/ VOLUME 69 1982 1/ OF THE v Uieeovm BoTANtSXB IBarben Library The Annals, published four times a year, contains papers, pri- marily in systematic botany, contributed from the Missouri Botan- ical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the Annals. Editorial Committee Nancy Morin, Editor Missouri Botanical Garden Marshall R. Crosby Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden & St. Louis University Peter Goldblatt Missouri Botanical Garden Colophon This volume of the Annals of the Missouri Botanical Garden has been set in VIP Times Roman. The text is set in 10 point type while the figure legends and literature cited sections are set in 8 point type. The volume has been printed on 70# Centura Gloss, an acid-free paper designed to have a shelf-life of over 100 years. Centura Gloss is manufactured by the Consolidated Paper Company. Photographs used in the Annals are reproduced using 300 line screen halftones. The binding used in the production of the Annals is a proprietary method known as Permanent Binding. The Annals is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A. Missouri Botanical Garden 1983 ISSN 0026-6493 Announcements The Dennis Stanfield Award 234 The 1982 Jesse M. Greenman Award 234 Atsatt, Peter R. & Phillip Rundel. Pollinator Maintenance vs. Fruit Production: Partitioned Reproductive Effort in Subdioecious Fuchsia lycioides - — Baker, H. G. & I. Baker. Starchy and Starchless Pollen in the Onagra- ceae — Baker, I. (See H. G. Baker & I. Baker) - Berry, Paul E. (See Dennis E. Breedlove, Paul E. Berry & Peter H. Raven) — — Berry, Paul E. The Systematics and Evolution of Fuchsia Sect. Fuchsia (Onagraceae) Boufford, David E. The Systematics and Evolution of Circaea (Ona- graceae) Breedlove, Dennis E., Paul E. Berry & Peter H. Raven. The Mexi- and Central American Species of Fuchsia (Onagraceae) except for 199 748 748 209 1 804 can Sect. Encliandra 209 755 Carlquist, Sherwin. Wood Anatomy of Onagraceae: Further species; root anatomy; significance of vestured pits and allied structures in Di- cotyledons Coney, Peter J. Plate Tectonic Constraints on the Biogeography of Mid- dle America and the Caribbean Region Cross, Aureal T. & Ralph E. Taggart. Causes of Short-term Sequential Changes in Fossil Plant Assemblages: Some Considerations Based on a Miocene Flora of the Northwest United States 676 Eyde, Richard H. Evolution and Systematics of the Onagraceae: Floral 432 Anatomy 735 Gentry, Alwyn H. Neotropical Floristic Diversity: Phytogeographical Connections Between Central and South America, Pleistocene Climatic Fluctuations, or an Accident of the Andean Orogeny? 557 Goldblatt, Peter. Corm Morphology in Hesperantha (Iridaceae, Ixi- oideae) and a Proposed Infrageneric Taxonomy 370 Goldblatt Peter. Notes on Geissorhiza (Iridaceae): The Species in ' 379 Madagascar Goldblatt, Peter. A Synopsis of Moraea (Iridaceae) with New Taxa, Transfers, and Notes 351 Gomez P., Luis D. The Origin of the Pteridophyte Flora of Central America 548 Grayum, Michael H. (See Barry E. Hammel & Michael H. Grayum) 420 Hammel, Barry E. & Michael H. Grayum. Preliminary Report on the Flora Project of La Selva Field Station, Costa Rica 420 Huft, Michael. Pseudocroton tinctorius Muell. Arg., a Synonym of Cap- Is indica (L.) Fawc. & Rendle 426 pari Humphries, C. J. Vicariance Biogeography in Mesoamerica 444 Keating, Richard C. The Evolution and Systematics of Onagraceae: Leaf Anatomy 770 418 Kenton, Ann (See Ching-I Peng & Ann Kenton) Kral, Robert & Lyman B. Smith. Xyris apureana Krai & Smith, A New Species of Xyris (Sect. Nematopis) from Venezuela 412 Kral, Robert. Xyris ni^rescens Kral, a New Species of Xyris (Sect. Nematopus) from Costa Rica 415 McDade, Lucinda A. Three New Species of Aphelandrn ( Acanthaceae) from Central America 402 McLaughlin, Siivi.n P. A Revision of the Southwestern Species of Amsonia (Apocynaceae) - 336 Martin, Helene A. Changing Cenozoic Barriers and the Australian Paleobotanical Record - 625 Mohlenbrock, Robert H. Illinois Convolvulaceae in the Missouri Bo- tanical Garden Herbarium 393 Mohlenbrock, Robert H. Illinois Solanaceae in the Missouri Botanical Garden Herbarium and Biographical Sketches of Some Collectors .... 382 Morin, Nancy R. Biological Studies in Central America: The Twenty- Eighth Annual Systematics Symposium 431 Peng, Ching-I & Ann Kenton. Chromosome Number of Byblis lini- flora Salisb. (Byblidaceae) 41° Prance, Ghillean T. A Review of the Phytogeographic Evidences for Pleistocene Climate Changes in the Neotropics _ - 594 Raven, Peter H. (See Dennis E. Breedlove, Paul I Berry & Peter H. Raven) 209 Raven, Peter H. Nomenclatural Corrections in the Genus Camissonia (Onagraceae) „ 995 Robbrecht, Elmar. The Identity of the Panamanian Genus Drcssleriopsis All (Rubiaceae) 4 -' Rundel, Phillip (See Peter R. Atsatt & Phillip Rundel) 199 Savage, Jay M. The Enigma of the Central American Herpetofauna: Dis- persals or Vicariance? 464 Smith, Lyman B. (See Robert Krai & Lyman B. Smith) 412 Solomon, James C. The Systematics and Evolution of Epilobium (Ona- graceae) in South America 239 676 Taggart, Ralph E. (See Aureal T. Cross & Ralph E. Taggart) Whitmore, T. C. Wallace's Line: A Result of Plate Tectonics 668 FTHE LUME69 1982 NUMBER /J / . J ' \VI/ ^ / f —^ ^* - " FUCHSIA NIGRICANS LINOEI STUDIES IN FUCHSIA CONTENTS The Systematics and Evolution of Fuchsia Sect. Fuchsia (Onagraceae) Paw/ E. Berry Pollinator Maintenance vs. Fruit Production: Partitioned Reproductive Effort in Subdioecious Fuchsia lycioides Peter R. Atsatt & Phillip Rundel 199 The Mexican and Central American Species of Fuchsia (Onagraceae) except for Sect. Encliandra Dennis E. Breedlove, Paul E. Berry & Peter H. Raven 209 The Dennis Stanfield Award 234 The 1982 Jesse M. Greenman Award 234 Index Additional copies of this special issue are available from the Missouri Botanical Garden. See page I for ordering information. VOLUME 69 1982 NUMBER 1 OF THE M. M botany contributed from the Missouri Botanical Gard de the Garden will also be accepted. Authors should ditor for information concerning arrangements for pub- the Annals. Editorial Committei Nancy Morin, Editor Missouri Botanical Garden Gerkit Davidse Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden 6- St. Louis University Peter Goldbla t t Missouri Botanical Garden Published four times a year by the Missouri Botanical Garden. St. Louis, Missouri 63110. ISSN 0026-6493 For subscription information contact the Business Office of the Annals, P.O. Box 368, 1041 New Hampshire, Lawrence. Kansas 66044. Subscription price is $45 per volume U.S., Canada, and Mexico, $50 all other countries. Four issues per volume. Second class postage paid at Lawrence, Kansas 66044 OF THE ID H m VOLUME 69 1982 NUMBER 1 THE SYSTEMATICS AND EVOLUTION OF FUCHSIA SECT. FUCHSIA (ONAGRACEAE) 1 Paul E. Berry 2 Abstract Nine sections are currently recognized in the genus Fuchsia, consisting of approximately 100 species. Morphological and biogeographical evidence indicates a Paleogene origin of the genus in temperate South America, with two early lines diverging to New Zealand and North America. Within South America, sects. Quelusia and Kierschlegeria have remained in the southern part of the con- tinent, while the two largest sections, Fuchsia and Hemsleyella, have developed in the tropical Andes. 1 Copies of this special issue of the Annals of the Missouri Botanical Garden can be obtained by sending $7.50 per copy to Department Eleven Studies in Fuchsia, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. This material is based upon research supported by the National Science Foundation under Grants No. DEB 78-05969 (to P.B.) and DEB 78-23400 (to Peter H. Raven) and is published with support of National Science Foundation Grant No. DEB-8122087. The Foundation provides awards for research and education in the sciences. The awardee is wholly responsible for the conduct of such research and preparation of the results for the publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author and do not necessarily reflect the views of the National Science Foundation. A grant-in-aid from Sigma Xi helped to support the initial stages of the field work. The color illustrations and some of the figures were prepared by the Unidad de Medios Audiovisuales, Universidad Simon Bolivar, Caracas, Venezuela. The Universidad Simon Bolivar also generously covered the costs of the color plates, through the Decanato de Investigaciones. This support is gratefully acknowledged. This study was undertaken as part of my doctoral thesis studies at Washington University, St. Louis, Missouri. I am especially thankful to Dr. Peter H. Raven for his continual interest, encour- agement, and support throughout the realization of this study. The assistance of the staff and tech- of the Missouri Botanical Garden greatly facilitated my work and is strongly appreciated. I extend special thanks to Dr. Stephen S. Tillett (Herbario Ovalles, Facultad de Farmacia, Universidad Central de Venezuela) for providing advice and laboratory facilities during extended stays in Caracas, to Dr. Julian A. Steyermark (Instituto Botanico, Caracas) for his ongoing help and encouragement, and to Dr. Peter Hoch (Missouri Botanical Garden) for carefully reviewing the final draft of my thesis. My companions W. Wagner, T. P. Ramamoorthy, and J. Solomon provided helpful discussion and assistance in the preparation of this paper. The illustrations were expertly done by Eduardo Perez; Gabriela Aedo assisted with maps and diagrams. Many persons generously gave of their time and knowledge to assist me in the field work. I would specifically like to mention the late Dr. Luis Ruiz-Teran (Merida, Venezuela); also, Dr. Cesar Vargas C. (Cuzco, Peru), Dr. Ramon Ferreyra (Museo de Historia Natural Javier Prado, Lima, Peru), Dr. Abundio Sagastegui (Universidad Nacional de Trujillo, Peru), Ing° Alberto Ortega (Universidad Nacional, Quito, Ecuador), Dr. Fernando Ortiz (Universidad Catolica, Quito, Ecuador), Linda Albert de Escobar (Comision Fulbright, Quito, Ecuador), Lie. Olga Benavides (Universidad de Narino, Pasto, Colombia), Ing° Eugenio Escobar (Universidad del Valle, Palmira, Colombia), Dr. Luis Mar- Ann. Missouri Bot. Gard. 69: 1-198. 1982. 0026-6493/82/000 1 -0 1 98/$ 1 9.95/0 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 This paper treats the systematics of Fuchsia sect. Fuchsia, which includes 61 species, ten of them newly described here. Despite the large number of species compared to other sections in the genus, sect. Fuchsia appears to be a natural grouping of species. These species all have morphological variations on a common theme, namely shrubs with tubular, reddish flowers pollinated mostly by hummingbirds. The main differences between species occur in the shape, length, or coloration of the floral tube and in the position of flowers in the leaf axils or in different kinds of racemose inflorescences. The close relationship between species in sect. Fuchsia is indicated by their high level of inter- fertility with numerous naturally occurring hybrids and by chromosome numbers that are invariate or on just two levels. From over 100 new chromosome counts in populations of sect. Fuchsia, 37 species are now known as diploids (n = 11), five as tetraploids, and one species has both diploid and tetraploid populations. Chromosomal pairing in interspecific hybrids was normal in all cases examined. Naturally occurring interspecific hybrids were found in 17 different combinations, and, in two in- stances, populations of probable hybrid origin appear to be more successful than their local parental populations. Altitudinal stratification of species along mountain slopes is a major isolating mechanism between different species of sect. Fuchsia. Though sympatry is common, species that differ substantially in floral tube length and coloration may diminish or prevent interspecific pollination by hummingbirds. Sect. Fuchsia possesses a particularly flexible reproductive system, combining limited vegetative reproduction with a long, essentially aseasonal flowering period in which modal outcrossing is prev- alent, but autogamy also occurs. The cool, montane habitats of the Andes presently occupied by sect. Fuchsia have arisen largely in the Neogene. The Andean chain actually is composed of a number of distinct structural units, each one differing somewhat in its orogeny and floristic/faunistic composition. Distributional patterns of species in the different structural units suggest that sect. Fuchsia has undergone a recent differentia- tion closely tied to the major elevation of the Andes in the Pliocene and to the glacial events and climatic fluctuations that took place during the Pleistocene. The recent diversification of most species in the section is further supported by the morphological and cytological similarities noted above, as well as by the large numbers of species in the Andes compared to other areas where Fuchsia occurs, and the small geographic ranges of most species there. The ability to hybridize and recombine genetic material, the presence of self-compatibility allowing establishment from a single seed after dispersal, and the recent uplift and climatic fluctuations of the Andes have probably been the key factors in the differentiation of species in Fuchsia sect. Fuchsia. Fuchsia is a large, distinctive genus that lacks close affinities with any other group in the Onagraceae and therefore is classified as a monogeneric tribe. Its approximately 100 species are mostly mesophytic shrubs that grow in the Andes from Colombia and Venezuela to Tierra del Fuego, the southeastern coastal mountains of Brazil, Hispaniola, and the mountains of Mexico and Central Amer- ica, with a small disjunct section in New Zealand and Tahiti. On the basis of chromosome number and morphology (Kurabayashi et al., 1962), floral anatomy (Eyde & Morgan, 1973), wood anatomy (Carlquist, 1975), and leaf architecture (Hickey, 1980), Fuchsia appears to be one of the least specialized genera in the family. In its red, tubular, bird-pollinated flowers and fleshy, bird-dispersed fruits, however, Fuchsia as it exists today is clearly specialized. Both Fuchsia and Ludwigia, a genus that is also relatively unspecialized and represents a phylo- cano-Berti (Universidad de Los Andes, Merida, Venezuela), and Dr. Gary L. Smith and Milciades Mejia (Jardin Botanico Rafael Moscoso, Santo Domingo, Dominican Republic). 1 thank the curators of the following herbaria for allowing me to examine material under their care: A, AAU, B, BH, BM, BR, BREM, CAS, CGE, CM, COL, CUZ, DS, DUKE, F, FHO, G, GH, GOET, ISC, JBSD, K, LAM, LE, LG, LIL, LL, LOJA, M, MA, MER, MERF, MICH, MO, MPU, MSC, MU, MY, MYF, NA, NY, O, OS, OXF, P, PH, POM, PR, PSO, Q, QCA, RSA, S, SIU, TCD, TEX, U, UC, UPS, US, USM, VALLE, VEN, W, WU, and Z. Finally, my deep thanks go to my wife Eva and to Danny, who endured long separations so that this paper could be completed. 2 Departamento de Biologia de Organismos, Universidad Simon Bolivar, Apartado 80659, Ca- racas 1080, Venezuela. 1982] BERRY— FUCHSIA SECT. FUCHSIA 3 - genetic line distinct from the remainder of the family, have their center of distri- bution in South America, which may have been the center of origin for the family (Raven & Axelrod, 1974). In light of these relationships, an understanding of the evolutionary modes in Fuchsia will be fundamental to the overall understanding of evolution in the Onagraceae. In the past 15 years, detailed systematic studies have been completed for a number of the small sections of the genus. Sect. Encliandra (six spp., Central America and Mexico) was revised by Breedlove (1969), sect. Skinnera (four spp., New Zealand and Tahiti) by E. Godley and P. H. Raven (unpublished), and finally sect. Schufia (two spp., Central America and Mexico) and the new sections Ellobium (three spp., Central America and Mexico) and Jimenezia (one sp., Costa Rica and Panama) by Breedlove et al . ( 1 982) . These sections , some of which present interesting developments such as male sterility and wide disjunctions in range, represent less than a fifth of the total number of species in the genus. The great majority of species of Fuchsia are concentrated in Andean South America, and about three-fifths of the total belong to the section treated here. Fuchsia sect. Fuchsia was selected for the present revision because of its comparatively large size, with over 60 closely related species, and because it can be expected to provide a model of evolution widely applicable to other groups of the poorly studied tropical Andes. Emphasis was placed primarily on field ex- amination of populations and morphological analysis of herbarium specimens from the major collections of the world. In this manner, the amount of hybrid- ization and the principal isolation mechanisms operating between species could be evaluated along with morphological characters to determine more accurately species limits within the section. In addition, chromosome counts were obtained for 43 of the 61 species recognized in sect. Fuchsia. Concurrently with this study, an analysis of leaf flavonoids (J. Averett et al., in prep.) as well as a detailed, ultrastructural pollen survey of the entire genus (J. Nowicke et al., in prep.; J. Praglowski et al., in prep.) are nearing completion, and systematic studies are underway in the remaining three sections from South America, Quelusia, Kier- schlegeria and Hemsleyella (P. E. Berry and T. P. Ramamoorthy, in prep.). Origin of the Genus and Subgeneric Relationships AGE AND RELATIONSHIPS OF FUCHSIA Despite the unequivocal familial limits of the Onagraceae, no clear relation- ships are apparent between the seven tribes in the family. Although no tribe presents a series of features that can be considered ancestral to the rest, Lud- wigia, the sole genus of the Jussiaeeae, is the most distinct and clearly represents an early offshoot of the family (Raven, 1979a). The other six tribes are therefore more closely related to one another than any one is to Ludwigia. The monogeneric tribe Fuchsieae is the tribe with the largest assemblage of unspecialized features, including the absence of interxylary phloem and unspecialized chromosomes and placentation (Eyde & Morgan, 1973). The order Myrtales, to which the Onagraceae belong, is clearly a southern hemisphere group (Raven & Axelrod, 1974). Ludwigia has its least specialized 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 1. Main historical subdivisions of Fuchsia. Author deCandolle (1828) Sect. Quelusia Breviflorae Macrostemonae Longiflorae Sect. Skinnera Endlicher (1840) a. Encliandra 2 Brebissonia Lyciopsis b. Fuchsia 1 Kierschlegeria Fuchsia Schufia c. Skinnera' 1 Hemsley (1877) "American species having petals" Tropical Andean species Brazilian species South Andean species Mexican and Central American species "American species destitute of petals" "New Zealand species" Munz(1943) Sect. Sect. Sect. Sect. Quelusia Fuchsia Kierschlegeria Skinnera Sect. Hemsleyella Sect. Sect. Schufia Encliandra Equivalent Modern Sections 1 Encliandra, Kierschlegeria Fuchsia (part), Quelusia, Schufia Fuchsia (part), Hemsleyella, Ellobium Skinnera Encliandra (part) Encliandra (part) Kierschlegeria Fuchsia, Quelusia Schufia Skinnera Fuchsia Quelusia (part) Quelusia (part), Kierschlegeria Encliandra, Schufia, Ellobium Hemsleyella Skinnera Quelusia Fuchsia, Ellobium (part) Kierschlegeria Skinnera Hemsleyella, Ellobium (part) Schufia Encliandra i Includes only groups or species known at the time of publication. 2 Groups of undesignated rank, but subsequently taken up as sections by Walpers (1843), Ben tham & Hooker (1867), and Baillon (1877). members and greatest species diversity in South America, as well as all of its self-incompatible species. Fuchsia likewise has by far its greatest concentration of species in South America and had reached New Zealand by the latest Oligocene (Mildenhall, 1980). Because of these relationships, it has been hypothesized that the Onagraceae originated in South America (Raven & Axelrod, 1974; Raven, 1979a). All of the remaining tribes of the family, however, have their centers of diversity in the northern hemisphere, and fossil pollen of Ludwigia is known from the northern hemisphere from the early Paleogene onward (Eyde & Morgan, 1973). Thus very early dispersal between the two hemispheres clearly occurred. The Onagraceae originated by the close of the Cretaceous (Raven & Axelrod, 1974); earliest fossil remains assignable to genera are from the Paleocene for Ludwigia and the Oligocene for Circaea. The earliest fossil record for Fuchsia 1982] BERRY— FUCHSIA SECT. FUCHSIA 5 Figure 1. Native distributions of the South American sections of Fuchsia. 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 T 75" !.:;!;!i ,; 'i Sect. Fuchsia Sect Jimenezia ^ > »0* - Sects Encliandra. Schufia. and Ellobium 105* _i 95 _1_ 85 Figure 2. Native distributions of the sections of Fuchsia outside of South America, not in- cluding sect. Skinner a, which is endemic to New Zealand and Tahiti. is pollen from the uppermost Oligocene of New Zealand (Couper, 1960; Milden- hall, 1980), which gives us a minimum age for the genus. Because of the posses- sion of a number of advanced characters, the New Zealand species of Fuchsia are almost certainly derived from American ancestors, so the genus probably evolved considerably earlier. Bird pollination is a dominant theme in Fuchsia and is present in both New World and Old World groups; the fruits are also adapted to bird dispersal. Specialized flower- visiting birds probably did not evolve until the Eocene (Sussman & Raven, 1978), but it is uncertain whether the com- mon ancestor of Fuchsia would have been bird-pollinated or not. At any event, plants similar to the existing species of Fuchsia probably could not have evolved prior to the Eocene. The most likely hypothesis, therefore, is that the genus originated in the Eocene or Oligocene in South America. SUBGENERIC TREATMENTS OF FUCHSIA Attempts to recognize several related genera in the tribe Fuchsieae, such as Spach's (1835) treatment, have met with little acceptance because of the existence of intermediate taxa linking the most divergent species and because of the lack of fundamental differences in the basic features of the group. In contrast, a num- ber of infrageneric groups have traditionally been recognized as sections or as groups of undesignated rank. A synopsis of the different subgeneric treatments of Fuchsia until Munz's (1943) generic monograph is listed in Table 1. DeCandolle (1828) divided the genus into sect. Skinnera, from New Zealand, and sect. Quelusia, from America. He based his subdivisions of sect. Quelusia on the degree of staminal exsertion and the relative length of the floral tuhe and 1982] BERRY— FUCHSIA SECT. FUCHSIA 7 Table 2. The sections c )f Fuchsia and the ir native geographical distribution. Estimated number Section of Species Distribution 1. Quelusia 5 SE coastal Brazil, Chile & Argentina 2. Fuchsia 61 Tropical Andes, Hispaniola 3. Ellobium 3 Mexico and Central America 4. Hemsleyella 14 Tropical Andes 5. Kierschlegeria 1 Central coastal Chile 6. Schufia 2 Mexico and Central America 7. Jimenezia 1 Panama and Costa Rica 8. Encliandra 6 Mexico and Central America 9. Skinner a 4 New Zealand and Tahiti TOTAL: 97 sepals. His subdivisions represent what now seems to have been an unnatural division of the presently recognized sections. Endlicher (1840) made a few im- provements over DeCandolle's treatment, such as the segregation of the distinc- tive Schufia group and the separation of Kierschlegeria from Encliandra. Despite Hemsley's (1877) informal style of presentation, his arrangement is remarkably close to our modern sectional concepts of the genus. Through his geographical criteria and familiarity with living, introduced species, Hemsley first recognized the integrity of the modern sects. Fuchsia, Hemsleyella, and, in part, Quelusia. Finally, Munz's (1943) generic revision ordered the previous diverse subgeneric concepts into a consistent, formal system consisting of seven sections. Recent study of the Mexican and Central American species of Fuchsia has introduced several changes into Munz's sectional classification (Breedlove et al., 1982). The currently recognized sections are listed along with their native distri- bution and species numbers in Table 2, and maps of their distributions appear in Figures 1 and 2. Key to the Sections of Fuchsia la. Petals lacking or much reduced, if present, less than one third the length of the sepals and with floral tubes 5-15 mm long; leaves generally alternate. 