SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 84 

Revision of Pearcea 

(Gesneriaceae) 

Lars Peter Kvist 

and Laurence E. Skog 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1996 
ABSTRACT 

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

Contributions to Botany, number 84,47 pages, 22 figures, 1 table, 1996.-Seventeen species of 

Pearcea (Gesneriaceae) are recognized, nine of which are new: Pearcea bella, P. bilabiata, P. 

cordata, P. fuscicalyx, P. glabrata, P. gracilis, P. grandijolia, P. intermedia, and P. strigosa. Of 

the remaining eight species, two were originally described in Pearcea, one is transferred herein 

from Kohleria, and five come from Parakohleria, a genus placed in synonymy herein. Pearcea 

occurs mainly on the eastern Andean slopes of Colombia (5 species), Ecuador (12 species), Peru 

(6 species), and Bolivia (1 species), and one species also occurs on the western Andean slopes 

in Ecuador. The largest number of species appears to occur in Ecuador, although additional 

species may occur in less explored parts of Peru. Pearcea differs from related and similar genera 

in the Gloxinieae by having fleshy capsules dehiscing by two valves and exposing a sticky seed 

mass. All species are herbs, usually with bright red flowers, and are usually found in the shady, 

humid forest understory, often near small streams. Hybridization, local speciation, and 

polymorphic variation may complicate the species delimitation. The variable Pearcea sprucei 

occurs throughout nearly the entire range of the genus (and includes the only accepted variety, 

var. parvzji’ora), but most species are rare and local. 

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 Pearcea (Gesneriaceae) / Lars Peter Kvist and Laurence E. Skog. 

p. crn. - (Srnithsonian contributions to botany ; no. 84) 

Includes bibliographical references (p. 46) and index. 

1. Pearcea-Classification. 1. Skog, Laurence E. 11. Title. 111. Series. 

QKl.S2747 no. 84 [QK495.G4] 581 s-dc20 [583’.81] 96-28456 

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

National Standard for Permanence of Paper for Printed Library Materials 239-48-1984. 
Contents 

Page 

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

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

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

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

Distinguishing Characters ................................ 4 

Geographical and Altitudinal Distribution ....................... 7 

Variation ......................................... 15 

Polymorphic Variation ............................... 16 

Introgression .................................... 16 

Speciation ...................................... 16 

Evolution ......................................... 16 

Pearcea Regel ...................................... 17 

Key to the Species and Varieties of Pearcea Regel ................. 18 

1 . Pearcea ahunda (Wiehler) L.P. Kvist & L.E. Skog. new combination ..... 20 

2 . Pearcea bella L.P. Kvist & L.E. Skog. new species ............... 20 

3 . Pearcea bilabiata L.P. Kvist & L.E. Skog. new species ............. 22 

4 . Pearcea cordata L.P. Kvist & L.E. Skog. new species ............. 22 

5 . Pearcea fuscicalyx L.P. Kvist & L.E. Skog. new species ............ 25 

6 . Pearcea glabrata L.P. Kvist & L.E. Skog. new species ............. 25 

7 . Pearcea gracilis L.P. Kvist & L.E. Skog. new species ............. 27 

8 . Pearcea grandijolia L.P. Kvist & L.E. Skog. new species ........... 29 

9 . Pearcea hispidissima (Wiehler) L.P. Kvist & L.E. Skog. new combination . . 29 

10 . Pearcea hypocyrtiflora (J.D. Hooker) Regel ................... 31 

11 . Pearcea intermedia L.P. Kvist & L.E. Skog. new species ............ 31 

12 . Pearcea purpurea (Poeppig) L.P. Kvist & L.E. Skog. new combination .... 32 

13 . Pearcea reticulata (Fritsch) L.P. Kvist & L.E. Skog. new combination .... 33 

14 . Pearcea rhodotricha (Cuatrecasas) L.P. Kvist & L.E. Skog. 

new combination ................................. 35 

15 . Pearcea schimpjii Mansfeld ........................... 35 

16 . Pearcea sprucei (Britton) L.P. Kvist & L.E. Skog. new combination ...... 37 

16a . Pearcea sprucei var . parviflora (Rusby) L.P. Kvist & L.E. Skog. 

new status .................................. 39 

16b . Pearcea sprucei var . sprucei ........................ 40 

17 . Pearcea strigosa L.P. Kvist & L.E. Skog. new species ............. 40 

Appendix 1: Numerical List of Taxa .......................... 43 

Appendix 2: List of Exsiccatae ............................. 44 

Literature Cited ..................................... 46 

Index to Scientific Names ................................ 47 

Ecology ......................................... 4 

iii 

Revision of Pearcea 

(Gesneriaceae) 

Lars Peter Kvist 

and Laurence E. Skog 

Introduction 

During our research in preparation of the revision of 

Kohleria Regel (Kvist and Skog, 1992), we began to study a 

group of species that until recently was placed in Kohleria. We 

concluded that these plants did not belong in that genus and 

postponed our revision of them until we had finished the study 

of Kohleria. The result of this revision is presented herein. We 

expand the genus Pearcea Regel from only two species to 17 

species, including the ones previously referred to Kohleria (and 

to Parakohleria by Wiehler in 1978). 

ACKNOWLEDGMENTS.-we greatly appreciate the assistance 

of Ximena Londono during field work and for her collections in 

Colombia; Alice Tangerini, who prepared the excellent 

illustrations; Leslie Brothers, who helped in manuscript and 

plate preparation; Linda Trimmer, George Mason University 

Biology Department, for scanning electron micrographs; 

Marjorie Prochaska for help in proofreading; and the herbaria 

listed below for the loan of their specimens for study and/or 

help and encouragement during herbaria visits. 

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 

Suzanne Liebers Erickson Danish Exchange Fund of the 

Smithsonian Institution, a Smithsonian Short-term Visitor 

Award, as well as a Redoctoral Fellowship that allowed him to 

undertake research at various times at the National Museum of 

Natural History, Smithsonian Insitution. The second author 

acknowledges the support of the National Capital Area Chapter 

Lars Peter Kvist, Forestry Unit, Royal Veterinary and Agricultural 

University. Thontaldsensvej 57 DK-1877 Frederiksberg C. (Copenhagen), Denmark. Laurence E. Skog, Department of Botany, NHB166, Smithsonian Institution, Washington, D.C. 20560-OOOI. 

Reviewers: Susanne Renner, Department of Biology, University of 

Missouri at St. Louis, St. Louis, Missouri, 63211; James F. Smith, 

Department of Biology, Boise State University, Boise, Idnho, 83725. 

of the American Gloxinia and Gesneriad Society and the 

Research Opportunity Fund of the National Museum of Natural 

History, Smithsonian Institution, which provided travel funds 

to visit herbaria and to undertake field work in South America. 

Finally, we appreciate the comments of the outside reviewers, Susanne Renner and James F. Smith, which added 

immeasurably to the paper; however, the responsibility for the 

opinions expressed and any errors remain with the authors. 

MATERIALS AND METHODS.-The revision is based on 

studies of herbarium material (451 collections from the 

following herbaria: A, AAU, AMAZ, BH, BM, BUS, C, CGE, CM, COL. 

MANCH, MO, MOL. MPU, NA, NY, P, PH, QCA, QCNE, S, SEL, TRT. 

TULV, U, UC, US, USM, VEN, W, WAG, WU. WIS) and living plants 

collected from the wild and grown in the greenhouses of the 

Department of Botany, National Museum of Natural History, 

Smithsonian Institution. In addition, the following nine species 

were studied in their natural habitats in Colombia, Ecuador, and 

Peru: Pearcea abunda, P. bella, P. hispidissima, P. intermedia, 

P. purpurea, P. reticulata, P. schimpfii, P. sprucei, and P. 

strigosa. 

DAV, E, ECON, F, FI, FI-W, G, GB, GH, HBG, IBE, K, L, LAM, M, MA, 

Taxonomic History 

Pearcea was established by Regel (1867) based on P. 

hypocyrtiflora, described earlier the same year as Gloxinia 

hypocyrtiflora by J.D. Hooker. Regel named the genus in honor 

of Richard William Pearce (-1835-1868), who collected 

plants in South America for the horticultural firm of James 

Veitch and Sons. The Veitch company supplied plants to the 

Royal Botanic Gardens, Kew, from which the drawing and 

description of the original species were made. It was not until 

1936 that a second species, P. schimpfii, was added to the genus 

by Mansfeld. Both of these species from Ecuador have peculiar 

basally inflated corollas with a narrowed throat. This urceolate 

corolla shape was considered by earlier authors as the main 

character distinguishing Pearcea from other genera in the tribe 

Gloxinieae Fritsch. Pearcea bellu, which is newly described in 

1 
2 

this work, also has an urceolate corolla. In addition, we herein 

expand Pearcea with 14 other species having funnelform, 

ventricose, and bilabiate corollas, and we propose the main 

uniting and distinguishing character for the genus to be the 

unusual fruit dehiscence. The rest of this section deals with the 

taxonomic history of these 14 species and with the rationale for 

referring them to Pearcea. We initially studied this material, 

and became increasingly aware of its affinities with Pearcea, 

during our revision of Kohleria (Kvist and Skog, 1992). 

The funnelform-flowered Pearcea species formerly were 

members of other genera. The first two of these species were 

described from Peru as Rhytidophyllum purpureum and R. 

melastoma (both now Pearcea purpurea) by Poeppig in 1840. 

Later Hanstein ( 1865) and Bentham ( 1876) transferred them to 

Moussonia Regel and Isoloma (Bentham) Decaisne, respectively. The next two of the funnelform-flowered species were 

described from Bolivia in 1896 as Isoloma sprucei (now 

Pearcea sprucei) by Britton and Isoloma parviflorum (now P. 

sprucei var. parviflora) by Rusby. Three more species were 

added to Isoloma by Kuntze (1898) and Rusby (1900), but 

these now are all synonyms of P. sprucei var. parviflora. 

Isoloma (Bentham) Decaisne (1848, not Isoloma J. Smith 

(1841)) is a synonym of Kohleria that was mainly used by 

British workers in the 19th Century. Several other synonyms of 

Kohleria were in common use, and the Central American genus 

Moussonia (Regel, 1847, 1848) also was confused with 

Kohleria. Numerous species were added to all of these genera, 

but from near the turn of the century (e.g., Fritsch, 1893-1894), 

the whole group usually was treated as Kohleria sensu lato, 

which then included nearly 100 species names and several 

sectional names. New (Pearcea) species were referred to 

Kohleria (and a few to Diastema Bentham), but they were not 

recognized as a subgroup in Kohleria s.1. In 1913 Fritsch 

described Kohleria reticulata and K. jamesonianu from 

Ecuador (now Pearcea reticulata and P. sprucei, respectively), 

Kohleria weberbaueri from Peru (now Pearcea purpurea), and 

Diastema galeopsis from Bolivia (now Pearcea sprucei var. 

parviflora). In 1935 Cuatrecasas added three Ecuadorian 

species to Kohleria (all now synonyms of Pearcea sprucei) as 

well as Diastema rhodotrichum (now Pearcea rhodotricha). 

The complex and confusing taxonomic history of Kohleria 

s.1. has already been discussed (Kvist and Skog, 1992). Here it 

is sufficient to repeat that Wiehler (1975) realized that Kohleria 

s.1. consisted of three natural groups: Moussonia, restricted to 

Mexico and Central America, Kohleria sensu stricto, which 

ranges from Mexico to Peru and the Guianas with Colombia 

being the country having the highest number of taxa (Kvist and 

Skog, 1992), and a third group ranging along the eastern 

Andean slopes of Ecuador and Peru. In 1978 Wiehler described 

this third group as the new genus Parakohleria, with 14 

species. Three species were new, and the others were 

transferred from Kohleria (6), Moussonia (2), Diastema (2), 

and Pearcea (1). It is remarkable that Wiehler transferred 

Pearcea schimpfii, described by Mansfeld (1936), to ParSMITHSONIAN CONTRIBUTIONS TO BOTANY 

akohleria in spite of the urceolate corollas that supposedly 

characterize Pearcea. We find no major floral differences 

between the two “old” Pearcea species, P. hypocyrtiflora (J.D. 

Hooker) Regel and P. schimpfii, although they can be separated 

from each other by leaves of different shapes and patterns of 

coloration and venation. 

In a later publication, Wiehler (1 983) gave some characters 

delimiting the monotypic genus Pearcea from Parakohleria 

and including but not mentioning Pearcea schimpfii. He argued 

that Pearcea (i.e., P. hypocyrtiflora) differs by having (1) 

epedunculate 1-2 flowered inflorescences, (2) bilobed stigmas, 

and (3) an inability to hybridize with Parakohleria. The first 

argument does not hold because two of our new species, the 

urceolate-flowered Pearcea bella and the funnelform-flowered 

P. fuscicalyx, always have epedunculate inflorescences. Some 

specimens of the variable P. sprucei are, as well, epedunculate, 

and plants of P. hypocyrtijlora occasionally have up to 5 mm 

long peduncles. The second argument is doubtful because the 

stigmas are only weakly bilobed in P. hypocyrtijlora (also the 

case in P. schimpfii) and are more similar to the usual capitate 

stigmas of Parakohleria than to the bilobed stigmas of 

Kohleria. The third argument is not documented with hybridization data available to us. It is not known if anyone has tried 

to cross Pearcea schimpfii with any other species. We 

concluded therefore (Kvist and Skog, 1992) that the species 

with strongly inflated corollas heretofore placed in Parakohleria and Pearcea belong to Pearcea, and then we took a 

closer look at the funnelform-flowered species placed in 

Parakohleria. 

Wiehler (1978, 1983) segregated Parakohleria from Kohleria based on the following characters: a capitate rather than a 

bilobed stigma, the usual absence of scaly rhizomes, the basally 

more strongly bent corolla tube, a more congested inflorescence, and the five free floral nectary glands. Only the stigma 

seemed to afford a clear distinction, but while doing field work 

we discovered that the fruits also are distinctive. Kohleria 

species have either dry capsules with wind-dispersed seeds that 

split from the apex with two valves (Figure Ic) or fleshy 

capsules that split with one slit from the apex to the base 

(Figure ld,e). Parakohleria (= Pearcea) species have fleshy 

capsules that split with two valves from the apex to the base 

(Figure la). We concluded that the segregation of Parakohleria from Kohleria was justified, but we became aware that the 

funnelform-flowered Parakohleria species were not well 

delimited from the Pearcea species with urceolate flowers. 

The discovery of Pearcea intermedia strengthened the 

affinity of Parakohleria with Pearcea. This species has the 

funnelform corollas (Figure 20g) typical of Parakohleria and 

has upper leaf surfaces with a characteristic pattern of green 

blotches surrounded by yellow green areas along the primary, 

secondary, and reticulate tertiary veins. The urceolate-flowered 

Pearcea schimpfii (Figure 20) has leaves with a pattern so 

similar that it is impossible to distinguish the two species 

vegetatively, but their leaves set them apart from all other 
NUMBER 84 3 

FIGURE 1.-Capsule dehiscence in Peurceu and in some other genera in tribe Gloxinieae: u, Peurceu ubunda, 2 

fleshy valves that split from apex to base and reflex completely; b, Diusremu rucemiferum Bentham, 2 apical 

valves with 1 of 2 slits continuing to base, c, Kohleriu ullenii Standley & L.O. Williams, 2 apical valves; d, 

Kohleriu longiculyx L.P. Kvist & L.E. Skog, 1 longitudinal slit from apex to base, e, Kohleriu inaequulis 

(Bentham) Wiehler, 1 longitudinal slit from apex to base;f, Monopyle mucrocurpu Bentham, 1 longitudinal slit 

from middle of capsule toward apex and base. u from Kvisr 60451 (A AU); b from Steyermurk & Rabe 97441 (US); 

c from Liesner 2106 (US); 153 (AAU);e from Londono & Kvisr 190 (AAU); f from d from Lundonio & Kvisr 153 (AAU);e from Londono & Kvisr 190 (AAU); f from 

Churchill er ul. 4735 (US). 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

species. The trichomes of both species also consist of unusually 

large, flattened, yellow green cells (Figure 206). Similar 

trichomes, although with smaller cells, cover the two other 

urceolate-flowered species, P. hypocyrtijlora and P. belfa 

(Figure 13). and the funnelform-flowered P. fuscicafyx 

(Figure 15), P. gfabrata (Figure 16), and P. rhodotricha. 

The affinity also is strengthened by the fact that two other 

species have intermediate corollas. Pearcea befla (Figure 13) 

has urceolate corollas, but they are not as inflated as in the two 

original Pearcea species, P. hypocyrtijlora and P. schimpjk’ 

(Figure 20). and they are not strikingly different from the 

ventricose corollas of P. abunda (see photo in Kvist, 1991:22). 

In addition, the same range of corolla variation can be found in 

the distantly related New World Gesneriaceae genera Gasteranthus Bentham, Drymonia Martius, and Alfopfectus Martius 

(Kvist, 1991). As a result, the merger of Parakohferia with 

Pearcea does not result in an unusually variable genus. 

Pearcea apparently differs from Kohferia by having 11 

rather than 13 haploid chromosomes. Wiehler (1 978) wrote that 

the haploid numbers of Kohleria and Parakohferia both are 13 

whereas n = 11 in Moussonia, but no documentation for counts 

on Parakohleria was provided. During the past few years, 

Royce Oliver, cytologist in the Department of Botany, 

Smithsonian Institution, repeatedly tried to count the chromosomes of P. abunda. It proved exceedingly difficult because the 

chromosomes are small. Twice he succeeded and found 11 

chromosome pairs. This is interesting because this number can 

set Pearcea apart from Kohferia and Diastema (Table l), and 

it is unexpected because all other South American genera of the 

Gloxinieae that have been examined have a haploid number of 

13 (except for a few tetraploids with 26). Smaller numbers 

(10-12) have been reported only from Central American/ 

Mexican genera (Skog, 1984). Additional counts from Pearcea 

are needed to substantiate or refute the base number of 11 for 

the genus. 

Distinguishing Characters 

In Table 1 some features of Pearcea are compared with those 

of nine other genera in the Gloxinieae in order to discuss their 

differences and affhities. Gloxinieae currently includes 23 

genera (excluding Parakohleria) and approximately 200 

species. 

The following suite of characters distinguishes Pearcea from 

related and similar genera. The plants are terrestrial herbs, 

usually with a conspicuous indumentum, occasionally with 

stolons but usually without scaly rhizomes. The flowering 

shoots have frondose subtending leaves supporting congested 

cymes that are usually pedunculate, but some are epedunculate 

and some have only one or two flowers (Figure 2a,d-f). The 

corollas are bent downward at the base, usually red (sometimes 

orange or yellow but never white), and 0.9-2.7 cm long. The 

nectariferous disc consists of five (or rarely four) free glands. 

The stigma is capitate, stomatomorphic, or only weakly 

bilobed, and the fruit is a fleshy capsule, up to 8 mm long, 

dehiscing apically with two reflexing valves disclosing the seed 

mass. 

The urceolate-flowered Pearcea species cannot be confused 

with species of the other genera in the Gloxinieae, although the 

funnelform-flowered species occasionally may be confused 

with Kohleria and Diastema (Table 1). In the absence of fruits, 

the conspicuously bilobed stigma sets species of Kohferia 

apart, and most species of Kohferia also have considerably 

longer tubular corollas. Diastema species mostly have white or 

whitish flowers, which are never found in Pearcea (Table 1). 

and the plants are low-growing herbs. 

The inflorescences of Pearcea also distinguish the genus 

from Diastema, as well as from some small-flowered Kohferia 

species. The inflorescence morphology of Kohleria and related 

genera was discussed by Kvist and Skog (1992) (Figure 2). 

Diastema has flowering shoots with single flowers (probably 

much reduced cymes) supported by subtending leaves reduced 

to bracts (Figure 2c), whereas the subtending leaves that 

support the cymes of Pearcea are never reduced to bracts 

(Figure 2u). Two small-flowered Kohferia species (K. 

spicata (Kunth) Oersted and K. stuebefiana Fritsch) that are 

sympatric with Pearcea in Ecuador and possibly in Colombia 

have flowering shoots in which the subtending leaves gradually 

reduce to bracts toward the apex (Figure 2b, frondose-bracteose 

following Troll, 1964). A similar inflorescence with small red 

flowers is found in Gloxinia sylvatica (Kunth) Wiehler, which 

is sympatric with Pearcea species in Peru and Bolivia. 

The cymes of Pearcea are similar to those described by 

Weber (e.g., 1973) as “pair flowered cymes,” in which regular 

flowers are accompanied by an additional bractless flower in 

frontal-median position, and are similar to those found in many 

Kohferia species (Kvist and Skog, 1992) (Figure 2d-f). The 

cymes of Kohferia, however, never develop more than 10 

flowers (Figure 25 seven regular flowers and three bractless 

flowers), whereas no fixed maximum seems to exist for 

Pearcea. The highest numbers that have been counted are 

around 20 in P. reticufata (Figure 19). Only two species, P. 

befla (Figure 13) and P. fuscicafyx (Figure 1% apparently 

always have epedunculate inflorescences, and only the first 

consistently has solitary flowers in the leaf axils. 

Ecology 

Many terrestrial New World Gesneriaceae grow in fairly 

bright and open places, such as tree-fall gaps or along trails and 

in other disturbed places. Most disappear when the forest is cut 

completely, but some, e.g., many species of Kohferia (Kvist 

and Skog, 1992), Heppieffu Regel (Kvist, 1990). and Gloxinia 

L’Hbritier (pers. obs.), appear to thrive in deforested, degraded 

areas. Species of Pearcea are forest plants that can grow in 

fairly well-lit places, but many also are found in deep shade, 

e.g., in ravines along small streams. They share these dim, 

humid, moss-covered slopes with other genera of Gesneriaceae, 
NUMBER 84 5 

b 

le 

P If 

FIGURE 2.--lnflorescences of Pearcea, Kohleria, and Diasremu: a, flowering shoot with single flowers supported 

by frondose subtending leaves; b, flowering shoot with single flowers supported by frondose-bracteose 

subtending leaves gradually reduced to bracts (some species of Kohleria); c, flowering shoot with single flowers 

supported by bracteose subtending leaves reduced to bracts (Diasremu); d, cyme with 3 regular flowers and 1 

front-flower; e, cyme with 5 regular flowers and 1 front-flower; f, cyme with 7 regular flowers and 3 

front-flowers. (ad-ffound in both Pearcea and Kohleria.) 

such as Gasteranthus, Napeanthus Gardner, Diastema, Monopyle Moritz ex Bentham, and even a few species of Kohleria 

(Kvist and Skog, 1992), but with few other flowering plants 

(Kvist, 1991). There are occasional reports on herbarium labels 

that specimens of Pearcea were epiphytic. None of the species, 

however, seem to be true epiphytes, but they may grow in moss 

cushions covering tree trunks in montane forests. 

