73 

warszewiczii (Regel) L.H. Bailey, 70 

var. minus-hirsutum (W.J. Hooker) Planchon, 66 

Isoloma J.D. Smith, 38 

Koellikeria Regel. 36 

erinoides (A.P. de Candolle) Mansfeld, 36 

Kohleria Regel, 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 13, 14, 15, 16, 19, 20, 21, 

27,28,30,31, 32, 35,36,37,38,39,44, 46, 52, 57.60, 62,66,68 

section Calycostema (Hanstein) Fritsch, 3 

section Cryptoloma (Hanstein) Fritsch, 3 

section Eukohleria Fritsch, 3 

section Isoloma (Bentham) Fritsch, 3 

section Moussonia (Regel) Fritsch, 3 

section Sciadocalyx (Regel) Fritsch. 3 

section Tydaea (Decaisne) Fritsch, 3 

subgenus Brachyloma (Hanstein) Hanstein, 3, 51 

alleniistandley & L.O. Williams, 1, 3, 4, 7, 8, 11, 13, 16, 19, 20, 25, 38, 

anonyma (Hanstein) Hanstein, 66 

amubilis (Planchon & Linden) Fritsch, 1,2, 3.4, 8, 13, 19, 20,30,38,44, 

39,42,43, 44 

45,47,60,72 

var. amubilis, 5, 16,41,45 

var. bogofensir (iVichoIson) Kvist & L. Skog, 2, 9, 13, 14, 16, 38, 41, 

45,46,75 

anisophylla (Fritsch) Wiehler, 3,70 

avilana Cuatrecasas, 73 

baezana Cuatrecasas. 73 

behnickii Fritsch, 74,75 

bella C. Morton, 74, 75 

bogotensis (Nicholson) Fritsch, 8, 45.46 

brachycalyx Fritsch, 8, 50, 51 

chiapensis T.S. Brandegee, 60 

collina T.S. Brandegee, 73 

colorata (Hanstein) Hanstein, 66 
82 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cordlfolia (Hanstein) Hanstein, 66 