2a. Floral tube 5-15 mm long; gynodioecious or dioecious; leaves usually present when flowering; New Zealand and Tahiti sect. Skinnera 2b. Floral tube 17-170 mm long; hermaphroditic; often flowering when leafless; South America. sect. Hemsleyella lb. Petals present, usually conspicuous and more than one third the length of the sepals, if shorter, with floral tubes 20-130 mm long; leaves generally opposite or whorled. 3a. Antepetalous stamens reflexed and included in the floral tube. 4a. Flowers axillary, dioecious or subdioecious; berries with 6-35 seeds; Mexico to Panama. sect. Encliandra 4b. Flowers in terminal racemes or panicles, hermaphroditic; berries with 35-80 seeds; Costa Rica and Panama. sect. Jimenezia 3b. Stamens all erect and exserted beyond the floral tube. 5a. Small trees; flowers numerous, erect in well-developed di- or trichotomously branched panicles. sect. Schufia 5b. Shrubs, climbers, or epiphytes, rarely arborescent; flowers spreading or pendant and axillary, racemose, or in few-branched panicl 6a. Petiole bases persisting as woody, spinose knobs on the stem; subdioecious; floral tube 3-10 mm long and sepals reflexed; berries with 14-30 seeds. sect. Kierschlegeria 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 6b. Petioles dehiscing flush with the stem and never spinose; hermaphroditic; floral tube 10-130 mm long or, if shorter, sepals not reflexed; berries with 30-250 seeds. 7a. Floral tubes usually shorter than the sepals, these ± connate at the base; stamens exserted well beyond the sepals; petals convolute after anthesis; flowers axillary; Southern Andes and southeast Brazil (introduced else- where). sect. Quelusia 7b. Floral tubes usually longer than the sepals, if not, with sepals free to the base; stamens generally not exserted beyond the sepals, petals not over- lapping at anthesis; flowers axillary, racemose, involucrate, or paniculate; Central Andes to Mexico and Hispaniola. 8a. Leaves mostly opposite, broad, elliptic-ovate or cordate; tubers pres- ent, or plants epiphytic, if not, with green petals; nectary a smooth, unlobed band lining the base of the tube; Central America and Mex- ico. sect. Ellobium 8b. Leaves opposite to whorl ed, variously shaped but generally not cor- date at the base; tubers lacking; plants terrestrial, the petals never green; nectary annular and mostly free from the base of the tube or with prominent lobes adnate to it; South America and Hispaniola (introduced elsewhere). sect. Fuchsia INTERSECTIONAL RELATIONSHIPS AND EVOLUTIONARY LINES IN FUCHSIA As discussed previously, Fuchsia probably originated in South America. The genus almost invariably occurs in cool, mesic climates and is centered in tropical uplands or temperate forest below the frost zone. The three main ecogeographic areas of South America occupied by Fuchsia are 1) cloud forests of the tropical Andes, 2) Nothofagus forest in the temperate Andes, and 3) montane forest in southeastern Brazil. The floristic relationships of these three areas have been pointed out by numerous authors, including Gerth (1941), Smith (1962), and Vuil- leumier (1969), based on such diverse taxa as Araucaria, Drimys, Berberis, Le- pechinia, Podocarpus, Ilex, Chusquea, and Perezia. These elements are consid- ered to be remnants of ancient, widely distributed temperate forests across southern South America (Simpson, 1973, 1979). Paleontological, paleoclimatic, and geo- logical evidence indicates that a generally warm and equable climate prevailed throughout South America in the early Tertiary, with tropical rainforest vegeta- tion extending considerably farther south than at present (see reviews in Vuilleu- mier, 1969; Simpson 1973, 1979; Haffer, 1974; Solbrig, 1976; Duellman, 1979). Fossil evidence of fully tropical wet forest is known from the Eocene in Argentina at 38°S, with subtropical woods and pampa occurring at 42°S. A marked cooling and drying trend began in the Oligocene and Miocene, however, with the result that temperate elements of the Antarcto-Tertiary Geoflora migrated farther north, replacing the tropical flora of Chile and Argentina. In the Oligocene, austral forests of Nothofagus, Araucaria, and Laurelia were present as far north as 30°S in Argentina (Jeannel, 1967). Subsequently the continuity of this continuous forest was broken up by the rain shadows created by the rising Andes and by the spread of the Tertiary-Chaco Geoflora (Solbrig, 1976). The fragmentation of the once widespread southern forests continued, and by the Pliocene most of the Notho- fagus forests east of the Andes had disappeared completely. Given this background, we may hypothesize that the ancestral stock of Fuch- sia evolved within the southern temperate forests of South America during the Eocene or Oligocene. Several main lines of the genus probably diverged in the 1982| BERRY— FUCHSIA SECT. FUCHSIA 9 late Oligocene and Miocene, when the austral forests began to contract and frag- ment, and the Andes began to be uplifted substantially. One of the earliest off- shoots of Fuchsia was sect. Skinnera. As indicated by pollen fossils (Mildenhall, 1980), Fuchsia was present in New Zealand, where there are three very distinct species of sect. Skinnera, by the late Oligocene. It probably spread across Ant- arctica, contrary to the earlier view of Raven, who postulated a long distance dispersal route across the Pacific (Raven, 1972a, p. 242). At that time, Fuchsia had not been reported from New Zealand prior to the middle Miocene, and it was not understood that more or less direct opportunities for migration across Antarctica persisted until the Miocene (Raven, 1979b). The fourth species of sect. Skinnera, F. cyrtandroides, is endemic to the Pacific island of Tahiti, which is of volcanic origin and less than two million years old (Dymond, 1975). This species undoubtedly arrived there from New Zealand by long distance bird transport, since the entire Tahitian flora is adapted to long distance dispersal, and nearly 40% of the plant species there are adapted to internal transport by birds (Carlquist, 1967). Members of sect. Skinnera are probably better adapted to long distance transport than are other sections of the genus because they have the smallest and most numerous seeds (up to 450 per fruit) in the genus. Other distinctive characters reflect the early divergence of sect. Skinnera. Initial results of a survey of leaf flavonoids in the genus indicate that it is the only section with what appears to be sulfated flavonoids (J. Averett et al., in prep.). It also includes the most widely divergent growth forms in the genus — procum- bent creepers, lianas, and tall trees. A strong reduction of petals occurs in the section, and it has smooth, band-type nectaries like those of sects. Hemsleyella and Ellobium and beaded viscin threads on the pollen as in all South American sections except sect. Kierschlegeria. Another offshoot probably spread early in the history of the genus to North America by long distance dispersal, most likely in Paleogene time. Other families that show this pattern of an early South to North American migration include Cactaceae, Loasaceae, Nyctaginaceae, and Zygophyllaceae (see Raven & Ax- elrod, 1974, pp. 627 and 628 for more complete lists). This early offshoot of Fuchsia that reached North America eventually differentiated into the modern sects. Encliandra, Jimenezia, and Schufia. Though each one of these sections is now quite distinct from each other, common ancestry is suggested by the follow- ing shared characters: smooth viscin pollen threads (though a few intermediate types occur in sect. Encliandra), small flowers adapted to insect pollination in several species, and similar lobed-adnate nectaries. Other derived characters are the presence of male sterility in sects. Encliandra and Schufia, antepetalous stamens reflexed into the floral tube in sects. Encliandra and Jimenezia, and low 'fi Mexico Isthmus of Tehuantepec (Breedlove, 1969; Breedlove et al., 1982). Section Ji- menezia, clearly closely related to, but more primitive than, sect. Encliandra, is monotypic and endemic to a small area of Costa Rica and Panama. The remaining sections represent the majority of the species in the genus; they probably arose from a common ancestor as the austral temperate forests of South America became increasingly limited in area and the Andes were uplifted 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 during the Miocene and Pliocene. Sections Quelusia, Fuchsia, Hemsleyella, and Ellobium are all basically hummingbird-pollinated, hermaphroditic, and have sim- ilar beaded viscin pollen threads. They differ in characters such as nectary type, petal size, ploidy level, pollen pore number, and habit, however. Because sect. Quelusia inhabits areas in southeastern Brazil and Chile- Argentina that are more directly related to the ancient austral temperate forests than are the areas of the remaining sections, we may infer that its relatively short floral tubes and exserted stamens are more primitive characters than the long tubes characteristic of the other sections. Simpson (1973) presented extensive biogeographical evidence that the present day South American Nothofagus forests are refuges for many taxa that may have been present and have changed little since the early to mid-Ter- tiary. Section Quelusia is entirely polyploid and may have retained certain prim- itive characters for this reason. As Simpson pointed out for other taxa, relatively little differentiation seems to have occurred in the Nothofagus forests compared to related species elsewhere. A single widespread species of Fuchsia, F. magel- lanica, is found in these forests today. In contrast, the remaining species of sect. Quelusia in southeastern Brazil have radiated strongly there, as have many other plant groups (see Smith, 1962, and Plowman, 1979, for examples). Section Kierschlegeria consists of a single species, Fuchsia lycioides, that grows in the summer-dry coastal vegetation of central Chile. This is the only species in the genus that occupies a seasonally dry habitat, and it has evolved a series of xeromorphic adaptations such as small, deciduous leaves, spinose leaf bases, and thick seed coats. The habitats where it occurs did not exist before the Pleistocene, when the Andes reached their present elevation and the cold Hum- boldt Current brought on greatly increased aridity to the Pacific coast of south- central South America (Raven, 1973; Simpson, 1975a; Solbrig, 1976). Notwith- standing this, it probably began to evolve its peculiar constellation of characters earlier in local pockets of aridity at the margins of the tropics. The lobed-adnate nectaries, triporate pollen grains, and tetraploidy make a common origin with sect. Quelusia seem likely, especially in view of the geographical proximity of these groups. Fuchsia lycioides is dioecious (Atsatt & Rundel, 1982) and its low ovule number and conceivably even its smooth viscin threads might be specializa- tions related to this aspect of its breeding system. It seems possible that the three- pored pollen characteristic of these sections and present inall genera of Onagraceae other than Fuchsia, might be a shared primitive characteristic of sects. Quelusia and Kierschlegeria, and not directly related to their polyploid nature. Sections Fuchsia and Hemsleyella are sympatric and restricted to the moist slopes of the tropical Andes, except for two disjunct species of sect. Fuchsia on Hispaniola, in the West Indies. The ancestors of these two sections probably migrated northward from the austral temperate forests in South America into new, cool, montane habitats that began to develop on the eastern slopes of the central Andes as they began to rise during the Miocene. Large areas of cool Andean forest probably did not exist until the Pliocene, however, when the major uplift of the Andes occurred. Section Fuchsia occurs in aseasonal habitats and is petaloid and mostly diploid; most species have a particular annular nectary (see Figs. 42 and 43). Section Hemsleyella, in which polyploidy is more frequent, is apetalous and has evolved a series of seasonal adaptations linked to a predom- 1982] BERRY— FUCHSIA SECT. FUCHSIA 11 inantly epiphytic habit. Members of this section usually live on rocks or trees that are subject to much higher water stress than plants of sect. Fuchsia that grow in moist soil nearby. Most species of sect. Hemsleyella have developed adaptations such as thick, tuberous stems and dry season flowering and leaf drop. Serf inn F.Unhium was recently segregated from sects. Fuchsia and Hems- Mexico northern affinities and is morphologically intermediate between sects. Fuchsia and Hems- leyella, having petals as in sect. Fuchsia and band nectaries as in sect. Hemsle- yella. A clear specialization trend occurs in the section from a nontuberous, mostly aseasonal species with large petals to a strongly seasonal, tuberous, epi- phytic, and nearly apetalous species. Unlike the sympatric sections Schufia and Encliandra, the most generalized species of sect. Ellobium occur from Costa Rica to Mexico, while the more specialized ones are restricted to Mexico. This distribution pattern and the probable young age of the related sects. Fuchsia and Hemsleyella indicates that sect. Ellobium reached Central and North America by an independent dispersal event from South America. Most likely, the ancestors of sect. Ellobium reached the high mountains of Costa Rica once the Pliocene land connection between North and South America was established, then differ- entiated farther north into seasonally drier habitats. Summarizing, the most primitive existing sections of Fuchsia may be Quelusia and Kierschlegeria, of temperate habitats in southern South America. The ances- tors of sect. Skinnera reached New Zealand from temperate South America prior to the close of the Oligocene. The common ancestor of the North American sections Jimenezia, Encliandra, and Schufia likewise may have been derived from antecedents that occurred in the Paleogene of temperate South America and reached similar habitats in North America. Generalized features for the genus may be sought by comparing characteristics common to these main groups. The differentiation of the more modern and related sections Fuchsia, Hemsleyella, and Ellobium in the Andes and, ultimately, North America, seems definitely to have been a Neogene phenomenon linked with the uplift of the Andes and the diversification of suitable habitats there, as well as with floral specialization to hummingbird pollination. The rest of this paper deals with the systematics and evolution of sect. Fuchsia and is aimed at analyzing how this particular line of the genus was able to differentiate so extensively in a relatively short time span. The patterns of evolution in this section should serve to illustrate compa- rable patterns in many other groups that inhabit the same areas, and that have been derived from diverse sources. Ecology and Geographical Distribution ecology Habitat. Plants of Fuchsia sect. Fuchsia typically occupy moist, semidis- turbed habitats of cool, montane forest. In the Andes this vegetation zone is commonly referred to as " cloud forest," but locally other terms are used such as "yungas" (Bolivia), "ceja" (Peru and Ecuador), and "montana" (northern Andes). Lauer (1979) calls this formation the "tropical upper montane forest." 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figures 3-4. Habitats of Fuchsia sect. Fuchsia in the Andes. — 3. Habitat of F. venusta, on the edge of a patch of cloud forest at 2,400 m in Dept. Tolima, Colombia. Arrow points to the position of flowering branches of F. venusta, found growing as a liana here. It also occurs as an erect or scandent shrub. — 4. Habitat in which F. venusta (Berry 3455), F. nigricans (Berry 3454), and a hybrid between them (Berry 3453) were found growing together, between La Carbonera and La Azulita, 2,220 m, Edo. Merida, Venezuela. 1982] BERRY— FUCHSIA SECT. FUCHSIA 13 According to the Holdridge life zone system, the habitats of Fuchsia correspond to the humid, very humid, or pluvial montane or low montane life zones (Ewel et al., 1976). Three of Cuatrecasas' (1958) vegetation zones in Colombia are inhabited by Fuchsia, the upper part of the Subandean Forest (ca. 1 ,000-ca. 2,400 m). the Andean Forest (ca. 2.400-ca. 3.400 m). and the Subparamo (ca. 3,400-ca. 000 formations in much of Central America (oak-pine, mixed broadleaf forests), the cloud forests of the tropical Andes are essentially continuous on the eastern slopes from 12°N to 27°S latitude and are difficult to separate into distinct phys- iognomic units (Simpson, 1979). The two main ecophysiological requirements of sect. Fuchsia are ample light and high, constant soil moisture. As shown by the long, wide vessel elements in the xylem and the lack of interxylary phloem (Carlquist, 1975), Fuchsia is clearly adapted to strongly mesic conditions. Because of their moderately high light requirements, plants of sect. Fuchsia are rarely found in tall or heavily shaded forests; the few plants that I have seen in such habitats were climbers that had reached the canopy or a light break in the forest (Fig. 3). Although seeds of Fuchsia germinate in the light, the inability of most species to withstand even moderate water stress prevents them from growing in open or strongly disturbed sites such as clearings or pastures. Many species of sect. Fuchsia are found growing in wet soil or near moving water, and they are typically found in thickets formed along road cuts or forest edges. In areas where human interference has 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 5. Altitudinal zonation of Fuchsia along a transect in Prov. La Vega, Dominican Re- public. been minimal, Fuchsia is found in naturally disturbed sites that can be fairly widespread on the steep slopes of wet forest, including treefalls, landslides, and streamsides. Larger populations are found in more open areas, however, so that a limited amount of human disturbance such as occasional clearings and road cuts seems to promote the growth and expansion of populations of most species in sect. Fuchsia (Fig. 4). A few species occur outside of the cloud forest belt, either near its upper or lower limits or in drier formations. The Fuchsia petiolaris species group occurs at high elevations near the upper cloud forest limits and sometimes even extends into the paramo (Cleef, 1979). Fuchsia vulcanica is found at 4,000 m in northern Ecuador and southern Colombia, but Fuchsia macrophylla extends down into the subtropical forest at 1,200 m in central and southern Peru. Fuchsia triphylla, an endemic on Hispaniola, is exceptional in that it occasionally grows in open fields and also occurs as low as 1,000 m. Both F. loxensis and F. dependens are found in the high, rain-shadowed central valley of Ecuador, where they are usu- ally found as mostly erect shrubs in hedgerows. Fuchsia denticulata is the only species in the section that inhabits the dry Pacific slopes of central Peru; it lives there in moist canyons mostly as an erect shrub, whereas the same species is a sprawling shrub or climber in cloud forests on the eastern slopes. The only widely naturalized species in the section, the arborescent F. boliviana, is tolerant of open habitats with moderate moisture stress, occupying a wider range of climatic conditions than most other species in the group. 1982] BERRY— FUCHSIA SECT. FUCHSIA 15 Figure 6. Altitudinal zonation of Fuchsia along a transect on the west slopes of the Sierra Nevada de Merida, Edo. Merida, Venezuela. Altitudinal separation of species. Many species within sect. Fuchsia have different elevational tolerances, and the altitudinal stratification of species on the same mountain system is one of the principal isolating mechanisms operating between species. Hybrids are most often found where the altitudinal limits of partially sympatric species overlap. Figures 5 to 1 1 illustrate several examples of the successive altitudinal replacement of species on the same mountain range, which is found throughout the range of the section. The figures are based on data from field studies and herbarium specimens. Phenology. Data from collection dates of herbarium specimens and field observations of populations of several species indicates that very little periodicity occurs in the flowering or appearance of new leaves in sect. Fuchsia. All the species are evergreen, and senescent leaves gradually fall off the older stems and are replaced by new leaves on the young axillary or terminal shoots. Individuals or local populations of F. holiviana, F. dependens, F. nigricans, and F. venusta are known to flower throughout the entire year. Though annual temperature variation in the Andean cloud forests is slight, especially compared to the daily fluctuations, precipitation is seasonal, with one 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 2200 M 1 R petiolaris 2 F. hartwegii 3 F crassistipula 4 F boliviano (naturalized) 5 F venusta Figure 7. Altitudinal zonation of Fuchsia along a transect on the east slopes of the Cordillera Central in Dept. Tolima, Colombia. or two relative dry seasons in different latitudes (Simpson, 1979). Definite growth cycles and periodicity in leaf and flower production occur in the sympatric sect. Hemsleyella. These plants, however, are mostly epiphytic or grow on rocks, where they are subject to much greater water stress, especially in drier periods. Members of sect. Fuchsia rarely experience a water deficit because they are terrestrial and grow in semi-shaded sites with high year-round soil moisture. Section Fuchsia employs a fc 'steady-state' ' flowering strategy (Gentry, 1974), whereby relatively few flowers are produced over an extended time period (usu- ally a month or two). This strategy is adapted to pollinators with fixed foraging patterns such as bees and hummingbirds. It is important to note, however, that although pollination efficiency may be heightened by this strategy, temporal sep- aration of flowering periods can not be an operative reproductive isolation mech- anism in sect. Fuchsia. GEOGRAPHICAL DISTRIBUTION Geological background of the Andes. The Andes are a classic example of a mountain range that has a volcano-plutonic origin along a convergent plate margin (Sillitoe, 1974). As outlined in recent reviews of the Andean orogeny (Hammen, 1974; Haffer, 1974; Sillitoe, 1974; Simpson, 1975b, 1979; Flenley, 1979) three events in the history of the Andes have been of key importance to the develop- 1982] BERRY— FUCHSIA SECT. FUCHSIA 17 3300 2900 2500 Figure 8. Altitudinal zonation of Fuchsia along a transect crossing the Cordillera Central in Depts. Cauca and Huila, Colombia. ment of the native flora there. First, the final and major uplift throughout the Andes, accounting for an average 2,000-3,000 m increase in elevation, occurred toward the end of the Pliocene. As Hammen (1979) states, this implies that, at least in the northern Andes, the history of the Andean forest vegetation zone dates from the upper Miocene or lower Pliocene, while the paramo flora can only be traced back to the mid- to upper Pliocene. The youth of the high Andean flora is further demonstrated by the absence of endemic plant families in the paramos and by the low number of endemic paramo genera (Cleef, 1979). Second, geomorphological and tectonic evidence indicates that, rather than having a longitudinally continuous orogeny, the Andes are actually composed of a number of separate, longitudinal segments (Fig. 12). The boundaries between these segments usually coincide with major changes in overall geology, strike, or even width of the Andes, and each segment represents a different piece of litho- sphere that was individually subducted at probably different rates. The particular history of each segment affected its faunistic and floristic composition, and anal- yses such as those of Simpson (1975b) and Duellman (1979) have shown that distributions of Andean forest species tend to coincide with the basic geological discontinuities between structural units. Finally, the Andes, as well as lowland tropical South America, were strongly affected during the Pleistocene by the same glacial cycles that drastically altered the climates of the temperate regions of the world. One to four major glaciations 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Meters 3800 3000 2200 1400 1 2 3 4 5 o GO F macrostigma 6 F. loxensis F. scabriuscula 7 F. vulcanica F sessilifolia 8 F. pallescens F. sylvatica 9 F. oriental'* F ampliata 10 F. putumayensis Figure 9. Altitudinal zonation of Fuchsia along a transect crossing the Ecuadorian Andes in Prov. Pichincha and Napo. Figure 10. Altitudinal zonation of Fuchsia along a transect crossing the Cerro de Calla-Calla, Dept. Amazonas, Peru. 1982] BERRY— FUCHSIA SECT. FUCHSIA 19 w 3200 2600 2000 1400 M 1 F. austromontana 2 F. af f. denticulata 3 F. sanctae - rosae 4 F. tincta 5 F.vargasiana 6 F. tincta x sanctae -rosae Figure 1 1 . Altitudinal zonation of Fuchsia along a transect on the eastern slopes of the Peruvian Andes in Dept. Cuzco, Prov. Paucartambo. The vertical dashed line indicates the high degree of sympatry in the vicinity of Buenos Aires. have been detected on high peaks of the tropical Andes, and detailed palyno- logical evidence from the northern Andes (Hammen, 1974) shows that a maximum temperature depression of 7°C and a lowering of the upper elevation vegetation zones by 1 ,200-1 ,500 m occurred during recent glacials (Fig. 13). Glacial episodes were often accompanied by much drier periods than at present. The implications of these climatic fluctuations are complex and different for plants of different altitudes and humidity regimes. For instance, paramo organisms, as well as those adapted to dry, lowland habitats, experienced periods of range extension and secondary contact during the glacials. Moist lowland taxa, on the other hand, underwent isolation and differentiation during glacial episodes. During intergla- cials such as at present, the dry lowland and high paramo organisms became restricted to island-like enclaves. The preceding events regarding the recent history of the Andes have several important biogeographical and evolutionary implications for sect. Fuchsia. First, assuming that the ancestors of sect. Fuchsia were adapted to cool temperate 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 12. Major structural units of the Andes where Fuchsia sect. Fuchsia occurs. Adapted from Simpson (1979) and Sillitoe (1974). 