Nearly all exposed-habitat New World Gesneriaceae belong 

to the same tribe as Pearcea, the Gloxinieae. Plants of the 

genera in the Gloxinieae mostly possess scaly rhizomes (Kvist 

and Skog, 1992, fig. 9) or less commonly tubers (Sinningia 

spp.), enabling them to survive seasonal dry periods to resprout 

from the rhizomes or tubers when conditions improve. 

Considering that species of Pearcea usually grow in permanently humid places, it is no surprise that scaly rhizomes are 

absent. We have never seen them on plants in their native 

habitats, but occasionally Pearcea abunda and P. rhodotricha 

develop small scaly rhizomes in cultivation. Many or all 

species develop stolons in their natural habitats and thereby 

often grow in small clonal groups. 

The most weedy of the Pearcea species seems to be P. 

reticdata, which grows to 1.5 m tall and is common in the Rio 

Pastaza valley of Ecuador. There the species generally grows in 

disturbed, secondary shrub forests but not in areas that are 

completely exposed. Some smaller plants also may grow in 

shade. Pearcea reticulata lacks scaly rhizomes, but on many of 

its underground stems we found red, round “bulbs” up to 2 cm 

in diameter. Dissection indicated these to be galls. 

Pearcea species, like many other New World Gesneriaceae, 

are probably hummingbird pollinated, because the conspicuous 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Character 

Life-form 

lblTeShial 

Epiphytic 

Climbing 

Understory 

Exposed 

Scaly rhizomes 

Absent 

Present 

Frondose 

Bracteose 

Flowers in cymes 

Maximum > 2 

Maximum 1-2 

Red 

white 

Different 

Filaments/anthers 

Coiling/coherent 

Not coiling/free 

Anthers/stigma 

Included 

Excluded 

Habitat 

Flowering shoots 

Corolla color 

Nectary 

4-5 free glands 

Annular 

Absent 

Capitate 

Weakly bilobed 

Bilobed 

Fleshy, 2-valved 

Fleshy-dry, 1 slit 

Dry, 2-valved 

Dry, 4-valved 

Globose 

Ellipsoid 

n=ll 

n=13 

Stigma 

Capsule 

Seeds 

Chromosome number 

Number of species 

TABLE 1 .-Comparison between Pearcea and nine other genera in the Gloxinieae. The frequency of characters 

is indicated as follows: ++++ = Always; +++ = Mostly; ++ = Frequently; + = Occasionally; (+) =Rarely; 

? = Possibly or unknown; - = Never. 

Pearcea Achimenes Capanea Diastema Gloxinia Heppiella Koellikeria Kohleria Monopyle Moussonia 

++++ 

? 

- 

+++ 

+ 

++++ - 

++++ - 

+++ 

+ 

+++ 

- 

(+) 

++++ - 

++++ 

(+) 

++++ - 

- 

+++ 

+ 

- 

++++ - 

- 

- 

+++ 

+ 

++++ 

? 

17 

++++ - 

- 

+ 

+++ 

- 

++++ 

++++ - 

- 

++++ 

? 

? 

+++ 

++++ - 

++++ - 

- 

++++ - 

++ 

+++ 

+ 

- 

- 

++++ - 

- 

++++ 

++++ 

- 

22 

+ 

+++ 

+++ 

(+) 

+++ 

++++ - 

++++ 

- 

++++ - 

(+) 

- 

+++ 

++++ - 

- 

++++ 

++++ - 

- 

++++ - 

- 

- 

- 

- 

++++ 

- 

++++ 

- 

++++ 

2 

++++ 

- 

- 

+++ 

(+) 

? 

++++ 

- 

++++ 

- 

++++ 

(+) 

+++ - 

++++ 

- 

++++ 

- 

++++ - 

- 

- 

++++ - 

- 

++++ 

- 

- 

++++ 

- 

- 

++++ 

-5 

++++ - 

- 

++ 

++ 

? 

++++ 

+++ 

+ 

- 

++++ 

++ 

++ 

+++ 

++++ - 

++++ - 

- 

+++ 

++ 

++++ - 

- 

- 

+ 

+++ - 

+ 

+++ 

- 

++++ 

-12 

+++ 

+ 

++ 

- 

++++ 

++ 

+++ 

++++ - 

+++ 

+ 

++++ - 

- 

- 

++++ 

++ 

+++ 

- 

++++ - 

++++ 

- 

- 

- 

- 

+++ 

++ 

- 

++++ 

- 

++++ 

4 

++++ 

- 

- 

? 

+++ 

? 

++++ 

- 

++++ 

- 

++++ 

- 

++++ - 

++++ - 

++++ - 

- 

++++ - 

++++ - 

- 

- 

- 

++++ - 

- 

++++ 

- 

++++ 

1 

++++ - 

- 

++ 

+++ 

++ 

+++ 

+++ 

+ 

+++ 

+ 

++++ 

- 

- 

++++ - 

++++ - 

+++ 

+ 

- 

- 

- 

++++ 

- 

++ 

+++ - 

++ 

+++ 

- 

++++ 

17 

++++ - 

- 

+++ 

(+I 

? 

++++ 

- 

++++ 

+++ 

++ 

- 

++ 

+++ 

++++ - 

++++ - 

- 

- 

++++ 

++++ 

- 

- 

- 

++++ - 

- 

++++ - 

- 

++++ 

-5 

++++ 

? 

? 

+++ 

- 

++++ - 

++++ - 

++++ 

? 

++++ - 

- 

++++ 

- 

++++ 

- 

- 

++++ 

- 

++++ 

- 

- 

- 

- 

++++ - 

? 

+++ 

++++ - 

-10 

and usually red flowers conform to the hummingbird pollina- were observed in southern Ecuador, but although hummingtion-syndrome (e.g., see Howe and Westley, 1988:llO-114). birds visited neighboring plants of other Gesneriaceae, such as 

Unfortunately, no reports of successful observations of species of Besferia Linnaeus and Drymonia, they were never 

pollination have been found. Populations of Pearcea sprucei seen to visit Pearcea. 
NUMBER 84 7 

The flowers are protandrous and, after pollen is shed, the 

filaments coil and drag the decaying stamens back into the 

flower while the style elongates (see photo of Pearcea 

reticulata in Kvist, 1991:16). The same phenomenon occurs in 

flowers of Kohleria but not in Heppiella (Table 1). Another 

similarity with Kohleria, and difference from Heppiella, is the 

presence of tiny glandular trichomes on the limb, usually in the 

throat (particularly where the anthers are located while they 

shed the pollen), and often on the filaments and style. The 

function of the glandular trichomes is unknown, but according 

to Steiner (1985). similar trichomes in Drymonia serrulata 

(Jacquin) Martius exude substances that help the pollen to 

adhere to bees visiting the flowers. The trichomes in the 

corollas of Pearcea, however, probably serve a different but as 

yet unknown function. 

Apart from the species with urceolate (3), ventricose (2), and 

bilabiate (1) corollas, the species of Pearcea have fairly similar 

flowers; the color patterns and indumentum may vary, but the 

sizes and shapes are not that different. The corollas are 

probably not adapted to different specialized pollinators; if the 

primary pollinators are hummingbirds, the same birds can be 

expected to visit flowers of various Pearcea species because 

they are attracted by colors and shapes rather than by scents. 

Until recently it was believed that Pearcea had capsules with 

wind-dispersed seeds as are found in exposed-habitat species of 

Kohleria, Heppiella, Gloxinia, Sinningia, and others. These 

capsules dehisce only from the apex into two valves (Figure lc) 

and persist in this stage during few to several weeks or months 

while they dry out and the seeds are blown away. But as wind 

dispersal of seeds does not work well in the calm forest 

understory, we suspect there may be some other dispersal 

mechanism operating in Pearcea. In fact, field studies showed 

that Pearcea fruits instead split from the apex all the way to the 

base, with two fleshy valves reflexing to expose a glutinous 

seed mass (Figure la), which is quite different from the 

exposed-habitat species of the Gloxinieae. No seed dispersers 

were observed during daylight hours, but the glutinous seed 

masses disappeared overnight (perhaps eaten by nocturnal 

invertebrates). Within a day the degrading capsules hang slack 

below the inflorescences (P. abunda, P. sprucei, and P. 

reticulata, see photo of the latter in Kvist, 1991:19). Considering that the length of the ovoid capsules rarely exceeds 7 mm, 

it is no surprise that this dehiscence was neither detected from 

herbarium sheets nor reported earlier in the literature. Even the 

similarly opening but longer (to several cm) fleshy capsules of 

the distantly related genera Alloplectus and Drymonia were 

long overlooked. 

Seeds of Pearcea (Figure 3a-c; see Beaufort-Murphy, 1983, 

pl. 5 1 b,c) are subglobose or irregular. The understory species of 

Kohleria with unilaterally splitting capsules have similar seeds, 

whereas the exposed-habitat wind-dispersed Kohleria species 

have narrowly ellipsoid seeds with longitudinal, transverse, 

rather regular striations (Figure 3d; see Kvist and Skog, 1992, 

figs. 5, 6). There seems to be a general correlation in the 

Gloxinieae between irregularly subglobose seeds and understory habitats (e.g., in plants of Diastema and Monopyle) and 

between regularly ellipsoid seeds, exposed habitats, and wind 

dispersal (Table 1; Kvist, 1991; Kvist and Skog, 1992). 

From a field trip in 1987, we brought material of at least 

seven Pearcea species into cultivation. They were not difficult 

to keep alive; small cuttings planted in wet sphagnum moss 

maintained in plastic bags survived but often hardly grew for 

months or years, and in their native habitats plants probably 

also survive until conditions improve. Pearcea abunda, P. 

bella, and P. sprucei eventually flowered in cultivation but 

none developed fruits, and this may indicate that they, like 

Kohleria, require cross-pollination, Pearcea will probably 

continue in cultivation as a specialty for growers with a 

particular interest in unusual Gesneriaceae. Plants of Pearcea 

are attractive pot-plants, but sunny windows are too different 

from dark rainforest ravines; the plants will not thrive unless 

they are protected from strong light and dry air. The most hardy 

of the species in cultivation seem to be P. abunda and P. 

rhodotricha. 

Pearcea also seems to have been ignored in its native range. 

No vernacular names have been recorded, whereas Kohleria is 

well known to the rural population (Kvist, 1987), and only one 

folk use of Pearcea has been recorded. The Coah Amerindians 

in the Amazon basin of Ecuador use Pearcea abunda for 

medicinal purposes (see that species). 

Geographical and Altitudinal Distribution 

The range of the entire genus (Figures 4, 5) is nearly 

identical to the range of the commonest species, Pearcea 

sprucei, which is found on the eastern Andean slopes and 

adjacent lowlands from the Colombian department of Caqueta 

in the north through Ecuador and Peru to the department of 

Cochabamba in northwestern Bolivia (Figure 6). Pearcea 

sprucei also is the only representative of the genus on the 

western slopes in Ecuador and very locally in the upper Rio 

Magdalena watershed in the Colombian department of Huila. 

All other Pearcea species are much more local in distribution: 

P. abunda (Figure 11) is distributed from southern Colombia to 

northern Peru, P. hispidissima (Figure 9) and P. reticulata 

(Figure 10) occur in Colombia and Ecuador, and 13 species 

are endemic to a single country (Figures 7-12). 

The numbers of species known from Ecuador, Peru, 

Colombia, and Bolivia are twelve, six, five, and one, 

respectively. Figure 5 shows how many species are known 

from departments or provinces of these countries. The largest 

number of species is found in the northern and central 

Ecuadorian Amazon region, particularly the Upano-Zamora 

valley (with 8 species). In addition, there seems to be a second 

speciose center in the Rio Huallaga valley of central Peru (with 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 3.Seeds of Pearcea and Kohleria (scale length= 100 pm): a, Pearcea hispidissima; h, Pearcea 

fuscicalyx; c, Pearcea rhodotricha; d, Kohleria allenii. Micrograph a of Lug0 3931 (08): b of Schunke 10366 

(MO); c of Lug0 5402 (US); d of Liesner 2106 (US). 

6 species). Pearcea is apparently rare between these areas, but 

this may reflect lower collecting activity in southern Ecuador 

and northern Peru. Most Peruvian collections have come from 

fairly few areas, e.g., close to Tingo Maria in the department of 

Huanuco, particularly from along the road east to Pucallpa. 

Extensive areas on the eastern Andean slopes in Peru have not 

been botanically explored, and current collecting activity is low 

due to political problems. It is therefore likely that additional 

species occur in Peru. At the present time, however, the 

morphological variation of Pearcea seems largest in Ecuador, 
NUMBER 84 9 

for example, the broadest range in corolla shapes occurs there. 

We define the altitudinal vegetation zones as lowland 

rainforest (below 700 m), lower montane forest (700-1500 m), 

montane forest (1500-2500 m), and cloud forest (above 2500 

m). These zones proved to be helpful in describing the 

altitudinal distribution of Columnea Linnaeus (Kvist and Skog, 

2 

0 

2 

4 

6 

0 

10 

12' 

14 

16. 

18. 

2 80 70 76 74 72 70 60 66 

4 

2 

,O 

'2 

,4 

-6 

-0 

.10 

.1 2 

.14 

-1 6 

-1 0 

FIGURE 4.-Distribution of Pearcea in Colombia, Ecuador, Peru, and Bolivia, and areas where species are most 

common. 
10 

d 

C 

2 

4 

6 

a 

10 

12 

14 

18, 

18 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

4 

2 

0 

2 

4 

6 

8 

10 

12 

14 

16 

18 

FIGURE 5.-Departments/provinces in Colombia, Ecuador, Peru, and Bolivia in which species of Pearcea have 

been collected, and number of species occurring in each departmentlprovince. 
NUMBER 84 11 

2 

0 

2 

4 

6 

8 

10 

12 

14 

16, 

la 

2 80 78 76 74 72 70 68 66 

S I 

1 

1 

1 

.-d 

1 ,-d-d . ,-' ,\ 

,.---- 1 I' \, f, I I I I 1 1 1 1 

80 78 76 74 72 70 68 66 62 

10 

12 

14 

16 

18 

FlGURE 6.-Distribution of Pearcea sprucei var. sprucei and var. parviflora (inset) in Colombia, Ecuador, Peru, 

and Bolivia. 
12 

2 

0 

2 

4 

2 

0 

2 

4 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

82 80 78 76 74 72 70 

I 

FIGURE 7.-Distribution of Pearcea bellu, P. hypocyrtiflora, P. intermedia, and P. schimpfii in Colombia and 

Ecuador. 

4 

2 

0 

2 

4 

t 

? 

D 

2 

4 

FIGURE 8.-Distribution of Pearcea bilabiatu, P. glabrara, and P. rhodotricha in Ecuador. 
NUMBER 84 13 

0 Pearcea hispidissima 

FIGURE 9.-Distribution of Pearcea cordato and P. hispidissima in Colombia and Ecuador. 

4 

2 

0 

2 

4 

-2 

-0 

-2 

-4 

RGURE 10.-Distribution of Peorcea gracilis and P. reticulato in Colombia and Ecuador. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

78 76 74 72 70 82 ao 

I 

Pearcea abunda 

;- P. grandifolia 

A P. fuscicalyx 

'-- -\ 

-1':- i 

,, e' -. ..---_ '\ 

2 

0 

2 

4 

6 

a 

10 

FIGURE 1 I.--Distribution of Pearcea obundo, P.fuscicolyx, and P. grandifolia in Colombia, Ecuador, and Peru. 

1993) and also are useful for Pearcea. The lower montane 

forest is the zone richest in Gesneriaceae in the Andes, and 

Pearceu also is concentrated here, in numbers of both species 

and specimens. 

Twelve species of Pearcea are found mainly or only in lower 

montane forest, two (Pearcea hypocyrtiflora and P. rhodotrick) are known only from lowland rainforest, and three (R 

cordura, P. gracilis, and P. intermedia) are known from 

montane forest. Cloud forest is penetrated only by the common 

P. sprucei, which ranges from 300 m to 2800 m elevation, but 

it is rare (or rarely collected) above 2000 m; most P. sprucei 

collections come from lower montane forest. In the lowland 

rainforest, P. sprucei is restricted to the most humid areas up to 

the base of the mountains (Figure 6). Pearcea abunda has been 

collected much more distantly from the Andes in the Peruvian 

department of breto (Figure 11). 
NUMBER 84 15 

I 8 

FIGURE 12.-Distribution of Pearcea purpurea and P. srrigosa in Peru. 

6 

8 

10 

12 

14 

16 

Variation 

Pearcea (including Parakohleria) is probably the least 

known and understood genus among the 23 genera now 

referred to the Gloxinieae (Wiehler, 1983). partly because its 

members occur in relatively inaccessible areas and habitats, and 

partly because species delimitation in the group is very 

problematic. In analyzing the intraspecific variation in Pearcea, it became clear to us that the species of Pearcea are 

polymorphic, may hybridize, and rapidly diverge morphologically in isolation. This causes taxonomic problems because one 

can either merge the more common and widespread entities or 

segregate numerous narrowly distributed endemics based on 

very few specimens. We have chosen to preserve some entities 

as separate taxa despite some intermediate specimens, and we 

accepted rare species only if they were represented by a 

sufficient number of specimens with stable characters. Pearcea 

cordata and P. grandifolia are known from only one and two 

specimens, respectively, but both are clearly set apart by 

features otherwise unknown in the genus and are not likely to 

be hybrids or extreme variations. 

It should be noted that we rarely find any apparent 

first-generation hybrids. In spite of that, hybridization seems 

evolutionarily important because the morphological variation 

of several species is best explained by backcrossing and 

introgression with other species. As in Kohleria (Kvist and 

Skog, 1992), hybridization seems to weaken the species limits, 

although none of the 17 accepted Pearcea species seems to 

have evolved by interspecific hybridization. It should also be 

noted that the term “population” is used below, although we 

mostly discuss morphological characters of herbarium specimens. Thus, herbarium collections of individual species from 

particular regions and altitudes represent the populations. The 

same applies to sympatric species. 

Examples of variation patterns that may reflect the polymorphic variation, introgression, and local speciation are briefly 

described below, but variation within individual species is 

described in more detail in the section on taxonomy. 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

POLYMORPHIC VARIATION.-Within populations of a species, individuals may differ strongly in certain features but be 

consistent in most others, as can be seen in the following 

examples: 

Pearcea purpurea. In the center of the range of the species in 

the Peruvian department of Huanuco, most specimens have 

ovate-lanceolate leaves, but a few (e.g., Poeppig 1632, 

Matthews 150) have longer, narrowly lanceolate leaves. 

Pearcea rhodotricha. 'Qpical specimens from the province 

of Napo, Ecuador, have ovate leaves and red corollas with a 

black apex, but two collections (Lug0 S. 3503, 3562) have 

lanceolate leaves and entirely red flowers. 

Pearcea sprucei. In Ecuador the majority of the specimens 

have rugose leaves, whereas this feature is otherwise uncommon except in the departments of Huanuco and nearby Ucayali 

(Figure 21c) in central Peru. Purplish leaves also are common 

in Ecuadorian populations and in some parts of Peru (Figure 

214, but reflexed calyx lobes are found only in Ecuador, 

where nearly one-half of the specimens exhibit this character. 

Reflexed calyx lobes also occur in the variety parvflora from 

Bolivia and southern Peru. Nearly glabrous upper leaf surfaces 

in this species are known only from the Peruvian department of 

Pasco (e.g., Gentry & Smith 35851), whereas a villous, rather 

than sericeous, leaf indumentum is found in Ecuador (and in the 

variety parvflora). 

INTRoGRESSION.-PlaIItS from the region(s) where a particular species seems to be common are, for the most part, fairly 

stable and distinct, whereas plants from the periphery of a 

species' range may show features of abundant sympatric 

species. Three examples follow: 

Pearcea reticulata. From the north in the Rio Pastaza valley 

(provinces of lhngurahua and Pastaza) of Ecuador, where the 

species is common, through the province of Nap0 and to the 

department of Putumayo in adjacent Colombia, the average 

size of the flowers is reduced and is more similar to the smaller 

flowers of the sympatric P. sprucei. To the south, one of the 

two collections from the province of Morona-Santiago (Steyermark 53639) resembles P. sprucei even more closely in having 

smaller flowers and shorter peduncles. 

Pearcea purpurea. This species is fairly common in the 

department of Huanuco, Peru, the apparent geographic center 

of the species' range. Here it is distinguished by having long 

peduncles, ovate and mostly reflexed calyx lobes, and a villous, 

purple indumentum. To the south in the departments of Pasco 

(a few collections) and Junin (two collections) and to the north 

in San Martin (one collection), the calyx lobes are erect and 

usually lanceolate, similar to those of P. sprucei, and the 

peduncle length is reduced from an approximate average of 8 

cm to 3 cm, but it still exceeds the pedicel in length. This in 

combination with the villous indumentum typical for P. 

purpurea sets these plants apart from sympatric P. sprucei 

populations. 

Pearcea strigosa. Qpical collections come from the 

Peruvian departments of Huanuco and adjacent Ucayali. 

Collections from Cuzco also have the strigose indumentum and 

subulate calyx lobes usual for the species, but they lack the 

sericeous corolla trichomes extending beyond the limb and 

have lanceolate leaves, similar to P. sprucei, rather than 

narrowly lanceolate leaves as are usually found in P. strigosa. 

SPECIATION.-Peripheral and isolated populations sometimes evolve distinct characteristics, diverging both from plants 

from the area(s) where the species is more common and from 

sympatric species. Examples can be found in Pearcea ahunda 

and P. sprucei. 

Pearcea abunda. The southernmost collection of this 

species, from the department of San Martin in Peru (Mug 

4216), differs by having considerably larger flowers (1.9 cm 

long rather than maximally 1.5 cm long) and a sparser 

indumentum than is typical for the species. It might even 

represent a separate species. 