deppeana (Schlechtendal & Chamisso) Fritsch, 73 

diosfemoides Kvist & L. Skog, 8, 9, 13, 19, 20, 36, 37, 41, 46, 47 53 

digitalinora (Linden & Andre) Fritsch, 8, 70, 72 

dubia Fritsch, 74,75 

elegans (Decaisne) Loesener, 73 

elongata (Kunth) Hanstein, 65 

elongata Regel, 65,73 

eriantha (Bentham) Hanstein, 50.70 

ferruginea Fritsch, 54, 55 

filisepala Standley. 73 

fruticosa TS. Brandegee, 65, 73 

x gigantea (Planchon) Fritsch, 74 

grandifora Kvist & L. Skog, 4, 8, 9, 19, 20, 30, 41,47, 48, 49, 55 

x hansteinii (Ortgies) H.E. Moore, 74 

x heeriana H.E. Moore, 74 

x hillii (Regel) H.E. Moore, 74 

hirsuta (Kunth) Regel, 1,2,3,4,7, 8, 11,13, 16, 19,20,24,30, 31,32,33, 

var. lasiantha (Zuccarini) Fritsch, 73 

var. pedunculata V.S. Brandegee) C. Morton, 73 

38, 39, 46, 49, 50, 51, 59,62,63, 65, 66, 68, 72 

var. hirsuta, 6,9, 13, 14, 29, 32,33,38,42, 50, 51, 

var. longipes (Bentham) Kvist & L. Skog, 24,29, 32,33, 38, 42, 50, 

51,59,65 

hirsutissima C. Morton, 73 

hondensis (Kunth) Hanstein, 1.2, 3, 4,6, 8, 11, 13, 14, 16, 19,20,23, 30, 

hookeri (Regel) Hanstein, 66 

ignorata (Kunth) Regel, 2, 3, 38, 60, 62, 63 

inaequalk (Bentham) Wiehler, 1, 2, 3, 4, 8, 11, 13, 16, 19,20, 27,31, 32, 

31,38,40, 52, 53, 67 

33,34,35, 36, 37,38, 39, 47, 49,53,55, 63 

var. inaequalis, 5.9, 13.33, 34.36, 37, 42.53.54 

var. lindenii (Hanstein) Kvist & L. Skog, 2, 3, 33, 34, 36, 37, 39, 42, 

var. ocellata (W.J. Hooker) Kvist & L. Skog, 2, 13.33, 34, 36,37,41, 

54 

53,54,55 

incana Klotzsch & Hanstein, 60,63 

jamesoniana Fritsch, 73 

kalbreyeri Fritsch, 55 

karsteniana (Hanstein) Hanstein, 51 

krameriana (Lehmann) Hanstein, 60 

lanata Lemaire, 51, 59, 60 

lanigera Fritsch, 50, 51 

Iehmannii Fritsch, 54 

x lenneana (Ortgies) H.E. Moore, 74 

leucomallos (Hanstein) Hanstein, 50 

lindeniana (Regel) H.E. Moore, 73 

lindenii (Hanstein) Hanstein, 54 

linkiana (Kunth & Bouche) Hanstein. 60,63 

1ongicalyxKvist & L. Skog, 1,4, 8, 11, 13, 16, 19, 20, 42,47,49,55 

longiflora (Kunth) Hanstein, 50 

longifolia (Lindley) Hanstein, 60 

longipedunculata Hanstein, 50,51 

longipes (Bentham) Hanstein, 51 

x luciani (Andre) Fritsch, 74 

magnifica (Planchon & Linden) H.E. Moore, 54 

mollis (Kunth) Hanstein, 50 

maculata C. Morton, 59,74,75 

martensii Fritsch, 73 

moritziana (BouchB & Hanstein) Hanstein, 50 

neglecta Kvist & L. Skog, 1,4,6,9, 11, 13,20,26,41, 56, 57.61 

ocellata (W.J. Hooker) Fritsch, 54 

x ortgiesii (Honorum ex Planchon) H.E. Moore, 74 

var. petiolaris (Bentham) C. Morton, 60 

papill~sa (Oersted ex Hanstein) Fritsch. 73 

var. pendula C. Morton, 73 

var. sericea Fritsch. 73 

var. solitaria C. Morton, 73 

patentipilosa (0. Kuntze) Schumann, 73 

pedunculata TS. Brandegee, 73 

peruviana Fritsch, 1, 3, 4,6,7, 8, 10, 13,20,24, 33, 38, 42, 49,57, 58,59 

var. pallida Fritsch, 57,59 

petiolaris (Bentham) Hanstein, 60 

picta (W.J. Hooker) Hanstein, 45, 66 

pilosa (Oersted) Hanstein, 66 

platylomata (J.D. Smith) Wiehler, 68, 70 

x pulchra (HCrincq) H.E. Moore, 74 

regelii Hanstein, 73 

reticulata Fritsch, 73 

pycnosuzygium (J.D. Smith) BadiUo, 50 

rhodomallos (Hanstein) Hanstein, 50 

x rossiana (Ortgies) H.E. Moore, 74 

rubricaulis (Kunth & Bouche) Hasskarl, 50 

rugata (Scheidweiler) Kvist & L. Skog, 1, 2, 6.7, 8,9, 13, 16.20, 25, 30, 

38,40,44, 51, 59, 60, 63 

rupestris Seemann, 60,63 

rupicola Standley & L.O. Williams, 73 

sancti-josephi Cuatrecasas, 73 

saxicola TS. Brandegee, 73 

scabrida Fritsch, 55 

schiedeana (A.P. de Candolle) Hanstein, 60 

seemannii (W.J. Hooker) Hanstein, 2, 60 

septentrionalis D. Denham, 73 

serrulata C. Morton, 73 

skutchii C. Morton & D. Gibson, 73 

spicata (Kunth) Oersted, 1, 2. 3, 4, 5, 6, 7, 10, 13, 16, 19, 20, 22, 30, 31, 

, 

34,36,38,40, 53, 57, 60, 61,62,63, 65, 67 

var. hispida Fritsch, 60, 63 

var. wageneri (Regel) Fritsch, 60 

sprucei (Britton) Fritsch, 73 

straussiana Fritsch, 50, 51 

stricta (Oersted) Hanstein, 66 