1 = Nudo de Pasto. 2 = Amotape zone, the northern limit of the Central Andes and a major tectonic segment boundary, aligned to the west with the Carnegie Ridge and the Gulf of Guayaquil. To the north a marked change in the geology of the Andes occurs, with a line of recent stratovolcanos. 3 = Huancabamba deflection. 4 ■ Pisco or Abancay deflection, marked by a major step in the coastline, a shallowing of the Peru-Chile trench, and the offshore Nazca Ridge. 1982] BERRY— FUCHSIA SECT. FUCHSIA 21 Meters 50004 21-13.000 BP VERY COLD ♦ DRY 13- 10.000 BP COLD + HUMID 6-4.000 BP MAXIMUM WARM + WET PRESENT WARM + S EMIHUMID 4000- 3000 2000 1000 Paramo (tree to snow line) Subandean forest Andean forest Lower tropical forest — savanna Figure 13. Effects of recent climatic fluctuations upon the vegetation zones in the Cordillera Oriental of Colombia. Adapted from Hammen (1974) and Lauer (1979). habitats as are almost all the extant species in the genus, they could not have arrived in the tropical Andes until the Miocene, and major opportunities for the colonization of Andean forest habitats probably did not arise until the early Plio- cene. Second, the glacial periods of the Pleistocene were probably times of range extension and secondary contact for populations of Fuchsia that would be com- paratively isolated at interglacials such as at present. Though the drier climatic affect 500 for Andean forest in the coldest glacials) would cause a great increase in its horizontal distribution, making possible migration and secondary contact of taxa across areas presently occupied by lowland vegetation. Third, a comparison of the species compositions of the different Andean structural units where Fuchsia sect. Fuchsia occurs should enable us to analyze the nature and directions of some of the migrations that have taken place in the section. >/ Fuchsia is a common element of the cloud forest stern slones of the Andes from northern Colombia northern northern Peru. The geology and physiognomy of the different struc 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 tural units of the Andes have been summarized by Simpson (1975b) and Duellman (1979). The structural units where Fuchsia sect. Fuchsia occurs are shown in Figure 12. Two basic changes from Simpson's (1975b and 1979) system are in- troduced, however. First, a small unit called the Amotape-Huancabamba zone is added because both the Amotape zone in southern Ecuador (as treated in Sillitoe, 1974) and the Huancabamba deflection in northern Peru constitute clearly denned tectonic boundaries where major changes in the overall geology of the Andes occur (Sillitoe, 1974). It is therefore misleading to extend the structural units of the Northern Andes south of the Amotape zone to the Huancabamba deflection or to extend those of the Central Andes north of the Huancabamba deflection into Ecuador. Though several authors considered the Huancabamba deflection to be the main north-south migration barrier in Andean organisms (Vuilleumier, 1971; Simpson 1975b, 1979; Duellman, 1979), this is least true for cloud forest or- ganisms. The 21 species of amphibians and reptiles that occur both north and south of the Huancabamba deflection are primarily cloud or wet forest inhabitants on the eastern slopes of the Andes (Duellman, 1979). In sect. Fuchsia and perhaps in other cloud forest organisms, it is perhaps more appropriate to treat the entire Amotape-Huancabamba zone as the transitional area between the species of the Northern and Central Andes. The second change is the separation of Simpson's (1975b, 1979) "Cordillera Oriental" of the Central Andes into two units, the Cordillera Central to the north and the Cordillera Oriental to the southeast. This aids our analysis of species distributions by producing three units in the Central Andes that are comparable in size and extent to the units of the Northern Andes. Technically and geologi- cally, however, there are ample reasons for this division as well. The dividing line of the two units corresponds to the Pisco or Abancay deflection (Fig. 12), which is marked on the western side by the offshore Nazca ridge, a major step in the coastline, and a shallowing of the offshore Peru-Chile trench. Along the boundary in the mountains to the east, there is a marked northward narrowing of the cordilleras, a change in direction of the Eastern Cordillera, and important changes in the Mesozoic paleogeography (Sillitoe, 1974). Figure 14 summarizes the species and species group distributions in the dif- ferent structural units of the Andes where sect. Fuchsia occurs. The following discussion deals with the species relationships of each unit. Reference is also made to species distribution maps (Figs. 55-67) in order to explain patterns ob- served within and between the structural units. Northern Andes Nevada de Santa Marta. This is a small, isolated volcanic range and the northernmost segment of the Andes. It has undergone a complex geological history, but the Sierra Nevada de Santa Marta was underwater until the Miocene and apparently experienced its major upheaval in the Quaternary (Simpson, 1975b). The sole species of sect. Fuchsia that occurs there is F. magdalenae, which is endemic to the Andean forest and subparamo of this small, but high, mountain massif. General floral characters place this species closest to the Peruvian-cen- tered F. denticulata species group, but it is very unusual in the section because 1982] BERRY— FUCHSIA SECT. FUCHSIA 23 HISPANIOLA 2/2 1 SANTA MARTA 1/1 4 CORDILLERA CENTRAL 23/2 10 8 CORDILLERA ORIENTAL 10/1 8 NORTHERN ANDES 29/7 9 AMOTAPE- HUANCABAMBA ZONE 10/2 6 1 CORDILLERA OCCIDENTAL 5/0 5 4 ^ CORDILLERA CENTRAL 22/14 8 1 "* CENTRAL ANDES CORDILLERA ORIENTAL 10/6 5 29/20 10 Figure 14. Schematic representation of the species composition of Fuchsia sect. Fuchsia in the major structural units of the Andes (see also Fig. 12). The first number of each pair separated by a slash is the total number of species in that unit; the second number is the number of species found only in that unit (= endemics). The bold face number below each of the above pairs is the number of species groups present in that unit. Numbers of species shared by two units appear between the arrows connecting different units. of its tetraploidy with biporate pollen and its smooth, non-annular nectaries. Although suitable habitats for Fuchsia are found less than 40 km away across the Cesar depression in the Sierra de Perija of the Cordillera Oriental, the only species of the genus that occurs there is F. gehrigeri, which bears no close 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 relationship to F. magdalenae and is found elsewhere only in the Merida Andes of Venezuela. Mayr (1963) has noted that peripheral isolates of many genera show one or more of the following characters: they usually occur in a small geographical area, have low absolute population sizes, and differ from the main body of the species in several, often unique morphological or other characters, criteria that apply well to F. magdalenae. Cordillera Oriental. This unit is characterized by a comparatively low num- ber of species but a high number of species groups, low endemicity, and strong east-west and north-south connections to neighboring Cordilleras. As shown by van der Hammen's detailed geological and palynological studies of the Cordillera Oriental of Colombia (Hammen, 1974) this unit was uplifted to nearly its present level in the mid- to late Pliocene, followed by a long sequence of glacials and interglacials during the Pleistocene. Through the examination of lacustrine sediments, van der Hammen and coworkers were able to document recent climatic extremes. During the Last Glacial, at 21,000 years BP, the climate around the high plains of Bogota became extremely cold and dry, with a ca. 7°C 200-1.500 Middle interglacial when the temperature became 2°C warmer and the vegetation zones several hundred m higher than at present. The average effects of these climatic changes on the altitudinal distribution of different vegetation zones are illustrated in Figure 13. Considering the magnitude of the vegetation zone fluctuations during the Pleis- tocene, migrations of Fuchsia across the Magdalena and Cauca River valleys to the Cordilleras Central and Occidental could have occurred almost without in- terruption or via bird dispersal across short lowland barriers. At present, there is only a 35 km gap between the 1,000 m contours of the Cordillera Central and the Cordillera Oriental at 5°N Latitude, which is the approximate latitude of Bogota and where F. hirtella, F. petiolaris, and F. venusta are found in both cordilleras. These species are all more common in the Cordillera Oriental than in the Cordillera Central, and they most likely migrated in a westerly direction to attain their present distribution there. The distribution patterns of the large paramo genera Espeletia and Puya give strong evidence of the same pattern of a Cordillera Oriental source area with subsequent migrations to the Cordillera Central (Cuatrecasas, 1979). The recent origin of these paramo genera (Hammen, 1979) further supports the idea that the Pleistocene was an important period for plant migrations in the Andes. southern Macizo northernmost corridor that spread into the three different units that diverge northwards from it. Several species of Fuchsia have present day distributions that indicate they used this corridor to reach the southern part of the Cordillera Oriental from the south. These include Fuchsia sessilifolia (Fig. 63), F. scabriuscula (Fig. 56), and F. cuatrecasasii (Fig. 58). The first of these species probably used the Nudo de 1982] BERRY— FUCHSIA SECT. FUCHSIA 25 Pasto as a corridor to spread northwards into all three units of the Northern Andes. Fuchsia verrucosa (Fig. 65), which is centered in the Cordillera Oriental, has a distribution indicating that it migrated southwards along the eastern edge of the Nudo de Pasto to reach its present locations in the Cordillera Central in southern Colombia. In the north, the Cordillera Oriental splits into two ranges near the Colombian- Merida Merida the low Tachira depression, which constitutes a secondary dispersal barrier within this unit. Six species occur just on the Colombian side of the barrier, and the only endemic species in this unit, F. gehrigeri, is found only in the Merida Andes and the Sierra de Perija. In summary, the distribution patterns of this unit show close ties to the ad- jacent Cordillera Central, with all but one species and numerous species groups shared between them. The presence of so many species in common and their relative lack of differentiation in the different cordilleras suggests that they have not been separated for very long or that they have been in secondary contact, and it is shown how Pleistocene climatic changes could easily account for migra- tions or secondary contact of species between the two cordilleras. Cordillera Central. This unit has the highest representation of species and species groups, yet the next lowest proportion of endemism of any structural unit. This is largely due to the central position of this unit in the Northern Andes and its close proximity to three adjacent units. The more gently sloping eastern flanks of the Cordillera Central offers a greater variety of habitats than the com- paratively steep slopes of the Cordillera Occidental. Twelve species are common to both the Cordillera Central and the Cordillera Occidental. The high interandean valleys of Ecuador, the Nudo de Pasco, and the high plateau separating the Cauca and Patia Rivers are major corridors for species exchange between these two units, and they all must have greatly facil- itated migration of upper montane species during Pleistocene glacials. The Fuchsia petiolaris species group, which is strongly centered in the Cor- dillera Central, has complex patterns of variability between populations and is the source of considerable taxonomic confusion. In Ecuador, populations of this species group are found on many of the high, semi-isolated peaks that are found in both cordilleras (Fig. 59). Since members of this group typically occur at high elevations near tree line, they were probably more strongly affected by Pleisto- cene events than other groups in the section; Sauer (1971) recorded three main glaciations in Ecuador. The F. dependens species group also developed strongly in a relatively small area centered in the Cordillera Central in southern Colombia (Fig. 66). Cordillera Occidental. Seventy-five percent of the species in this unit are shared with the Cordillera Central, while only three species are endemic there. Fuchsia sylvatica occurs on the Pacific slopes in Ecuador and is vicarious with F. nigricans, which is found in the same cordillera but north of the Patia River val- ley in southern Colombia. Fuchsia macrostigma, however, occurs on both sides of the Patia River, which is a secondary migration barrier within this unit. Two of the 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 endemic species, F. cinerea and F. polyantha, are very localized and occur on the edge of the Nudo de Pasto. Amotape-Huancabamba Zone Despite its relatively small area (Fig. 12), this unit has the same number of species as the much larger Cordillera Oriental of the Northern Andes. This reflects its key position bordering four different structural units, as well as the equatorially based center of species diversity in the section. Fuchsia glaberrima is the only species that occurs both north and south of this unit. The Amotape-Huancabamba Zone thus constitutes the major transitional zone between the species of the Northern and Central Andes. The species found within this unit, however, are a mixture from all four adjacent structural units and include two endemic species. This area is still insufficiently collected because of its inaccessibility, and future explorations will be certain of finding novelties in Fuchsia. One of the most distinctive species in the section, F. steyermarkii, occurs there and appears to be most closely allied to species from the Cordillera Central in northern Peru. Central Andes Cordillera Occidental. The cloud forest belt on the western side of the Andes ends near the Peruvian-Ecuadorian border. As a result, most of the Cordillera Occidental in Peru is too dry for Fuchsia, and four of the five species present in this unit are only found north of 8°S Latitude. These species inhabit local patches of mesic forest that is more a mixed scrub than true cloud forest. Fuchsia ayava- censis is the only species that is largely restricted to this unit, and it forms part of the northern F. petiolaris species group. Three of the species in the unit are shared with the Cordillera Central of Peru and probably were dispersed across the Maranon River valley from the east. At present, there are only approximately 25 km separating the closest populations of F. mathewsii on either side of the Rio Maranon. The genus Ascidiogyne (Compositae) provides another interesting example of this trans-Maranon disjunction (Cuatrecasas, 1979). This unique genus has only two species, one in the high "jalca" of Dept. Amazonas in northern Peru, and the other across the Maranon valley to the west in Dept. Cajamarca. Fuchsia denticulata is an exceptional species in this unit because it is found between 9°S and 13°S Latitude in isolated pockets of mesic vegetation on the otherwise arid Pacific slopes of Depts. Lima and Ancash. This species is wide- spread on the eastern slopes of the Andes, however, from Dept. Huanuco in Peru to Dept. Cochabamba in Bolivia. This disjunct pattern was previously discussed by Simpson (1975b, 1979) for other mesic Andean species. Simpson (1975a) pos- tulated that the high parts of the Cordillera Occidental and its upper western slopes experienced heavier precipitation during Pleistocene glacial periods than at present. At these times, east-west migrations of mesic montane plant taxa were made possible across the i4 nudos" or low connecting paths between the Pacific and Amazonian slopes of the Peruvian Andes. Fuchsia denticulata is a clear example of migration directly across the Andes via the Nudo de Pasco, at the 1982] BERRY— FUCHSIA SECT. FUCHSIA 27 latitude of Lima. There are fewer than 100 km (air distance) now separating the closest populations of F. denticulata on the western slopes above Lima and on the eastern slopes in Dept. Junin (Fig. 61). Besides its presence in cloud forests of the Cordillera Central in Junin, F. denticulata also occurs in higher interandean valleys such as the valley of Comas, which has an intermediate habitat type between the eastern "ceja" and the small mesic pockets restricted to protected canyons on the western slopes. This distribution pattern suggests how species with wide ecological tolerances may have been able to cross the Nudo de Pasco under more humid conditions than exist at present. Once F. denticulata reached the Pacific slopes, it probably spread north (and south?) in what may have been a continuous moist forest belt during Pleistocene glacials (Simpson, 1975a, 1979). Other species such as Arracacia incisa Wolff. (Umbelliferae) show the same pattern as F. denticulata, and Simpson (1975b) gave several examples of species that are presently disjunct between the western slopes above Lima and the eastern slopes further south near Cuzco. These latter species may have used a more southern pass, the Nudo de Vilcanota. Cordillera Central. This unit has almost as high a species diversity as the Cordillera Central of the Northern Andes, but two fewer species groups and a much higher proportion of endemic species. It is much more dissected and is in contact with fewer units than the northern Cordillera Central. The adjacent Cor- dillera Occidental is too dry to allow major opportunities for the establishment of new populations of Fuchsia, and strong migratory barriers are present to the north in the Huancabamba depression and to the south in the Rio Apurimac valley. Though there are proportionately fewer species groups present in this unit than in most others, at least half of those groups present are restricted or strongly centered here. All four species of the F. simplicicaulis species groups are endemic to the Cordillera Central, as are all but one species each of the F. macrophylla , F. boliviana, and F. decussata species groups. Speciation has probably been facilitated by the presence of several semi-isolated ranges such as the Cordillera Azul on the Huanuco-Loreto border. Five very localized endemics are concen- trated in Dept. Amazonas of northern Peru. Cordillera Oriental. As in the Cordillera Oriental of the northern Andes, ten species of Fuchsia occur in this unit, but a lower number of species groups and a much higher proportion of endemic species are found here. The Cordillera Peru northern Peru livia, showing the continuity of distribution of most species in this unit across the outer slopes of the Titicaca Basin. Fuchsia boliviana just barely extends north of the Rio Apurimac into the Cordillera Central, and it may only be naturalized there. Fuchsia tincta and F. vargasiana are restricted to one or two valleys in southern Peru, but are very closely related to the Bolivian F.furfuracea. Fuchsia macrophylla and F. decussata are both centered in the Cordillera Central and northern northern 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Hispaniola Two species of sect. Fuchsia are widely disjunct from the rest of the species in the section and are endemic to the Caribbean island of Hispaniola. A similar Andean-Hispaniola disjunction of montane species has recently been reported by Cuatrecasas ( 1 979) for the genus Lae stadia (Compositae) , which has four species in the Andean paramos, one in high elevation Costa Rica, and one on Hispaniola. The ancestors of the fuchsias on Hispaniola probably reached the island from South America by long distance bird dispersal, a mechanism that has been pro- posed to account for other disjunctions in the genus, such as F. cyrtandroides on Tahiti. Judging from the lack of clear affinities of either of the two Hispaniola species to any species group on the South American mainland, their arrival to the island does not appear to be recent. It is furthermore unclear whether F. triphylla and F. pringsheimii were derived from a common ancestor or whether they arrived separately, by a "double invasion" in the sense of Mayr (1963). Supporting the possibility of a common ancestor, both species have the combi- nation of tetraploidy with biporate pollen, which is unusual in the genus. In addition, they are presently isolated altitudinally throughout their range, except in certain contact areas where partly fertile hybrids are formed. On the other hand, the two species differ widely in morphological characters (Table 19). Fuch- sia triphylla is morphologically similar to the main body of Andean species of sect. Fuchsia, but F. pringsheimii has several floral characters not clearly related to any section in the genus, such as particular non-annular nectaries and very large, emarginate petals. If a double invasion did occur, then F. pringsheimii is most likely the descendant of the earlier immigrant, and its unique assemblage of characters may even be derived from the early offshoot of the genus that had reached Central America. Summary of Distribution Patterns A small number of taxa in Fuchsia sect. Fuchsia show wide disjunctions in range from the main body of species in the Andes or from other members of their species groups. These include species such as