Pearcea sprucei. The northernmost populations of this 

species, in the department of Caqueta, Colombia, differ by 

having peduncles usually several times longer than the pedicels 

and by having smaller flowers than more southern plants do, 

but the transition to populations in northern Ecuador is gradual 

(Figure 21a). A similar transition is seen in the southern part of 

the species' range, from long-pedunculate populations in Puno, 

Peru, and Bolivia, to short peduncles in northem Cuzco, Peru 

(Figure 21a). In addition, the former plants also have 

unusually large and long leaves with a sparse indumentum. 

Between the ranges of the two long-pedunculate P. sprucei 

populations in Colombia, Bolivia, and nearby Peru, there occur 

three other species with peduncles longer than 5 cm: P. 

reticulata in Ecuador, P. grandfolia in northern Peru, and P. 

purpurea in central Peru. Consequently, throughout nearly the 

entire range of Pearcea from southern Colombia to Bolivia, 

there are no sympatric long-pedunculate species. These species 

are probably close relatives (apart from the rare P. grandvolia), 

and the greatest taxonomic problem has been this complex. 

A further complication in Bolivia and southern Peru is that in 

addition to the long-pedunculate Pearcea sprucei plants, there 

sympatrically grow plants with short peduncles, to only 2 cm 

long (Figure 2 la), that also differ from the long-pedunculate 

specimens by having ovate leaves (Figure 21b), a villous 

indumentum (Figure 21g), and mostly reflexed calyx lobes 

(Figure 21e). These plants are placed in the variety 

parviflora of P. sprucei because var. sprucei and var. 

parviflora appear to be distinct in Bolivia and southern Peru 

(Figure 2la,b,d,e,g), the characters that separate the varieties appear to coalesce in more northern collections from the 

department of Cuzco in Peru, and some P. sprucei var. sprucei 

collections from high elevations in southern Ecuador are 

similar to var. parviflora (see further discussion under P. 

sprucei, below). 

Evolution 

Pearcea belongs to the tribe Gloxinieae, which consists of 

-1 8 genera that are found mostly in open habitats and that have 
NUMBER 84 17 

bivalved capsules (Figure lc) and wind-dispersed seeds, as 

well as some five genera found in the calm, humid forest 

understory that have evolved fleshy capsules adapted to animal 

dispersal. These latter genera are Pearcea (Figure la), fruits of 

which split into two fleshy valves from the apex to the base, 

Diastema (Figure lb), which also has fruits splitting with two 

apical slits, only one of which continues to the base, Kohleria, 

in which six of 17 species (Figure ld,e) split with one slit from 

capsule apex to base, and Monopyle (Figure If) and Solenophora Bentham, which split from the middle of the capsule 

rather than from the apex. In Monopyle, but apparently not in 

Solenophora, the slit continues to the apex. Most genera of the 

Gloxinieae are thus adapted to exposed habitats predominantly 

in areas with a seasonal climate. In such areas occur members 

of the tribe that, according to Kvist and Skog (1992). may be 

more primitive, such as Achimenes Persoon, Eucodonia 

Hanstein, and most species of Gloxinia. Later, a few groups, 

including Pearcea, apparently adapted independently to the 

forest understory, each switching to animal dispersal but 

developing different types of capsules. Pearcea and the other 

understory genera therefore seem to be advanced in the 

Gloxinieae (Kvist, 1991), although further studies are needed if 

one is to conclude that all the understory genera are, in fact, 

advanced. 

In the Gloxinieae the delimitation of species seems particularly difficult in the forest understory genera. In Kohleria 

(Kvist and Skog, 1992) the most common understory species, 

K. inaequalis, was as difficult to circumscribe as the most 

widespread and common species of Pearcea, P. sprucei, 

whereas the open-habitat species of Kohleria were more easily 

circumscribed. The species of the understory genera Diastema 

and Monopyle also are difficult to delimit. This may indicate 

that local speciation is particularly rapid in the understory 

groups, possibly due to the relatively narrow dispersal of 

animal-dispersed seeds in the understory habitat. In the 

wind-dispersed species, long-distance dispersal of the tiny 

seeds may counteract local speciation, e.g., Kohleria spicata 

has probably wind-dispersed to Isla del Coco 500 km 

southwest of the Costa Rican coast, and this very common 

species shows little variation throughout its range from Mexico 

to Peru. The plants on Isla del Coco, however, belong to a 

salt-influenced ecotype growing on rocks and otherwise found 

on islands along the Pacific coast of Panama and Costa Rica 

(Kvist and Skog, 1992). 

Hybridization followed by backcrossing and introgression 

seems to complicate taxonomic and phylogenetic studies in 

many genera of Gesneriaceae in northwestern South America. 

Other examples are Cremosperma Bentham, Reldia Wiehler, 

Kohleria, Columnea (Kvist and Skog, 1988, 1989, 1992, 

19931, and Pearcea. There are good indications, particularly 

from Kohleria, that this reflects extensive introgression or even 

reticulate evolution. 

Pearcea Regel 

Pearcea Regel, 1867:388. [Qpe: P. hypocyrriflora (J.D. Hooker) Regel 

Parakohleriu Wiehler, 1978:5. [m: P. abunda Wiehler = Pearcea abunda 

(Gloxinia hypocyrrifora J.D. Hooker).] 

(Wiehler) L.P. Kvist & L.E. Skog.] 

Terrestrial herbs, rarely epilithic or appearing epiphytic, 

stems decumbent to erect, usually terete in live material but 

becoming bisulcate or subquadrangular when dry, (5-)25- 100 

(-150) cm long, diameter 1-5(-10) mrn basally, branched or 

unbranched, stolons occasional, scaly rhizomes rare, but 

adventitious roots often present, basally often glabrescent or 

pilose to sericeous or villous apically. Leaves opposite, equal, 

subequal, or rarely unequal in each pair, blades usually 

lanceolate or ovate-lanceolate, less commonly ovate-elliptic, 

obovate, or oblanceolate, (3-)5-18(-25) x (1-)2-8(-10) cm, 

base oblique, cuneate, attenuate, or obtuse, apex acute or 

acuminate, margin senulate, less commonly serrate or nearly 

entire, above green or less commonly purplish, occasionally 

rugose, often pilose or hirsute, less commonly velutinous, 

strigose, or glabrous, below lighter green or purple, velutinous, 

pilose, or sericeous; lateral veins (5-)7-lo(-12) per side, often 

conspicuous and light green to yellow contrasting with darker 

green or purple areas between veins; petioles (0.2-)1-3(-6) 

cm long, Inflorescences of congested cymes of 1-12(-20) 

flowers in upper leaf axils, often purplish, sericeous, villous, or 

hispid; usually with peduncles to 2 cm long, less commonly 

epedunculate or with peduncles to 20 cm long; bracts mostly 2, 

but occasionally with up to 4 additional smaller bracts, each 

bract sessile, 2-25 mm long, ovate, lanceolate, or subulate; 

pedicels 0.2-5 cm long. Flowers protandrous; floral tube 

(combined bases of perianth and androecium) conic, green, 

yellow green, or purple, usually villous; calyx 3-13 mm long, 

the 5 lobes basally connate for 1-4 mm, mostly erect, less 

commonly reflexed, equal, triangular, ovate, lanceolate, or 

subulate with margin entire, both sides often sericeous or 

villous. Corolla bent downward at base, narrowly to broadly 

funnelform, subventricose, or ventricose, less commonly 

urceolate, rarely bilabiate, 0.9-2.7 cm long, base occasionally 

saccate, glabrous and yellow, tube red, often yellow basally and 

ventrally, rarely entirely yellow, outside villous, sericeous, or 

pilose, occasionally glandular-puberulent, rarely only puberulent, inside lighter red, cream, or yellow, often with darker 

dots, frequently puberulent and/or glandular-puberulent, throat 

often constricted, limb subregular, rarely bilateral, of 5 (rarely 

4) ovate to rotund lobes, outside red, inside red or yellow, often 

with darker dots, usually glandular-puberulent at least along 

margin; stamens 4, didynamous, included, subincluded, or 

rarely exserted, filaments 5-25 mm long, coiling in later 

female flowering stage to pull anthers deeper into flower, 

mostly glabrous, less commonly puberulent, rarely glandularpuberulent, cream or yellowish, anthers cream or yellowish, 

oblong, 1.2-2 x 1-1.5 mm, coherent at apices and sides, thecae 

distinct, dehiscing by longitudinal slits; 1 dorsal staminode, 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1-4 mm long, usually present; nectariferous disc fleshy, 

yellow, and glabrous, usually of 5 free glands, each 0.5-1 mm 

tall and 0.5- 1.5 mm wide at base, rarely with 2 glands placed 

closely together, with 4 glands, and/or with 1-2 of glands 

bilobed; ovary '/~~/4 inferior, pilose to sericeous, style 7-25 

mm long, cream or yellow white, mostly puberulent-pilose, 

especially basally, and glandular-puberulent, especially apically, stigma included or subincluded, rarely exserted, stomatomorphic, rarely capitate or weakly bilobed. Fruit a fleshy 

capsule, globose-ovoid, 4-7(-10) mm long, diameter 3-6(-8) 

mm, dehiscing apically and loculicidally into 2 valves, both 

slits apparently always continuing to base, valves reflexing 

completely and exposing sticky seed mass; seeds very 

numerous, irregularly subglobose or rarely ellipsoid, 0.2-0.4 

mm long, shape and size varying considerably even in a single 

capsule, surface irregularly striate, tawny, brown, or black. 

Key to the Species and Varieties of Pearcea Regel 

1. Leaves cordate; inflorescences and stems with trichomes to 8 mm long and retrorse; corolla yellow [plants 

from Ecuador]. .............................. -4. P. cord&, new species 

1. Leaves rarely cordate and only in combination with urceolate corollas; inflorescences and stems with 

trichomes only to 5 mm long and never retrorse; corolla red or orange [plants from throughout range 

of genus]. 

2. Leaves to 27 x 15 cm; peduncles 13-20 cm long, villous-hate [plants from Peru] ......... 

2. Leaves never exceeding 22 x 10 cm; peduncles rarely exceeding 8 cm, if longer then sericeous. 

.................................... 8. P. grandgolia, new species 

3. Corolla obviously bilabiate (upper lobes 7-9 mm) [plants from Ecuador] .............. 

3. Corolla urceolate, ventricose, or funnelform, rarely shortly bilabiate (upper lobes to 3 mm). 

.................................... 3. P. bikbiatu, new species 

4. Corolla urceolate; peduncles often absent, if present then shorter than pedicels; inflorescences with 

5. Corolla 11-13 mm long; upper leaf surface dark green along primary and secondary veins 

[plants from Colombia] ...................... .2. R bells, new species 

5. Corolla 15-25 mm long; upper leaf surface light green to yellow along veins [plants from 

Ecuador]. 

6. Leaves ovate, seldom cordate or oblong, both surfaces pilose, often purplish with yellow 

veins; calyx lobes 1-2 mm long .................. 10. R hypocyrtifrora 

6. Leaves elliptic to lanceolate, upper surface glabrous, green with lighter areas along primary, 

secondary, and tertiary veins surrounding darker blotches, lower surface veins with 

conspicuous trichomes of up to 10 large, flat cells; calyx lobes 3-8 mm long ...... ....................................... 15. R schhpfl 

4. Corolla funnelform or ventricose; peduncles rarely absent and frequently longer than pedicels; 

7. Leaves unequal or equal in each pair, upper surface usually glabrous (if sparsely pilose then with 

unequal leaves), lower surface with conspicuous villous or sericeous indumentum at 

primary, secondary, and tertiary veins, or lower surface pilose (then with unequal leaves); 

peduncles usually absent, if present then shorter than pedicels; floral tube with dense 

indumentum often contrasting with nearly glabrous calyx lobes. 

8. Leaves unequal in each pair, lower surface pilose; stems and inflorescences with densely 

hispid or villous 3-5 mm long trichomes; bracts and calyx lobes subulate [plants from 

Colombia and Ecuador] .............. 9. I? hispidissima, new combination 

8. Leaves equal or subequal in each pair, lower surface glabrous except on veins; stems and 

inflorescences usually with appressed trichomes; bracts and calyx lobes ovate or 

lanceolate. 

9. Leaves mostly ovate or elliptic, serrate, surface below green, above green with darker 

blotches surrounded by yellow green areas along primary, secondary, and tertiary veins 

[plants from Ecuador] ................. 1.1. P. intermedia, new species 

9. Leaves mostly lanceolate, sermlate, or subentire, usually purplish, particularly below 

between conspicuous yellow green veins. 

1-3(-5) flowers. 

inflorescences with 1-1 2(-20) flowers. 
NUMBER 84 19 

10. Calyx lobes glabrous; inflorescences usually pedunculate, pedicels 1-1.5 cm long 

10. Calyx lobes hispid; inflorescences epedunculate, pedicels 1.5-4.5 cm long [plants from 

7. Leaves subequal or equal in each pair, not glabrous; peduncles usually longer than pedicels; 

floral tube and calyx lobes usually with similar indumentum. 

Margin of leaves ciliate with 3-5 mm long tichomes; stems and inflorescences with 3-5 mm 

long purple trichomes; apical 5 mm of corolla, including limb, usually black, rarely 

11. Margin of leaves not ciliate or with trichomes only to 2 mm long; stems and inflorescences 

rarely with dark villous indumentum, trichomes not exceeding 3 mm long; corolla never 

black apically. 

12. Upper leaf surface stigose; calyx lobes lanceolate-subulate, strigose; corolla either 

minutely puberulent or sericeous with tricliomes extending -2 mm beyond limb. 

13. Peduncles longer than pedicels; calyx lobes 3-5 mm long; corolla glabrous except for 

13. Peduncles shorter than pedicels; calyx lobes 5-12 mm long; corolla sericeous with 

tichomes extending -2 mm beyond limb [plants from Peru] ............. 

Upper leaf surface velutinous, puberulent, pilose, or hirsute; calyx lobes ovate or 

lanceolate, pilose, villous, or sericeous; corolla pilose, villous, or sericeous with 

trichomes not extending beyond limb. 

14. Peduncles (if present) shorter than pedicels; corolla funnelform, limb not entirely red, 

diameter of throat 2 4 mm; calyx lobes lanceolate and about twice as long as wide at 

base (1 6. P. sptucei, new combination). 

15. Leaves usually ovate, never conspicuously rugose; stems, inflorescences, and veins 

usually with whitish villous indumentum [plants from Bolivia and Peru] 

15. Leaves usually lanceolate, often rugose; stems, inflorescences, and veins usually 

sericeous with yellow green or purplish indumentum [plants from throughout 

range of genus] .................... 16b. P. sprucei var. sprucei 

14. Peduncles at least as long as pedicels (occasionally shorter at flowering onset but 

elongate to exceed pedicels); corolla funnelform or venticose, limb entirely red or 

with darker dots or markings, diameter of throat 3-1 1 mm; calyx lobes lanceolate and 

about twice as long as wide at base or mostly ovate and about as long as wide at base. 

16. Limb red, rarely yellow, without markings, corolla ventricose, 9-15 mm long and 

3-4 mm wide in throat (to 19 mm long, 7 mm wide in specimens from San Martin 

Department, Peru) [plants from Colombia, Ecuador, and Peru] .......... 

16. Limb yellow or red with darker markings, corolla funnelform to venticose, 

(12-)15-25 mm long and (3-)5-11 mm wide in throat. 

17. Calyx lobes lanceolate and about twice as long as wide at base, never reflexed in 

central Peru but occasionally so in Ecuador and northern Peru; inflorescences 

and calyces sericeous with green to purplish indumentum [plants from 

throughout range of genus] ............ 16b. P. sprucei var. sprucei 

17. Calyx lobes mostly ovate and about as long as wide at base, often reflexed; 

inflorescences and calyces sericeous with yellow green indumentum or villous 

with grayish green or purple brown indumentum. 

Stems, inflorescences, and calyces with grayish or purple brown villous 

indumentum; 2 upper lobes of limb red, 3 basal lobes yellow with darker spots 

or markings centrally or along margin; peduncle 3-7 cm long [plants from 

Peru] ................... 12. P. purpurea, new combination 

18. Stems, inflorescences, and calyces sericeous with yellow green indumentum; 

entire limb yellow with darker dots; peduncle 4-15 cm long [plants from 

Colombia and Ecuador] ......... 13. P. reticukzta, new combination 

[plants from Ecuador] ................. .6. P. ghbrata, new species 

Peru] .......................... 5. P. fuscicalyx, new species 

11. 

entirely red [plants from Ecuador] ........ 14. P. rhodotricha, new combination 

minute puberulence [plants from Ecuador] ....... 7. P. gracib, new species 

.............................. 17. P. strigosa, new species 

12. 

..................... 16a. P. sprucei var. pant$ora, new status 

.......................... 1. P. abunda, new combination 

18. 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1. Pearcea abunda (Wiehler) L.P. Kvist & L.E. Skog, 

new combination 

FIGURES la, 11 

Parakohleria abunda Wiehler, 19785, 7, pl. 1A. [1Lpe: plants cultivated at the 

Marie Selby Botanical Gardens, Sarasota, FL, originally collected by 

Madison et al. (3194) from Ecuador, prov. Morona-Santiago, Wiehler 77129 

(SEL, holotype).] 

Stems to 100(-150) cm long, occasionally branched, yellow 

green and villous to lanate apically. Leaves subequal in each 

pair, blades ovate-lanceolate, 7- 15(-22) x 2.5-7(- 10) cm, 

rarely purplish, above hirsute, below pilose with villoussericeous veins, margin semlate. Inflorescences with 3- 12 

flowers, yellow green to purple, villous; peduncles 23-5 cm 

long; bracts lanceolate, 4- 10 mm long; pedicels 0.5-1.5 cm 

long. Floral tube villous-lanate; calyx lobes often reflexed, 3-4 

mm long, ovate; corolla 0.9-1.5(-1.9) cm long, tube ventricose, diameter 5-9(-12) mm medially and 3-4(-7) mm in 

throat, bright red, less commonly intense yellow, outside 

sericeous, inside usually glandular-puberulent, lobes 1-2 mm 

long, red, rarely with darker purple lower lobes; filaments 

occasionally sparsely puberulent, style puberulent and/or 

glandular-puberulent. Capsule subglobose, 3-5 mm long, 

diameter 3-5 mm; seeds broadly ellipsoid, -0.4 mm long, 

brown, striate. Chromosome number n = 11 (counted by Royce 

Oliver, Department of Botany, Smithsonian Institution, from 

Kvist 60451 (vouchers in AAU, QCA, and US). 

DISTRIBUTION.-Pearcea abunda occurs in the Ecuadorian 

Amazon lowland and adjacent areas of Colombia and Peru 

(Figure 11). The species is locally common in Ecuador, but 

only the cited specimens have been seen from Peru and 

Colombia. The Peruvian collection from Loreto is farther from 

the Andean slopes than any other collection of Pearcea (Figure 

11). Pearcea abunda mostly occurs below 500 m elevation but 

reaches elevations up to loo0 m in the Ecuadorian province of 

Morona-Santiago. 

REPRESENTATIVE COLLECTIONS.-COLOMBIA. Putumayo: 

Towards Puerto Asis, between Rio Chipayacu and Finca 

Pedrogoso, Ewan 16734 (US); Umbria, Klug 1659 (BM, F, GH, 

ECUADOR. Morona-Santiago: Taisha, Rio Pangui, 10 km 

S-SE of military camp, Brandbyge & Asanza 32112 (AAU, 

QCA, us); Mendez, 600-800 m, Camp 902E (NY, S, US); Macas, 

Kvist 60451 (MU, QCA, US); 60452 (AAU, QCA, QCNE, 

us). Napo: Aningu along Rio Napo, Besse et af. 1601 (SEL), 

1602 (SEL); Rio Aguarico, Durenio,Ceron 344 (NY). Pastuza: 

10 km E of Puerto Sarayacu, Sarayaquillo, Lug0 S. 4018 (GB, 

SEL), 5443 (GB, us). 

PERU. Loreto: Rio Huallaga, Lewis et al. 11309 (USM); 

Valencia. Rio Comentes near Platanoyacu, McDaniel & 

Marcos 11139 (us). Sun Martin: Upper Rio Huallaga, Juan 

Jui, Klug 4216 (A, BH, BM, F, K, MO, NY, S, U, UC, US, USM). Total 

of 27 wild collections seen. 

DISTINGUISHING FEATURES.-The combination of hirsute 

leaves, villous inflorescences with peduncles longer than 

MO, NY, US). 

pedicels, ovate and mostly reflexed calyx lobes, ventricose and 

red (rarely yellow) corollas with a narrow throat (see 

illustration in Kvist, 1991:22), and an entirely red/purple limb 

without darker spots or markings sets Pearcea abunda apart 

from all other species in the genus. 

VARIATION.-The sole collection from the Peruvian department of San Martin (Klug 4216) differs by having considerably 

larger flowers (corolla 1.9 cm long, diameter 1.2 mm medially 

and to 0.7 mm in the throat) and by having longer (to 22 cm), 

relatively narrower leaves. It may represent a new species 

closely related to P. abunda, but additional collections are 

necessary before this can be determined. 

NOTES.-In northwestern Ecuadorian Ammonia, the Cofan

Amerindians crush and boil the leaves of Pearcea abunda and 

drink the decoction for colds. They also bum the leaves and 

apply the ash to sores in the mouth, according to Ceron 344. 

2. Pearcea bella L.P. Kvist & L.E. Skog, new species 

FIGURES 7, 13 

Pearceae hypocyrtijlorae (J.D. Hooker) Regel affinis, sed 

corollis 11-13 mm longis urceolatis, floribus in foliorum 

axillis solitariis, foliis in paginis superioribus secus vena 

atro-viridibus differt. 