strigosa C. Morton, 74 

stuebeliana Fritsch, 3, 4, 6.7, 8, 13, 20.26, 30, 38, 40, 61, 63, 64, 65 

tetragona Oersted, 60 

tomentosa Hanstein, 60, 63 

trianae (Regel) Hanstein, 1,2.3,6, 7,8, 11, 16,20.29,30,31,32,40,49, 

50, 51,65, 66. 72, 74, 75 

triflora (W.J. Hooker) Regel, 2, 52 

tubifora (Cavanilles) Hanstein, 1, 2, 3, 10, 11, 13, 16, 19, 20, 23, 30,31, 

urticifolia (Rusby) Fritsch, 74 

ventricosa (Hanstein) Hanstein, 50 

vestita (Bentham) Hanstein, 50 

villosa (Fritsch) Wiehler, 1,2,3,4,8, 11, 13, 16, 19,20,30,36,38,44,45, 

38.40.52, 61, 62, 63, 66,67,68 

47, 68 

var. villosa, 3,5, 10, 13, 41, 68, 69,70 

var. anisophylla (Fritsch) Kvist & L. Skog, 3,4,4 1.68, 70 

viminalis TS. Brandegee, 74 

violacea Fritsch, 72 

wageneri Regel, 60 

warszewiczii (Regel) Hanstein, 1, 2, 6, 7, 8, 10, 11, 13, 14, 16, 19, 20, 26, 

weberbaueri Fritsch, 74 

30,31,37 38, 40, 47, 49,51,53,54,66, 70, 72, 75 

Kohlerieae Fritsch, 3 

Lompornk calolaema, 19 

Liefzia Regel, 44 

Locheria magnifica Planchon & Linden, 54 
NUMBER 79 83 

Monopyle Moritz ex Bentham, 8,33,38 

Moursonia Regel, 1,3, 11, 13, 36,73 

macrocarpa Bentham, 19 

deppeana (Schlechtendal & Chamisso) Hanstein, 3,73 

elegam Decaisne, 73,74 

elongata Auctorum, 3 

fruticosa (TS. Brandegee) Wiehler, 73 

hirsutissima (C. Morton) Wiehler, 73 

rupicola (Standley & L.O. Williams) Wiehler, 73 

septentrionalis (D. Denham) Wiehler, 73 

serrulata (C. Morton) Wiehler, 73 

skutchii (C. Morton & D. Gibson) Wiehler, 73 

strigosa (C. Morton) Wiehler, 74 

trtj7ora (Martens & Galeotti) Hanstein, 73 

virninalis (T.S. Brandegee) Wiehler, 74 

Nematanthus erianthus Bentham, 2.70 

Omiltemia filisepala (Standley) C. Morton, 73 

Paliavana Vandelli, 44 

Parakohleria Wiehler, 1, 3, 8, 11, 13, 19, 33, 36, 38 

abunda Wiehler, 19 

awilana (Cuatrecasas) Wiehler, 73 

baezana (Cuatrecasas) Wiehler, 73 

jamesoniana (Fritsch) Wiehler, 73 

parvifora (Rusby) Wiehler, 73,74 

purpurea (Poeppig) Wiehler, 73 

sancti-josephi (Cuatrecasas) Wiehler, 73 

sprucei (Britton) Wiehler, 73 

weberbaueri (Fritsch) Wiehler, 74 

Pearcea Regel, 3 

hypocyrtiflora (W.J. Hooker) Regel. 73 

Reldia Wiehler, 4. 33 

Rhylidophyllwn awiculatwn W.J. Hooker, 4 

Sciadocalyx Regel, 2,3, 38 

digataliflora Linden & Andre, 2.70 

x luciani Andre, 74 

warszewiczii Regel, 2. 38.70, 75 

Sinningka Nees, 11, 33, 38, 73 

incarnata (Aublet) D. Denham, 73 

Sinningiae Fritsch, 3 

Smithiantha 0. Kuntze, 36 

Solenophoreae Fritsch, 3 

Synepileana Baillon. 3, 39 

Tydaea Decaisne, 2, 3.38 

granatensis Baillon, 3, 39, 54 

amabilis Planchon & Linden, 45 

ceciliae Andre, 45 

x decaisniana Regel, 74 

decaisniana Hortorum ex Fritsch, 74.75 

elegans (Decaisne) Planchon & Linden, 74 

gigantea Van Houtte ex Voss, 74 

x gigantea Planchon, 74,75 

grandiflora Hortorum Behnick ex Fritsch, 74 

x hansteinii Ortgies, 74 

hillii Regel, 74 

X hybnda Honorum ex H.E. Moore, 74 

karsteniana Hortorum ex Hanstein, 74 

x lenneana Ortgies, 74 

var. viridis Ortgies, 74 

var. rutilans Ongies, 74 

hdeniana Regel, 74 

lindenii Regel ex Andre, 74 

magnifica (Planchon & Linden) Hanstein, 54 

x meyendorffii Regel, 74 

var. rubra Regel, 74 

ocellata (W.J. Hooker) Regel, 2, 54 

var. confluens Regel, 54 

var. fonnosa Regel, 54 

var. picta Regel, 55 

x ortgiesii Honorum Van Houtte ex Planchon, 74 

pardina Linden & Andre, 55 

picta (Bentham ex W.J. Hooker) Decaisne, 2, 38, 45, 74 

x pulchra Herincq, 74 

var. karsteniana Hortorum ex Hanstein, 74 

var. argyroneura Herincq, 74 

var. chIoroneura Herincq, 74 

var. concolor Herincq, 74 

var. multiflora Regel, 74 

var. argentea Herincq, 74 

var. chlorophylla Herincq, 74 

var. erythrichina Herincq, 74 

x robert-le-diable Van Houtte, 74 

x rossiana Ortgies, 74 

var. concolor Ortgies. 74 

var. lucida Ortgies, 74 

x pyramidalis Regel, 74 

x regelii Herincq, 74 

venosa Lescuyer, 45 

warszewiczii Regel, 54 

Vanhoutrea Lemaire. 36, 38