F. magdalenae in the Sierra Ne- vada de Santa Marta and F. triphylla and F. pringsheimii on Hispaniola. These species show fundamental differences from each other and from the main body of the section in several morphological and cytological features. In this way, they can be considered peripheral isolates in the sense of Mayr (1963) and may rep- resent early offshoots of the section that were isolated at the margins of the range. In the contiguous structural units of the Andes where the bulk of the species in sect. Fuchsia are concentrated, a major change in the species and species group composition occurs in southern Ecuador and northern Peru in a transitional area called the Amotape-Huancabamba zone. Only one species and four species groups occur both north and south of this area. Consequently, it has been possible to compare the distribution patterns of the species in the Northern Andes with those of the Central Andes. As seen in Figure 14, the Northern and Central Andes each have 29 species and nearly the same number of species groups. The structural units with the greatest number of species are the Cordillera Central of the Northern Andes and 1982] BERRY— FUCHSIA SECT. FUCHSIA 29 the Cordillera Central of the Central Andes. While we cannot place the center of diversity of the section in either one of these two areas, it is clearly located on the eastern slopes of the Andes in Ecuador and northern Peru. From there, we find a gradual reduction in species number to the north and south in a pattern analogous to the distribution of hummingbird species (Skutch, 1973). Just over half (32 of 61) of the species in sect. Fuchsia are restricted to a single structural unit of the Andes or to the island of Hispaniola. As might be expected of easily dispersed plants in the cloud forest belt, this proportion is lower than that normally observed in paramo plants, where high degrees of local endemism are common (Simpson, 1975b; Cuatrecasas, 1979). The proportion of endemics (= species limited to a single structural unit) in sect. Fuchsia varies significantly, however, between the Northern and Central Andes. In the Northern Andes, with 24% endemics, migration between structural units has been facili- tated by the proximity of the three Cordilleras and the numerous corridors that would be available in glacial periods of lowered vegetation zones. Accordingly, four species in the Northern Andes are present in all three Cordilleras, and twelve are found in two Cordilleras. In contrast, the Central Andes have 69% of the species endemic, with only one species present in all three Cordilleras and five in two cordilleras. The Central Andes have much stronger migration barriers because of the greater spatial isolation of the Cordilleras Central and Oriental, and the aridity of the Cordillera Occidental. The key migratory corridor in the Northern Andes has been the Nudo de Pasto (Macizo Colombiano). The three northern cordilleras merge at this massif, and many species have used it to disperse latitudinally or longitudinally into the different structural units. A total of 14 species are present along the Nudo de Pasto (as defined by Duellman, 1979), which is the richest local species concen- tration of sect. Fuchsia in the Andes. Although the Cordillera Oriental of the Northern Andes is poorer in species number than the adjacent Cordillera Central, it has been an important source area of several species present in the northern part of the Cordilleras Central and Occidental. At least three species have migrated westwards across the Magdalena valley, and one or two migrated further west across the Cauca valley into the Cordillera Occidental. In Ecuador, the opportunities for migration between the two cordilleras have been numerous, and it is difficult to determine for most species in which direction they may have spread. Most species groups of the Central Andes are restricted to a single structural unit, but three migration paths between different structural units were identified, 1) the western migration of two or three species across the Maranon River in north- ern Peru, 2) the western migration of Fuchsia denticulata across the Andes to the Pacific slopes via the Nudo de Pasco, and 3) the southern expansion of two species from the Cordillera Central into the Cordillera Oriental. Secondary barriers exist within many structural units that have also been important filters to migration. For example, only three often species in the Cordi- llera Oriental of the Northern Andes occur on both sides of the Tachira depression in western Venezuela. Similarly, the Patia River valley in southwestern Colombia and the Huallaga and Mantaro Rivers in central Peru have also acted as selective barriers within their respective units. 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 Reproductive Biology FLORAL BIOLOGY The maturation sequence of flowers seems to be similar in all species of sect. Fuchsia. Anthesis usually occurs in the early morning, at which time the stigmatic surface becomes wet with a sticky exudate and is presumably receptive (Raven, 1979a). Anther dehiscence is usually delayed from several hours to one or two days, depending on the species, but the stigma remains receptive in all species until after the anthers open; protogyny is ubiquitous but does not preclude self- pollination. Flowers last an average of three to four days from anthesis to abscis- sion. A spatial separation of 2-20 mm usually exists between the stamens and the exserted stigma, further reinforcing outcrossing. Notable exceptions, however, are F. boliviano, F. nigricans, and F. verrucosa, in which at least some populations have the stigma in close contact with the upper staminal whorl. In other species, if no pollinator visits are made before anther dehiscence, the stigma-anther separa- tion is usually small enough that wind or other mechanical factors are sufficient to cause self-pollination. Self-pollination is favored because the pollen is held to- gether in elastic strings by their viscin threads and because the stigma is situated below the anthers in the hanging flowers that characterize sect. Fuchsia. As far as known, all species of Fuchsia are self-compatible, which is consid- ered an advanced character state in the family (Raven, 1979a). Fuchsia is the only genus in the family in which marked protogyny occurs. Although Raven (1979a, p. 578) reports protogyny in Circaea, it is barely pronounced and there- fore not really comparable to the protogyny found in certain sections of Fuchsia. Controlled crosses in both native and naturalized populations of F. boliviano show that this species is self-compatible and largely autogamous. The high degree of autogamy is probably exceptional in the section and undoubtedly helps to account for the widely naturalized distribution of F. boliviano at present. In this species, protogyny is very poorly developed, and the anthers and stigma are in close contact; naturalized populations at El Junquito, Venezuela, were found to self-pollinate in bud. Male sterility is an outcrossing device that is found in the Onagraceae only in certain sections of the genus Fuchsia. All the known sections with male sterility are small and have distributions that are peripheral to the main Andean- Brazilian center of the genus. One possible case of gynodioecy has now been found in sect. Fuchsia, however. Certain populations of F. hartwegii from several different localities in Colombia include male-sterile individuals. Other populations of the same species apparently consist entirely of hermaphroditic plants. A similar sit- uation has recently been documented in F. paniculata (sect. Schufia) of Central America (Breedlove et al., 1982). Further field work and breeding studies are needed to clarify the situation of male sterility in F. hartwegii, but its confirmation would constitute the fifth independent occurrence of male sterility in the genus. POLLINATION Table 3 shows the features of sect. Fuchsia that strongly associate it with hummingbird pollination. In addition, the beaded viscin threads, which enable 1982] BERRY— FUCHSIA SECT. FUCHSIA 31 Table 3. Characters of Fuchsia sect. Fuchsia adapted to hummingbird pollination. Hummingbirds i Fuchsia sect. Fuchsia Distribution: General behavior/ phenology: Pollination-related characters: New World, highest diversity in the tropical Andes. Tropical species are non-migratory, territorial, diurnal. Poor sense of smell, preference for red coloration, long bill and suck- ing tongue, hovering approach, high energy requirement, and fre- quent daily feeding. Tropical Andes, Hispaniola Plants long-lived with diurnal anthe- sis, long-lived flowers (av. 3-4 days), and long flowering season. Flowers odorless, reddish, tubular, divergent to pendulous, copious nectar production. 1 From Grant & Grant, 1968; Percival, 1969; Raven, 1972a. hundreds of grains to be removed in a single bundle, appear to be of adaptive value in bird pollination (Percival, 1969; Skvarla et al., 1978). The hummingbirds feed on the nectar that is produced at the base of the floral tubes, and, unlike many groups of insects, they remain active in the rain and at cool temperatures, an obvious advantage for montane cloud forest groups such as Fuchsia (Heinrich & Raven, 1972). Only incidental observations were made on pollination in Fuchsia during the course of field work. Numerous unidentified hummingbird species were seen visiting different species of Fuchsia sects. Fuchsia and Hemsleyella, and no non- avian visitors were observed. On Hispaniola, there are two species of Fuchsia that occur at different altitudes and three resident species of hummingbirds. Chlo- rostilbon swainsonii, which is the "large highland species" of Lack's (1976) niche theory of West Indian hummingbirds, feeds exclusively below the canopy (Lack, 1976) and has been photographed visiting F. triphylla (Fig. 32). Lack's k 'large lowland species," Anthracothorax dominicus, is actually present in the highlands there, but is restricted to the tree canopies. The third "small species" completing the hummingbird fauna of the island is Mellisuga minima, which occurs at all elevations. But because it is the second smallest bird in the world it is therefore very unlikely to pollinate the moderately long-tubed F. triphylla and F. prings- heimii. The hybrids found between these two species of Fuchsia in areas where their altitudinal limits are in close proximity can almost certainly be attributed to visits by Chlorostilbon, which ranges throughout the elevational ranges of both species and occupies the understory niche almost exclusively. Naturalized populations of F. holiviana were observed over a several day period in March 1979, at El Junquito and Colonia Tovar, Venezuela. Two differ- ent populations were aggressively defended and visited throughout the day by the long tailed sylph, Aglaiocercus kingi. None of the visits were legitimate pol- linations, however, because the hummingbird consistently pierced the base of the tube and robbed the flowers of their nectar. At higher elevations in the Colonia Tovar, the same species of Fuchsia was visited by the booted racket-tail, Ocrea- tus underwoodii, which appeared to be making legitimate visits without piercing the tube. 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 ® ® \0jjL. r i n Figures 15-16. Meiotic chromosomes in Fuchsia sect. Fuchsia, metaphase I. — 15. F. loxensis, 11, from Berry 3133 (MO). — 16. F. verrucosa, a tetraploid, n = 22, from Berry 3662 (MO). In several cases two closely related species differ mainly in floral tube length and coloration, yet occur together or in close proximity without the presence of intermediate plants. Fuchsia poly ant ha, which has red flowers 34-43 mm long, * grows with F. sessilifolia, which has pale pink tubes 13-20 mm long. A similar pattern is found between F. tine t a and F. vargasiana, and between F. macro- phylla and F. macropetala. This pattern suggests that hummingbird species are species-specific pollinators of Fuchsia, the specificity linked to tube lengths and color patterns, or that flower dimensions mechanically act against interspecific pollination. In hawkmoth-pollinated species of Oenothera, Gregory (1964) found that short-tubed flowers were only effectively pollinated by short-tongued hawk- moths, but long-tubed species were able to use long-tongued as well as short- tongued hawkmoths as effective pollinators. Further observations on the pollination system of sect. Fuchsia are needed and could analyze specific problems such as the degree of specificity between hummingbirds and Fuchsia species, the relation of bill length to tube length, the importance of territoriality of hummingbirds in pollen dispersal, and the possible presence of insect pollinators. VEGETATIVE REPRODUCTION Trailing stems or broken stem pieces of Fuchsia root readily in the moist humus that is usually found in thickets where most species of the genus occur. Large colonies of F. holiviana that reproduce mainly in this manner were seen in Venezuela (naturalized) and in Bolivia (native). The same process also gives rise in other species to dense thickets of plants that probably arose from a single individual. In 1979 I returned to the precise locality of a hybrid between F. gehrigeri and F. nigricans that had first been collected in 1958 and found what was most probably the same population. Regrowth from viable hybrid seed cannot be discounted, but the uniformity of the population and the large number of vigorous, young shoots make it more likely that vegetative reproduction has maintained this colony for over 20 years. 1982] BERRY— FUCHSIA SECT. FUCHSIA 33 DISPERSAL Fuchsia is the only genus of Onagraceae with fleshy fruits, clearly an advanced character within the family. The berries are plump and generally reddish at ma- turity, with a high sugar content, and are clearly adapted to dispersal by birds. Berries are well suited for long distance dispersal, and the distribution of widely disjunct species such as F. cyrtandroides on Tahiti and F. triphylla on Hispaniola have been explained by internal transport by birds (Carlquist, 1967). A similar long-distance dispersal event has been postulated for the early estab- lishment of the Mexican and Central American sections of the genus. On a local scale, bird dispersal is an effective means of spreading seeds to new habitats, and it probably has been instrumental in the extension of many species into different structural units of the Andes. Cytology Compared with those of other Onagraceae, the chromosomes of Fuchsia are relatively large and unspecialized, contract evenly in the course of mitosis, and show poor differentiation into heterochromatic and euchromatic portions (Ku- rabayashi et al., 1962). Translocation systems common in the tribe Onagreae and aneuploidy prevalent in Lopezieae and in Clarkia are not found in Fuchsia, which retains the basic chromosome number of the family, x = 11. Reports of chromosome numbers in the native species of Fuchsia previous to the beginning of the present study in 1977 were mostly diploid (n = 11), except for two tetraploid species in sect. Quelusia, Fuchsia lycioides (sect. Kierschle- geria), and a single individual of sect. Encliandra (Warth, 1925; Haque, 1952; Chaudhuri, 1956; Beuzenberg & Hair, 1959; Kurabayashi et al., 1962; Huynh, 1965; Breedlove, 1969). The chromosome numbers of only three of the 61 species now recognized in sect. Fuchsia had been reported, all diploid, but since vouchers apparently were not preserved, these reports are of little value. Recent studies conducted at the Missouri Botanical Garden have shown sect. Quelusia (eight spp. , Brazil and the southern Andes) to be polyploid, with three tetraploid and one octoploid species known; tetraploid species have been found in the Andean sect. Hemsleyella (P. Berry and T. P. Ramamoorthy, unpublished). Because these sec- tions are considered to be rather closely related to sect. Fuchsia, an extensive sur- vey was undertaken in sect. Fuchsia to determine if polyploidy had played any sig- nificant role in the broad diversification of this section. Methods. Gametic counts were obtained from pollen mother cells in young, field collected buds fixed in 3 parts absolute ethanol : 1 part glacial acetic acid for 70% rm 2-4 hydroxyquinoline, then fixed as above in acetic alcohol for 10 min. to 6 hr. The roots were hydrolyzed in 1 N HC1 for 20 min. at 60°C and stained for 12-24 hr. in aceto-carmine or lacto-propionic orcein; they were then squashed and heated in equal parts glycerine and 45% acetic acid. Results. One hundred new counts were obtained for populations of Fuchsia 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 4. Chromosome numbers in Fuchsia sect. Fuchsia. 1 Taxon // In Collection Data or Reference F. abrupta I. M. Johnston F. ampliata Benth. F. austromontana I. M. Johnston F. ayavacensis Humboldt, Bonpland & Kunth F. boliviana Carriere F. canescens Benth. F. caucana P. Berry F. ceracea P. Berry F. cinerea P. Berry F. corollata Benth. von F. crassistipula P. Berry F. denticulata Ruiz & Pa- von F. aff. denticulata Ruiz & Pavon 11 11 11 II F. cf. andrei I. M. Johnston 11 11 11 11 11 11 11 11 11 II 11 11 11 11 11 22 22 22 11 11 F. corollata x Icaucana 22 (probable hybrid) F. corymbiflora Ruiz & Pa- 1 1 11 11 11 11 11 22 22 44 22 22 22 Peru, Junin: 58 km E of Satipo, B3078 (MO) Peru, huanuco: above Chinchao, Mathias & Taylor 4033 (RSA) 2 Ecuador, imbabura: 8 km W of Laguna Cuicocha, B3I69 (MO) Ecuador, pichincha: 9 km W of Chillogallo, B3234 (MO) Peru, amazonas: 42 km of E of Pedro Ruiz, B3628 (MO) Peru, cuzco: 8 km E of Abra de Acanaco, B2594 (MO) Peru, cuzco: 58 km from Ollantaytambo to Quilla- bamba, B3035 (MO) Peru, piura: 35 km E of Canchaque, B3630 (MO) , B3552 (MO) Colombia, antioquia: Alto de Minas, Escobar WOO (MO) Colombia, tolima: 29 km W of Fr Mexico, chiapas: Chamula, Breedlove 80/8 (DS) Peru, cuzco: Urubamba, B256I (MO) Peru, apurimac: Ampuy, Stork, Horton & Vargas 10595 (UC) 2 Peru, huanuco: 5 km SE of Carpish, Stock & Hor- ton 9921 (UC) 2 Colombia, cauca: 31 km E of Totoro, B3578 (MO) Colombia, cauca: 34 km E of Totoro, B3579 (MO) Colombia, narino: 24 km E of Pasto, B3252 (MO) Peru, huanuco: 5 km W of Carpish pass, B308I (MO) Ecuador, carchi: Tufino, B3I47 (MO) Ecuador, carchi: ridge between Tulcan and El Car- melo, B3159 (MO) Ecuador, imbabura: Hacienda Curubi, B3I73 (MO) Eucador, imbabura: 1 1 km SW of Otavalo to Mojan- da, B3I75 (MO) Colombia, cauca: 11 km E of Totoro, B3575 (MO) Colombia, cauca: 4 km W of Gabriel Lopez, B3576 (MO) Colombia, cauca: 23 km E of Purace, B3584 (MO) Peru, huanuco: Carpish, B3082 (MO) Colombia, tolima: 40 km W of Fresno, B3553 (MO) Bolivia, la paz: road to Chulumani, Albert de Esco- bar 1305 (TEX) Peru, cuzco: ca. 70 km from Quillabamba to Ollan- taytambo, B2573 (MO) Peru, cuzco: 72 km from Quillabamba to Ollantay- tambo, B3042 (MO) Peru, cuzco: ca. 65 km from Quillabamba to Ollan- taytambo, B3048 (MO) Peru, ayacucho: 41 km E of Tambo, B3050 (MO) Peru, cuzco: Km 131 of Cuzco-Paucartambo-Pilcopa- ta road, B2598 (MO) 1982] BERRY— FUCHSIA SECT. FUCHSIA 35 Table 4. Continued. Taxon n F. depend ens Hook. F. ferreyrae P. Berry F. fontinalis J. F. Macbr F. gehrigeri Munz F. gehrigeri x nigricans F. gehrigeri x Ivenusta F. hartwegii Benth. F. hirt el I a Humboldt, Bon- c pland & Kunth F. hirt el I a x venusta F. lehmannii Munz F. loxensis Humboldt, Bon- pland, & Kunth F. macrophylla I. M. Johnston F. magdalenae Munz F. mathewsii J. F. Macbr F. nigricans Linden 1 1 2/i 44 Collection Data or Reference Colombia, narino: 13 km S of Pas to, B3145 (PSO) Ecuador, carchi: 4 km W of El Carmelo, B3164 (MO) Peru, juni'n: 68 km W of Satipo, B3073 (MO) Peru, amazonas: 62 km from Balsas to Leimebam- ba, B3608 (MO) Venezuela, merida: 43 km W of El Aguila to Pifian- go, B3138 (MO) Venezuela, merida: 9 km above Santo Domingo to Apartaderos, B3139 (MO) Venezuela, merida: Km 19-20 from Apartaderos to Santo Domingo, B3599 (MO) Venezuela, tachira: 6 km E of Zumbador to Queni- quea, B3297 (MO) Venezuela, trujillo: Visiin, above Las Mesitas, B3129 (MO) Venezuela, tachira: 3.5 km E of Zumbador, B34I4 (MO) Colombia, putumayo: W of Sibundoy, B3255 (PSO) Colombia, valle: above El Guayabo, Palmira to Taco, B3568 (MO) Colombia, cauca: 21 km E of Piendamo, B3572 (MO) Colombia, cundinamarca: 23 km from Pacho to Zi- paquira, B3538 (COL) Colombia, cundinamarca: Km 34 of old Bogota-Fu- sagasuga road, B3543 (MO) Colombia, cundinamarca: Km 36 of old Bogota-Fu- sagasuga road, B3543-B (no voucher) Colombia, cundinamarca: Km 39 of old Bogota-Fu- sagasuga road, B3547 (MO) Ecuador, zamora-chinchipe: 27 km E of Loja, B3200 (MO) Ecuador, zamora-chinchipe: below El Retorno, Mathias & Taylor 5200 (RSA) 2 Ecuador, azuay: above Sayausi, B3185 (MO) Ecuador, loja: 5 km S of Saraguro, B3192 (MO) Ecuador, pichincha: Panamerican Hway S of Quito, B3233 (MO) Peru, ayacucho: 58 km E of Tambo, B3053 (MO) (Wright, 1978a) Cultivated plants from Colombia, SE slopes Sierra Nevada de Santa Marta, Trombachuca Valley, Wright (RDG) 3 Peru, cajamarca: 39 km W of Celendin to Cajamar- ca, B3602 (MO) Peru, amazonas: 42 km E of Balsas to Leimebamba, B3603 (MO) Peru, amazonas: 51 km E of Balsas to Leimebamba, B3606 (MO) Peru, amazonas: 24 km above Leimebamba to Bal- sas, B3612 (MO) Venezuela, trujillo: Agua Negra, 8 km SW of El Batatal, B3093 (MO) Venezuela, trujillo: 9 km SW of El Batatal, B3095 (MO) 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 4. Continued. Taxon F. nigricans x venusta F. orientalis P. Berry F. pallescens Diels F. petiolaris Humboldt, Bonpland & Kunth F. pilosa Fielding & Gard- ner F. polyantha Killip F. pringsheimii Urban F. rivularis J. F. Macbr. F. sanctae-rosae Kuntze F. sessilifolia Benth F. sylvatica Benth F. tincta I. M. Johnston F. triphylla L. F. vargasiana Munz pland & Kunth n 11 22 c.z ii 1 22 22 11 F. venusta Humboldt, Bon- 11 11 c.ll In 22 44 Collection Data or Reference c.22 Venezuela, trujillo: 12 km from San Rafael to Tru- jillo, B3104 (MO) Venezu , merida: above La Mucuy, Berry in 1980 (MO) Venezuela, merida: La Carbonera, B3451 (MO) Venezuela, merida: between La Azulita and La Carbonera, B3453 (MO) Ecuador, zamora-chinchipe: 24 km E of Loja, B3199 (MO) Ecuador, napo: 10 km from Baeza to Cosanga, B3246 (MO) Colombia, cauca: Km 41 W of Uribe, B3570 (MO) Colombia, norte de santander: 7 km above Pam- plona to Bucaramanga, B3533 (MO) Colombia, cundinamarca: Cerro El Tablazo, B3539 (MO) Colombia, cundinamarca: E slopes Paramo de Choachi, B3542 (MO) Colombia, tolima: 47 km W of Fresno, B3558 (MO) Colombia, tolima: 48 km W of Fresno, B3559 (MO) Peru, amazonas: 9 km E of Molinopampa to Men- doza, B36I8 (MO) Ecuador, carchi: 55 km W of Tulcan to Maldonado, B3154 (MO) Dominican Republic, la vega: Valle Nuevo, B3707 (MO) Dominican Republic, la vega: Valle Nuevo, B3709 (MO) Peru, amazonas: 9 km E of Molinopampa to Men- doza, B3617 (MO) Bolivia, la paz: Chulumani, Albert de Escobar 1303 (TEX) Peru (Huynh, 1965) * Ecuador, carchi: El Carmelo, B3I63 (MO) Colombia, huila: Km 25 from Pitalito to Mocoa, B3592 (MO) Ecuador, imbabura: 25 km W of Laguna Cuicocha, B3171 (MO) Ecuador, pichincha: 25 km W of Chillogallo to Chi- riboga, B3237 (MO) Peru, cuzco: Km 131 of Cuzco-Paucartambo-Pilcopa- ta road, B2597 (MO) Source unknown, cultivated at Berkeley, California, 49.823 (UC) 2 Dominican Republic, la vega: El Convento, 10 km S of Constanza, B370I (MO) Dominican Republic, la vega: ca. 1 km above El Convento, B3702 (MO) Peru, cuzco: Km 132 of Cuzco-Paucartambo-Pilcopa- ta road, B3000 (MO) Venezuela, merida: El Paramito, above La Carbo- nera, B3450 (MO) Colombia, cundinamarca: below Zipacon, B3540 (MO) Colombia, cundinamarca: Km 39 of old Bogota-Fu- sagasuga road, B3548 (MO) 1982] BERRY— FUCHSIA SECT. FUCHSIA 37 Table 4. Continued. Taxon n In Collection Data or Reference F. verrucosa Hartweg F. vulcanica Andre F. wurdackii Munz 22 c. 20-22 22 22 22 c.22 11 44 44 Venezuela, tachira: 6 km E of Zumbador to Queni- quea, B34I7 (MO) Colombia, cundinamarca: Km 34-35 to old Bogota- Fusagasuga road, B3535 (MO) Venezuela, tachira: 5 km E of Las Porqueras to Pregonero, B3662 (MO) Ecuador, azuay: 13 km from Cuenca above Sayausi, B3184 (MO) Ecuador, azuay: 17 km from Cuenca above Sayausi, B3I87 (MO) Ecuador, azuay: 18 km from Cuenca above Sayausi, B3188 (MO) Ecuador, azuay: 8 km S of Cumbe, B3I90 (MO) Ecuador, napo: Papallacta, B3245 (MO) Peru, amazonas: vicinity of Leimebamba, Hutchi- son & Wright 4888 (UC) 2 1 Numbers preceded by B were collected by the author. 