Erect stems 10-20 cm long, basal diameter 2-3 mm, 

unbranched but with elongated rhizomes and adventitious 

roots, villous with yellow brown trichomes, shoots somewhat 

dorsiventral; internodes 0.5-2 cm long. Leaves subequal to 

unequal in each pair, blades oblique, ovate, 3-7 x 1.5-3.5 

cm, base attenuate-cuneate, apex acute-acuminate, margin 

sermlate, above with dark green areas along primary and 

secondary veins, weakly rugose, appressed pilose, below 

lighter green, pilose, villous along veins, lateral veins 5-6 per 

side; petiole 1-3 cm long. Inflorescences of solitary flowers in 

upper leaf axils, villous with yellow green trichomes; epedunculate and ebracteate; pedicels 3-5 cm long. Floral tube conic, 

1.0-1.5 mm long, villous; calyx 3-4 mm long, villous, lobes 

erect, triangular, 1-1.5 mm long and 1.5-2 mm wide at base, 

apex acute; corolla 11-13 mm long, tube urceolate, 9-12 mm 

long, diameter 7-9 mm medially and 1.5-2.5 mm in throat, 

outside red, ventrally and basally yellow red, villous, inside 

yellow, puberulent, limb with lobes subequal, rotund, 1-1.5 

mm long and 1.5-2 mm wide at base, red, inside puberulent, 

with acute-obtuse apex; stamens included, filaments 5-6 mm 

long, puberulent, anthers -1.2 x 1 mm; staminode 5-6 mm 

long; nectariferous disc of 5 glands, all -0.5 mm high and 0.5 

mm wide at base; ovary 2/3 inferior, sericeous, 2-2.5 mm long, 

diameter 1-1.5 mm, style included, 7-9 mm long, puberulent 

and sparsely glandular-puberulent apically, stigma stomatomorphic. Fruit and seeds not seen. 

TYPE.-COLOMBIA. Caquetal: Eastern slopes of the Andes 

along the road to Florencia, Quebrada Delicias, 750 m, 9 Feb 

1987, Londonio & Kvist 136 (COL, holotype; AAU, TULV, US, 

isotypes). 
NUMBER 84 21 

FIGURE 13.-Peurceu bellu: a, habit and inflorescences; b, indument of upper leaf surface; c, flower showing 

corolla mouth and anthers; d. lateral view of flower; e, laterally opened corolla showing androecium;f, lateral 

view of flower with corolla removed; g, calyx, nectary, and gynoecium. All from Cuurrecusus 9161 (F). 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ETYMOLOGY.-The specific epithet, bella, comes from the 

Latin bellus, meaning pretty. 

DISTRIBUTION.-The species is known only from the eastern 

slopes of the Andes in the Colombian department of Caqueta 

between 600 m and 1500 m elevation (Figure 7). Dense 

populations arising from creeping rhizomes were observed on 

shaded banks and slopes near small streams (Londono & Kvist 

136,141). 

OTHER COLLECTIONS.- COLOMBIA. Caqueta: Sucre, 

banks of Rio Hacha, Cuatrecasas 9161 (COL, F, us); road to 

Florencia, Londonio & Kvist 141 (AAU, US). Total of three 

collections seen. 

DISTINGUISHING FEATURES.-The approximately 1 cm long, 

urceolate corollas easily identify Pearcea bella. In addition, 

this species is unique in the genus by having solitary flowers in 

the leaf axils. Pearcea bella is not spectacular, but the pattern 

of lighter and darker green on the upper leaf surfaces and the 

small, urceolate, red corollas make populations of this species 

attractive. 

3. Peurceu biltabiata L.P. Kvist & L.E. Skog, new species 

FIGURES 8, l5f 

Species insignis limbo corollae valde bilabiato, lobis 

superioribus 7-9 mm longis, a speciebus cogniti notis bene 

distincta. 

Stems 20-100 cm long, basal diameter to 5 mm, frequently 

branched, glabrescent basally to sericeous apically; internodes 

1-5 cm long. Leaves equal or subequal in each pair, blades 

mostly oblique, ovate to lanceolate, 4-lo(-15) x 1.5-4(-6) 

cm, base attenuate-cuneate, apex acute-acuminate, margin 

serrate-serrulate, above dark, dull pen, glabrous or sparsely 

appressed pilose particularly along veins, rarely rugose, below 

lighter green, velutinous, lateral veins 6-9 per side; petioles 

0.5-2 cm long, Inflorescences of 2-7 flowers in upper leaf 

axils, yellow green or purple, sericeous or villous; peduncles 

0.5-2(-4) cm long; bracts 2, lanceolate-subulate, 5-12 x 1-2 

mm; pedicels 1.5-4.5 cm. Floral tube conic, 2-4 mm long, 

green or purple, villous; calyx 6-11 mm long, villous, lobes 

erect, lanceolate, 5-9 mm long and 1-2 mm wide at base, apex 

acute-acuminate; corolla 16-24 mm long, tube 9-15 mm long, 

diameter 3-7 mm, outside red, sericeous, inside yellow, 

glabrous, limb bilabiate, red outside, yellow and glandularpuberulent inside, lobes rotund, lower 3 with dark dots, upper 

lobe (= 2 dorsal connate lobes) 4-7 mm long (to base of lateral 

lobes and 7-9 mm long to base of basal lobe), lateral lobes 

1-2.5 mm long, basal lobe 2-3.5 mm long and reflexed; 

stamens exserted, filaments 17-25 mm long, glabrous, anthers 

-1.8 x 1.5 mm; staminode 2-4 mm long; nectariferous disc of 

5 glands, all -1 mm high and -1 mm wide at base; ovary 2/3 

inferior, 3-4 mm long, diameter 3-4 mm, sericeous, style 

exserted, 17-25 mm long, sparsely glandular-puberulent or 

glabrous, stigma stomatomorphic. Capsule globose, 4-5 mm 

long, diameter 4-5 mm; seeds subglobose-ellipsoid, 0.25-0.35 

mm long, brown black, irregularly striate. 

TYPE.-ECUADOR. Moronu-Santiago: Road from Sigsig to 

Gualaquiza, between Oroyaco and El Porton. 22 Jul 1948, 

Scolnik 1490 (US, holotype). 

ETYMOLOGY .-The specific epithet refers to the bilabiate 

form of the corolla. 

DlsTRIBuTloN.-Apparently a rare endemic, Pearcea bilabiata is restricted to the Ecuadorian province of MoronaSantiago (Figure 8). The species occurs at 800-1500 m 

elevation in the Upano-Zamora valley between the Andean 

slopes to the west and the Cordilleras de Condor and Cutucui to 

the east. In the Upano-Zamora valley at least seven Pearcea 

species occur, including P. bilabiata; the others are P. abunda, 

P. glabrata, P. gracilis, P. hispidissima, P. sprucei, and P. 

reticulata. 

OTHER COLLECTIONS.-ECUADOR. Morona-Santiago: 

Cordillera de Cutucui, Mendez-Morona road, km 19, Dorr & 

Valdespino 6297A (US); Indanza-Limon (General Plaza), 

Hurling & Andersson 12755 (GB); new road from Mendez to 

Morona, Hirtz et al. 7130 (MO); 20 km S of San Juan Bosco, 

Madison et al. 7553 (SEL); Rio Calagras, Luer et al. 11870 (US); 

Cordillera de Cutucui, Mendez-Morona road, van der Weg& 

Palacios 10408 (US). Total of seven collections seen. 

DISTINGUISHING FEATURES.-The bilabiate corollas set 

Pearcea bilabiata apart from all other species in the genus, 

although the species is difficult to distinguish vegetatively. 

4. Pearcea corduta L.P. Kvist & L.E. Skog, new species 

FIGURES 9,14 

Species insignis foliis cordatis, inflorescentiis et caulis 

trichomatibus ad 8 mm longis hispidis retrorsis, corollasque 

flavis, a speciebus cognitis notis bene distincta. 

Stems -50 cm tall, 5 mm or more in diameter, unbranched, 

with the petioles, inflorescences, and flowers all with conspicuous, 5-8 mm long, yellow brown, hispid, retrorse trichomes, 

each consisting of -10 cells; internodes -5 cm long. Leaves 

subequal in each pair, blades cordate, 7-12 x 6-8 cm, margin 

serrate, above dull green, somewhat rugose, hirsute, below 

lighter yellow green, pilose, lateral veins 6-8 per side; petiole 

3-7 cm long. Inflorescences of -5 flowers; peduncles -2 cm 

long; bracts 2, ovate-lanceolate, 8-15 x 3-5 mm; pedicels 

0.8-1.5 cm long. Floral tube conic, -3 mm long; calyx -8 mm 

long, lobes lanceolate, -6 mm long, apex acute; corolla -15 

mm long, tube funnelform, diameter to 6 mm in throat, outside 

yellow, inside, particularly limb, puberulent and glandularpuberulent, lobes subequal, rotund, 2-3 mm long and 2-3 mm 

wide at base, inside glandular-puberulent; stamens included, 

filaments - 12 mm long, puberulent and glandular-puberulent, 

anthers -2 x 1 mm; nectariferous disc of 5 glands, -0.8 mm 

high; ovary -2/3 inferior, -4 mm long, diameter -2 mm, pilose, 
NUMBER 84 23 

- /// , 

5mm 

FIGURE 14.-feurceu cordola: a. habit and inflorescences; b, indument of upper leaf surface; c, lateral view of 

flower, d, laterally opened corolla showing andmcium; e, calyx, nectary, and gynoecium. All from Holm-Nielsen 

el at. 26512 (AAU). 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 84 25 

FIGURE 15 (opposite page).-f eorcea fuscicalyx: a, habit and inflorescences; 

b, indument of lower leaf surface; c, lateral view of flower; d, laterally opened 

corolla showing androecium; e, calyx, nectary, and gynoecium. feorceo 

bilabiata: J lateral view of flower. a,b from Schunke 5116 (us); c-e from 

Schunke 3111 (US); f from Scolnik 1490 (US). 

style included, -10 mm long, basally pilose and glandularpuberulent apically, stigma stomatomorphic. Fruits and seeds 

not seen. 

TYPE.-ECUADOR. Napo: Rio Borja, 1740 m, 21 Sep 1980, 

Holm-Nielsen et al. 26512 (AAU, holotype). 

ETYMOmY.-The specific epithet refers to the cordate leaf 

bases. 

DISTRIBUTION.-Montane forest in the Ecuadorian province 

of Nap0 (Figure 9). According to the label (of the single 

collection), Pearcea cordata was growing in a swamp forest at 

1740 m elevation. 

DISTINGUISHING FEATURES.-Pearcea cordata is the only 

species with cordate leaves and a hispid, retrorse indumentum 

(the latter character probably also distinguishes P. cordata 

from all other Ecuadorian Gesneriaceae). For these reasons, we 

have described Pearcea cordata despite the fact that only one 

poor collection (with two flowers) has been seen. It is also very 

unusual for a Pearcea species to have yellow rather than red or 

orange red flowers, but yellow flowers occasionally occur in P. 

abunda and P. sprucei. 

5. Pearcea fuscicalyx L.P. Kvist & L.E. Skog, new species 

FIGURES 36, 11, 15a-e 

Differt a Pearceae glabratae L.P. Kvist & L.E. Skog lobis 

calycum hispidis, inflorescentiis epedunculatis, pedicellis 1.5- 

4.5 cm longis. 

Stems often decumbent, 20-50 cm long, basal diameter to 3 

mm, usually unbranched, frequently with stolons and adventitious roots, often purplish, sericeous; internodes 1-3(-5) cm 

long. Leaves equal in each pair, blades weakly oblique, 

lanceolate, 6-15 x 1-3 cm, base cuneate, apex acuteacuminate, margin sermlate, above dark green with lighter 

veins, glabrous, below purplish at least centrally on both sides 

of primary vein, glabrous apart from sericeous veins, lateral 

veins 8-9 per side; petiole 1-3 cm long. Inflorescences of 1-3 

flowers, violet or purple; hispid; epedunculate; bracts 2, 

lanceolate-subulate, 6-12 x 1-2 mm; pedicels 1.5-4.5 cm 

long. Floral tube conic, 1.0-1.5 mm long, dark purple, hispid; 

calyx 6-8 mm long, hispid, lobes erect, lanceolate, 4-6 mm 

long and 1-2 mm wide at base, apex acute-acuminate; corolla 

18-24 mm long, tube funnelfom, subventricose, 16-20 mm 

long, diameter 8-12 mm medially and 6-9 mm in throat, 

outside red, pilose, inside glabrous or sparsely glandularpuberulent, limb with lobes subequal, rotund, 2-3 mm long 

and 3-4 mm wide at base, apex obtuse-truncate, outside red, 

densely glandular-puberulent, inside lighter red, sparsely 

glandular-puberulent; stamens subincluded, filaments 15- 18 

mm long, glabrous, anthers -1.8 x 1.2 mm; staminode 3-4 

mm long; nectariferous disc of 5 glands, all -0.7 mm high and 

1.0 mm wide at base; ovary 3/4 inferior, 4-5 mm long, diameter 

3-4 mm, sericeous, style subincluded, 15-18 mm long, 

glandular-puberulent and puberulent, stigma stomatomorphic. 

Capsule globose, 5-6 mm long, diameter 5-7 mm; seeds 

ellipsoid, 0.25 x 0.35 mm long, brown, striate. 

TYPE.-PERU. Huainuco: Loncio Prado, Jacintillo on left 

bank of Rio Monzon, 170 m, 3 Nov 1975 Schunke V. 5116 (us, 

holotype; COL, F, G (2). GH, IBE, K, NY, P, USM, isotypes). 

ETYMOLOGY.-The specific epithet refers to the purple-. 

hispid floral tube. and calyx. 

DISTRIBUTION.-Pearcea fuscicalyx is so far known only 

from near Tingo Maria in the department of Huanuco in thRioo 

Huallaga valley of Peru (Figure 11). All collections come from 

elevations around 700 m. Three other species of Pearcea, P. 

purpurea, P. sprucei, and P. strigosa, occur in the same area 

(Figures 6, 12). The first two occur at the same elevations as 

P. fuscicalyx, whereas the latter is found only at considerably 

higher elevations (in Huanuco). 

OTHER COLLECTIONS.-PERU. Huainuco: Tingo Maria, 

Allard 20838 (US), 21856 (US), 22233 (us), 22287 (BH, US); 

junction of Rio Monzon and Rio Huallaga, Asplund 12684 (s); 

trail along Rio Monzon near Rio Huallaga at Tingo Maria,

Croat 21261 (MO); across Rio Huallaga from Tingo Maria, 

Gentry et al. 15941 (F, MO, US); SW of Tingo Maria, road to 

Jacintillo, Schunke V. 3111 (F, US); left bank of Rio Monzon, 

Jacintillo, Schunke V. 10366 (MO, us). Total of 10 collections 

seen. 

DISTINGUISHING FEATURES.-The combination of narrowly 

lanceolate, glabrous (apart from some hairs on the veins below) 

leaves with a purple lower surface, epedunculate inflorescences 

with up to three flowers, pedicels approximately 3 cm long, and 

purple-hispid floral tubes sets Pearceafuscicalyx apart from all 

other species of Pearcea. 

6. Pearcea gkbrata L.P. Kvist & L.E. Skog, new species 

FIGURES 8,16 

Species insignis laminis foliorum glabris lanceolatis paginis 

inferioribus purpureis, marginibus subintegris, tubis floralibus 

sericeis, lobis calycum reflexis a speciebus cognitis notis bene 

distincta. 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 84 27 

FIGURE 16 (oppposite page).-feurceu glubmfu: u, habit and inflorescences; 

b, indument of lower leaf surface: c, lateral view of flower; d, laterally opened 

corolla showing androecium; e, flower with corolla removed; f, calyx, nectary, 

and gynoecium. All from Hurling 3756 (S). 

Stems to 1 m long, basal diameter to 3 mm, frequently 

branched, yellow green, sericeous; internodes 1 -6(-10) cm 

long. Leaves subequal in each pair, blades oblique, lanceolate, 

less commonly elliptic, (2-)4-10 x 1-3 cm, base cuneate, 

apex acute-acuminate, margin subentire, above dull green, 

glabrous, below usually purplish, glabrous apart from the 

strigose veins, lateral veins 7-9 per side; petioles 0.2-2 cm 

long. Inflorescences of 1-3(-5) flowers, yellow green, 

sericeous; peduncles 0.2- 1.5 cm long; bracts 2, lanceolatesubulate, 4-8 x 1-2 mm; pedicels 1-1.5 cm long. Floral tube 

conic, 1.0-1.5 mm long, yellow green, sericeous; calyx 4-6 

mm long, basally sericeous, lobes reflexed, ovate, 2-4 mm 

long and 1.5-3 mm wide at base, glabrous, apex acute; corolla 

15-20 mm long, tube funnelform, subventricose, 12-18 mm 

long, diameter 5-9 mm medially and 4-8 mm in throat, 

outside red, pilose, inside glabrous, limb with lobes subequal, 

reflexed, rotund, 2-3 mm long and 2-3 mm wide at base, red, 

margin frequently glandular-puberulent, apex obtuse-truncate; 

stamens subincluded, filaments 10-1 8 mm long, glabrous, 

anthers -1.5 x 1.0 mm; staminode -3 mm long; nectariferous 

disk of 5 or rarely 4 glands, all -0.6 mm high and 0.8 mm wide 

at base; ovary 3/4 inferior, 4-5 mm long, diameter 3-4 mm, 

sericeous, style subincluded to exserted, 15-20 mm long, 

puberulent, stigma stomatomorphic. Fruit and seeds not seen. 

TYPE.-ECUADOR. Morona-Santiago: Arapicos, 850 m, 2 

Apr 1987, Lug0 S. 5923 (US, holotype; GB, isotype). 

ETYMOLOGY.-The specific epithet refers to the glabrous 

upper leaf surface. 

DISTRIBUTION.-Apparently Pearcea glabrata is a locally 

endemic inhabitant of the central eastern Andean slopes of 

Ecuador (Figure 8), where it has been collected between lo00 

m and 1500 m elevation. 

OTHER COLLECTIONS.-ECUADOR. Pastaza: Mera, Harling 3756 (S); Mera-Shell Mera road, km 5, Ollgaard et al. 

35583 (AAU). Tungurahua: Colonia Regina, 5 km W of Rio 

Verde, Lug0 S. 1894 (GB, SEL); Banlos-Mera, km 35, alongRio

Cashaurco, Ollgaard & Balslev 9268 (AAU, NY). Province 

Unknown: Pearce sen. (K). Total of six collections seen. 

DISTINGUISHING FEATURES.-The glabrous (apart from the 

veins below), lanceolate leaves, usually with a purple lower 

surface, and nearly entire margin distinguish Pearcea glabrata. 

The contrast between the sericeous floral tube and the glabrous, 

reflexed calyx lobes also characterizes the species. 

NOTES.- According to the label of one of the specimens, 

(Ollgaard et al. 35583) the species occasionally grows as an 

epiphyte. 

7. Pearcea gracilis L.P. Kvist & L.E. Skog, new species 

FIGURES 10, 17 

Foliis lanceolatis, hirsutis, viridibus, marginibus subintegris, 

inflorescentiarum pedunculis quam pedicellis plus quam duplo 

longioribus, bracteis et lobis calycum subulatis, corollis 

subglabris. 

Stems to 120 cm long, basal diameter to 3 mm, frequently 

branched, yellow green, sericeous; internodes 2-5 cm long. 

Leaves equal or subequal in each pair, blades weakly oblique, 

lanceolate, 4-10 x 1-2.5 cm, base cuneate, apex acuteacuminate, margin subentire, above green, strigose, below paler 

green, hirsute with sericeous veins, lateral veins 6-8 per side; 

petiole 0.3-1.2 cm long. Inflorescences of 2-4(-7) flowers, 

yellow green, sericeous; peduncles 1.5-4.5 cm long; bracts 2, 

subulate, 2-10 x 0.5-1.5 mm; pedicels 0.3-1.5 cm long. 

Floral tube conic, 1.0-1.5 mm long, yellow green, sericeous; 

calyx 4-6 mm long, strigose, lobes erect, lanceolate-subulate, 

3-5 mm long and 1-1.5 mm wide at base, apex acuminate; 

corolla 13- 18 mm long, tube funnelform, subventricose, 

12-16 mm long, diameter 4-6 mm medially and 3-6 mm in 

throat, outside red, with only tiny puberulent and frequently 

also glandular-pubemlent trichomes, inside glabrous, limb with 

lobes subequal, rotund, 1-2.5 mm long and 2-3.5 mm wide at 

base, red, outside densely glandular-puberulent, inside 

glabrous, apex obtuse-truncate; stamens included, filaments 

5-15 mm long, glabrous, anthers -2 x 1.5 mm; staminode 

3-4 mm long; nectariferous disc of 5 or rarely 4 glands, all 

-0.5 mm high and 0.8 mm wide at base; ovary 2/3 inferior, 

3-4 mm long, diameter 2-3 mm, sericeous, style subincluded, 

12-18 mm long, glabrous, stigma stomatomorphic. Capsule 

globose, 6-8 mm long, diameter 5-7 mm, dehiscing by 2 

valves; seeds subglobose-ellipsoid, 0.2-0.3 mm long, brown 

black, irregularly striate. 

TYPE.-ECUADOR. Morona-Santiago: 10 km S of San 

Juan Bosco, 1570 m, 27 Jan 1987, Gentry et al. 30865 (QCA, 

holotype; K, MO, SEL, isotypes). 

ETYMOLOGY.-The specific epithet refers to the slender 

stems of this species. 

DlSTRIBUTION.-Pearcea gracilis is known from the eastern 

Andean slopes of Ecuador between 1500 m and 2000 m 

elevation (Figure 10). 

OTHER COLLECTIONS.-ECUADOR. Morona-Santiago: 

Trail from Sevilla de Or0 to Mendez, Tambo Pilas, Camp 761E 

(US); road Limon (General Plaza)-Gualaceo, Hurling & Andersson 12685 (GB, SEL); 8 km S of San Juan Bosco, Madison 

& Besse 7534 (SEL). Total of four collections seen. 

DISTINGUISHING FEATURES.-The combination of lanceolate, hirsute, green leaves with nearly entire margins, inflorescences with peduncles at least twice as long as the pedicels, 

subulate bracts and calyx lobes, and glabrous corollas (apart 

from tiny puberulent trichomes) distinguishes Pearcea gracilis. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 84 29 

FIGURE 17 (opposite page).-Pearcea gracilis: a, habit and inflorescences; b, 

indument of upper leaf surface; c, lateral view of corolla; d, laterally opened 

corolla showing androecium; e, flower with corolla and androecium removed; 

f, calyx, nectary, and gynoecium; g, enlargement of calyx, nectary, and ovary; 

h, capsule; i, seeds. a,b from Harling & Andersson 12685 (GB); c-i from 

Madison et al. 7534 (SEL). 

8. Pearcea grandifolia L.P. Kvist & L.E. Skog, new species 

FIGURES 11, 18 

Foliis magnis 27 x 15 cm, indumento caulium inflorescentiarumque villoso-lanato, pedunculis 13-20 cm longis, sed lobis 

calycum longioribus quam latioribus aequantibus. 

Stems rising 30-50 cm tall, basal diameter to 1 cm, 

unbranched, yellow green or reddish, villous basally to lanate 

apically; internodes 3-4 cm long. Leaves equal in each pair, 

blades weakly oblique, ovate-obovate, 13-27 x 8-15 cm, 

base attenuate to obtuse, apex acute-acuminate, margin serrate, 

above green, pilose with villous veins, below lighter green, 

pilose, lateral veins 10-12 per side; petiole 2-6 cm long. 

Inflorescences cymes of 6- 15 flowers, villous-lanate; peduncles 13-20 cm long (but shorter at beginning of flowering); 

bracts 2-4, lowest 2 ovate-elliptic, 12-25 x 6-10 mm, other 

bracts smaller and lanceolate-subulate; pedicels 1-4 cm long. 

Floral tube conic, -5 mm long, villous; calyx 8-12 mm long, 

villous-pilose, lobes erect, ovate, 7-10 mm long and 4-7 mm 

wide at base, apex acute; corolla 23-27 mm long, tube 

funnelform, diameter to 15 mm in throat, outside red with 

villous white indumentum, inside glandular-puberulent, limb 

with rotund lobes, 2-3 mm long and 4-6 mm wide at base, 

glandular-puberulent inside; stamens included, filaments -20 

mm long, glandular-puberulent, anthers -2 x 1.5 mm; nectariferous disc of 5 glands, each -1.5 mm high and 1 mm wide; 

ovary */3 inferior, -5 mm long and 3 mm wide, villous, style 

included, -1 2 mm long, pilose, glandular-puberulent apically, 

stigma stomatomorphic. Capsule globose, -10 mm long, 

diameter -8 mm; seeds several hundreds, -0.3 x 0.2 mm, 

ellipsoid, obliquely striate, brown. 

Sun Martin: Prov. Mariscal Caeres, Quebrada de Santa Rosa de Cachiyacu (road to Progreso), 600 m, 19 

Jul 1974, Schunke V. 7558 (us, holotype; MO, isotype). 

ETYMOLOGY.-The specific epithet refers to the large 

leaves. 

DISTRIBUTION.-Known only from lower montane forest 

(500-1000 m) in the Peruvian department of San Martin 

(Figure 1 1). 

Sun Martin: Prov. Mariscal 

Caceres, District Campanilla, “Carretera marginal de la selva,” 

7 km N of Pulcache, Plowman & Schunke V. 1161 1 (K). Total 

of two collections seen. 

DISTINGUISHING FEATURES.-The very large leaves easily 

distinguish Pearcea grandifolia even vegetatively. The lanate 

TYPE.-PERU. 

OTHER COLLECTION.-PERU. 

indumentum of the stems and inflorescences also is distinctive, 

and only P. reticulata may have equally long peduncles (but 

that species does not have calyx lobes that are longer than the 

width at base). We have described P. grandifolia despite the 

limited material because the species is so unusual that we 

believe it cannot be a hybrid or a variant of some other species. 

9. Pearcea hispidissima (Wiehler) L.P. Kvist 8z L.E. Skog, 

new combination 

FIGURES 3a, 9 

Parakohleria hispidissimo Wiehler, 1978:6, 7, pl. 1B. [”)ye: Ecuador, Pas-, 

Stems to 100 cm long, 3-8 mm in diameter, occasionally 

branched, densely hispid with yellowish or purplish 3-5 mm 

long trichomes. Leaves unequal in each pair, rarely subequal, 

larger blade of pair ovate-elliptic, 8-18 x 3-9 cm, margin 

entire to sermlate, above dull green, glabrous, below lighter 

green, pilose; smaller blade of pair ovate-rotund, 0.8-3(-7) X 

0.5-1 3-5) cm. Inflorescences of 1-4(-6) flowers, yellow 

green, villous; peduncles to 2.5 cm long, occasionally absent; 

bracts subulate, 6-8 mm long; pedicels 1.5-4 cm long. Floral 

tube yellow green, villous; calyx lobes subulate, 5-8 cm long, 

villous; corolla funnelform, subventricose, 16-21 mm long, 

tube diameter 7-11 mm medially and 5-9 mm in throat, red, 

outside pilose-villous, inside glabrous, lobes 1- 1.5 mm long, 

red filaments glabrous; style glandular-puberulent and/or 

puberulent. Capsule globose, 5-7 mm long, 5-6 mm wide; 

seeds ellipsoid, 0.8 x 0.4 mm, brown, striate. 

DISTRIBUTION.-Pearcea hispidissima has been collected at 

600-1200 m elevation on the eastern Andean slopes in 

Ecuador, and most collections come from the region of Rio 

Bobonaza east of Puyo, the provincial capital of Nap0 (Figure 

9). The single Colombian collection comes from very humid 

montane forest at 1900 m elevation close to a steep cliff. 

REPRESENTATIVE COLLECTIONS.-COLOMBIA. Caqueta: 

Florencia-Guadalupe road, Londono & Kvist 158 (AAU, COL, 

TULV, US). 

ECUADOR. Moronu-Santiago: 9 km N-NW of Taisha, 

Brandbyge & Asanza 31 943 (AAU). Napo: Hollin-Loreto 

road, km 17, Hurtado & Shiguango 1656 (MO). Pastaza: 

Veracruz, Indillama, Lug0 S. 28 (GB, SEL), 1083 (GB, SEL); 

Banios-Mera road, km 35,Olllgaard et al. 9270 (MU). Total of 

20 collections seen. 

DISTINGUISHING FEATURES.-The strongly anisophyllous 

leaves and the dorsiventral shoots distinguish Pearcea hispidissima. In addition, the species is characterized by having mostly 

ovate leaves with glabrous upper surfaces and nearly entire 

margins, villous inflorescences, and subulate calyx lobes. 

VARIATION.-one specimen (Asplund 9523) is unusual in 

having nearly equal leaf pairs. The single collection from the 

department of Caquea in Colombia is similar to the Ecuadorian 

specimens. This similarity is unusual in Pearcea because plants 

Puyo-Veracruz road, km 4, Wiehler 71 76 (SEL, holotype).] 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 84 31 

FIGURE 18 (opposite page).-Pearcea grandifolia: (1, habit and inflorescences; 

b, indument of lower leaf surface; c, lateral view of corolla; d, laterally opened 

corollashowing androecium; e, calyx, nectary, and gynoecium (front calyx lobe 

removed); f, calyx, nectary, and young capsule; g, seeds. All from Schunke 

7558 tUSl. 

from the periphery of a species’ range usually have some 

distinct features (see “Variation” preceding key, above). 

TYPIFICATION.-According to Wiehler (1978), isotypes are 

located in K, NY, SEL, and US, but we could not find them and 

possibly they were never distributed. 

10. Pearcea hypocyttiflom (J.D. Hooker) Regel 

FIGURE 7 

Pearcea hypocyrtiflora (J.D. Hooker) Regel, 1867:388. Gloxinia hypocyrtiflora J.D. Hooker, 1867, pi. 5655. Isoloma hypocyrtiflorwn (J.D. Hooker) 

Bentham, 18761002 (“hypocyrtiflora”).-Hoehne, 1970323, pl. 161. [w: Based on cultivated material, originally collected by Jameson in 

eastern Ecuador, Rov. Napo, Archidona, Jameson 786 (K, lectotype selected 

Erect stems 5-20 cm long, unbranched, but with long 

rhizomes and adventitious roots, yellow brown, villous. Leaves 

subequal to unequal in each pair, blades ovate, less commonly 

cordate or oblong, 2.5-17 x 2-10 cm, rugose, above and 

below appressed pilose, often purplish, primary and secondary 

veins often yellowish and contrasting with darker green or 

purplish areas between veins. Inflorescences of 1-2 flowers, 

villous, yellow green to purple; peduncles 0-0.5 cm long; 

bracts ovate, sometimes to 5 mm long but usually absent or 

caducous; pedicels 4-7 cm long. Floral tube villous; calyx 

lobes erect, rotund-ovate, 1-2 mm long; corolla tube urceolate 

(see photograph by Dunn in Anonymous, 1991:15), 16-23 mm 

long, diameter 8-15 mm medially and 2-3 mm in throat, 

outside red, villous, inside puberulent or glandular-puberulent, 

lobes 1-2 mm long, red; filaments puberulent; style puberulent 

or glandular-puberulent, stigma weakly bilobed. Capsule and 

seeds not seen. 

DISTRIBUTION.-Pearcea hypocyrtiflora is currently known 

from an area -50 km long near the Nap0 river, below 700 m 

elevation in the Ecuadorian province of Nap0 (Figure 7). 

REPRESENTATIVE COLLECTIONS.-ECUADOR. Napo: 

Bna-Puyo, Besse et al. 1671 (SEL); Shinguipino, 8 km SE of 

Bna, Grub6 et al. 1503 (K); Ahuano at Rio Nap0 25 km E of 

Puerto Napo, Lug0 S. 235 (GB, SEL); 8 km downstream from 

Puerto Misahualli, Palacios et al. 484 (AAU, US); Santa Rosa 

along Rio Napo, Prance 26587P (SEL). Province Unknown: 

spruce s.n. (K). Total of 18 collections seen. 

DISTINGUISHING FEATURES.-Pearcea hypocyrtiflora 

shares urceolate corollas only with P. bella and P. schimpfii. 

Pearcea bella differs by having corollas -1 cm rather than -2 

crn long and by having leaves with darker areas along the 

lateral veins on the upper leaf surfaces rather than between 

them. Pearcea schimpfii differs by having glabrous rather than 

herein; BM, E. MPU, iso~ectotypes).] 

appressed pilose and rugose upper leaf surfaces and elliptic 

rather than ovate leaves. Vegetatively, however, the often large, 

ovate, purplish, and yellow-veined leaves distinguish P. 

hypocyrtijlora from all other species (see illustrations in 

Batcheller, 1992:19; Boggan, 1990:9; Striepens, 1991:16). 

TYPIFICATION.-NO specimen labeled as holotype has been 

seen. Possibly there were no specimens of the species preserved 

while it was under cultivation at the Royal Botanic Gardens, 

Kew, where it was drawn by W. Fitch and described by J.D. 

Hooker. The Kew plants came from Messrs. Veitch who grew 

them from a specimen collected in Ecuador by Richard Pearce. 

Hooker (1867) cites a Jameson collection, which he also 

annotated with the figure number from Curtis’s Botanical 

Magazine. Daniel Oliver, keeper at Kew at the time, also noted 

on the same sheet that the Jameson collection was the “same or 

(very close ally) sent by Messrs. Veitch 4/67. [He] called it 

Hypocyrta brevicalyx & sent it to Mr. Fitch to figure.” Oliver 

later added to his note: “This name [i.e., Hypocyrta brevicalyx] 

gives place to Gloxinia hypocyrtiflora, Bot. Mag.” The name 

Hypocyrta brevicalyx was used by Henry James Veitch in the 

plant list in Gardener’s Chronicle of 4 May 1867 for a plant 

exhibited at the Royal Horticultural Show in April 1867 

(Veitch, 1867). This name was mentioned by Regel when he 

transferred Gloxinia hypocyrtiflora to his new genus Pearcea, 

but by the typography in the Regel paper, it appears that Regel 

did not consider Hypocyrta brevicalyx to be a name that had 

any standing. The name does appear, however, in Index 

Kewensis (Jackson, 1895) where it was relegated to being a 

synonym of Pearcea hypocyrtijlora. We believe, as Regel did, 

that the name Hypocyrta brevicalyx has no standing beyond 

referring to a horticultural novelty. 

11. Pearcea intermedia: L.P. Kvist & L.E. Skog, new species 

FIGURES %20g 

Differt a P. sprucei (Rusby) L.P. Kvist & L.E. Skog foliis 

ovatis glabris pagina superioribus flavi-viridibus reticulatis 

ordinatiis, a P. schimpfii Mansfeld corollis subventricosis 

differt. 

Stems frequently decumbent, to 50 cm long, basal diameter 

to 4 mm, apparently unbranched but with adventitious roots, 

basally glabrescent to apically villous (apart from glabrous, 

violet nodes), trichomes yellow brown with -10 large, flat 

cells; internodes 2-5 cm long. Leaves equal in each pair, blades 

oblique, ovate-elliptic-obovate, 4-10 x 2-5 cm, base obtuseattenuate-cuneate, apex acute, margin serrate, above with 

regular pattern of green blotches surrounded by yellow green 

areas along primary, secondary, and, particularly, reticulate 

tertiary veins, glabrous, below grayish green, glabrous apart 

from villous yellow-green veins, lateral veins 8-10 per side; 

petiole 2.5-4 cm long. Inflorescences of 1-4 flowers, yellow 

brown, villous; peduncles 0.3-0.8 cm; bracts 2, lanceolate, 

15-25 x 3-6 mm; pedicels 2-4 cm long. Floral tube conic, 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1.0-1.5 mm long, yellow brown, villous; calyx 7-10 mm long, 

villous, lobes erect, lanceolate, 5-8 mm long and 2-3 mm 

wide at base, apex acute-acuminate; corolla 16-20 mm long, 

tube funnelform, subventricose, 16-1 8 mm long, diameter 

7-12 mm medially and 5-10 mm in throat, outside red, 

sericeous, inside yellow, occasionally glandular-puberulent, 

limb with lobes subequal, rotund, 1-2 mm long and 2-4 mm 

wide at base, inside glandular-puberulent and with darker dots, 

apex obtuse-truncate; stamens subincluded, filaments 13- 18 

mm long, glabrous basally to pubescent medially and glandular-puberulent apically, anthers -2 x 1.5 mm; staminode 1-3 

mm long; nectariferous disk of 5 glands, all -0.8 mm high and 

0.8 mm wide at base; ovary -3/4 inferior, 3-4 mm long, 

diameter 2-3 mm, sericeous, style subincluded, 14-18 mm 

long, glandular-puberulent, stigma stomatomorphic. Fruits and 

seeds not seen. 

TYPE.-ECUADOR. Napo: Baeza-Tena road, km 40, 1900 

m, 29 Mar 1979, Lojtnant & Molau 11494 (AAU, holotype; GB, 

isotype). 

ETYMOLOGY.-The specific epithet refers to this species’ 

leaves being similar to those of the urceolate-flowered P. 

schimpfii but having funnelform corollas similar to those found 

in the majority of Pearcea species. 

DISTRIBUTION.-Pearcea intermedia is known from the 

region near Baeza on the northeastern Ecuadorian Andean 

slopes, between 1700 m and 1900 m elevation (Figure 7). 

OTHER COLLECTIONS.-ECUADOR. Napo: Baeza-Tena road, 

S slope of Cordillera de Guacamayos above Jondachi, Hurling 

& Andersson 16357 (GB, us); Baeza-Tena, Holm-Nielsen et al. 

16277 (MU); Napo-Lago Agrio road, Kvist 60386 (MU, QCA, 

US); Baeza-Lago Agrio road, km 16, Santa Rosa, MacBryde 

720 (US); road to Tena, km 45 from N, Webster 23238 (DAY 

us); Baeza-Lago Agrio road, km 46, Wiehler & Masterson 

79300 (SEL). Total of seven collections seen. 

DISTINGUISHING FEATURES.-The subventricose corollas of 

Pearcea intermedia are similar to those of several other 

Pearcea species, e.g., the common P. sprucei. Vegetatively, the 

species is strikingly similar to P. schimpfii, which, however, 

differs by having urceolate rather than subventricose corollas. 

Both species have ovate, glabrous leaves with a reticulate 

yellow green pattern surrounding green blotches on the upper 

surface, and the trichomes of the stem apex and inflorescences 

in both consist of unusually large, flat cells. Pearcea schimpfii 

has also been collected in the province of Napo, but in different 

parts of that province (Figure 7) and at different altitudes. 

12. Pearcea purpurea (Poeppig) L.P. Kvist & L.E. Skog, 

new combination 

RGURE 12 

Rhyridophyllum purpureum Poeppig, 18406, pl. 206 [“Rytidophyllum”]. 

Moussonia purpurea (Poeppig) Hanstein, 1865:289. Isoloma purpureum 

(Poeppig) Bentham, 1876: 1002. Parakohleria purpurea (Poeppig) Wiehler, 

1978% [w: Peru, Hubuco, Cuchero, Poeppig 1632 (W, holotype; BM. F, 

G. P, w (3). isotypes).] 

Rhyridophyllum melasroma Poeppig, 1 Ea.7 r‘Rytidophyllum’7. Moussonio 

melastoma (Poeppig) Hanstein, 1865:288. Kohleria melastoma (Poeppig) 

Cuatrecasas. 1935:258 [reprint p. 551. Parakohleria melastoma (Poeppig) 

Wiehler, 1978% [m: Peru, Huanuco, Pampayaco, Poeppig 1239 (w, 

holotype)]. 

Kohleria weberbaueri Fritsch, 191 3:430. Parakohleria weberlmueri (Fritsch) 

Wiehler, 1978:lO. 1%: Peru, Junfn, Tarma, E of Huacapistana, Weberbauer 2109 (B, holotype, no longer extant: MOL, lectotype, selected herein).] 

Stems 40-100 cm long, occasionally branched, grayish 

villous. Leaves nearly equal in each pair, blades ovate, less 

commonly lanceolate, 4-15(-22) x 2-8 cm, margin serrulate, above dull green, velutinous, less commonly hirsute, 

below grayish green, velutinous, veins pilose-villous below. 

Inflorescences with (2-)5- 12(- 16) flowers, grayish green to 

purplish, villous; peduncles 3- 10 cm long; bracts 2-4(-6), 

lanceolate, 2 lowest 0.5-1.5 cm long, others smaller; pedicels 

0.5-2.5 cm. Floral tube grayish green, villous; calyx lobes 

usually reflexed, ovate, rarely erect, lanceolate, 2-6 mm long; 

corolla 15-22 mm long, tube funnelform, subventricose, 

diameter 7-12 mm medially and 3-9 mm in throat, red, 

outside pilose-sericeous, inside glandular-puberulent, lobes 

2-3 mm long, 2 upper lobes red, 3 lower lobes yellow with 

brown area near margin or brown dots centrally; filaments 

glabrous; style glandular-puberulent and/or puberulent. Capsule subglobose, 4-6 mm long, diameter 4-5 mm; seeds ovoid, 

-0.5 mm long, brown black, striate. 

DISTRIBUTION.-Pearcea purpurea occurs in central Peru, 

where most collections come from the department of Huanuco, 

fewer from Pasco, two from Junin and one from San Martin 

(Figure 12). It has been found at 600-2400 m elevation. 

REPRESENTATIVE COLLECTIONS.-PERU. Junin: TarmaChanchamayo border, 45 km S of San Ramon, Rio ’Mumayo 

drainage, Rondayacu, Foster & Smith 9161 (F, MO, US). Huanuco: Tingo Maria, Asplund 12073 (s); 5 km S of Tingo Maria, 

Schunke V. 3262 (COL, F, G, us, USM); Tingo Maria-Huanuco 

road, km 8, Skog et al. 5147 (US, USM). Pasco: 30 km SW of 

Oxapampa, Rio Paucartambo, Madison 955 (F, GH); SW of 

Oxapampa on road to Maria Teresa and Llaupi, Foster 7602 (F, 

MO, US, USM); Oxapampa-Villa Rica road, km 7 SE from 

Miraflores crest, Foster 9112 (F, MO, US). Sun Martin: 

Infantes 5524 (HBG). Department Unknown: Matthews 150 

(FI-W, G, P (2)), 2012 (BM, CGE, E, K (2)). Total of 33 COlleCtiOnS 

seen. 

DISTINGUISHING FEATURES-PearCea purpurea is most 

similar to P. reticulata and to some populations of P. sprucei 

that also have inflorescences with peduncles several times 

longer than the pedicels. Both differ from P. purpurea by 

having a sericeous rather than a villous indumentum on the 

inflorescences as well as on the veins of the lower leaf surfaces. 

Although some populations of P. sprucei have a villous 

indumentum, these have peduncles only to 2 cm long (see 

“Variation,” in P. sprucei). The corollas of Pearcea purpurea 

are notable for their contrasting upper red lobes and the three 

lower yellow lobes with brown areas either near the margin or 

centrally. 
NUMBER 84 33 

VARIATION.-The type collection of Pearcea purpurea has 

lanceolate leaves instead of the ovate leaves usually found in 

this species, but the type is otherwise similar to the specimens 

of the bulk of the collections. Only the collections of Matthews 

have similar lanceolate leaves. 

Specimens from the department of Pasco (e.g., Foster 9112) 

often have lanceolate calyx lobes similar to those of P. sprucei 

but have villous stems and inflorescences as in other specimens 

of P. purpurea, as well as the brown area on the three lower 

corolla lobes. This variation probably reflects introgression 

between P. purpurea and P. sprucei (see “Variation,” preceding key). 

lected a neotype for Kohleria weberbaueri because, at that 

time, no additional specimens of the type, Weberbauer 2109, 

had been found to lectotypify the species. Recently, a duplicate 

of the type was discovered at MOL. This specimen of 

Weberbauer 2109 is selected as lectotype and supersedes the 

neotypification. 

NOMENCLATURE AND TYPIFICATIoN.-Wiehler (1978) se13. Pearcea reticulaca (Fritsch) L.P. Kvist & L.E. Skog, 

new combination 

FlGURES 10, 19 

Kohleriu rericuluru Fritsch, 1913:428. [w: Ecuador, ’hngurahua, BaClos, 

spruce 5841 (W, holotype; BM. CGE, E. G. K. NY. P, WAG, isotypes).] 