2 Counted by P. H. Raven from cultivated progeny. 3 A separate count of In = 44 was obtained by Ching-I Peng from seeds sent to the Missouri Botanical Garden by J. O. Wright. (see Table 4), comprising 43 species and five probable hybrids. Of these, 37 species yielded only diploid counts, five only tetraploid counts, and one species both diploid and tetraploid counts. Fuchsia vulcanica yielded four tetraploid counts and one possible diploid count. One of the putative hybrids, Berry 3584 (MO), is tetraploid, forming 22 bivalents at metaphase I, and appears to be an allotetraploid derived from diploid populations of F. corollata and F. caucana. Four other hybrids were diploid and had normal meiotic pairing of 1 1 bivalents at metaphase I, although chromosomal bridges and fragments were observed in some cells at anaphase I. Since only limited field collected material was available for the hy- brids, these results should be considered tentative and their interpretation doubt- ful. Two of the six tetraploid or partly tetraploid species in sect. Fuchsia are disjunct and confined to the island of Hispaniola. One of these, F. pringsheimii, has nectaries and flowers uncharacteristic for sect. Fuchsia', its affinities are unclear, but it may share a common ancestor with F. triphylla, which is also endemic to Hispaniola but much more similar to other species of sect. Fuchsia. Of the South American species, F. verrucosa and F. magdalenae are also con- sidered anomalous in sect. Fuchsia, because of their non-annular nectaries, as well as the extremely short floral tubes in F. verrucosa. The only species really typical of the main body of Andean species that are tetraploid are F. vulcanica and F. corollata. These are the only two of all the tetraploid species in the section that commonly have a large, though variable, proportion of triporate pollen grains. The other four species have entirely biporate grains, as do the other diploid species in the section. In the context of other polyploids that have been studied in the family Onagraceae (P. H. Raven, pers. comm.), this suggests that F. vul- canica and F. corollata might have become tetraploid recently, and that the other 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 four tetraploid species may be of more remote origin and have become stabilized. In F. corollata, both diploid and tetraploid populations are known, and the tet- raploidy doubtless originated within the species. All species in sect. Quelusia and the single species of sect. Kierschlegeria are polyploid and have entirely triporate (or rarely four-porate) grains. Sympatry and Interspecific Hybridization The evaluation of sympatry and interspecific hybridization in sect. Fuchsia is hampered by our still fragmentary knowledge of the population structure and distribution of many species. Despite many months of field studies and over 20,000 km of road travel in the Andes, I was not able to find living plants of 12 of the rarer species in the section. The broad latitudinal distribution of the section, the enormous stretches of cloud forest on the eastern slopes of the Andes that are virtually inaccessible, and the heavy rains that often cut off areas that do have roads are all factors that contribute to the difficulty of making field obser- vations. The amount of known sympatry (discussed for each species in the sys- tematic treatment) and of known or suspected interspecific hybridization (pre- sented in Fig. 17) is therefore a conservative indication of the amount that likely occurs in nature. Figure 17 shows that evidence for interspecific hybridization was found in 23, or just over a third, of the species in sect. Fuchsia, and species such as F. nigricans and F. sanctae-rosae were found to hybridize with three or four different species. Evidence for individual hybrid plants is discussed under the suspected parent species in the systematic treatment except for the example of F. nigricans in the following discussion. The main criterion used for recognizing naturally occurring hybrids was morphological intermediacy between the presumed par- ents, but decreased pollen stainability and local ecological factors such as habitat disturbance and the sympatric occurrence of the parental species were also con- sidered to be correlated with hybrid origin in many cases. Pollen viability was estimated using the double staining malachite green/acid fuchsin/orange G stain of Alexander (1969), and stainability was found to be usually 90-100% in most species of Fuchsia. In interspecific hybrids it is usually considerably lower, but some suspected hybrids were almost fully fertile, while others were plants that had totally aborted pollen grains. Normal meiotic pairing was observed in several hybrids, and the sterility observed is not thought to be related to chromosomal reorganization. In most cases, however, morphological interme- diacy and pollen stainability of less than 70% are considered strong evidence of hybrid origin. These criteria obviously limit the recognition of most hybrids to F/s, since backcross individuals are much harder to detect. Although it is usually much easier to recognize living hybrid plants than dried ones, a number of hybrids were recognized from herbarium specimens, especially between morphologically well differentiated species. Because of low pollen stain- ability and the presence of unmistakable features of two distinct species, the type specimen of Fuchsia caracasensis (Fielding & Gardner, 1844) was recognized as a hybrid between F. nigricans and F. gehrigeri, both of which are known to grow near the type locality. Fuchsia colombiana (Munz, 1946) and F. platypetala (Johnston, 1939) were also probably based on hybrid individuals. 1982] BERRY— FUCHSIA SECT. FUCHSIA 39 1 decuss • (•) Naturally occurring hybrids 2 ferrey (•) • in sect. Fuchsia 3 fontin • • • h san-ros • • • • 8 nigric • • • • • 17 putum ?•••(•)• 21 abrupt (g) 23 coroll ?J\ caucan (•) • 26 vulcan • • • 27 ampliat ..••••-••(§)• 28 venust ••••(§) * 30 gehrig ••••(§)••••••(#)• 34 dentic •••(§) • ■ 35 austro (§) • Uk sessil ••••(§) 46 tincta •••(§) 47 furfur •••<§) 52 mathew • • (§) 5^ hirtel <§) 57 canesc • •••••••(•)•••••••••••• 59 triphy • ••••••••••••••••••••• 60 prings ® • Figure 17. Naturally occurring hybrids in Fuchsia sect. Fuchsia. Most hybrids found in the field were rare and localized, and the parental populations were usually large and at least partly sympatric. Hybrids between F. triphy lla and F. pringsheimii, F. venusta and F. hirtella, and F. venusta and F. gehrigeri occur in a narrow belt where the different altitudinal limits of the species coincide (Figs. 5 and 6). Although human disturbance such as that associated with road cuts usually creates more local habitats for Fuchsia and thus increases the possibility for hybridization, hybrids of F. venusta and F. gehrigeri were also found in primary forest. A high incidence of natural hybridization is expected in sect. Fuchsia for several reasons. First, the species of sect. Fuchsia are modally outcrossing and hummingbird-pollinated. Second, there is a high degree of sympatry between species, but remarkably little ecological differentiation. For example, F. cuatre- casasii, F. scabriuscula, F. sessilifolia, and F. verrucosa were all found growing together in the same thicket south of Pitalito on the border of Depts. Huila and Cauca, Colombia, in July 1979. Finally, the thousands of hybrids artificially pro- duced in the 19th and 20th centuries attest to the ease of hybridization in the 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 5. Floral characters of Fuchsia nigricans. F. venusta, and a presumed hybrid. F. nigricans Hybrid F. venusta (Berry 3454) (Berry 3453) (Berry 3455) Inflorescence Subracemose Subracemose Subracemose to racemes to racemes 20 cm long 12 cm long Pedicel length 3-5 mm 9-18 mm 25-28 mm Ovary length 10-11 mm 9-13 mm 7-9 mm Floral tube length 19^21 mm 30-37 mm 50-60 mm Floral tube color Lavender pink Red pink Orange red Sepal length 8 mm 14-16 mm 16-18 mm Petal length 8-10 mm 15-17 mm 1&-20 mm Petal color Dark purple Medium purple Orange red Fruit length 1 £-20 mm 14-15 mm ca. 12 mm genus (see Porcher, 1858; Hemsley, 1876; and Reiter, 1941). Although most of these hybrids were from other sections in the genus, F. dominiana (van Houtte, 1854) was an early hybrid between F. macrostigma and F. denticulata. Towards the end of the 19th century, the tetraploid F. triphylla was used in many inter- specific crosses (Wright, 1978b). Numerous sympatric populations have been found with no evidence of hy- bridization, however. For example, a dense patch of F. pringsheimii was found growing along the edge of a potato field with scattered plants of F. triphylla at La Nuez, on the border of La Vega and Peravia Provinces, Dominican Republic, in December 1979. Despite the large sympatric populations and the fact that both of these species form hybrids on other parts of the island, no intermediates be- tween these two distinctive species could be found. Only F. triphylla was flow- ering at the time, and it is possible that F. pringsheimii has a different flowering period at this locality. Fuchsia tincta and F. vargasiana are apparently closely related diploid species that occur side by side and flower together in southern Peru, yet no intermediate plants were detected. The same situation occurs with F. macrophylla and F. macropetala in central Peru. It is possible that hybrids between these species do occur but are not easily recognizable or were not flow- ering at the time the populations were examined. On the other hand, there may be prezygotic isolation mechanisms such as pollinator discrimination, especially in instances in which the sympatric species differ widely in flower size, but these are factors that have yet to be examined. Despite the rarity of most hybrids, extensive hybrid populations were ob- served in two combinations. Suspected hybrids between Fuchsia corolla ta and F. caucana are numerous and widespread in southern Colombia. They occur in geographically intermediate areas between observed populations of the presumed parents, but at higher altitudes and in harsh subparamo habitats. One of the probable hybrids was tetraploid. This indicates the possibility of well established hybrid populations that have occupied a new habitat and that might have origi- nated as allotetraploids. A second case concerns probable hybrid swarms of F. ampliata and F. vulcanica in Prov. Cotopaxi, Ecuador. Very variable local popula- tions have been found in heavily disturbed roadside areas, and these outnumber the presumed parental populations. 1982] BERRY— FUCHSIA SECT. FUCHSIA 41 Table 6. Pollen stainability of putative hybrids of Fuchsia nigricans and representatives of sympatric parental populations. Collection F. nigricans (Berry 3454) Hybrid (Berry 3453) F. venusta (Berry 3455) F. nigricans (Bernardi 653) Hybrid (Bernardi 652) F. cf. venusta (Bernardi 664) Other hybrids F. nigricans x F. venusta Ricardi & Hernandez 5722 Bernardi s.n. F. gehrigeri (Berry 3128) Hybrids x F. nigricans: Berry 3129 Aristeguieta & Medina 3670 Linden 368 (OXF) F. nigricans (Foster g & Core 21550) Hybrid (Fosberg 21556) Hybrid (Fosberg 21558) F. sessilifolia (Fosberg 21560) F. nigricans x F. putumayensis: Robinson 138 i Estimated due to poor stain differentiation. Stainability 81.8% (500 grains) 34.0% (500 grains) 81.0% (500 grains) 81.6% (1000 grains) 28.5% (1000 grains) 69.0% (1000 grains) 4.0% (500 grains) less than 10% i 88.3% (600 grains) 5.3% (300 grains) 8.6% 21.3% (500 grains) (400 grains) 84.0% (500 grains) 28.6% (500 grains) 0.2% (1000 grains) 94.7% (1000 grains) 0.6% (500 grains) The relative rarity of most hybrids found indicates that unless large or nu- merous populations are carefully examined, many hybrids will be overlooked, and thus that the likelihood of finding hybrid individuals is strongly dependent on the sampling intensity. The Venezuelan Andes was the only area that was intensively studied. Four species are sympatric there, F. gehrigeri, F. nigricans, F. venusta, and F. verrucosa, and all possible hybrid combinations were found except those involving F. verrucosa. Since this is the only tetraploid species in the group and is distantly related to the others, it is probable that postzygotic barriers purpl als, canescent pubescence, and elongate, tapered ovaries. Because of these fea- tures, hybrids involving this species are relatively easy to detect. These features, together with characters such as the sympatric occurrence of the parent species, field observations in some cases, and reduced pollen stainability (Table 6), were used to compile the following list and discussion of putative hybrids with this species. Putative hybrids involving Fuchsia nigricans Fuchsia nigricans x F. venusta. Berry 3453 (MO, VEN): Merida rphologically 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 7. Comparison of morphological characters in Fuchsia nigricans, F. gehrigeri, and pre- sumed hybrids. Hybrid (Aristeguieta F. gehrigeri Hybrid & Medina F. nigricans (Berry 3128) Subglabrous to (Berry 3129) 3670) (Average Values) Stem pubescence Densely canescent Canescent Densely canescent strigillose to strigillose number of 5-7 10-12 13 12-16 secondary veins Leaves/node 3-4 2-3 3 3 Pedicel length 25-35 mm 9-12 mm 8-15 mm 3-10 mm Floral tube length 5 1-54 mm 30-33 mm 30-35 mm 14-22 mm Petal length 14-15 mm 10-12 mm 13-14 mm 5-10 mm Petal color Red Purple Dark red Dark purple Stigma exsertion 9^10 mm 2 mm 9 mm 0-2 mm above anthers Fruit shape Subglobose Cylindrical — Cylindrical Seed size (L x W) 2-2.5 x 1.2 mm 1.5-2 x 1 mm (mostly aborted) 1.5 x 0.8 mm collection are shown in Table 5 and in Figure 33, and the reduction in pollen stainability is evident in Table 6. Both parental species occurred side by side at this locality in wet roadside thickets (Fig. 4), though F. nigricans was locally more abundant. The hybrid showed strong vegetative growth and was also inter- formation Meiotic preparations from Berry nearby . (NY): Venezuela, Edo. Merida, Monte Zerp m, 21 June 1953. Both parental species were collected in the vicinity by the same same mm) Other specimens with hybrid characters similar to those described above and with low pollen fertility include Bernardi s.n. (NY; Venezuela, without locality, i 6119 (MER; Venezuela, Edo. Merida, Selva La Hernandez 5772 (MER; Venezuela, Edo. Merida 300-2.500 Fuchsia nigricans x F. gehrigeri. Medina Mesitas Aug. 1958. Berry 3129 (MO, VEN) was collected at the same site in Sept. 1978. These collections probably come from the same population, which formed a dense thicket about 5 m 2 in a sunny hollow with extensive vegetative reproduction occurring by stem shoots. Just 10 m away, several plants of F. gehrigeri were growing in shade along the forest edge. No plants of F. nigricans were seen in the vicinity, but they undoubtedly occur in the area. The presumed hybrids are clearly morphologically intermediate between the indicated species (Table 7), and their pollen stainability is in both cases less than 10% (Table 6). Buds of Berry 3129 were examined cytologically and had 11 bi- valents at metaphase I, with occasional bridges and fragments at anaphase I. Though ripe fruits were produced in Berry 3129, most of the seeds were aborted. 1982] BERRY— FUCHSIA SECT. FUCHSIA 43 Table 8. Comparison of morphological characters in Fuchsia nigricans, F. sessilifolia, and presumed hybrids. Leaves/node Leaf shape Blade length (max.) Petiole length Leaf margin number of secondary veins Floral tube color Petal color Inflorescence F. nigricans (Fosberg & Core 21550) Hybrid Hybrid {Fosberg 21556) (Fosberg 21588) F. sessilifolia (Fosberg 21560) 2-3 3-4 3-4 Broadly elliptic- Oblanceolate-ob- Elliptic obovate 9.5 cm ovate 15 cm 13 cm 1 5-20 mm 8-10 mm Subentire to den- Serrulate 6-15 mm Denticulate ticulate 10-11 17-18 13-16 Dull crimson Dark purple Green red to crimson Crimson to pur- ple Crimson Purple 4 Narrowly elliptic- lanceolate 18 cm 5-11 mm Serrate to serrulate 15-19 Green Red Spreading, race- Drooping, pani- Drooping, sub- Drooping, panicu- mose culate paniculate late Two other probable hybrids with F. gehrigeri are Benitez de Rojas 1887 ( VEN ; Venezuela, Edo. Trujillo, road from Bocono to Trujillo) (OXF; Venezuela, Edo. Merida, between Mendoza 1843). This latter collection was used as the type specimen of F. caracasensis Fielding & Gardner (Sert. PI. t. 29. 1844) but has highly inviable pollen (Table 6) and a series of intermediate characters between F. nigricans and F. gehrigeri such as long floral tubes, elongate ovaries, unequal leaf bases, and oblong leaves. The same collection number at P was used to describe F. nigricans, but this and the du- plicates at BM, G, K, LE, TCD, US, and W are clearly different, with high pollen stainability and no indication of intergradation with F. gehrigeri. Fuchsia nigricans x F. sessilifolia. Fosberg 21556 (RSA, US): Colombia, Choco, 7 km N of Carmen de Atrato, bank above trail, 2,675 m, 6 March 1944, "calyx varying from greenish red to crimson, corolla from crimson to purple." Also Fosberg 21558 (RSA, US): 8 km N of Carmen de Atrato, steep bank above purpl lifolia) but colors not intergrading." Both of these collections have highly reduced pollen stainability and characters clearly intermediate between F. nigricans and F. ses- silifolia (Tables 6 and 8). The variability and intermediate flower color noted above is also indicative of their hybrid origin. Fuchsia nigricans was collected along the same trail at 2,500 m, and F. sessilifolia grew together with Fosberg 21588. Although the flower size and shape of the two parental species are very similar, the very different leaves and flower color offer excellent indicators for analyzing presumed hybrids. The combination of such distinctive characters as the subsessile, lanceolate leaves of F. sessilifolia and the purple petals of F. nigricans leave little doubt as to the parentage of the presumed hybrids. 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 The largely triporate pollen grains found in Fosberg & Core 21550 (F. nigricans) are noteworthy. The four diploid counts of F. nigricans from Venezuela were made from plants with biporate grains. Triporate grains in Fuchsia are some- times indicators of polyploidy, so there may be tetraploid populations of this taxon in Colombia. If so, this would perhaps explain the extremely low pollen stainability of the presumed hybrids between these rather closely related species. Fuchsia nigricans x F. putumayensis. Robinson 138 (COL, K, US): Colom- bia, Dept. Valle, Rio Bravo, northwest of Darien, 4,200 ft., 28 July 1962. This plant has almost entirely inviable pollen (Table 6). The purple petals and elongate ovaries indicate genetic input from F. nigricans. The subnitid, sulcate-nerved upper leaf surface is typical of F. putumayensis, however, as are the divergent, slender petals. The rachis is too long for that species, though, and the leaves are often ternate as in F. nigricans. Both presumed parent species have been col- lected in the general area, and the low altitude indicated on Robinson's label is consistent with the parentage of F. putumayensis, which grows at lower altitudes 200-1.800 Colombia. All of the preceding evidence supports a high degree of interfertility in most species of sect. Fuchsia and indicates that natural interspecific hybridization is a common, but not ubiquitous, phenomenon between sympatric species. When hybrids do occur, they are usually rare and localized, but two examples were given that illustrate the evolutionary potential of populations of hybrid origin that have become better adapted to certain habitats than the parent species. The increased genetic recombination afforded by interspecific hybridization has prob- ably been of considerable adaptive significance in sect. Fuchsia during the late Tertiary uplift of the Andes and especially in the Pleistocene, when new and disturbed habitats were formed as a result of severe climatic fluctuations. The Evolution of Fuchsia sect. Fuchsia One of the main questions addressed by this study is how and why so many more species evolved in Fuchsia sect. Fuchsia than in other sections of the genus. Through the evidence presented in the preceding sections, it is possible to attempt answers to this question. We can begin by distinguishing the importance of extrinsic or physical factors from ones related to the plants themselves. Prior to the Neogene, the tropical Andes were probably too low for fuchsias to occur there. In the context of angiosperm evolution, the Andes are newly uplifted mountains, and they are furthermore composed of a number of distinct structural units. Starting in the Miocene, and especially during the Pliocene, they were strongly uplifted from the surrounding lowland tropical areas to give rise to a wide variety of new habitats over a very broad geographical area. Cool, montane forest habitats ("cloud forest" or "Andean forest") first appeared and increased in extension, followed by open, high elevation habitats ("puna" and "paramo"). In this way, large expanses of new, cool habitats in the tropics became available for adaptation by lowland, tropical taxa, or for colonization by previously cool-adapted organisms 1982] BERRY— FUCHSIA SECT. FUCHSIA 45 from temperate areas. The latter possibility was shown to be the case in Fuchsia, which had a probable Paleogene origin in temperate South America. The other key extrinsic factor in understanding plant evolution in the Andes is the series of strong climatic changes that occurred as a result of Pleistocene glaciations, whose magnitude in the tropics went largely unrecognized until the last two decades. As discussed in Raven (1980), drastic climatic shifts (such as those caused by the uplift of the Andes and by Pleistocene glaciations) produce new or fundamentally altered environments, thereby setting the stage for rapid speciation. This can also be described as an increase in the ecological opportunity for the establishment of new immigrants or new adaptive gene combinations (Mayr, 1963). The ways to exploit and adapt to these changes in the physical conditions of the Andes are seen in the different reproductive strategies that the organisms inhabiting them have evolved. Fuchsia is a good colonizer as a result of its self- compatibility and dispersal by birds. This is probably how the genus migrated into the tropical Andes and reached isolated islands such as Tahiti and Hispaniola. Especially when new Andean forest habitats were developing, major jumps within the tropical Andes may have occurred by long-distance dispersal. Populations established in this way may have formed the nuclei of the different species groups that we recognize today, and which are usually centered in a specific geographic area. Bird dispersal is an important factor in local migrations as well, such as those discussed previously between the different structural units of the Andes. Certain ecological adaptations were favorable to the diversification of sect. Fuchsia in the Andes. Of the two sections of the genus now present in the tropical Andes, sect. Hemsleyella became adapted to seasonal conditions and developed a basically epiphytic or lithophytic habit, whereas sect. Fuchsia is more gener- alized and remains more or less aseasonal in mesic, terrestrial habitats. Section Fuchsia has over four times as many species as sect. Hemsleyella over roughly the same geographical range. Because members of sect. Fuchsia are restricted to mesic habitats where more or less constant moisture is available, they have not radiated strongly into dif- ferent ecological zones. Nonetheless, many species have become adapted to dif- ferent altitudinal tolerances and can remain spatially isolated on the same moun- tain range. Montane cloud forests are dynamic communities where natural disturbances such as landslides, tree falls, clearings, and water courses abound. These are the main areas where sect. Fuchsia occurs, though man-made distur- bances have recently added a new dimension to this system. The particularly flexible breeding system of sect. Fuchsia is well adapted to such disturbed hab- itats; plants are loosely growing, mostly scandent shrubs that are modally out- crossing but self-compatible and have a relatively high and constant flowering rate, yet they are capable of considerable vegetative reproduction. These are adaptations that would have been particularly advantageous during the Pleisto- cene, when habitat disturbances were increased by the glacial cycles. Adaptation to hummingbird pollination has been another important factor in the evolution of sect. Fuchsia. Hummingbirds can maintain their flower-visiting activity throughout the year in cool, wet habitats whereas insects are far more 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 " -\ N •' V - * ? t <. * i \ ; ■ V CQ On ^0> 13 o CO 2 E _2 "5 e£ 55 OJ & C/3 U si O 3 O 3 0> 00 2 «5 2* O Q 05 u 4J E CO ■^ .2 r s o c §1 E 54 e u N m at 3 c/5 3 H 5 •is ov 2 ^ O o v X) P E qjd g 1982] BERRY— FUCHSIA SECT. FUCHSIA 41 Figures 20-22. Illustrations of Fuchsia sect. Fuchsia. — 20 (Upper half). Fuchsia abrupta I. M. Johnston, from Berry & Aronson 3077 (MO), Junin, Peru. Scale: floral tubes of largest open flowers are 45 mm long.