Stems to 150 cm long, frequently branched, yellow green, 

apically sericeous-lanate. Leaves nearly equal in each pair, 

blades ovate-lanceolate, 9-19 x 4-7 cm, margin semlate, 

above dull green, hirsute, below lighter green, pilose, primary, 

secondary, as well as tertiary veins sericeous and distinctly 

yellowish. Inflorescences with 5- 12(-20) flowers, yellow 

green, sericeous-lanate; peduncles 4- 15 cm long; bracts 

usually 6, ovate-lanceolate, 2 lowest 0.5-2.5 cm long, others 

smaller; pedicels 0.5-2.5 cm long. Floral tube yellow green, 

villous; calyx lobes ovate, 3-4(-6) mm long and 3-4 mm 

wide, sericeous; corolla (17-)20-25 mm long, tube funnelform, subventricose, diameter (7-)12-17 mm medially and 

5-11 mm in throat, red, outside villous, inside glandularpuberulent, lobes 2-3 mm long, yellow with darker dots (see 

photograph in Kvist, 1991: 16); filaments glabrous; style 

puberulent. Capsule broadly globose, 3-5 mm long, diameter 

4-6 mm; seeds broadly ellipsoid, -0.5 mm long, black, striate. 

DIsTRlBuTIoN.-Pearcea reticulata occurs on the eastern 

Andean slopes of Ecuador and in the adjacent Colombian 

department of Putumayo (Figure 10). The species is common 

in the Rfo Pastaza valley of Ecuador, where it often grows in 

secondary vegetation (pers. obs.), but it appears to be much 

rarer both to the south and to the north. Only the two cited 

collections are known from Colombia. Pearcea reticulata has 

been collected at 500-1600 m elevation but is usually found at 

900-1400 m. 

REPRESENTATIVE COLLECTIONS.-COLOMBIA. Putumayo: 

Between Mocoa and Sachamates, Cuatrecasas 11402 (COL, F, 

US); Mocoa, Schuftes & Smith 2054 (COL, F, GH, US). 

ECUADOR. Moronu-Santiago: Sangai National Park, km 17 

N of Macas, Bohlin et al. 1485 (GB). Napo: Baeza-Lago 

Agrio, km 49, Balslev & Madsen 10284 (AAU, F, MO, NY, SEL 

(2)). Pastaza: Mera, Lug0 S. 1125 (GB, SEL); Banios-Mera 

trail, Mexia 6824 (F, uc, us). Tungurahua: Trail between 

Machay and Rio Mapoto, Penland & Summers 112 (F, GH, US); 

Rfo Negro, Hurling & Andersson 17238 (GB, US); Pastaza 

Valley, Kvist 60315 (AAU, COL, GB, MO, NY, QCA, QCNE. US). 

Total of 53 collections seen. 

DISTINGUISHING FEATURES.-The long peduncles distinguish Pearcea reticulata from all other species except P. 

grandifolia from northern Peru, P. purpurea from central Peru, 

and populations of P. sprucei found in southern Peru and 

Bolivia and in the department of Caqueta in Colombia. Pearcea 

grandifolia (Figure 18) differs from P. reticulata (Figure 19) 

by having much larger leaves, P. purpurea differs by having 

smaller flowers and having generally a grayish green rather 

than a yellow green indumentum, and P. sprucei differs by 

having lanceolate calyx lobes rather than ovate lobes. Plants of 

P. reticulata are usually taller and stouter than any other 

sympatric species, and its size combined with the conspicuous 

yellow green indumentum distinguishes the species even 

vegetatively. 

VARIATION.-Pearcea reticulata specimens from the Rio 

Pastaza valley in Ecuador cannot be confused with any other 

species of Pearcea. To the north and to the south, however, the 

specimens of P. reticdata gradually become more similar to P. 

sprucei and have smaller flowers, shorter peduncles, and a 

sparser, but yellow green, indumentum. These specimens are 

nevertheless distinct from P. sprucei (see “Variation,” preceding key). 

Acosta-Solfs 7432 is a remarkable collection with inflorescences and calyces typical for Pearcea reticulata but with 

flowers 3-3.5 cm long, similar to those found in Kohleria 

hirsuta (Kunth) Regel. The plant appears to be an intergeneric 

hybrid between those two species. In Ecuador K. hirsuta has so 

far been collected only a few times in the province of 

Morona-Santiago (Kvist and Skog, 1992). The exact provenance of Acosta-Solis 7432 is unclear, but the specimen comes 

either from Morona-Santiago or from Zamora-Chinchipe 

further to the south. 

Wiehler (1976) reported that he hybridized Kohleria 

reticufata with Kohleria hirsuta. We have not seen material of 

the collection he used for this cross, but it came from the Rfo 

Pastaza valley near Banos where Pearcea reticulata is very 

common, and it is consequently very likely that it actually was 

this species that he crossed with Kohleria hirsuta. He reported 

zero percent pollen stainability, indicating the hybrid was 

completely sterile (he also reported zero percent pollen 

stainability for a hybrid between Parakohleria weberbaueri 

(apparently Pearcea sprucei var. parviflora) and Kohleria 

spicata). Later (Wiehler, 1983) he established a generic name, 

x Paleria, for hybrids between Parakohleria and Kohleria. 
NVINOSHUMS SNOLLn9INLN03 CAI, AWL09 
NUMBER 84 35 

FIGURE 19 (opposite page).-Pearcea reticulata: a, habit and inflorescences; b, 

indument of lower leaf surface; c, inflorescence; d, lateral view of flower, e, 

laterally opened corolla showing androecium;f, calyx, nectary, and gynoecium 

(front calyx lobes removed). a,b from Harling 3197 (S); c-f from Dodson & 

Thien 1926 (Ws). 

14. Pearcea rhodotricha (Cuairecasas) L.P. Kvist & 

L.E. Skog, new combination 

FIGURES 3c, 8 

Diastema rhodotrichum Cuatrecasas, 1935252 [reprint p. 491, pl. 20. 

Parakohleria rhodotricha (Cuatrecasas) Wiehler, 1978:7, 8, pl. 1C. [w: 

Ecuador, Napo, E Santa Rosa ad Cotapino, kern 496 (MA, lectotype).] 

Stems to 50 cm long, unbranched, villous with 3-5 mm long 

brown red trichomes. Leaves frequently subequal to unequal in 

pairs, blades ovate, rarely lanceolate, 6-12(-18) x 2-6(-8) 

cm, margin serrate, with conspicuous 3-5 mm long trichomes, 

above dull green, appressed pilose, below yellow green to 

purple, pilose. Inflorescences of (1-)4-8(-12) flowers, purplevillous; peduncles 0.5-3 cm long, occasionally absent; bracts 

lanceolate, less commonly ovate, 5-8(-12) mm long; pedicels 

1.5-2.5 cm. Calyx purple, villous, lobes lanceolate, 3-7 mm 

long; corolla 11-15 mm long, tube funnelform, subventricose 

to ventricose, diameter 5-6 mm medially and 3-4 mm in 

throat, red and usually black on the upper -5 mm including 

limb, rarely entirely red, outside villous, inside sparsely to 

densely glandular-puberulent, lobes 1-1.5 mm long, usually 

black; filaments 8-10 mm long, puberulent, rarely nearly 

glabrous; style 8-11 mm long, puberulent and glandularpuberulent. Capsule globose, -3 mm long, diameter -3 mm; 

seeds subglobose, -0.3 mm long, black, striate. 

DISTRIBUTION.-Pearcea rhodotricha is known from areas 

below 600 m elevation in the Ecuadorian Amazon region. Most 

of the collections have been made near the Nap0 and Bobonaza 

rivers (Figure 8). 

REPRESENTATIVE COLLECTIONS.-ECUADOR. Napo: 

herto Misahualli, Besse el al. 68 (QCA, SEL); San Jose, Isern 

596 (MA); Lago Agrio, Lug0 S. 3147 (GB, SEL); Lago Agrio-El 

Chaco road, Lug0 S. 3503 (GB, SEL); Lago Agrio-Baeza road, 

km 16, Santa Cecilia, Lug0 S. 3562 (GB, SEL); new road toward 

Rio Arajuno, Neil1 6556 (us). Pastaza: Rio Bobonaza, 8 km 

NW of Sarayacu, Lug0 S. 5327 (GB, US); Rio Bobonaza, Spruce 

4962 (BM, K (2), W). Total of 24 collections seen. 

DISTINGUISHING FEATURES.-Pearcea rhodotricha is easily 

distinguished from all other Pearcea species by having leaf 

margins with conspicuous trichomes to 5 mm long. Other 

characteristic features are the ovate, serrate leaves, the 

purple-villous indumentum of stems and inflorescences, and 

the red and apically black corollas (see illustration in Ertelt, 

199 1 :2 1). 

VARIATION.-TWO collections (Lugo S. 3503, 3562) from 

Nap0 Province differ from other specimens of Pearcea 

rhodotricha by having lanceolate instead of ovate leaves and by 

having red rather than redblack corollas. A third collection 

(Lugo S. 3147) has ovate leaves and entirely red corollas. These 

plants, however, are considered as variants of Pearcea 

rhodotricha because they are similar in other aspects, e.g., by 

having conspicuous trichomes on the leaf margins. 

A collection (Hirtz et al. 3116) from the province of 

Morona-Santiago in southern Ecuador has characters intermediate between Pearcea rhodotricha (e.g., similar although 

shorter trichomes and a similar leaf shape and inflorescence) 

and P. sprucei (e.g., rugose leaves and flowers without a black 

apex). Because this intermediate collection was made 200 km 

south of the southernmost previously known occurrence of P. 

rhodofricha and at 1500 m elevation, whereas P. rhodotricha 

has so far never been collected above 700 m, this collection 

may represent a new species. 

15. Pearcea schimpfii Mansfeld 

FIGURES 7, 200-f 

Pearcea schimpfii Mansfeld, 1936 149. Parakohleria schimpfii (Mansfeld) 

Wiehler, 1978:9. [w: Ecuador, Pastaza, W of Mem, Schimpff 704 (B, 

holotype, no longer extant; GH, lectotype selected by Wiehler (1978); G. M. 

MO, TRT, isolectotypes).] 

Terrestrial or epilithic stems to 70(-100) cm long, glabrous 

to pilose near apex, trichomes yellow brown, of -10 large, flat 

cells. Leaves usually nearly equal in each pair, blades elliptic, 

less commonly lanceolate, 4-12 x 2-6 cm, margin serrate, 

upper surface often with conspicuous, regular pattern of green 

blotches surrounded by reticulate yellow green areas, glabrous. 

Inflorescences of 1-3(-5) flowers, yellow brown to purple, 

villous; peduncles 0-2 cm long; bracts lanceolate, 7-15 mm 

long; pedicels 1-2.5 cm. Floral tube yellow brown, villous; 

calyx lobes erect, ovate, 3-8 mm long; corolla urceolate, 

15-22 mm long, tube diameter 8-17 rnm medially and 2-3 

mm in throat, red to orange red, outside sericeous-villous, 

inside densely puberulent and often glandular-puberulent, 

lobes 1.5-2.0 mm long, red; stamens included, filaments 9- 18 

mm long, puberulent and often glandular-puberulent; style 

included, 8-12 rnm long, densely puberulent and glandularpuberulent, stigma weakly bilobed. Capsule subglobose, -4 

mm long, diameter -5 mm; seeds not seen. 

DISTRIBUTION.-PearCea schimpfii is known only from a 

small area between 1 100 m and 1500 m elevation on the eastern 

Andean slopes in Ecuador (Figure 7). Most collections come 

from the Rio Pastaza valley, where the species was observed 

growing on shady, humid cliffs along streams in forest-covered 

ravines (Kvisf 60319). 

REPRESENTATIVE COLLECTIONS.-ECUADOR. MoronaSantiago: 27 km SE of San Juan Bosco, Gentry et al. 30921 

(MO, SEL). Napo: Hollin-Loreto road, km 17, Hurtado & 

Shiguango 1638 (us), 1751 (MO). Pastaza: Mera, sandy 

shore of Rio Mangayacu, Asplund 18673 (s); Colonia Jaitiva, 7 

km N of Mera, Harling & Andersson 16916 (GB, US). Tungurahua: Pastaza valley between Banos and Cashuroc, Hitchcock 
36 TANY 
NUMBER 84 37 

RGURE 20 (opposite page).-Pearcea schimpfi: a, habit and'inflorescences; b, 

indument of lower leaf surface; c, upper leaf surface; d, flower; e, laterally 

opened flower showing androecium; f, calyx, nectary, and gynoecium (front 

calyx lobes removed). Pearcea intermedia: g, flower. a-c from Asplund 18673 

(S); d-f from Lug0 417 (GB); g from Hurling & Andersson 16357 (GB). 

21852 (GH, NY, US); Pastaza valley, Kvist 60319 (MU, NY, QCA, 

QCNE, us). Total of 20 collections seen. 

DISTINGUISHING FEATURES.-The combination of urceolate 

corollas and adaxially glabrous, serrate leaves distinguishes 

Pearcea schimpfi (Figure 20a-f; illustration in Kvist, 

1991 :21) from all other species. Pearcea intermedia has similar 

vegetative characters but has ventrimse corollas (Figure 20g). 

NOTES.-Mansfeld (1 936) named the species for Heinrich 

Schimpff who collected in South America in the 1920s and 

1930s. Unfortunately, Mansfeld misspelled Schimpff s name, 

which he had spelled correctly when he honored the same 

collector with a species of Columnea in 1934. 

16. Pearcea sprucei (Britton) L.P. Kvist & L.E. Skog, 

new combination 

RGURES 6,21 

Isoloma sprucei Britton, 1896:97. "Qpe: Bolivia, La Paz, Mapiri, Bang I541 

(NY, holotype; BM, E, F, G (2). GH (2), K. NY. PH, US (2). isotypes).] 

Stems 30-100(-150) cm long, rarely to frequently 

branched, yellow green or purplish, sericeous, rarely villous. 

Leaves equal to subequal in each pair, blades weakly to 

conspicuously oblique, lanceolate, less commonly ovate, 

elliptic or oblanceolate, (2-)5-12(-16) x (1-)2-4(-6) cm, 

base usually attenuate, less commonly obtuse or cuneate, 

margin sermlate, rarely serrate, apex acute to acuminate, often 

rugose, above dark green to purplish, often with conspicuous 

primary and secondary veins, velutinous, pilose or hirsute, 

rarely nearly glabrous, below often purple, velutinous-pilose 

with sericeous veins. Inflorescences with 1-8(-18) flowers, 

sericeous, less commonly villous; peduncles 1-2(-9) cm long, 

rarely epedunculate; bracts 2(-6), lanceolate or ovate, 2 lowest 

0.5-2 cm long, other bracts smaller; pedicels (0.5-)1.5-2.5 

(-4.5) cm long. Floral tube villous; calyx lobes erect or less 

commonly reflexed, lanceolate, less commonly subulate, rarely 

ovate, (2-)5-11 mm long; corolla (12-)15-23 mm long, tube 

funnelfonn, subventricose, diameter 6- 1 1 mm medially and 

4-10 mm in throat, red, often yellow ventrally and basally, 

rarely entirely yellow, outside pilose-villous, inside occasionally sparsely glandular-puberulent, lobes 1.5-3 mm long, often 

yellow with red purple dots, rarely entirely red; filaments 

glabrous, rarely sparsely puberulent; style often puberulent 

and/or glandular-puberulent, rarely glabrous. Capsule 

subglobose, 3-5 mm long, diameter 3-7 mm; seeds ovoid, 

0.5-0.7 mm long, brown black, striate. 

DISTRIBUTION.-Pearcea sprucei is by far the most widespread and commou species in the genus ranging along the 

eastern Andean slopes from the southern Colombian department of Caqueta to the department of Cochabamba in Bolivia 

(Figure 6). It is the sole member of the genus extending into the 

inter-Andean valleys, to the western slopes of southern 

Ecuador, and to the upper Rio Magdalena watershed in the 

department of Huila in southern Colombia. Here, however, P. 

sprucei has only been collected at 2000 m elevation, close to 

the ridge of the eastern Colombian Cordillera that separates the 

Rio Magdalena watershed from the Amazon basin. The range 

of P. sprucei is nearly identical to the range of the entire genus 

(Figures 4, 6). and consequently P. sprucei is sympatric with 

nearly all other Pearcea species (Figures 7-12). Only P. 

abunda seems also to occur in areas where P. sprucei has not 

yet been found, because this species ranges considerably farther 

to the east in Amazonian lowland rainforest (Figure ll), 

whereas P. sprucei is restricted to more humid lowland 

rainforest along Andean slopes (Figure 6). 

Most collections of Pearcea sprucei come from the 

provinces of Nap0 and Pastaza in eastern Ecuador (Figure 6). In 

Peru, most collections come from the region near Tingo Marla 

in the department of Huanuco, from the adjacent northwestern 

part of Ucayali, and from Cuzco farther to the south. Pearcea 

sprucei, however, as well as the entire genus, seems rare or 

rarely collected in northern Peru (Figures 4-6). 

On the western Andean slopes of Ecuador, P. sprucei has 

been found in the southern and central provinces of Chimborazo, Canar, and El Oro (Figure 6), but not north to Pichincha, 

Imbabura, or Carchi. The species may have spread from the 

eastern to the western Andean slopes fairly recently and may 

not yet have reached the extensive humid forests farther to the 

north. Collections of P. sprucei from the inter-Andean valleys 

of the south-central provinces of Canar and Azuay (Figure 6) 

strongly suggest that the dispersal across the Andes occurred 

here. This also is likely because the Andean Cordilleras are 

lower here and are less of a barrier than in northern Ecuador. 

Pearcea sprucei has been collected at 300-2800 m elevation 

but seems rare above 2000 m. While collecting on the eastern 

Andean slopes along the border of Ecuador and Colombia, we 

encountered the first P. sprucei populations at 2300 m (Kvist 

60251). Herbarium studies provided evidence that 2300 m is 

usually the upper limit throughout much of its range. The 

highest records of P. sprucei are from Ecuador and come from 

2600 m and 2800 m elevation from the western Andean slopes 

and interAndean valleys, respectively. 

DISTINGUISHING FEATURES.-Most Pearcea sprucei collections, particularly those from Ecuador and central Peru, are 

easily recognized by having most of the following characters: 

leaves lanceolate with a conspicuous and often pilose indumentum, blades more or less rugose, with conspicuous secondary 

veins, dark green upper surfaces and purplish lower surfaces, 

inflorescences also purplish and covered with an appressed 

indumentum, and peduncles shorter than pedicels. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Peduncles 

> 2cm 

1 a 

1-6) 

Blades 

ovate 

1 

b 

Blades 

rugose 

L 

C 

Blades 

purple 

d 

Calyx lobes 

reflexed 

L 

1 

1 

1 

e 

Bract width Inflorescen- 

> 5mm 

1 

z 

c f 

ces villous 

1 

1 

9 

Morphological forms (morphs) of var. sprucei and 7) var. parviflora 

Number of 

collections 

13 

62 

20 

16 

15 

14 

18 

FIGURE 2 1 .-Frequencies of 7 characters in 7 morphological forms of P earcea sprucei. 1-6 show var. sprucei; 

7 shows var. pawflora. Amount of hatching in each rectangle is relative to frequency of character state in each 

morph: a, presence of peduncles longer than 2 cm; b, leaves with ovate blades; c, leaves with rugose blades; d, 

leaves with purple-colored blades particularly on lower surface: e, calyx lobes reflexed (rather than erect); f, 

inflorescences in which largest bracts are more than 5 mm wide; g, inflorescences, stems, and veins of the lower 

leaf surface with patent, usually villous indumentum rather than appressed and usually senceous indumentum. 

Populations of Pearcea sprucei, however, may be confused 

with several other species because specimens may have only 

some of the above-mentioned attributes. Short-peduncled P. 

sprucei populations may be confused with species such as P. 

hispidissima, P. rhodotricha, and particularly P. strigosa (see 

those species). In most cases apparent reasons for the similarity 

may include hybridization and introgression (see “Variation,” 

preceding key). 

Long-peduncled populations of Pearcea sprucei also may be 

confused with P. purpurea and P. reticulata, which both 

develop peduncles more than 5 cm long and longer than the 

pedicels, but they differ from P. sprucei by having ovate rather 

than lanceolate calyx lobes and in several other characters. In 

addition, they are not sympatric with the long-peduncled 

populations of P. sprucei, which are found in Colombia, 

Bolivia, and nearby Peru (Figure 21a). whereas P. purpurea 

(Figure 12) and P. reticulata (Figure 10) occur in central 

Peru and Ecuador, respectively. Unfortunately, both species 
NUMBER a4 39 

may hybridize with P. sprucei, causing identification problems 

in both Ecuador and Peru (see P. purpurea, P. reticulafa, and 

“Variation,” preceding key). 

VARIATION.-PeUrCea sprucei is extremely variable, partic - 

ularly in the vegetative characters, although the obvious and 

even the more subtle details of the corolla characters remain 

fairly stable throughout. Apparent reasons for the vegetative 

variation are discussed herein (see “Variation,” preceding key). 

Populations from upper montane forests and cloud forests 

often have shorter stems and internodes, smaller leaves, and a 

denser and more villous indumentum. The latitudinal and 

longitudinal variation, however, is much more pronounced than 

the altitudinal variation. 

Figure 2 1 illustrates the frequencies of different characters 

throughout the range of Pearcea sprucei. Seven morphologically distinctive forms or morphs are recognized (numbered 1 

to 7). In some areas these morphs are separated by natural 

bamers, such as mountain ranges higher than -2500 m, e.g., in 

central Peru (morphs 4 and 5 and morphs 5 and 6). In other 

areas there is no obvious physical barrier, e.g., between 

Colombia and northern Ecuador (morphs 1 and 2), and in 

southern Ecuador (morphs 2 and 3). In both these areas the 

transitions seem more gradual. In addition, the plants in 

southern and western Ecuador and in northern Peru (morph 3) 

are less distinctive as a group than are the other six morphs 

shown in Figure 21. 

In Bolivia and southern Peru there are two sympatric 

Pearcea sprucei morphological forms (morphs 6 and 7 in 

Figure 21). Britton (1896) and Rusby (1896) described these as 

Isoloma sprucei and Isoloma parviflorum, respectively. This 

was quite understandable because the morphs appear to be 

consistently distinct in Bolivia; the former (morph 6) has long 

peduncles, a sparse indumentum, and lanceolate blades, and the 

latter (morph 7) has short peduncles, a villous indumentum, 

and ovate blades. Herein they are treated as varieties (Figure 6) 

because the morphs are distinct in Bolivia but not farther to the 

north in central Peru, and because all the attributes that 

characterize them also can be found in other P. sprucei 

morphological forms, e.g., pedunculate inflorescences in 

Colombia (shared between morphs 1 and 6 in Figure 21) and 

ovate leaves, particularly in southern Ecuador (shared between 

morphs 3 and 7). 

NOMENCLATURE AND TYPIFICATION.-ISO~O~~ sprucei and 

Isoloma pawiflorum were described in 1896 in the same 

publication by Rusby and by Britton, respectively. Both were 

based on “atypical” Pearcea sprucei collections. Intermediate 

specimens are found to the north in Peru (Figure 21). There are 

consequently neither nomenclatural nor taxonomic arguments 

to decide which is the more appropriate epithet to use for the 

most common species in the genus Pearcea. We have selected 

P. sprucei because this epithet has been used most frequently in 

identifying collections of this species from Ecuador and Peru. 

16a. Pearcea sprucei var. parvijlora (Rusby) L.P. Kvist & 

L.E. Skog, new status 

FIGURES 6,21 

Isoloma parviflorum Rusby, 1896:97. Diastema pawiflorum (Rusby) Fritsch, 

1913:406.-Hoehne, 1970175, pl. 87. Parakohleria parviJlora (Rusby) 

Wiehler, 1978% [Type: Bolivia, La Paz, Mapiri, Bang 1563A (NY, 

holotype). duplicates distributed as Bang 1565 (G. GH (2). K, MO. NY. US (2), 

isotypes). J 

lsoloma patentipilosum 0. Kuntze, 1898:242. Kohleria patentipilosa (0. 

Kuntze) Schumann, 1900:386. [’I)pe: Bolivia, Cochabamba(?), Yungas, 

Kuntze s.n. (NY, holotype).] 

Isoloma flexuoswn Rusby, 1900.28. [Type: Bolivia, Cochabamba(?), Yungas, 

Rusby 2424 (NY, holotype; US, isotype).] 

Isoloma urticifolium Rusby, 1900:28. Kohleria urticifolia (Rusby) Fritsch, 

1913:430. [’Qpe: Bolivia, La Paz, Mapiri, Ruby 2422 (NY, holotype; BM. 

Diastema galcopsis Fritsch. 1913:405. [Type: Bolivia, Cochabamba(?), 

Yungas, Bang 544 (NY, lectotype selected by Wiehler, 1978; BM, E. F. G. GH. 

Stems frequently branched, villous with white to yellow 

green trichomes. Leaf blades ovate, (2-)4-lo(-15) x 1-5 

cm, margin serrate, rarely and only weakly rugose, rarely 

purplish, veins villous below. Inflorescences with 1-5 flowers, 

often on short shoots, villous, yellow green to white; the largest 

bracts to 6 mm wide; peduncles to 2 cm long. Calyx lobes often 

reflexed, lanceolate; corolla red, 1.5-2.5 cm long, outside 

villous. 

DISTRIBUTION.-Pearceu sprucei var. parviforu has been 

collected from southem Peru (Cuzco, Puno) and adjacent 

Bolivia (Cochabamba, La Paz) (Figure 6). Most collections 

come from elevations between 500 m and lo00 m, but Pearcea 

sprucei var. parwiforu has been collected as high as 2300 m 

elevation. 

REPRESENTATIVE COLLECTIONS.-BOLIVIA. La Puz: Mapiri region, San Carlos by Sarampium, Buchtien 1348 (GH, NY, 

US (2)); Nor Yungas, 14 km SW of Yolosa-Chuspipata road, 

km 14, Solomon et al. 12085 (US). Cochabamba: Chapare, 

road to Villa nnari, Luer et al. 4902 (SEL). 

Cuzco: Quispicanchi, Quincemil-San hrenzo road, 

km 3, Moore et al. 8576 (BH (2), US); Quispicanchi, between 

Inambari and Forteleza, Vargas 16536 (US). Puno: San 

Gabon along Rio Gabon, Dillon et al. 1184 (MO, US); vicinity of 

Sangaban, Maas et al. 6079 (USM); Carabaya, Ollachea valley, 

Vargas 16124 (US (2)). Total of 25 collections seen. 

DISTINGUISHING FEATURES.-Pearcea sprucei var. parviflora is distinguished from sympatric var. sprucei populations 

by having a villous indument of whitish trichomes rather than 

a sericeous indument, particularly on the inflorescences and the 

veins of the lower leaf surfaces, by having peduncles no more 

than 2 cm long, ovate leaves with obtuse bases rather than 

lanceolate leaves with cuneate bases, mostly reflexed rather 

than erect calyx lobes, and serrate rather than sermlate margins 

of the leaves. Some high-elevation collections of var. sprucei 

from southern Ecuador are difficult to distinguish from var. 

GH. K, NY (3), PH, US (2). WIS, isotypes).] 

K, PH. US (2), WIS, iSOkCtOtypeS).] 

PERU. 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

parviflora plants, however. The Ecuadorian plants may have 

developed these characteristics independently of var. parviflora in southern Peru and Bolivia, and we refer them to var. 

sprucei. In contrast to populations of the true var. parviflora, 

the similar Ecuadorian populations are not well separated from 

sympatric more “typical” P. sprucei var. sprucei populations, 

and it is unlikely that the plants have dispersed from Bolivia to 

Ecuador or vice-versa (see “Ecology,” above). 

NOMENCLATURE AND TYPIFICATION.-RUSby (1 896:97) 

wrote following the original description of Isoloma parviflorum that the type, Bang 1563A, was “perhaps partly distributed 

as 1565,” but apparently all duplicates of Bang I563A were 

distributed as 1565, as no other specimens of the type number 

have been found. 

NOTES.-According to its label, Luer et al. 4902 was a 

collection of an epiphytic plant. 

16b. Pearcea sprucei var. sprucei 

FIGURES 6,21 

Isoloma sprucei Britton, 1896:97. Kohleria sprucei (Britton) Fritsch, 1913:429. 

Parakohleria sprucei (Britton) Wiehler, 1978:9. [Type: Bolivia, La Paz, 

Mapiri, Bang 1541 (NY, holotype; BM, E. F, G (2), GH (2), K. NY. PH. US (2), 

isotypes).] 

Kohleria jamesoniana Fritsch, 1913:429. Parakohleria jamesoniana (Fritsch) 

Wiehler, 1978:6. [”pe: Ecuador, Pastaza, Archidona, Jameson 236 (8, 

holotype, no longer extant); Ecuador, Napo, Canton Archidona, HollfnLoreto road, km 17, Hurtado & Shiguango 1608 (US, neotype, selected 

herein; MO, isoneotype).] 

Kohleria avilana Cuatrecasas, 1935256 [reprint 1935, p. 531, pl. 22, 23. 

Parakohleria avilana (Cuatrecasas) Wiehler, 1978:5. [’I@?: Ecuador, Avila, 

Isern 169 (MA, holotype; F, MA, isotypes).] 

Kohleria baezana Cuatrecasas, 1935:257 [reprint 1935, p. 541. Parakohleria 

baezana (Cuatrecasas) Wiehler, 1978:6. [Type: Ecuador, ’hngurahua or 

Pastaza, between Baeza and Archidona, Isern 180 (MA, holotype; F, fragment 

of holotype).] 

Kohleria sancti-josephi Cuatrecasas, 1935:258 [reprint 1935, p. 551. Parakohleria sancti-josephi (Cuatrecasas) Wiehler, 1978:8. [?Lpe: Ecuador, San 

Joe, Isern 504 (MA, holotype).] 

Parakohleria vinicolor Wiehler, 1978:9, pl. 1D. [me: Plants cultivated at the 

Marie Selby Botanical Gardens, Sarasota, FL, from specimens originally 

collected as Moore el al. 8387 in Peru, at Boqueron del Padre Abad between 

Aguaytia and Divisoria in Ucayali, Wiehler 75266 (SEL, holotype).] 

Stems rarely branched; inflorescences usually sericeous, 

often purplish, rarely villous with whitish trichomes. Leaf 

blades usually lanceolate with attenuate base, rarely ovate with 

obtuse base, frequently rugose and purplish. Inflorescences 

with 1-8(-18) flowers, mostly sericeous; largest bracts to 12 

mm wide. Calyx lobes usually erect, occasionally reflexed. 

DISTRIBUTION.-The range of P. sprucei var. sprucei is 

nearly identical with the range of the species (Figure 6). 

REPRESENTATIVE COLLECTIONS.-BOLIVIA. La Paz: Road 

to Tipuani, Buchtien 5567 (NY, US), 7487 (US (2)). 

COLOMBIA. Caqueta: Florencia-Guadalupe road, km 86, 

Londono & Kvist 106 (AAU, COL, TULV, us). Huila: Resina, 

Garcia-Barriga 18194 (COL, GH, us); Altamira-Florencia road, 

Rosa Blanca, Vogel 168 (US). Putumayo: San Francisco de 

Sibundoy, trail to Planada de Minchoy, Londono & Lozano 227 

(us). 

ECUADOR. Azuay: Cuenca-Colas de San Pablo highway, 

Boeke & Loyola 688 (AAu, NY, SEL, us). Canar: Above 

Rivera on road to Pindilig, Hurling & Andersson 23024 (GB, 

QCA, US), 23025 (GB, QCA, US). Chimbarazo: Guayllanac, 

Acosta-Solis 5284 (F), 5316 (F). El Oro: 11 km W of Pinas,

Dodson et al. 9020 (MO, SEL). Loja: Loja-Zamora road, km 

19, Hurling & Andersson 13640 (GB, SEL). MoronuSantiago: Sangai National Park close to Macas, Kvist 60440 

Hurling & Andersson 12676 (GB, SEL). Napo: Baeza-%na 

road, Cordillera de Guacamayos, Kvist 60356 (AAU, COL, GB, 

MO, NY, QCA, QCNE, US). Pastaza: Vicinity of hyo, Skutch 

4456 (A, F, K. MO, NY, US). Tungurahua: Pastaza river valley 

between Banos and Cashurco, Hitchcock 21866 (GH, NY, 

US). Zamora-Chinchipe: 10 km S of Zamora on road along 

Rio Jamboe, Hurling & Andersson 24031 (GB, US). 

PERU. Amazonas: Bagua, Mesones-Muro road below Montenegro, Hutchison & Wright 3685 (F, K, MO, NY, UC, US, 

USM). Ayacucho: Between Huanta and Rio Apurimac, Estrella, Killip & Smith 23059 (F, NY, US). Cuzco: Rio 

Mapituriani, tributary to Rio Apurimac, Wasshausen & 

Encarnacion 659 (K, NY, MO, US, USM). Huanuco: Leoncio 

Prado, Divisoria, Skog et al. 5121 (us, USM). Junin: Pichis 

trail, Porvenir, Killip & Smith 25926 (F, NY, US). Loreto: 

Wurdack 2381 (USM). Madre de Dios: Inambari, Chavez A. 

1160 (MO). Pasco: Cordillera Yanachaga, 8 km E of 

Oxapampa-Villa Rica road, Gentry & Smith 35851 (MO, 

us). San Martin: Lamas, Aloso de Alvarado, Cerro Blanco, 

road to Moyobamba, Schunke V. 6265 (NY, US). Ucayali: 

Coronel Portillo, Boqueron de Padre Abad, Schunke V. 3060 (F, 

171 wild collections seen. 

DISTINGUISHING FEATURES.-see the discussion of Pearcea 

sprucei var. parviflora. 

NOMENCLATURE AND TYPIFICATION.-The only species that 

fits the description of Kohleria jamesoniana (Fritsch, 1913) is 

Pearcea sprucei. A collection (Hurtado et al. 1608) from the 

same area and with similar short, reflexed calyx lobes and a 

similar indumentum has been chosen as a neotype to substitute 

for the holotype (Jameson 236), which is no longer extant. 

(AAU, COL, GB, MO, NY, QCA, QCNE, US); Gualaceo road, km 17, 

G, IBE, MO, NY, US, USM), 3061 (F, G, MO, NY, US, USM). TOM Of 

17. Pearcea shgosa L.P. Kvist & L.E. Skog, new species 

FIGURES 12,22 

Pearceae gracile L.P. Kvist & L.E. Skog affkis, sed 

pedunculis quam pedicellis brevioribus, lobis calycum 5- 12 

mm longis, corollis sericeis, trichomatibus -2 mm trans 

limbum extensis differt. 

Stems often decumbent, 30-80 cm long, basal diameter to 4 

mm, occasionally branched, basally pilose to sericeous and 

yellow green apically; internodes 1-9 cm long. Leaves equal in 

FlGURE 22 (opposite page).-Pearcea sfrigosa: a, habit and inflorescences; b, 

indument of lower leaf surface; c, lateral view of flower, d, laterally opened 

corolla showing androecium; e, calyx, nectary, and gynoecium;f, enlarged view 

of calyx, nectary, and gynoecium. a,b from Rimachi 4976 (us); c-ffrom 

Woytkowski 34523 (uc). 
NUMBER 84 41 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

each pair, blades weakly oblique, lanceolate, less commonly 

elliptic, 4-10 x 1.5-3.5 cm, base cuneate, rarely attenuate, 

apex acute-acuminate, margin nearly entire or sermlateundulate, above green, strigose, below (slightly) lighter green, 

glabrous apart from the sericeous-strigose veins, lateral veins 

7-9 per side; petioles 1-4 cm long. Inflorescences of (1-)3-7 

flowers in upper leaf axils, yellow green, sericeous; peduncles 

0.2-0.8 cm long; bracts 2, lanceolate-subulate, 10-20 x 2-4 

mm; pedicels 1-4 cm long. Floral tube conic, 1.5-2.5 mm 

long, yellow green, sericeous; calyx strigose, 6-13 mm long, 

lobes erect, lanceolate-subulate, 5-12 mm long and 1-2 mm 

wide at base, apex acute-acuminate; corolla 14-20 mm long, 

base saccate, tube funnelform, 12- 18 mm long, diameter 4-9 

mm medially and 3-9 mm in throat, red, outside sericeous, 

inside glabrous, limb with lobes subequal, rotund, 1.5-2.5 mm 

long and 2.5-3.5 mm wide at base, glandular-puberulent at 

least along margin, apex obtuse-truncate; stamens subincluded, 

filaments 10-1 8 mm long, often sparsely glandular-puberulent 

and puberulent, anthers -2 x 1.2 mm; staminode 2-3 mm 

long; nectariferous disc of 5 glands, all -0.6 mm high and 0.8 

mm wide at base; ovary -2/3 inferior, 2.5-3.5 mm long, 

diameter 2-3 mm, sericeous, style subincluded, 12-18 mm 

long, puberulent, stigma stomatomorphic. Capsule globose, 

4-5 mm long, diameter 5-6 mm, splitting into 2 valves from 

apex to base; seeds irregularly globose, 0.2-0.3 mm long, 

brown black, striate. 

TYPE.-PERu. Huanuco: Tingo Mm’a-Pucallpa road, La 

Divisoria, 1600 m, 22 Mar 1980, Rimachi Y 4976 (US, 

holotype; F, IBE, MO, USM, isotypes). 

ETYMOLOGY.-The specific epithet refers to the strigose 

upper leaf surfaces. 

DISTRIBUTION.-Most collections of Pearcea strigosa have 

come from along the Tingo Maria-Pucallpa road close to the 

border of the departments of Huanuco and Ucayali in Peru 

(Figure 12). Pearcea strigosa occurs at 1500 to 1600 m 

elevation on both sides of the ridge known as La Divisoria on 

the border of the departments. In addition, collections from the 

department of Cuzco in the south indicate that the species 

actually may be widespread in the mostly unexplored Peruvian 

front ranges of the Andes (Figure 12). Only one collection from 

Cuzco (Skog & Skog 5209). however, appears to be pure P. 

strigosa, and the remaining specimens may be the result of 

introgression with P. sprucei. These plants have wider 

ovate-lanceolate leaves with serrate-semlate margins but still 

have a strigose indumentum. They have been collected at 

400- 1800 m elevation. 

OTHER COLLECTIONS.-PERU, Cuzco: Quincemil-San Lorenzo road, km 17, Moore et al. 8584 (BH); Manu National 

Park, Paucartambo-Manu road between Pillahuata and Milcopata, Skog & Skog 5209 (US, USM); NE of Cuzco, Sandemun 

3681 (K); Paucartambo, Santa Isabel, Vargas 5165 (us); 

Quincemil toward Rio Negro, Vargas 15354 (us); San 

Pedro-Santa Isabel, Vargas 16843 (us); Paucartambo, Vargas 

17679 (WIS). Huanuco: Near La Divisoria between Tingo 

Maria and Pucallpa, Asplund 12603 (s); La Divisoria, -1 km 

from Ucayali, Gentry et al. 29595 (AMAZ, MO, US); Dtto. 

Hermilio Valdiah, Schunke V; 11455 (US); La Divisoria, 

Woytkowski 34523 (F, MO, UC). Ucayali: La Divisoria on the 

border between Huanuco and Ucayali, Ferreyra 1031 (GH, US, 

USM), 1072 (US, USM), 1643 (US, USM); Padre Abad, La 

Divisoria, close to Rio Chino, Schunke V; 9266 (MO, us), 9860 

(MO, u, US). Total of 17 collections seen. 

DISTINGUISHING FEATURES.-Pearcea strigosa is characterized by its lanceolate, yellow green or grayish green, strigose, 

and somewhat undulate leaves with nearly entire margins and 

by lanceolate calyx lobes and corollas that are covered with 

sericeous, red trichomes extending to 2 mm beyond the lobes of 

the corolla (Figure 22). Pearcea strigosa also has a conspicuous, gibbous corolla base, and the sericeous-strigose indument 

on the lower leaf surfaces is restricted to the veins. 
Appendix 1 

Numerical List of Taxa 

1. Pearcea abunda (Wiehler) L.P. Kvist & L.E. Skog 

2. Pearcea beffa L.P. Kvist & L.E. Skog 

3. Pearcea bilabiafa L.P. Kvist & L.E. Skog 

4. Pearcea cordafa L.P. Kvist & L.E. Skog 

5. Pearceafuscicafyx L.P. Kvist & L.E. Skog 

6. Pearcea gfabrafa L.P. Kvist & L.E. Skog 

7. Pearcea gracifis L.P. Kvist & L.E. Skog 

8. Pearcea grandijofia L.P. Kvist & L.E. Skog 

9. Pearcea hispidissima (Wiehler) L.P. Kvist & L.E. Skog 

10. Pearcea hypocyrfifora (J.D. Hooker) Regel 

11. Pearcea infermedia L.P. Kvist & L.E. Skog 

12. Pearcea purpurea (Poeppig) L.P. Kvist & L.E. Skog 

13. Pearcea reticulafa (Fritsch) L.P. Kvist & L.E. Skog 

14. Pearcea rhodofricha (Cuatrecasas) L.P. Kvist & L.E. Skog 

15. Pearcea schimpfi Mansfeld 

16. Pearcea sprucei (Britton) L.P. Kvist & L.E. Skog 

16a. Pearcea sprucei var. parviflora (Rusby) L.P. Kvist & L.E. Skog 

16b. Pearcea sprucei var. sprucei 

17. Pearcea sfrigosa L.P. Kvist & L.E. Skog 

43 
Appendix 2 

List of Exsiccatae 

(Numbers in parentheses following collection numbers refer to species numbers in taxonomic treatments and in Appendix 1.) 

Aco~ta-li~ll~, M., 5284 (16b); 5316 (16b); 7336 (16b); 7345 (16b); 7406 (16b); 

7414 (16b); 7426 (16b); 7432 (13). 

Allard, H., 20411 (16b); 20838 (5); 21658 (12); 21856 (5); 22101 (16b); 22233 

(5); 22287 (5). 

Asplund, E., 7831 (13); 8518 (13); 9522 (16b); 9523 (9); 10341 (16b); 12073 

(12); 12603 (17); 12684 (5); 18306 (13); 18336 (16b); 18673 (15); 19424 

(9); 19444 (16b); 20048 (13); 20077 (13). 

Balslev, H., et al., 2584 (16b); 10284 (13). 

Bang, M., 544 (16a); 1524 (16b); 1541 (16b); 1563A (16a); 1565 (16a). 

Beck, S., 3067 (l6a); 4918 (16b). 

Bennett, B., et al., 4176 (16b); 4180 (10); 4377 (16b); 4509 (14). 

Besse,L..etal.,68(14);77(13);599(16b); 1573(1); 1601 (1); 1602(1); 1671 

Boeke, J., et al., 688 (16b). 

Bohlin, J., et al., 709 (16b); 1485 (13); 1562 (15). 

Brandbyge, J., et al., 31943 (9); 32014 (1); 32112 (1). 

Bristol. M., 546 (16b). 

Buchtien, O., 1348 (16a); 1411 (16a); 5567 (16b); 7487 (16b). 

Camp, W., 761E (7); 902E (1); 14698 (1). 

Wet, P., et al.. 7598 (1). 

Ceron M., C., 344 (1); 1316 (14); 1456 (16b); 1560 (13); 1703 (10); 3582 (14); 

Ceron M., C., et al., 2098 (16b); 2937 (16b); 5600(10); 5604 (1). 

Chavez A., R., 1160 (16b). 

Croat, T., 21261 (5); 49522 (16b); 49713 (15); 49725 (16b); 57926 (12); 58554 

Croat, T, et al., 59046 (16b). 

Cuatrecasas, J.. 8727 (16b); 9154A (16b); 9161 (2); 11402 (13); 11428 (16b). 

DArcy, W., 14005 (16b). 

Davidse, G., et al., 5626 (16b). 

Davidson, C., 3461 (16b). 

Davis, E., 400 (13). 

Dillon, M., et al., 1184 (16a). 

Dodson, C.. et al., 906 (13); 1926 (13); 2071 (15); 2134 (16b); 9020 (16b); 

Dorr, L., et al., 6297A (3). 

Dudley, T., 10481 (16b); 11579 (16b). 

Espinoza, O., 16 (16b). 

Ewan, J., 16734 (1). 

Fagerlind, F., et al., 1082 (16b); 1277 (13). 

Femm R., 965 (12); 1031 (17); 1072 (17); 1598 (12); 1643 (17); 1941 (12); 

Foster, R., 2374 (16b); 7602 (12); 9112 (12). 

Foster, R., et al., 9161 (12). 

Garcia-Barriga, H., 18194 (16b). 