— 21 (Lower left). Fuchsia nigricans Linden, from Berry 3095 (MO), Trujillo, Venezuela. Scale: floral tubes of open flowers are 20 mm long. — 22 (Lower right). Fuchsia scabriuscula Benth., from Berry & Escobar 3178 (MO), Pichincha, Ecuador. Scale: floral tubes of open flowers are 23 mm long. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 limited in their activity. Like Fuchsia, hummingbirds have diversified mainly in the tropical Andes. Although the dynamics of hummingbird pollination in Fuchsia have not been examined, it is likely that at least in some species the differences in floral tube length restrict interspecific pollination by hummingbirds. Enough cytological data is now available to state that sect. Fuchsia is chro- mosomally homogeneous, and species tend to be largely interfertile. Six species are tetraploid or partly so, but these occur mostly in outlying geographical areas or have morphological features anomalous in the section. The presence of nu- merous naturally occurring hybrids, the near normal meiotic behavior of those hybrids examined, and the existence of at least two cases of widespread popu- lations of probable hybrid origin, indicate that interspecific hybridization on the homoploid level, rather than polyploidy, has probably played an important role in the evolution of the section. It is now recognized that the lack of internal barriers to hybridization in plants, especially in perennials, is more the rule than the exception, and that the main differences between plant groups occur in the nature and strength of external mechanisms that restrict interspecific hybridization (Raven, 1980). In other groups in the Onagraceae that have radiated very recently, such as Epilobium in New Zealand (Raven & Raven, 1976) and Oenothera in South America (Dietrich, 1978), interspecific hybridization has been the key feature in the differentiation of new taxa, but the different units now recognized as species have remained distinct mostly through predominant autogamy, strong radiation into widely different hab- itats, and the unique genetic system of complex structural heterozygotes in the case of Oenothera. Fuchsia sect. Fuchsia, on the other hand, has not been able to radiate much beyond the cloud forest zone, and it is primarily outcrossing, with specialized hummingbird pollination, rather than autogamous. Other cloud forest groups such as Monochaetum (Almeda, 1978) have reproductive systems similar to Fuchsia, and these are likely to prove quite common in other groups of woody Andean angiosperms. Interspecific Relationships in Fuchsia sect. Fuchsia The lack of any major morphological, cytological, or ecological differentiation in all but a few of the 61 species in sect. Fuchsia precludes a formal division of the species into series or other subsectional taxonomic categories. The mode of evolution in sect. Fuchsia appears to have been strongly reticulate, permitting only clusters of rather closely related species to be recognized. In general, it has not been possible to discriminate between more primitive or advanced species, and this may in part be due to the recent differentiation of the section. Munz (1943) informally divided sect. Fuchsia into two lines, species with long floral tubes and those with short floral tubes. Groups were circumscribed based on inflorescence structure. The shape of petals and floral tubes was used to further subdivide the species. Because a continuous range of floral tube lengths from 3 to 130 mm exists in the species of sect. Fuchsia, it is arbitrary to divide them into two groups on this basis. Pairs of closely related species are known, such other long-tubed. lifolia 1982] BERRY— FUCHSIA SECT. FUCHSIA 49 \v ; r Figures 23-26. Illustrations of Fuchsia sect. Fuchsia. — 23 (Upper left). Fuchsia ampliata Benth., from Berry & Escobar 3234 (MO), Pichincha, Ecuador. Scale: floral tube of open flower is 45 mm long. — 24 (Upper right). Fuchsia venusta Humboldt, Bonpland & Kunth, from Berry 3292 (MO), Tachira, Venezuela. Scale: floral tubes of open flowers are 48 mm long. — 25 (Lower left). Fuchsia harlingii Munz, from Berry & Escobar 3206 (MO), Loja, Ecuador. Scale: floral tubes of open flowers are 48 mm long. — 26 (Lower right). Fuchsia caucana P. Berry, from Berry 3590 (MO), Huila, Colombia. Scale: floral tube of open flower is 40 mm long. 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 ^■^li Figures 27-30. Illustrations of Fuchsia sect. Fuchsia.— 27 (Upper left). Fuchsia ferreyrae P. Berry, from Berry & Aronson 3073 (MO), Junin, Peru. Scale: floral tubes of open flowers are 26 mm long.— 28 (Upper right). Fuchsia verrucosa Hartweg, from Berry 3286 (MO), Tachira, Venezuela. Scale: ovary is 11 mm long.— 29 (Lower left). Fuchsia triphylla L., from Berry, Smith & Mejia 3701 (MO), La Vega, Dominican Republic. Scale: floral tube of open, lowermost flower is 36 mm long.— 30 (Lower right). Fuchsia pringsheimii Urban, Berry, Smith & Mejia 3711 (MO), La Vega, Dominican Republic. Scale: floral tubes are 27 mm long. 1982] BERRY— FUCHSIA SECT. FUCHSIA 51 i ""%« Figures 31-33. Illustration and photographs of Fuchsia sect. Fuchsia. — 31 (Left). Fuchsia ceracea P. Berry, from Berry & Aronson 3081 (MO), Huanuco, Peru. Scale: sepals of the left hand flower are 29 mm long. Note the tender, concave bracts sheathing the base of the flowers. — 32 (Upper right). The hummingbird Chlorostilbon swainsonii visiting Fuchsia triphylla. Both species are endemic to Hispaniola. Photograph by Donald Dod, taken between Constanza and Valle Nuevo, Prov. La Vega, Dominican Republic. — 33 (Lower right). Flowers of two sympatric species of sect. Fuchsia and their hybrid. Upper flower is from F. venusta {Berry 3455, MO), the lower flower is from F. nigricans (Berry 3454, MO), and the middle flower (Berry 3453, MO) is from a natural hybrid between the above species. Note the intermediate size and color of the hybrid flowers. The three plants were found growing together in roadside thickets between La Carbonera and La Azulita, at 2,200 m, Edo. Merida, Venezuela, in June 1979; their habitat is illustrated in Figure 4, and further evidence of the probable hybrid origin of Berry 3453 is given in the text. 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Whereas my key to the species of sect. Fuchsia closely follows Munz's scheme, a different approach has been used here to recognize species groups. Floral po- sition, tube length, and petal shape are used here, but these characters were analyzed in more detail and were supplemented by ecological, geographical, cy- tological, and vegetative characters. Instead of a dichotomous, linear system, 14 different species groups that are intended to reflect the evolutionary relationships between the species more realistically are proposed. These species groups are first approximations and should not be regarded as formal taxonomic categories. The order of the species groups is not necessarily meant to reflect any partic- ular evolutionary sequence, though the short-tubed, axillary-flowered species may be more primitive and are placed at the beginning. Those species of doubtful affinity are included with the closest probable species group, and those of un- known affinities are placed together in a final group. Each species group is fol- lowed by a brief summary of its principal characteristics. 1) Fuchsia decussata species group: F. decussata, F. ferreyrae, F. fontinalis, and F. sanctae-rosae (Fig. 55). Short-tubed, mostly axillary flowers with narrow petals. The first three are closely related and have divaricate or predominantly horizontal branching and denticulate leaves, but F.fontinalis has flowers tending towards a paniculate inflorescence. Fuchsia sanctae-rosae has thicker, less con- stricted floral tubes, nearly entire leaves, more erect branching and is more dis- tantly related to the first three species. Peru and Bolivia. 2) Fuchsia loxensis species group: F. loxensis, F. scahriuscula, F. steyermarkii (Fig. 56). A very loosely connected group with exclusively axillary flowers, short to medium floral tubes, and rather broad petals. Fuchsia loxensis is treated as a polymorphic species whose taxonomy is not fully understood; its round petals and large leaves suggest that it may be close to the F. petiolahs species group. Fuchsia scabriuscula has no clear relative and has very few flowers and opposite, reticulate leaves, unlike the other members of this group. The linear leaves of F. steyermarkii are unique in the section, but its flowers resemble those of F. loxensis more than any other species. Colombia and Ecuador. 3) Fuchsia nigricans species group: F. nigricans, F. sylvatica, F. pallescens, F. orientalis, and F. glaherrima (Fig. 57). Characterized by mostly short-tubed, axillary to racemose flowers with narrow petals, short pedicels, and cylindrical ovaries and fruits. The first three species are closely allied and each has petals that are considerably darker than the sepals. The transitional stages of axillary to racemose inflorescences are particularly evident in these species. The last two species are closely related and have definite racemes with persistent bracts and prominent stipules, but the flowers and leaves of F. orientalis link the two subgroups together. Venezuela to northern Peru. 4) Fuchsia macrophylla species group: F. macrophylla, F. macropetala, F. ova- lis, and F. pilosa (Fig. 56). The first two species are closely related and, with F. oralis, have lateral or axillary inflorescences rather than the usual terminal or subterminal ones (Fig. 40). Though F. pilosa has a terminal inflorescence, it is 1982] BERRY— FUCHSIA SECT. FUCHSIA 53 placed here because of the close similarity of leaves, flowers, and pubescence to Peru 5) Fuchsia putumayensis species group: F. putumayensis, F. lehmannii, F. an- drei, F. cuatrecasasii, and F. abrupta (Fig. 58). Short to long floral tubes with reddish orange flowers, and delicate, usually recurved petals. Flowers mostly densely packed in short, terminal racemes with lanceolate, recurved, or decidu- ous bracts. Pedicels usually divergent. Fuchsia abrupta is long-tubed and has elongate inflorescences; its affinities are unclear, but it is placed here because of its strongly divergent pedicels, delicate, recurved petals, and mostly opposite leaves, which are found in most of the above species. Colombia to central Peru. 6) Fuchsia petiolaris species group: F. petiolaris, F. corollata, F. caucana, F. ayavacensis, F. vulcanica, and F. ampliata (Fig. 59). Long-tubed, axillary flowers with various shaped, often pubescent petals. High altitude, often subparamo habitats. A well delimited group, but individual species are similar and poorly defined. Populations of F. vulcanica and F. corollata are tetraploid. Additional cytological and population sampling is needed, especially for F. vulcanica, F. corollata, and F. petiolaris. Southernmost Venezuela to northern Peru. 7) Fuchsia venusta species group: F. venusta, F. rivularis, F. gehrigeri, F. llew- elynii, F. scherffiana, and F. confertifolia (Fig. 60). Medium- to long-tubed flowers with narrow petals, in the upper axils or in incipient racemes with mostly long pedicels. The first two species are closely related and have a common climbing habit, crispate petals, subcoriaceous elliptic leaves, and slightly hairy petals. They are probably linked to the F. petiolaris species group through F. gehrigeri. Since the last three species are known only from a few fragmentary collections, their affinities are unclear. Fuchsia confertifolia has very distinctive reduced leaves, but they are subcoriaceous as in the first two species, and the more or less crispate petals and axillary to racemose inflorescence place it closest to this group. Fuch- sia llewelynii and F. scherffiana are probably closely allied and are placed here on the basis of their subcoriaceous leaves, long pedicels, and medium-long flow- ers. Venezuela to northern Peru. 8) Fuchsia denticulata species group: F. denticulata, F. austromontana, F. har- lingii, F. cochabambana, F. macrostigma, and F. magdalenae (Fig. 61). Long- tubed, axillary flowers with thick, firm, mostly cylindric floral tubes. Pedicels stout; petals variable, but often drying purplish; anthers large (3-6 mm long). A distinctive group in its solitary, stout flowers. The first three species are closely related. Fuchsia cochabambana has funnelform flowers grouped at the tips of branches, but they are more or less thick-tubed and have purple-drying petals as in the first two species. Fuchsia macrostigma has thick, axillary flowers, but long, curved, narrowly funnelform floral tubes with large, spreading petals and a massive quadrangular stigma. Though F. magdalenae seems clearly to belong to this group because of its strong resemblance to F. denticulata, it has an entirely different type of nectary from the rest of the species in sect. Fuchsia. It is also Marta Bolivia. 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 9) Fuchsia simplicicaulis species group: F. simplicicaulis , F. ceracea, F. coria- cifolia, and F. sanmartina (Fig. 62). Very rare, viny shrubs with long to very long floral tubes and petals generally (much) shorter than the sepals. The first two species are unique in the genus with involucres of short-pedicellate flowers and thin membranous, concave, sessile bracts. The latter two species have ra- cemose inflorescences but seem to be derived from, or at least closely related to the first two. The bracts of F. coriacifolia are smaller than those of the first two species, but they are still sessile and slightly concave; the basal flowers of the inflorescence are verticillate; the upper flowers are alternate. The inflorescences of F. sanmartina have mostly alternate flowers, and the bracts are lanceolate, short-petiolate, and mostly deciduous. Central Peru. 10) Fuchsia sessilifolia species group: F. sessilifolia and F. polyantha (Fig. 63). Two closely related species, one with short, bicolored flowers and the second with long, uniformly red flowers. Leaves quaternate, lanceolate, subsessile, nitid dark green. Flowers in well-branched terminal panicles. Colombia and Ecuador. Fuchsia sessilifolia shares several important characters with the F. nigricans species group. 11) Fuchsia tincta species group: F. tincta, F. furfuracea, and F. vargasiana (Fig. 62). A closely related group with opposite, ovate leaves and pilose pubes- cence; short, few-flowered, terminal, corymbiform racemes with long pedicels. Endemic to southern Peru and Bolivia. 12) Fuchsia boliviana species group: F. boliviana, F. corymbiftora, F. wurdackii, and F. mathewsii (Figs. 63 and 65). A loosely united group with large, pubescent leaves and terminal, long racemes or few-branched panicles of long- tubed flowers. Fuchsia boliviana is distinctive in its arborescent habit, wide ecological toler- ances, reflexed sepals, and early petal dehiscence. The leaves of these species are mostly opposite or ternate, which distinguishes them from the following group. Northern Peru to northern Argentina (excluding the naturalized range of F. bo- liviana). 13) Fuchsia dependens species group: F. dependens, F. hirtella, F. hartwegii, F. crassistipula, F. canescens, and F. cinerea (Fig. 66). Pubescent, mostly quater- nate leaves, narrow petals, flowers mostly paniculate and long-tubed. The first three species are very closely related, differing mainly in tube and petiole length. Fuchsia crassistipula is also allied to these species but has unusually thick, per- sistent stipules and frequently more than four leaves per whorl. Fuchsia canes- cens has thick floral tubes and intermediate axillary to racemose flowers. Though the flowers of F. cinerea are all axillary, the leaves and individual flowers are very similar to F. dependens, and it cannot be placed in any other species group. Colombia and Ecuador. 14) Anomalous species: F. triphylla, F. pringsheimii ', and F. verrucosa (Figs. 65 and 67). All three of these species are tetraploid and cannot be placed close to any of the preceding groups. Except for its disjunct distribution, low-growing habit, and suberect inflorescences, F. triphylla has no features in which it differs substantially from the Andean species. Fuchsia pringsheimii, like F. triphylla, is 1982] BERRY— FUCHSIA SECT. FUCHSIA 55 endemic to the island of Hispaniola. It has very small leaves, broad, obconic floral tubes, and large, emarginate petals unlike any other species in sect. Fuch- sia. It also possesses a non-annular, lobed nectary that is clearly atypical of the section. Although it does not closely resemble F. triphylla, the two do hybridize on Hispaniola, and they may have evolved from a distant, common ancestor. Fuchsia verrucosa also has an atypical non-annular nectary (Figs. 46 and 47) as well as an extremely reduced floral tube. Although it occurs in Venezuela and Colombia, well within the main range of sect. Fuchsia, it shows no clear affinities to any of the species in that group, and, unlike all other members of sect. Fuchsia, it has smooth viscin threads (J. Nowicke, pers. comm.). Morphology and Anatomy habit All species in sect. Fuchsia are basically perennial shrubs or lianas. Although two species, F. pallescens and F. triphylla, can flower while still remaining es- sentially herbaceous, woody-stemmed plants are more common. Fuchsia boli- viana is usually arborescent and can reach a height of 6 m with a stem diameter of up to 5 cm. A number of species such as F. glaberrima, F. pilosa, F. tincta, F. vargasiana, and F. verrucosa, are suberect and rarely exceed a height of 2-3 m. Many others, however, can surpass this height and then become scandent or climbing. Since plants of sect. Fuchsia usually grow in moist thickets, they can brush shrubs semicultivated sites, but in suitable areas such as streamside forests, they occur as lianas clambering well into the canopy of small trees. Fuchsia denticulata is a widely distributed species that grows as erect shrubs in the drier parts of its range but as scandent or climbing plants in cloud forest. A few species, such as F. venusta and F. rivularis, are especially prone to climbing and often have long, flexible stems trailing along the ground until suitable support such as a tree or tangle is found (Fig. 3). They reach heights of 10-15 m by means of long, flexuous- pendant branches. One species, F. ceracea, is known only as a liana with long, drooping branches. A few herbarium sheets of species such as F. petiolaris and F. nigricans record their habit as epiphytic, but all plants of sect. Fuchsia that I examined in the field were rooted in the ground. STEMS Carlquist's (1975) comparative analysis of Onagraceae wood anatomy showed that species of Fuchsia have relatively long and wide vessel elements, as expected of mesomorphic species. Unlike most Onagraceae, however, Fuchsia lacks in- terxylary phloem. This character state is undoubtedly primitive in the family and is shared only by Ludwigia and Hauya, two other relatively unspecialized genera (Carlquist, 1975). Within the genus, sect. Fuchsia lacks the tuberous-thickened stems frequently found in the related sects. Ellobium and Hemsleyella. The most useful external stem characters in sect. Fuchsia are the branching pattern, cross-sectional configuration, color, and bark texture on older stems. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Branching pattern. Most species are loosely branched near the base or distal to a relatively short main stem. The branches are usually ascending when young but then become spreading or drooping when they elongate and age. Fuchsia fontinalis and F. ferreyrae commonly have horizontally spreading secondary branches. The related F. decussata is unique in the section with its well developed branching system and short, strongly divaricate tertiary branchlets. The higher order branches of F. confertifolia are very much shortened and ascending. Fuch- sia scabriuscula is often low-growing with divergent to nearly prostrate branches. Those species with a climbing habit often have a flexuous main stem and long, poorly branched secondary branches that vary from divergent to drooping or pendant. A few species such as F. glaberrima, F. pilosa, and F. sessilifolia generally have few or no secondary branches. ■ Stem cross-section. The majority of species in sect. Fuchsia have terete branchlets, but some have branchlets that are noticeably ridged, which can be a diagnostic field character. The upper stems of F. crassistipula have a series of ridges and deep furrows that correspond in number to the petioles of the whorl of leaves above it, but the ridges begin to disappear towards the basal part of the internode. Another species with conspicuously ridged stems is F. cane sc ens , and a few species such as F. venusta lack the prominent ridges of the above species, but have trigonous or angled stems. Stem color and bark texture. Light green or reddish stems are typical of purpl purpl /< ferrugineous-pilose pubescence. Young stems of F. verrucosa are somewhat tu- berculate, and F. Ilewelynii has numerous spinescent projections on the stems and pedicels. Older, woody stems have three basic bark types. The most common is the sort of flaky, exfoliating bark typical of F. decussata and thick-stemmed species such as F. hartwegii, F. gehrigeri, F. loxensis, and F. boliviana. Also widespread is the finely fissured, grayish tan bark characteristic of F. andre'u F. hirtella, and F. putumayensis. Finally, a third smooth bark that splits off in a few long, wide strips occurs in F. harlingii, F. rivularis, and F. magdalenae. PUBESCENCE The few glabrous or nearly glabrous species in sect. Fuchsia are F. ceracea, F. polyantha, F. putumayensis, and F. cuatrecasasii. In the remaining species, the pubescence can be quite variable between populations, but it is usually con- stant enough to distinguish species with fundamentally different pubescence types. Trichomes in Fuchsia are all simple and one- to few-celled (R. Keating, pers. comm.). The basic pubescence types used in the species descriptions include puberulent (minute, erect hairs), canescent or incanous (dense, fine grayish white hairs), hirsute (erect, moderately stiff hairs ca. 1 mm long), hispid (stiff, erect hairs more than 1 mm long), pilose (soft, suberect hairs less than 1 mm long), strigose (± appressed hairs 0.5-1 mm long), strigillose (± appressed hairs less than 0.5 mm long), and villous (loose, soft hairs ca. 1 mm long). Additional color 1982] BERRY— FUCHSIA SECT. FUCHSIA 57 types described above. pruinose) LEAVES The leaves of sect. Fuchsia are simple, opposite or whorled, usually petiolate, stipulate, and pinnately veined. The secondary veins depart from the midvein at a right to acute angle and generally curve apically toward the margin, either joining the superadjacent secondary to form marginal loops and an intramarginal vein (brochidodromous type of Hickey, 1973) or diminishing toward the margin without forming marginal loops with the superadjacent secondary (essentially the eucamptodromous type of Hickey, 1973). The secondary veins are commonly more or less sulcate on the dorsal surface and prominent on the ventral side. The number of secondary veins is often a diagnostic leaf character, but comparison between species should be based on similar basal leaves. A diagnostic character of F. verrucosa is the clearly impressed brochidodromous venation. Fuchsia steyermarkii and F. coriacifolia are unusual in their almost inconspicuous sec- ondary veins. The secondary veins of F. caucana have an unusually acute angle of divergence, and they remain straight until nearly the margin of the leaf. Fuchsia scabriuscula and some populations of F. sylvatica have conspicuous higher order venation that gives a rugulose-reticulate appearance to the leaf surface. Leaf anatomy studies by R. Keating (1980, in prep.) have determined the following basic leaf structure in Fuchsia: one trace, unilacunar nodes; protruding midribs with a semicircular primary vasculature; blade with a thick, single-celled cuticle, single palisade and mesophyll layers; stomata present only on the ventral surface, and raphides in vertical bundles in the palisade layer. In dried specimens the raphide bundles appear to be parallel to the leaf surface. Stipules. The stipules in sect. Fuchsia lose much of their diagnostic value once the plants are dried. The commonest stipule type is thin, narrowly lanceo- late, not connate, and early deciduous (Fig. 34). Some species, however, have connate stipules or a mixture of separate and connate ones, making them inter- petiolar and reminiscent of those of the Rubiaceae (Fig. 35). By far the most distinctive stipule type in the genus is found in F. crassistipula : the stipules on the lower leaf nodes are large, persistent, incrassate, connate, recurved, and with a thickened midvein (Fig. 36). Other species with thick, prominent, and persistent stipules are F. canescens, F. glaberrima, F. oriental is, F. ovalis, and F. pilosa. Fuchsia abrupt a usually has large, connate stipules, but they are not thickened as in the above species. Fuchsia wurdackii has unusually long (3-5 mm), pale, chartaceous, blunt-tipped stipules. Leaf margin. The margins of Fuchsia leaves typically have glandular, ' 'fuch- sioid" teeth (Hickey, 1980). These teeth may be inconspicuous, however, so that the margin appears entire in species such as F. putumayensis, F. macrophylla, and F. cuatrecasasii. Other species in the section are remotely to markedly den- or serrulate. Strongly serrulate leaves char; lifolia, and the margins of F. confer tifoli curved. leaf characters. Mature 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 ► 1cm 1 cm »■ 1 cm i Figures 34-36. Main stipule types in Fuchsia sect. Fuchsia. — 34. The most common type of separate, lanceolate stipules, F. boliviano, from living material growing at El Portachuelo, D.F., Venezuela. — 35. Connate, interpetiolar stipules, F. verrucosa, Berry 3662 (MO). — 36. Incrassate, connate stipules of F. crassistipula, Berry 3553 (MO). 