Gentry, A., et al., 15941 (5); 28743 (13); 29595 (17); 30865 (7); 30921 (15); 

30975 (15); 35851 (16b); 41994 (16b); 60027 (14). 

Grisar, M., s.n. (16b). 

Grubb, P., et al., 85 (16b); 150 (10); 1432 (16b); 1503 (10). 

Gudino, E., et al., 1002 (16b). 

Haenke, T., 1961 (12). 

(10). 

3675 (16b). 

(1 6b). 

14102 (16b). 

4189 (12); 4205 (12); 6450 (12); 6750 (12); 18042 (16b). 

Harling,G.,941 (16b); lOOO(13); 1052(16b); 1126(1); 3146(13);3165(16b); 

3197 (13); 3350 (13); 3419 (13); 3756 (6): 13058 (16b). 

Haling, G., et al., 7649 (1); 9732 (13); 12676 (16b); 12685 (7); 12755 (3); 

13015 (15); 13640 (16b); 13817 (16b); 16208 (16b); 16355 (16b); 16357 

(11); 16430 (16b); 16826 (16b); 16916 (15); 17238 (13); 21410 (16b); 

23024 (16b); 23025 (16b); 23880 (16b); 24031 (16b). 

Heinrichs, E., 262 (13); 858 (13). 

Him, A., 4087 (16b). 

Hirtz, A., et al., 3116 (14); 3160 (15); 7130 (3). 

Hitchcock, A., 21773 (13); 21790 (13); 21852 (15); 21866 (16b); 21893 (15). 

Hodge, W., 6028 (16a). 

Holm-Nielsen, L., 16278 (16b). 

Holm-Nielsen, L., et al., 946 (16b); 4057 (16b); 16277 (11); 26512 (4). 

Hurtado, F., 1197 (16b). 

Hurtado, F., et al., 79 (16b); I17 (16b); 227 (16b); 422 (16b); 1608 (16b); 1619 

Hutchison, P., et al., 3685 (16b); 6052 (16b). 

Infantes, J., 5524 (12). 

Isem, J., 169 (16b); 180 (16b); 186 (10); 187 (10); 496 (14); 504 (16b); 596 

Jameson, W., 786 (10). 

Jaramillo, J., et al., 3058 (1); 3566 (10); 5631 (16b). 

Kennedy, H., 3892 (16b). 

Killip, E., et al., 22673 (16b); 22770 (16b); 23059 (16b); 25926 (16b); 26103 

(16b); 26111 (16b). 

Klug, G., 1659 (1); 4216 (1). 

Knight, D., 182 (16b); 346 (16b). 

Kuntze, O., s.n. (16a). 

Kvist, L., 60251 (16b); 60315 (13); 60319 (15); 60351 (13); 60356 (16b); 

60378 (16b); 60386 (11); 60419 (16b); 60440 (16b); 60451 (1); 60452 (1); 

60455 (1). 

(15); 1626 (16b); 1638 (15); 1656 (9); 1751 (15); 2786 (16b). 

( 14). 

Lehmann, F., 443 (13). 

Lewis et al., 11309 (1). 

Lotnant, B., et al., 11494 (1 1). 

Londono, X., et al., 106 (16b); 134 (16b); 136 (2); 141 (2); 154 (16b); 158 (9); 

Luer, C., et al., 1665 (13); 3293 (13); 4902 (16a); 11870 (3). 

Lugo, M., 260 (13). 

Lug0 S., H., 28 (9); 93 (13); 222 (14); 235 (10); 417 (15); 688 (13); 788 (16b); 

810 (15); 1083 (9); 1099 (9); 1125 (13); 1806 (13); 1894 (6); 2107 (10); 

2124 (I); 2163 (10); 2165 (14); 2189 (14); 2412 (13); 3147 (14); 3503 (14); 

3562 (14); 3931 (9); 4018 (1); 4417 (9); 4494 (16b); 4611 (9); 5060 (1); 

5243 (9); 5327 (14); 5402 (14); 5443 (1); 5923 (6); 7090 (14). 

227 (16b). 

Luteyn, J., et al., 4933 (16b). 

Maas, P.. et al., 6079 (16a). 

MacBryde, B., 133 (13); 720(11); 1550 (13). 

Madison, M., 955 (12); 10016-70 (16b). 

Madison, M., et al., 2521 (9); 2573 (9); 3194 (1); 7534 (7); 7553 (3); 7569 

Martin, R., 98 (1). 

Martinet, J., 1489 (12). 

(16b). 

44 
NUMBER 84 45 

Mason, H., 13937 (16b). 

Matthews, A., 150 (12); 2012 (12). 

McCarroll, D., 18 (16a). 

McDaniel, S., et al., 11 139 (1). 

Metcalf. R., 30659 (16a). 

Mexia, Y., 6824 (13); 6851 (16b); 6852 (16b). 

Miller, J.. et al.. 2197 (14). 

Molau. U., et el., 1291 (13); 1362 (16b); 2474 (16b). 

Moore, H., et al., 8387 (16b); 8546 (12); 8576 (16a); 8584 (17); 8622 (16b). 

Mom. R., et al., 5142 (16b); 5178 (16b); 5185 (16b). 

Neill, D., 6556 (14). 

Neill, D., et al.. 5824 (16b); 7239 (14); 8632 (16b). 

Nunez. P., 11953 (17). 

Ollgaard, B., et al., 9097 (16b); 9268 (6); 9270 (9); 35532 (13); 35583 (6); 

Palacios, W., 5610 (9). 

Palacios, W.. et al., 48 (16b); 49 (16b); 484 (10); 862 (14); 3743 (16b); 5155 

Pavon, J., s.n. (12). 

Pearce, R., s.n. (6). 

Penland, C., et al., 112 (13); 278 (15). 

Pennell, F., 13969 (16b). 

Plowman, T, et al., 4060 (1); 4937 (16b); 4986 (16b); 7403 (16b); 11611 (8). 

Poeppig, E., 1239 (12); 1632 (12). 

Poortmann, H., 319 (16b). 

Prance, G., 26587P (10). 

Prieto, F., Cup8 (9). 

Rimachi Y., M., 4976 (17). 

Ruiz, H.. et al., s.n. (9). 

Rusby, H., 2151 (166); 2422 (16a); 2424 (l6a). 

Sagastegui, A., et al., 7155 (16b). 

Sallier, 476 (10). 

Sandeman, C., s.n. (16b); 79 (13); 3681 (17); 5162 (12). 

Schimpff, H., 704 (1 5). 

Schultes, R., et al., 2054 (13). 

35849 (16b). 

(16b). 

Schunke V., J.. 3060 (16b); 3061 (16b); 3111 (5); 3262 (12); 5116 (5); 6263 

(16b); 6265 (16b); 7558 (8); 9266 (17); 9368 (16b); 9858 (16b); 9860(17); 

10366 (5); 11262 (16b); 11455 (17). 

Scolnik, R., 1490 (3). 

Skog, L., et al.. 5121 (16b); 5147 (12); 5209 (17). 

Shtch, A., 4456 (16b). 

Smith, D.. et al., 5578 (12); 5600 (12); 5788 (12); 8624 (12). 

Smith, E., 232 (12). 

Sobel, G., et al., 2443 (16b); 2457 (16b). 

Solomon, J., 9320 (16a); 9515 (16a). 

Solomon, J., et al., 12085 (16a). 

Soukup, J., 5254 (16b). 

Sparre, B., 17501 (10); 17508 (13). 

Spruce, R., s.n. (10); 4%2 (14): 5841 (13). 

Stein, B., et al., 2337 (16b); 2789 (16b). 

Steyermark, J., 53639 (13); 54231 (16b); 54682 (16b); 54863 (13); 54899 (15). 

Stork, H., et al., 9872 (16b). 

Sullivan, G., et al., 1142 (16b). 

Synge, P., 1 I (13). 

Vargas C., C., 5165 (17); 7399 (I6b); 11766 (16a); 15354 (17); 15380 (16a); 

16124 (16a); 16486 (l6a); 16536 (16a); 16843 (17); 16954 (16b); 17679 

(17). 

Vidal-Senege, M., s.n. (12); s.n. (16b); s.n. (16b). 

Vogel. S., 168 (16b). 

Wasshausen, D., et al., 659 (16b); 1250 (Ma). 

Weberbauer, A., 2109 (12). 

Webster, G., 23166 (13); 23238 (11). 

Webster, G., et al., 28449 (166); 28467 (14). 

Werff, H. van der et al., 9224 (16b); 10408 (3); 11210 (9). 

Wiehler, H., 7176 (9); 75266 (16b); 77129 (1). 

Wiehler, H.. et al., 79300 (1 1). 

Wiggins, I., 10935 (16b). 

Woytkowski, F., 156 (16b); 34496 (17); 34523 (17). 

Wurdack, J., 2381 (16b). 

zak, V., et al., 3143 (16b); 3558 (13); 3776 (16b). 

Zaruma, J., 376 (14). 
Literature Cited 

Anonymous 

1991. From ... Out of the Wild. Gesneriad Journal, 3(5)[SeptemberOctober]: 14-1 5. 

1992. Foliage Plants as Show Entities. Gloxinian, 42(6):18-19. 

1983. The Seed Surface Morphology of the Gesneriaceae Utilizing the 

Scanning Electron Microscope and a New System for Diagnosing 

Seed Morphology. Selbyana, 6220-422. 

Bentham, G. 

1876. Gesneriaceae. In G. Bentham and J.D. Hooker, Genera Plantarum, 

2:990-1025. London: Lovell Reeve and Co. 

Boggan, J. 

1990. Gardening in a Cold Basement. Gloxinian, 40(6):8-9. 

Britton, N. 

1896. Isoloma Sprucei Britton, sp. n. In H.H. Rusby, An Enumeration of 

the Plants Collected in Bolivia by Miguel Bang, 111. Memoirs of the 

Torrey Botanical Club, 6:97. 

Batcheller, F. 

Beaufort-Murphy, H. 

Cuatrecasas, J. 

1935. Plantae Isernianae, 1. Anales Ciencias Universidad de Madrid, 

4:245-259 [reprint pages 42-56]. 

Decaisne, J. 

1848. Isoloma Bentham (Kohleria Regel). Revue Horticole, 20(series 3, 

2):465. 

Ertelt, J. 

1991. The Bull’s-Eye Pollinator Attractant of Parakohleria rhodotricha. 

Gloxinian, 41 (2):21-22. 

Fritsch, K. 

1893-1894 Gesneriaceae. In A. Engler and K.A.E. Prantl, editors, Die 

narurlichen Pflanzenfamilien, 4(3b):133-185. Leipzig: Verlag 

Engelmann. 

1913 (“1914”). Beitrag zur Kenntnis der Gesnerioideae. Botanische 

Jahrhucherfur Sysrematik, 50392-439. [Date on title page is 1914; 

actually published in 1913.1 

Hanstein, J. 

1865. Die Gesneraceen des Khiglichen Herbarium und der Garten zu Berlin, nebst monographischer Uebersicht der Familie im Ganzen, 

11: Gattungen und Arten, Drittes Struk: Die Eugesnereen, Rhytidophylleen und Beslerieen. Linnaea, 34225-446. 

Hoehne, F. 

1970. lconografia das Gesneriaceas do Brasil. 521 pages, 260 plates. Sao 

Paulo: Instituto de Botanica. 

Hooker, J. 

1867. Gloxinia hypocyrriflora: Hypocyrta-flowered Gloxinia. Curtis’s 

Botanical Magazine, 93, plate 5655. 

Ecological Relationships of Plants. 273 pages. New York: Oxford 

University Press. 

1895. Index Kewensis. Volume 1, xvi + 1268 pages. Oxford: Clarendon 

Press. 

Howe. H., and L. Westley 

1988. 

Jackson, B.D. 

Kuntze, C.E.O. 

1898. Gesneriaceae. In Revisio Generum Plantarum, 3(2):240-242. 

Leipzig: Arthur Felix. 

Kvist. L. 

1987. Hunting Kohleria in Colombia. Gloxinian, 37(4):17-21. 

1990. Revision of Heppiella (Gesneriaceae). Systematic Botany, 15: 

1991. Parakohleria: Beautiful Herbs in Deep, Damp Shade. Gloxinian, 

720-735. 

41(4):16-22. 

Kvist, L., and L. Skog 

1988. The Genus Cremosperma (Gesneriaceae) in Ecuador. Nordic 

Journal of Botany, 8:259-269. 

1989. Revision of Reldia (Gesneriaceae). Nordic Journal of Botany, 

8501-61 1. 

1992. Revision of Kohleria (Gesneriaceae). Smirhsonian Contributions to 

Botany, 79: 83 pages. 

1993. The Genus Columnea (Gesneriaceae) in Ecuador. Allertonia, 

6~327-397.14 figures. 

Mansfeld, R. 

1934. VIII: Gesneriaceae Novae Andinae, 1. Repertorium Specierwn 

Novarum Regni Vegetabilis, 36: 120-1 24. 

1936. Gesneriaceae Austroamericanae Novae vel Rariores, 111. Repertorium Specierum Novarum Regni Vegetabilis, 41:145-150. 

Poeppig, E. 

1840. Gesneriaceae. In E.F. Poeppig and S.L. Endlicher, Nova Genera et 

Species Plantarum, 3: 1-36. Leipzig: F. Hofmeister. 

Regel, E. 

1847. Index Seminum in horto botanic0 Turicensi, anno 1847 collectorum, 

1847: 4 pages. 

1848. Ueber die Gattungen der Gesnereen. Flora. 31:241-252. 

1867. 11: Neue oder Empfehlenswerthe Zierpflanzen. Gartenfora, 16: 

Rusby, H. 

1896. An Enumeration of the Plants Collected in Bolivia by Miguel Bang, 

Ill. Memoirs of the Torrey BotanicalClub, 6:l-130. 

1900. An Enumeration of the Plants Collected by Dr. H.H. Rusby in South 

America, 1885-86, XXVIII. Bulletin of the Torrey Botanical Club, 

382-390. 

27:22-31. 

Schumann, K. 

1900. Neue Arten der Siphonogamen 1898. Just’s Botanischer Jahresberichr, 26323-396. 

Skog, L. 

1984. A Review of the Chromosome Numbers in the Gesneriacese. 

Selbyana, 7:252-273. 

Smith, J. 

1841. XXIII: Enumeratio Filicum Philippinarum; or asystematic Arrangement of the Ferns Collected by H. Cuming, Esq., F.L.S., in the 

Philippine Islands and the Peninsula of Malacca, between the Years 

1836 and 1840. JOUrMl of Botany (Hooker), 3:392-422. 

Steiner, K. 

1985. The Role of Nectar and Oil in the Pollination of Drymonia serrulata 

(Gesneriaceae) by Epicharis Bees (Anthophoridae) in Panama 

Biotropica, 12217-229. 

Striepens, A. 

1991. Pearcea hypocyrriflora. Gloxinian, 41(6): 16-1 7. 

Troll, W. 

1964. Die Infloreszenzen: Typologie und Stellung im Aufbau des Vegetationskorpers. Volume l, 615 pages. Jena: Gustav Fischer. 

Veitch, H. 

1867. New Plants for 1867 [advertisement]. Gardener’s Chronicle, 

1867[May 4]:458. 

Weber, A. 

1973. Die Struktur der Paarblutigen Partialinfloreszenzen der Gesneriaceen und Bestimmter Scrophulariaceen. Beitrage zur Biologie der 

Pflanzen, 49:429-460. 

Wiehler, H. 

1975. The Re-establishment of Moussonia Regel (Gesneriaceae). Selbyana, 1 :22-3 1. 

1976. A Report on the Classification of Achimenes, Eucodonia, Gloxinia, 

Goyazia, and Anetanthus (Gesneriaceae). Selbyana, 1 :374-404. 

1978. Parakohleria, a New South American Genus in the Gesneriaceae. 

Selbyana, 5:4- 10. 

1983. A Synopsis of the Neotropical Gesneriaceae. Selbyana, 61-219. 

46 
Index to Scientific Names 

(Currently accepted names are in italics.) 

Achimenes Persoon, 6.17 

Alloplectus Mattius, 4, 7, 17 

Besleriu Linnaeus, 6 

Capaneu Decaisne ex Planchon, 6 

Colwnneu Linnaeus, 9, 17,37 

Cremospermu Bentham, 17 

Diastemu Bentham, 2, 4, 5, 6, 7, 17 

galeopsis Fritsch, 2, 39 

parviflorum (Rusby) Fritsch, 39 

racemiferwn Bentham, 3 

rhodotrichum Cuatrecasas, 2.35 

serrulutu (Jaquin) Martius, 7 

Dtymoniu Martius, 4,6, 7 

Eucodoniu Hanstein, 17 

Casterunthus Bentham, 4,5 

Cloxiniu L’Heritier, 4,6, 7, 17 

hypocyrtifl0raJ.D. Hooker, 1, 1731 

sybuticu (Kunth) Wiehler, 4 

Gloxinieae Fritsch, 3,4, 5, 6.7, 15, 16, 17 

Heppiellu Regel, 4,6, 7 

Hypocyrta brevicalyx Hortorum ex Regel, 31 

Isoloma (Bentham) Decaisne, 2 

flexuosum Rusby, 39 

hypocyrtiflorum (J.D. Hooker) Bentham, 31 

parviflorum Rusby, 2, 39,40 

patentipilosum 0. Kuntze, 39 

purpureum (Poeppig) Bentham, 32 

sprucei Britton, 2, 37,39.40 

urticifolium Rusby, 39 

Isoloma J. Smith, 2 

Koellikeria Regel, 6 

Kohleriu Regel, 1,2,4, 5,6, 7, 8, 15, 17,33 

ullenii Standley & L.O. Williams, 3, 8 

avilana Cuatrecasas, 40 

baezana Cuatrecasas, 40 

hirsutu (Kunth) Regel, 33 

inuequalis (Bentham) Wiehler, 3, 17 

jamesoniana Fritsch, 2,40 

longiculyx L.P. Kvist & L.E. Skog, 3 

melastoma (Poeppig) Cuatrecasas, 32 

patentipilosa (0. Kuntze) Schumann, 39 

reticulata Fritsch, 2, 33 

sancti-josephi Cuatrecasas, 40 

spicatu (Kunth) Oersted, 4, 17.33 

sprucei (Britton) Fritsch, 40 

stuebeliuna Fritsch, 4 

urticifolia (Rusby) Fritsch, 39 

weberbaueri Fritsch, 2,32,33 

Monopyle Monk ex Bentham, 5,6,7,17 

Moussoniu Regel, 2,4,6 

macrocurpa Bentham, 3 

melastoma (Poeppig) Hanstein, 32 

purpurea (Poeppig) Hanstein, 32 

Nupeunthus Gardner, 5 

x Paleria Wiehler, 33 

Parakohlena Wiehler, 1.2.4, 15, 17,33 

abunda Wiehler, 17,20 

avilana (Cuatrecasas) Wiehler, 40 

baezana (Cuatrecasas) Wiehler, 40 

hispidissima Wiehler, 29 

jamesoniana (Fritsch) Wiehler, 40 

melastoma (Poeppig) Wiehler, 32 

parviflora (Rusby) Wiehler, 39 

purpurea (Pwppig) Wiehler, 32 

rhodotricha (Cuatrecasas) Wiehler, 35 

sancti-josephi (Cuatrecasas) Wiehler, 40 

schimpfii (Mansfeld) Wiehler, 35 

sprucei (Britton) Wiehler, 40 

vinicofor Wiehler, 40 

weberbaueri (Fritsch) Wiehler, 32,33 

PeurceuRegel, 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 14, 15, 

16, 17, 18,20,22,25,29, 31, 32,33, 35, 37, 

39 

ubundu (Wiehler) L.P. Kvist & L.E. Skog, 1, 3, 

4, 5.7, 14, 16, 17, 19, 20, 22, 25, 43 

bella L.P. Kvist & L.E. Skog, 1, 2, 4, 7, 12, 18, 

20,21,22, 31,43 

bilabiuta L.P. Kvist & L.E. Skog, 12, 18,22,25, 

corduta L.P. Kvist & L.E. Skog, 13, 14, 15, 18, 

fusciculyx L.P. Kvist & L.E. Skog, 2, 4, 8, 14, 

glubrutu L.P. Kvist & L.E. Skog, 4, 12, 19, 22, 

grucilis L.P. Kvist & L.E. Skog, 13, 14, 19.22, 

grandifolia L.P. Kvist & L.E. Skog, 14, 15, 16, 

hispidissima (Wiehler) L.P. Kvist & L.E. Skog, 

hypocyrtifloru (J.D. Hooker) Regel, 1, 2, 4, 12, 

inrermediu L.P. Kvist & L.E. Skog, 1, 2, 12, 14, 

purpurea (Poeppig) L.P. Kvist & L.E. Skog, 1, 

reticulafa (Fritsch) L.P. Kvist & L.E. Skog, 1,2. 

. 4.5.7. 13, 16, 19,22,29,32,33,35,38,39, 

43 

rhodotricha (Cuatrecasas) L.P. Kvist & L.E. 

Skog, 2,4, 5, 7,8, 12, 14, 16, 19, 35, 38.43 

schimpfii Mansfeld, 1, 2, 4, 12, 18, 31, 32, 35, 

37.43 

sprucei (Britton) L.P. Kvist & L.E. Skog, 1,2,6, 

7, 14, 16, 17, 19, 22, 25, 31, 32, 33, 35, 37, 

38, 39,40,42,43 

var. purvijloru (Rusby) L.P. Kvist & L.E. 

Skog, 2, 11, 16, 19, 33, 38, 39,40,43 

var. sprucei, 11, 16, 19,38,39,40,43 

strigosu L.P. Kvist & L.E. Skog, 1, IS, 16, 19, 

43 

22, 23, 25, 43 

19, 25.43 

25, 27.43 

27,29, 40.43 

18, 29, 31, 33,43 

1, 7, 8, 13, 18, 22, 29, 38.43 

14, 17, 18.20, 31,43 

18, 3 1,32, 37,43 

2, 15, 16, 19, 25, 32, 33, 38, 39,43 

25,38,40,42,43 

Reldiu Wiehler, 17 

Rhytidophyllum Martius 

melastoma Poeppig, 2, 32 

purpureum Poeppig, 2,32 

Sinningia Nees. 5, 7 

Solenophoru Bentham, 17 

47 

US GOVERNMENT PRINTING OFFICE: 1996 405-261/20015