1 to 27 cm. Fuchsia confertifolia has tiny, coriaceous leaves less than 15 mm long, and the leaves of F. decussata and F. pringsheimii rarely exceed 4 cm long. Species with leaves exceeding 10 cm long include F. macrostigma, F. macro- phylla, F. glaberrima, F. boliviano, and F. sessilifolia. Leaves of most species range from 3 to 10 cm long and 1.5 to 4 cm wide and are membranous. Fuchsia steyermarkii is a very distinctive species with linear leaves just 3 mm wide. A few species are readily distinguishable by their nearly sessile leaves (F. cocha- bambana, F. confertifolia, F. sessilifolia, and F. polyantha). Ovate leaves are unusual in the section and characterize the F. tincta species group as well as many plants of F. boliviana. Unusual leaf textures are found in F. coriacifolia (firm-coriaceous), F. ceracea (waxy-pruinose), and F. pallescens (thin membra- nous). A few species are commonly found that have leaves strongly purple-flushed on the under surface. The best examples of this unusual coloration are F. tri- phylla, F. tincta, and F. glaberrima. The number of leaves per node varies in many species, but it is quite constant in certain groups. Exclusively opposite leaves are found in F. corymbiflora, F. cuatrecasasii, F. putumayensis, F. tincta, and F. verrucosa. Predominantly qua- ternate leaves occur in the F. dependens and F. sessilifolia species groups. INFLORESCENCES Floral position and arrangement is one of the best diagnostic characters in sect. Fuchsia, and definite trends can be recognized in certain groups. The four basic types of floral arrangement in sect. Fuchsia are 1) axillary, 2) verticillate or involucrate, 3) racemose, and 4) paniculate (Figs. 37-41). Intermediate ar- rangements occur between all these types, especially between axillary and ra- cemose flowers. The position of the inflorescence is also important; most are terminally disposed, but the F. macrophylla species group has distinctly axillary racemes or panicles. 1982] BERRY— FUCHSIA SECT. FUCHSIA 59 Axillary flowers are considered the least advanced type, since they are the basic type found in the rest of the genus and family. Four sections of Fuchsia have flowers that are exclusively axillary (Quelusia, Encliandra, Skinnera, and Kierschlegeria), and only two small, probably advanced sections are exclusively racemose or paniculate (Jimenezia and Schufia). Section Hemsleyella is a special- ized section that has axillary and racemose flowers, these last derived from axillary flowers through the shortening of the terminal internodes (Berry, unpublished). In the three species of sect. Ellobium, the most generalized species has axillary flowers, the intermediate one has racemose flowers, and the most advanced species has lateral panicles (Breedlove et al., 1982). The strictly involucrate type of inflorescence (Fig. 38) is rare in sect. Fuchsia, found only in two species of the F. simplicicaulis species group. These species are unique in having sessile, concave bracts. The apparently derived F. coria- cifolia maintains the similar sessile (but only slightly concave) bracts; its inflo- rescence has elongated into a raceme with the basal flowers verticillate and the distal ones alternate. In F. simplicicaulis, a verticillately branched system of terminally arranged involucres sometimes occurs. Plants with axillary flowers that intergrade into terminal racemes are relatively common, especially in the F. nigricans species group. Rachis length is a diag- nostic character in species that have inflorescences. The F. putumayensis species group is characterized by very short, corymbiform racemes with narrow bracts and often divergent pedicels. Species such as F. boliviana, F. wurdackii, and F. pilosa have elongate racemes. The inflorescences of the F. dependens and F. sessilifolia species groups are almost entirely paniculate, and intermediate racemose to paniculate inflores- cences are found scattered in several species groups, as in F. oval is, F. boliviana, F. mathewsii, and F. fontinalis. FLORAL CHARACTERS Floral tube and sepals. In all species of Fuchsia there is an elongate floral tube above the inferior ovary. The tube length is measured from the top of the ovary to the rim of the tube on which the sepals, petals, and stamens are inserted. The length of the floral tube in the section varies from 3 mm (F. verrucosa) to 130 mm (F. ceracea), but the length is constant within fairly narrow limits for most species and is a valuable taxonomic character. Flowers in sect. Fuchsia are divergent to drooping, but never erect. Most species have floral tubes that are somewhat nodose or bulbous at the base around the nectary, creating a nectar reservoir inside the tube. This character varies with tube shape and there is little or no bulging at the base in more or less cylindric tubes such as are commonly found in the F. denticulata species group. On the — — 7 ^j ^ _ _- both species with marked constrictions in their lower portions. /< An important field character is the texture and thickness of the floral tube. Most fresh. The F. denticulata species group and a few other species such as F. ca- nescens and F. coriacifolia have firm-fleshy floral tubes with thick, spongy walls 1-2 mm thick when fresh. Thick tubes may well have evolved in connection with 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figures 37-41. Basic inflorescence types in Fuchsia sect. Fuchsia. — 37. Axillary flowers; pres- ent in several species groups. — 38. Involucrate flowers with sessile bracts; found only in the F. simplicicaulis species group. — 39. Terminal racemes; found in several species groups. — 40. Lateral racemes; found only in the F. macrophylla species group. — 41. Terminal panicles; typical of the F. dependens species group. 1982] BERRY— FUCHSIA SECT. FUCHSIA 61 protection against flower piercers. Tetragonous floral tubes are typical of F. ver- rucosa, and less markedly angled tubes are found in F. pringsheimii and part of the F. putumayensis species group. The sepals are valvate in bud and generally resemble the floral tube in texture and thickness. Presence and length of sepal tips in buds varies. Fuchsia pilosa, F. rivularis, and F. macros tig ma have free, spreading sepal tips in bud. At an- thesis, the angle of the sepals is suberect to divergent in most species, but both F. ampliata and F. boliviana are immediately recognizable because their sepals become fully reflexed soon after anthesis. Petals. Important petal characters are shape, size relative to the sepals, presence of pubescence, texture, surface, and angle at anthesis. Only a few species have pubescent petals, and in these cases the hairs are distributed on the outer side. Fuchsia petiolaris has the most consistently hairy petals, which vary from having a few villous hairs along the midnerve to having scattered or dense, pu- berulent hairs. Hairs can also be found on petals of some populations of F. gehrigeri, F. harlingii, F. venusta, and F. rivularis. The petal surface of F. boliviana, F. venusta, and F. rivularis is usually ridged, and the petal margin can be somewhat undulate or crispate. The petal margins of F. magdalenae are often irregular in the distal half, with a small mucronate tip. Fuchsia boliviana is the only species that has petals that regularly dehisce before the tube drops off. Unusual petal shapes include the trullate (trowel-shaped) petals in F. canescens, rhomboid petals in F. corollata, and large, obovate, emarginate petals in F. pringsheimii. Flower color. Flower color is an important field character, and species that difficult patterns have some basic red coloration, many species are characterized by orange, lav- ender, purple, or scarlet colors. Others have color gradations within the flower. Fuchsia nigricans, for example, has pinkish sepals and floral tubes but contrasting dark purple petals and filaments (Fig. 21). Several species, like F. denticulata, have green sepal tips (Fig. 19). This species usually has more or less orange petals that contrast with the waxy red or pink floral tube. Fuchsia ferreyrae has deep blue violet flowers, a color unique in the section (Fig. 27). Other distinctively (purpl lifolia purpl purpl flowers are fresh. Androecium. The eight stamens are all erect and biseriate, and the antesep- alous filaments are always longer than the antepetalous ones. In the species de- scriptions, the measurements of the antesepalous filaments are always listed first, followed by the antepetalous ones. Gvnoecium . The inferior ovary is four-locular with unspecialized axile placen- tation (Eyde & Morgan, 1973). The numerous ovules are anatropous and biseriate in each locule. Ovaries are either terete or quadrangular in transection. Style 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 pubescence can vary in certain species, so it is only of relative value. Stigmas are smooth-surfaced with a wet stigmatic surface (Raven, 1979a), and they vary in shape from clavate to capitate or obconic. The stigma always separates into four lobes, but these may be very slight as in Fuchsia denticulata, or they may consist of 4 large, prominent mounds as in F. macrostigma. Nectaries. Nectaries recently have been found to be a highly diagnostic character at the sectional level (Breedlove et ah, 1982). Section Fuchsia is char- acterized by having an annular nectary that is not found in any other section. This nectary is formed by a ringlike disc that surrounds the style at the base of the tube. The ring is mostly free from the tube or attached only at its base and can usually be dissected out of the flower intact; it is entire in some species or four-, eight-, or irregularly lobed in others (Figs. 42 and 43). All other sections have nectaries that are mostly or completely fused to the floral tube. Three species in sect. Fuchsia have anomalous nectary types, however. Fuchsia pringsheimii has an irregularly lobed nectary that is fully adnate to the floral tube (Fig. 44); its flowers are anomalous in sect. Fuchsia in other respects, and it is isolated on the island of Hispaniola. Fuchsia magdalenae seems to belong to the F. denticulata species group based on general floral characters, but its nectary consists of a smooth or slightly ridged band adnate to the tube, similar to the nectaries in sects. Hemsleyella and Ellobium (Figs. 45 and 48). Fuchsia verrucosa is yet another anomalous species in sect. Fuchsia, and its nectary is composed of four separate, antesepalous lobes adnate to the floral tube (Figs. 46 and 47). All three of these species are tetraploid, have biporate pollen, and are distinctive from other mem- bers of the section. FRUITS The berries of sect. Fuchsia vary in shape from cylindrical-fusiform to glo- bose. When ripe, they usually have a reddish coloration clearly related to dis- persal by birds. Seeds in sect. Fuchsia vary approximately twofold in size and are generally of little taxonomic value. Because they number from ca. 50 to 200 per fruit, the seeds are laterally compressed and irregularly oblong- triangular in shape, with considerable variation within the same fruit. Bright red seeds, rather than the typical tan brown ones, are sometimes found in fruits of F. caucana, F. scahriuscula, and F. sessilifolia. POLLEN Onagraceae pollen is probably the most distinctive of the angiosperms, be- cause of the viscin (exine) threads, protruding apertures, and atypical (for the dicots) structure of the exine. The unusual ultrastructure of Onagraceae has been analyzed and documented Figures 42-48. Nectary types in Fuchsia sects. Fuchsia and Hemsleyella. — 42-47. Sect. Fuch- sia. — 48. Sect. Hemsleyella. — 42-46, 48. Longitudinal sections. — 47. Cross-section through the floral tube above the nectary, viewed from above. — 42. Unlobed ring, F. denticulata, Berry 3065 (MO). 1982] BERRY— FUCHSIA SECT. FUCHSIA 63 * - 1 5mm 5mm i < 5mm »— —i h 5 mm 3mm i- 4 5 mm i 43. Four-lobed ring, F. boliviana, Berry 2592 (MO).^4. Irregularly lobed, adnate nectary of F. pringsheimii ', B*rry ?/ a/. J709 (MO).-^*5. Nectary of F. magdalenae, J. O. Wright (pickled material, MO), consisting of nectariferous tissue lining the floral tube with 4 poorly defined prolongations. Note general similarity to the nectaries of sect. Hemsleyella (see Fig. 48). — 46, 47. Separate, antesepalous , , «, -48 the base of the floral tube, F. juntas ensis, Berry 3638 (MO); this type of nectary is characteristic of sects. Hemsleyella, El labium, and Skinner a. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figures 49-51. SEM of pollen from Fuchsia sect. Fuchsia. (Provided by Joan W. No- wicke. )--«)-50. F. pilosa, Berry 3618 (MO).^49. Whole grain, equatorial view (?), x 1,550.— 50. Surface of exiqe with viscin threads, x 5,000. — 51. F. putumayensis , Berry 3562 (MO), viscin thread attachment, x 15,000. Note beaded thread type. in a series of publications by J. J. Skvarla and coworkers (Skvarla et al., 1975, 1976, 1978). The taxa illustrated in the nine plates by Skvarla et al. (1976) include 17 of the then recognized 18 genera and are representative of the family. In most dicotyledons, the exine is stratified and consists of an ektexine that includes tectum, columellae, and foot layer, and an endexine. In most Onagraceae pollen, however, the exine is distinctly bizonal in thin section, and consists of an external spongy layer and an internal solid one. The spongy layer is now considered to represent the ektexine and it may be more or less homogeneous as in almost all species of Fuchsia, or it may be coarsely granular on the proximal face, e.g., as found in Gongylocarpus (Skvarla et al., 1976: Plate 3F), or the ektexine may be differentiated into a tectum and columellae, with the latter being uniform in size and distribution, e.g., as in Camissonia arenaria (Skvarla and Nowicke, unpub- lished data). The inner solid layer is now considered to be the endexine and is remarkably similar in all taxa examined in thin section (Skvarla et al., 1976; Nowicke and Skvarla, pers. comm.). In Fuchsia (Figs. 49-54) the mature pollen is shed in monads, the grains are paraisopolar or heteropolar, and two-aperturate and bilaterally symmetrical (Figs. 49, 52, 53) or more rarely three-aperturate and radially symmetrical (Fig. 54). They are very large, and longest dimensions of 100 /xm are not uncommon. The ap- ertures are compound and protruding, the ectoaperture is porate or slightly ellip- tical (elongated horizontally), the endoaperture consists of a circular oval-shaped opening in the endexine, and the massive deposition around this opening makes the endoaperture very conspicuous in light microscopy. The sculpture of the ektex- ine surface consists of globular elements, ovoid elements, or more rarely elongated 1982) BERRY— FUCHSIA SECT. FUCHSIA 65 Figures 52-54. SEM of pollen from Fuchsia sect. Fuchsia. (Provided by Joan W. No- wicke.)— 52. F. putumayensis, Berry 3562 (MO), grain at center left illustrates the expanded distal pole or face, the prominent "ridge" visible on a number of grains reflects the deposition of the endexine, x500.— 53. F. loxensis, Berry 3185 (MO), views of grains at bottom right and center left are probably that of the distal pole but grains are partially collapsed, x500. — 54. F. vulcanic a, Berry 3190 (MO), almost all grains in this micrograph are 3-aperturate, but the sample included 2-aperturate ones also, x350. elements. The viscin threads are mostly segmented-beaded or segmented-spiral, or more rarely smooth. The vast majority of the Onagraceae, 590 of 675 species, have pollen with three apertures. However, in Fuchsia, 86 of 100 species have pollen with two apertures and in this respect the genus is unique within the family and a rarity in the dicotyledons. All species of the polyploid sections Quelusia and Kierschle- geria have three-aperturate pollen. However, the separation of two-aperturate pollen from three is by no means absolute. In sect. Fuchsia, known polyploid populations of F. vulcanica and F. corollata show varying proportions of two and three-aperturate pollen grains. In a sample of the polyploid species F. prings- heimii there were a few grains with three apertures, but the apertures were not equidistant from each other. The remaining three tetraploid species in sect. Fuch- sia, F. triphylla, F. verrucosa, and F. magdalenae, were entirely two-aperturate. In addition, a number of species known only as diploids, such as F. nigricans and F. venusta, have a small proportion of three-aperturate grains (see Brown, 1967). Although increase in aperture number often accompanies polyploidy (see above for sect. Quelusia, and Mosquin, 1966, for Epilobium), and this correlation clearly applies to sect. Fuchsia. It has been suggested for other onagraceous genera such as Clarkia that the occurrence of normal pore numbers in certain polyploid species and higher numbers in other polyploid species can be associated with older vs. more recent polyploidy (P. Raven, pers. comm.) Since a majority of the species of Fuchsia have segmented threads, including all of the generalized South American sections, Fuchsia, Hemsleyella, and Que- lusia, as well as the early disjunct sect. Skinnera, this type of thread can be 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 9. The distribution of viscin thread type, pollen pore number, and ploidy level in Fuch- sia. Gametic Chromosome Section Viscin Thread Type 1 Pore Number Number Quelusia Fuchsia Hemsleyella Ellobium Kierschlegeria Schufia Jimenezia Compressed beaded Compressed beaded Compressed beaded Compressed beaded Smooth Smooth Smooth 3 2,3 2,3 2 3 2 2 22,44 11, 22 11, 22 11 22 11 11 Encliandra Smooth, ropy, and transitional smooth to beaded 2 11 Skinnera Loosely beaded 2 11 1 From Skvarla et al., 1978; Nowicke et al., in prep assumed to be primitive in Fuchsia. Smooth threads were found in the three probably derived Mexican and Central American sections, Encliandra, Jimene- zia, and Schufia, as well as in the monotypic, anomalous sect. Kierschlegeria from coastal Chile. A general correlation found by Skvarla et al. (1978) was that segmented threads are associated with bird or moth pollination, whereas smooth, simple threads are associated with bee pollination. In Fuchsia, the sections with large hummingbird-pollinated flowers all have segmented threads, while all those with smooth threads have small flowers and may have at least some degree of insect pollination. The pollen of species in sect. Fuchsia is remarkably similar and for that reason has limited taxonomic value. A summary of the distribution of viscin thread types, pore number, and ploidy level is presented in Table 9. Systematic Treatment Taxonomic history of Fuchsia sect. Fuchsia. The earliest publication of Fuchsia was the description and illustration of "Fuchsia triphylla flore cocc^eo" by Charles Plumier in his Plantarum Americanarum Genera (1703). This species was found by the French missionary and naturalist during a stay in Haiti from 1689 to 1697 and was dedicated to Leonhart Fuchs, an important sixteenth cen- tury herbalist. The generic name Fuchsia was validated by Linnaeus (1737, 1753), using Plumier's plate as the type of Fuchsia triphylla L. The first group of South American species of sect. Fuchsia was discovered in Peru by the Spanish explorers Hipolito Ruiz and Jose Pavon between 1778 and 1788. They subsequently described six new species in the section (Ruiz & Pavon, 1802). A similar surge of collections and new species resulted from the explora- tions of Alexander von Humboldt and Aime Bonpland in the northern Andes from 1800 to 1802. Five new species were described from their collections in a later publication (Humboldt et al., 1823). As a result of the great horticultural interest that fuchsias generated in Europe in the early to mid- 1800s, a large number of new discoveries were made in South 1982] BERRY— FUCHSIA SECT. FUCHSIA 67 America. The biggest contribution came from specimens collected by Theodor Hartweg in Colombia and Ecuador in 1841 to 1843. In a series of fascicles begun in 1839 and called Plantae Hartwegianae, George Bentham described 12 new species of Fuchsia, 8 of which are currently recognized in sect. Fuchsia. Andrew Mathews made a set of valuable collections from northern Peru between 1830 and 1841, but no complete sets of his collections have been maintained. Fielding 1844) described 2 species from Mathe Macbride described 3 more species. Edouard Andre (1888) later followed much the same route in South America as Hartweg and managed to find several more novelties in sect. Fuchsia. Macbride Weberbauer Macbride them 26 new species in sect. Fuchsia. Only 11 of these names are currently recognized, however, which reflects the lack of a comprehensive understanding of the group by these authors. Regional floristic treatments of sect. Fuchsia have been made for Peru (Macbride, 1941) and for Ecuador (Munz, 1974). Philip Munz's (1943) generic revision of Fuchsia was the only attempt to examine the entire Munz 100 Unfortunately, Munz World War experience and understanding of South American geography limited the thor- oughness of his work. Munz Wurdack Cesar Vargas, and Julian Steyermark, as well as by a large number of other botanists including the author. The result of these collections has been a better geographical coverage of the areas in which Fuchsia occurs and a substantial increase in the number of specimens available, which has enabled us to redefine species limits and ranges in many taxa of the genus. Fuchsia Linnaeus, Sp. PI. 1191. 1753. type: Fuchsia triphylla L. Tilco Adanson, Fam. 2:498. 1763. type: Thilco Feuillee, J. Obs. Phys. Math. Bot. Amer. Merid. 3:64, pi. 47. 1725. "Thilco" or "Chilco" is the common name in Chile for Fuchsia magellanica Lamarck, and there is no doubt that this is what Feuillee illustrated and described. Skinnera J. R. & J. G. A. Forster, Charact. Gen. 57. 1771. type: S. excorticata J. R. & J. G. A. Forster = Fuchsia excorticata (J. R. & J. G. A. Forster) Linnaeus f. Quelusia Vandelli, Fl. Lusit. Brasil. 23, fig. 10. 1788. lectotype: Q. regia Vandelli ex Vello- zo = Fuchsia regia (Vandelli) Munz. Vandelli published the genus without naming a species. Munz (1943) designated Fuchsia magellanica as the lectotype, but that species occurs in Chile and Argentina, and Vandellfs publication was limited to Brazil. The illustration and description in Vandelli 's book are in good agreement with F. regia. Nahusia Schneevoogt, Icon. PI. Rar. 1:21. 1792. type: N. coccinea (Aiton) Schneevoogt = Fuchsia coccinea Aiton. Thilcum Molina, Saggio Chili, ed. 2:146, 286. 1810. type: T. tinctorium Molina = Fuchsia magella- nica Lamarck. Brehissonia Spach, Hist. Nat. Veg. Phan. 4:401. 1835, nom. rejic. type: B. microphylla (Humboldt, Bonpland & Kunth) Spach = Fuchsia microphylla Humboldt, Bonpland & Kunth. 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Kierschlegeria Spach, Hist. Nat. Veg. Phan. 4:403. 1835. type: K. lycioides (Andrews) Spach ■ Fuchsia ly chides Andrews. Lyciopsis Spach, Ann. Sci. Nat. Bot., ser. 2. 4:176. 1835. type: L. thymifolia (Humboldt, Bonpland & Kunth) Spach = Fuchsia thymifolia Humboldt, Bonpland & Kunth. Schufia Spach, Hist. Nat. Veg. Phan. 4:411. 1835. type: S. arborescens (Sims) Spach = Fuchsia arborescens Sims. Encliandra Zuccarini, Abh. Math. Phys. CI. Konigl. Bayer. Akad. Wiss. 2:335. 1837. type: E. par- viftora Zuccarini = Fuchsia encliandra Steudel. Kirschlegeria H. G. L. Reichenbach, Handb. 246. 1837, orth. var. = Kierschlegeria Spach. Myrinia Lilja, Fl. Sver. Suppl. 1:25. 1840, nom. rejic. type: M. microphylla (Humboldt, Bonpland & Kunth) Lilja = Fuchsia microphylla Humboldt, Bonpland & Kunth. Spachia Lilja, Tidning Tradg&dsskotsel allman Wextkultur 8:62. 1840, nom. illeg., non Spachea Jus- sieu. 1838. type: S.fulgens (DeCandolle) Lilja = Fuchsia fulgens DeCandolle. Ellobium Lilja, Linnaea 15:262. 1841, non Ellobum Blume. 1826, nom. rejic. Based on Spachia Lilja. 1840. Quilusa J. D. Hooker, J. Linn. Soc, Bot. 10:460. 1869, orth. var. = Quelusia Vandelli 1788. Erect to scandent shrubs, epiphytes, small- to medium-sized trees or pro- cumbent creepers. Stems swollen or with tuberous underground parts in some species. Leaves simple, alternate, opposite, or whorled, with small, generally deciduous stipules. Flowers hermaphroditic, gynodioecious, or dioecious; acti- nomorphic, pedicellate, axillary or in racemose, paniculate, or involucrate inflo- rescences. Floral tube cylindric to obconic, deciduous in fruit. Sepals 4, valvate. Petals 4 or 0, convolute or spreading at anthesis. Stamens 8, biseriate, the an- tesepalous ones usually longer than the antepetalous ones, all erect or the ante- petalous ones reflexed and included in the floral tube. Anthers oblong to reniform, bilocular, dorsifixed, longitudinally and introrsely dehiscent. Pollen shed singly, 2-4 porate, with beaded to smooth viscin threads. Nectary present at the base of the floral tube. Ovary 4-locular, ovules ca. 8-400, (uni-) bi- to multiseriate. Stigma usually exserted, capitate, globose, or clavate, 4-lobed or subentire. Fruit a berry; seeds 6-ca. 400 per fruit, triangular to compressed in cross-section, obovoid to ellipsoid or irregularly triangular in outline. Basic chromosome num- ber x = 11; gametic chromosome numbers n = 1 1, 22, and 44. Distribution (Figs. 1 and 2): Mostly cool, montane habitats in the Andes from Tierra del Fuego (55°S) to northern Colombia and Venezuela; in Central America and Mexico from central Panama to just north of the Tropic of Cancer (23°30'N); the coastal mountains of SE Brazil. Two species in Hispaniola, one species in central, coastal Chile; three species in New Zealand; one species in Tahiti. In the southern tip of South America and in coastal Chile, species descend to sea level. The genus consists of nine sections, as outlined with their respective numbers and geographical ranges in Table 2. The following systematic treatment covers the main Andean section, sect. Fuchsia. Fuchsia sect. Fuchsia Fuchsia sect. Quelusia (Vandelli) DeCandolle, sensu DeCandolle, Prodr. 3:36. 1828, pro parte. Fuchsia b. Fuchsia Zuccarini ex Endlicher, Gen. PI. 1193. 1840. Fuchsia sect. Fuchsia Zuccarini ex Walpers, Repert. Bot. Syst. 2:94, 1843. Fuchsia sect. Eufuchsia Baillon, Hist. PI. 6:467. 1877. Munz, Proc. Calif. Acad. Sci. IV. 25:15. 1943, pro parte. Fuchsia sect. Fuchsia; Munz, N. Am. Flora II. 5:4. 1965, pro parte. Erect, scandent, or climbing shrubs, subshrubs, or small trees. Leaves op- I982| BERRY— FUCHSIA SECT. FUCHSIA 69 posite or whorled. Flowers hermaphroditic, brightly (usually reddish) colored, divergent to pendant, axillary or arranged in racemose, paniculate, or involucrate inflorescences. Floral tubes longer than the sepals (except in F. verrucosa). Petals present and well developed, usually more than one-half as long as the sepals. Stamens erect, shorter than the sepals or slightly exserted beyond them. Nectary annular and mostly free from the floral tube, unlobed or shallowly 4-8-lobed, rarely an uneven band lining the base of the tube (in F. magdalenae) or composed of 4 or 8 prominent lobes adnate to the floral tube (in F. pringsheimii and F. verrucosa). Berry with ca. 50-250 seeds, the seeds compressed laterally and irregularly triangular-oblong in outline. Gametic chromosome numbers n = 1 1 , 22. Distribution (Figs. 1 and 2): Cloud forest of the tropical Andes from northern Argentina to Colombia and Venezuela; Hispaniola. Unless otherwise indicated, measurements used in the key and species de- scriptions are based on dried herbarium specimens. In some characters such as floral tube thickness, stipule dimensions, and petal dimensions, this may represent a substantial reduction in size compared to living material, so certain compen- sation must be allowed when using living specimens. The key is almost entirely based on floral and foliar characters; since flowering and vegetative growth in sect. Fuchsia are essentially aseasonal, specimens are almost always collected with both leaves and flowers, so that lack of plant parts used in the key is rarely a problem. Floral tube length is measured from the top of the ovary to the rim of the tube, where the stamens and petals are inserted. Measurements of the antesepalous staminal filaments always precede those of the (shorter) antepetal- ous ones, and the number of secondary leaf veins always refers to the number of each side of the midvein. The lack of major morphological differences and the overall similarity and variability in many species of sect. Fuchsia makes their separation by a simple key problematical, but a wide variety of characters are used when it is thought that the use of just one or two characters may be misleading. A number of dichotomies in the key are bridged by the variation present in certain species; in these cases, the species are keyed out under both entries. Literature citations in the synonymy of each species are limited to the major floristic or revisionary treatments of the genus and to those references in which illustrations of the species are provided. Since this paper covers a large number of taxa, some of them still poorly known, I have chosen not to recognize any taxa of subspecific rank. All the subspecific categories used by Munz (1943) were found to lie within the normal and continuous variation of their respective species. As with the other recent treatments in the family Onagraceae, considerably greater systematic knowledge of each species is required before subspecies are considered to be taxonomically meaningful units. Key to the Species of Fuchsia sect. Fuchsia la. Flowers axillary and subtended by normal or slightly reduced leaves, not tightly clustered at the branch tips, on short, lateral branches, or in definite inflorescences. 2 lb. Flowers tightly clustered at the branch tips, in definite racemes, panicles, involucres, or on short, lateral, raceme-like branches. 4 ' 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 2a. Floral tube 3-32 mm long. 3 2b. Floral tube 33-80 mm long. 13 3a. Floral tube 3-6 mm long, shorter than the ovary. (61) F. verrucosa 3b. Floral tube 10-32 mm long, longer than the ovary. 4 4a. Floral tube obconic, 9-16 mm wide at the rim; sepals 18-25 mm long; Hispaniola. (60) F. pringsheimii 4b. Floral tube cylindric to narrowly funnelform, 2-9 mm wide at the rim; sepals 6-17 mm long; South America. 5 5a. Branching well-developed and strongly divaricate, with short tertiary branchlets; leaves 15— 35(— 45) mm long, 7-15 mm wide; secondary veins 4-6 on either side of the midvein; central to southern Peru. (1) F. decussata 5b. Branching more or less lax and not strongly divaricate, short tertiary branchlets usually lacking; leaves generally more than 40 mm long, more than 15 mm wide; secondary veins 6-15 on either side of the midvein. 6 6a. Leaves opposite, very rarely ternate, the blade conspicuously rugulose; stems, leaves, and flowers with stiff, white, hispid pubescence; flowers 2-6 per branch. (6) F. scabriuscula 6b. Leaves not entirely opposite, whorls of 3 or 4 usually present. The blades not conspicuously rugulose; pubescence not hispid, or lacking; flowers generally 8 or more per branch. 7 7a. Young stems and leaves glabrous or occasionally lightly pubescent, the leaves sometimes villous on the adaxial midrib. 8 7b. Young stems and leaves canescent, strigose, or ferrugineous-pilose. 9 8a. Leaves firmly membranous, elliptic to oblanceolate, the margin entirely or obscure- ly denticulate; petiole 5-25 mm long; flowers uniformly bright orange or scarlet; Peru and Bolivia. (4) F. sanctae-rosae 8b. Leaves thinly membranous, elliptic-ovate, the margin gland-denticulate; petiole (15-)25-50 mm long; tube and sepals pale whitish-pink, the petals darker red to purple; Colombia and Ecuador. (10) F. pallescens 9a. Pedicels 3-10 mm long; ovary and berry narrowly cylindric-fusiform, the berry 13-25 mm long; young growth densely canescent. 10 9b. Pedicels (6-)8-32 mm long; ovary ellipsoid, the berry ellipsoid to subglobose, 9-18 mm long; young growth not densely canescent. 11 10a. Leaves mostly finely reticulate or rugulose; floral tube narrowly funnelform, 3.5-7 mm wide at the rim; sepals 10-13 mm long; petals red; berry tapered towards the apex, 13-17 mm long, 5-7 mm thick; Ecuador. (9) F. sylvatica 10b. Leaves not rugulose, mostly sulcate-veined; floral tube subcylindric, 3-6 mm wide at the rim; sepals 6-10 mm long; petals deep purple; berry barely tapered at the apex, 15-25 mm long, 5-10 mm thick; Colombia and Venezuela. (8) F. nigricans 11a. Leaf margin obscurely denticulate; stems canescent to finely pilose, hairs not reddish; ovary strongly tetragonous; petals oblong-elliptic to subrotund, (2— )3— 8 mm wide; Ecua- dor. (5) F. loxensis lib. Leaf margin distinctly denticulate or serrulate; stems mostly strigose-pilose, with white to reddish hairs; ovary terete or slightly angled; petals lanceolate to elliptic, 2-3 mm wide; Peru. 12 12a. Basal leaves mostly quaternate and considerably larger than the upper ones, not bluish-tinged; secondary veins 8-14 on either side of the midvein; flowers grouped towards the branch tips and subtended by somewhat reduced leaves; flowers pink to red, the floral tube 4-6 mm wide at the rim; Dept. Amazonas (Peru). (3) F . fontinalis 12b. Basal leaves mostly ternate and not much larger than the upper leaves, often bluish- tinged; secondary veins 6-10 on either side of the midvein; flowers subtended by normal leaves and not grouped at the branch tips; flowers red to blue violet, the floral tube usually 6-9 mm wide at the rim; Depts. San Martin to Junin (Peru). __ (2) F. ferreyrae 13a. Leaves sessile or subsessile, the petioles 0.5-3 mm long. 14 13b. Leaves not sessile or subsessile, the petioles more than 3 mm long. 16 14a. Leaves 10-50 mm wide, the margin denticulate or serrulate, not revolute; stems glabrous to puberulent; Bolivia. (37) F. cochabambana 14b. Leaves 2-6 mm wide, the margin subentire or revolute; stems ferrugineous-pilose; Ecuador and Peru. 15 1982] BERRY— FUCHSIA SECT. FUCHSIA 71 15a. Leaves elliptic-ovate, 8-12 mm long, 3-5 mm wide; pedicels 9-15 mm long; northern Peru. (33) F. confertifolia 15b. Leaves linear, 20-70 mm long, 2-3 mm wide; pedicels 16-20 mm long; southern Ecuador. (7) F. steyermarkii 16a. Petals broad, less than twice as long as wide, the apex rounded to broadly acute. 17 16b. Petals narrow, more than twice as long as broad, the apex narrowly to broadly acute. 28 17a. Sepals strongly reflexed soon after anthesis; petals erect; central Ecuador. _ (27) F. amp I iota 17b. Sepals suberect to divergent, not strongly reflexed after anthesis; petals spreading to divergent. 18 18a. Leaves entirely opposite, generally (narrowly) ovate; southern Peru. (48) F. vargasiana 18b. Leaves not entirely opposite, whorls of 3 or 4 usually present, mostly not ovate. „ 19 19a. Secondary leaf veins mostly 3-8 on either side of the midvein. 20 19b. Secondary leaf veins mostly 8-22 on either side of the midvein. 24 20a. Petals usually longer than the sepals, rhombic to broadly elliptic, the base attenuate or unguiculate; leaves nitid on the upper surface, usually drying glossy and more or less bullate, margin markedly glandular-serrulate; southern Colombia to central Ecuador. (23) F. corollata 20b. Petals shorter or nearly equal to the sepals, not rhombic or with an attenuate or unguiculate base; leaves generally not drying glossy and bullate on the upper surface. 21 21a. Petals elliptic-ovate, usually markedly shorter than the sepals, 5-6(-9) mm wide; ovary terete or subterete, 5-6 mm long; leaf margin generally conspicuously gland-serrulate; floral tube pink cerise to purple lavender, often more or less strongly dilated near the middle; petals usually considerably darker than the sepals; southern Colombia. (24) F. caucana 21b. Petals orbicular to broadly elliptic-(ob-)ovate, only slightly shorter than the sepals, (6— )7— 13 mm wide; ovary strongly tetragonous, 6-9 mm long; leaf margin subentire to gland-ser- rulate; floral tube orange to deep red, usually not strongly dilated near the middle; petals about the same color as the sepals. 22 22a. Leaves opposite or ternate, the margin gland-serrulate; pedicels 10-15 mm long; southern Ecuador. (36) F. harlingii 22b. Leaves 3-5-verticillate, margin subentire to denticulate; pedicels 5-55 mm long. 23 23a. Pedicels mostly somewhat tuberculate, 14-55 mm long; floral tube somewhat verrucose, 4-6 mm wide at the base, 10-12 mm wide at the rim, densely villous inside in the lower Va-Vi\ sepals thick-spongy, ca. 1.5 mm thick when fresh; petals usually drying purplish; berry 16-18 mm long, 8-11 mm thick; southern Peru to Bolivia. (35) F. austromontana 23b. Pedicels smooth, 5-25(-40)mm long; floral tube smooth, 3^4 mm wide at the base, 6-10 mm wide at the rim, pilose inside for most of the length; sepals membranous, less than 1 mm thick when fresh; petals not drying purplish; berry 11-15 mm long, 7-9 mm thick; southern Colombia and Ecuador. (26) F. vulcanica 24a. Floral tube 5-8 cm long, usually somewhat recurved; sepals 7-8 mm wide at the base; leaves mostly opposite, 6-27 cm long; Ecuador and Colombia. (38) F. macrostigma 24b. Floral tube 3.5-6 cm long, straight; sepals 3.5-7 mm wide at the base; leaves mostly in whorls of 3 or 4, 2-18 cm long 25 25a. Young stems and leaves glabrous or very lightly strigose; nectary non-annular, a smooth band 3-6 mm high lining the base of the tube; northeastern Colombia. (39) F. magdalenae 25b. Young stems and leaves markedly pubescent; nectary annular, 1-3 mm high and mostly free from the base of the tube; southern Colombia to Bolivia. 26 26a. Petals 8-11 mm long, 5.5-8 mm wide, considerably smaller than the sepals; floral tube 5-8 mm wide at the rim; stems and leaves canescent-strigose to hirtellous with mostly appressed hairs; leaves 6-18 cm long, 2.2-7 cm wide. (25) F. ayavacensis 26b. Petals 12-17 mm long, 8-13 mm wide, nearly as long as the sepals; floral tube 8-12 mm wide at the rim; stems and leaves subglabrous to densely pubescent, the hairs usually more or less erect; leaves 2.5— 8(— 10) cm long, 1-4 cm wide. 27 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 27a. Leaves mostly glabrous on the upper surface; floral tube 4-6 mm wide at the base, some- what tuberculate, the walls very firm and ca. 1.5 mm thick when fresh, the tube densely villous inside only in the lower X A-Vy, petals usually drying purplish; berry 16-18 mm long when ripe; southern Peru to Bolivia. (35) F. austromontana 27b. Leaves pubescent on the upper surface; floral tube 3-4 mm wide at the base, smooth, the walls not very firm, less than 1 mm thick when fresh, the tube pilose inside for most of its length; petals not drying purplish; berry 11-15 mm long; Ecuador. (26) F. vulcanica 28a. Leaf margin entire or more or less re volute, glandular teeth not visible without a lens. 29 28b. Leaf margin denticulate or serrate, glandular teeth clearly visible. 31 29a. Petals 9-11 mm long, smooth and not recurved; southern Ecuador. (32) F. scherffiana 29b. Petals 15-22 mm long, crispate-undulate and recurved when fresh. 30 30a. Secondary leaf veins 7-12 on either side of the midvein; leaves mostly ternate; sepal tips united in bud; Colombia and Venezuela. (28) F. venusta 30b. Secondary leaf veins 13-15 on either side of the midvein; leaves mostly quaternate; sepal tips mostly free and spreading in bud; northern Peru. (29) F. rivularis 31a. Leaves entirely opposite, generally (narrowly) ovate; northern Peru. (48) F. vargasiana 31b. Leaves not strictly opposite, whorls of 3 or 4 usually present, not ovate. 32 32a. Petals about as long or longer than the sepals. 33 32b. Petals distinctly shorter than the sepals. 38 33a. Petiole 3-9 mm long. 34 33b. Petiole 10-40 mm long. 36 34a. Secondary leaf veins 3-8 on either side of the midvein; Colombia and Ecuador (23) F. corollata 34b. Secondary leaf veins 9-15 on either side of the midvein; northern Peru. 35 35a. Leaves ternate or mostly quaternate; secondary veins 13-15 on either side of the midvein; margin entire to denticulate; stems smooth. (29) F. rivularis 35b. Leaves opposite or ternate; secondary veins 9-13 on either side of the midvein; margin dentate; stems verrucose. (31) F. Ilewelynii 36a. Floral tube slightly striated-tuberculate, hirtellous, 3.5-5.5 mm wide at the base, 1-2 mm thick and firm-spongy when fresh, dull orange; pedicels 5-13 mm long; southern Colombia. (57) F. canescens 36b. Floral tube smooth, glabrous to strigose, 2.5-4 mm wide at the base, less than 1 mm thick and not firm-spongy when fresh, bright red to pink; pedicels 12-40 mm long. 37 37a. Petals elliptic to rhombic, (5-)6-10 mm wide; leaves firm, 2-7 cm long, 0.7-3 cm wide, usually drying more or less bullate and glossy on the upper surface; petioles 3— 12(— 22) mm long; southwestern Colombia and Ecuador. (23) F. corollata 37b. Petals elliptic-lanceolate to slightly obovate, 4-8 mm wide; leaves thin, 3.5-12 cm long, 1.5-5 cm wide, dull and not bullate when dry; petioles 10—48 mm long; Venezuela and northeastern Colombia. (30) F. gehrigeri 38a. Floral tube subcylindric, (3-)4-8 mm wide at the base, the walls very firm and 1-1.5 mm thick when fresh; ovary 10-13 mm long; anthers 4-6 mm long; Peru and Bolivia. (34) F. denticulata 38b. Floral tube funnelform 2-4(-6) mm wide at the base, the walls not very firm, less than 1 mm thick when fresh; ovary 5-8 mm long; anthers 2-4 mm long; Colombia and Ecuador. 39 39a. Young branches angled, densely cinereous-pubescent; leaves mostly quaternate; floral tubes 5-6(-8) mm wide at the rim; sepals 3-4 mm wide at the base; Colombia- Ecuador border area. (58) F. cinerea 39b. Young branches terete, not densely cinereous; leaves mostly ternate; floral tubes (4— )6— 12 mm wide at the rim; sepals 5-9 mm wide at the base. 40 40a. Petals elliptic-ovate, 9-11 (-14) mm long, obtuse to cuneate at the apex, glabrous, purplish and usually darker than the sepals; sepals 5-6 mm wide at the base; southern Colombia. (24) F. caucana 40b. Petals lanceolate to lance-elliptic, (8-) 12-20 mm long, mostly acute at the apex, usually finely puberulent or villous on the dorsal surface, red and not much darker than the sepals; sepals mostly 6-9 mm wide at the base; central Colombia and Venezuela. (22) F. petiolaris 41a. Floral tubes 12-23 mm long. 42 41b. Floral tubes 24-130 mm long 58 1982] BERRY— FUCHSIA SECT. FUCHSIA 73 42a. Leaves sessile or subsessile, the petiole 0.5-5 mm long; secondary veins 18-25 on either side of the midvein. (44) F. sessilifolia Alb. Leaves not sessile or subsessile, the petiole more than 5 mm long; secondary veins 5-18 on either side of the midvein. 43 43a. Leaves entirely or mostly opposite. 44 43b. Leaves mostly 3-4-verticillate. 49 44a. Secondary leaf veins 3-8 on either side of the midvein; floral tube pale cream to pink, the petals much darker than the sepals. Ecuador and southern Colombia. (10) F. pallescens 44b. Secondary leaf veins 9 or more on either side of the midvein; floral tube red to orange, not pale or whitish, the petals not much darker than the tube and sepals. 45 45a. Stems and leaves densely pilose; leaves mostly ovate. 46 45b. Stems and leaves not densely pilose, subglabrous to puberulent or strigillose; leaves most- ly elliptic, not ovate. 47 46a. Flowers in a compact, terminal raceme 1.5-7 cm long; pedicels 18-30 mm long; sepals 7-9 mm long; southern Peru. (46) F. tincta 46b. Flowers in axillary racemes or panicles 5-10 cm long; pedicels 10-20 mm long; sepals 10-13 mm long; Central Peru. (15) F. ovalis 47a. Flowers in axillary racemes or on short side branches, the subtending leaves not strongly reduced or modified; Peru. (13) F. macrophylla 47b. Flowers in terminal racemes with strongly reduced, subsessile, linear to lanceolate bracts; Colombia and Ecuador. 48 48a. Racemes generally elongate, 5-16 cm long, the bracts persistent; pedicels 3— 7(— 12) mm long, usually not divergent; sepals 6-8 mm long; secondary leaf veins 12-17 on either side of the midvein. (11) F. orientalis 48b. Racemes generally compact, l-4(-10) cm long, the bracts mostly deciduous; ped- icels divergent, 8-25 mm long; sepals 8-11 mm long; secondary leaf veins 9-13 on either side of the midvein. (17) F. putumayensis 49a. Pedicels 2-10 mm long. 50 49b. Pedicels 10-40 mm long. 54 50a. Leaves mostly in whorls of 4; flowers generally paniculate or densely crowded on the ultimate branches; berries subglobose, 6-10 mm long. 51 50b. Leaves mostly in whorls of 3; flowers racemose, few racemes on each plant; berries oblong to cylindrical, 13-25 mm long. 52 51a. Leaves 4 or more per whorl, the margin slightly denticulate; stems canescent to hirtellous; floral tube 13-20(-24) mm long, 1.5-3 mm wide at the base; Colombia. ____ (55) F. hartwegii 51b. Leaves 4 or less per whorl, the margin markedly denticulate to serrate; stems canescent to densely pilose; floral tube 15-28 mm long, 3-4 mm wide at the base; northern Peru. (3) F. fontinalis 52a. Stems, leaves, and flowers densely pilose-hirsute; ovary 5-6 mm long; bracts nar- rowly lanceolate; northern Peru. (16) F. pilosa 52b. Stems, leaves, and flowers usually canescent or strigillose, never pilose or hirsute; ovary (6— )7— 1 1 mm long; bracts ovate to elliptic; Venezuela to Ecuador. 53 53a. Leaves usually finely reticulate or rugulose; floral tube narrowly funnelform, 3.5-7 mm wide at the rim; sepals 10-13 mm long; petals red; berries tapered towards the apex, 13-17 mm long, 5-7 mm thick; Ecuador. (9) F. sylvatica 53b. Leaves not finely reticulate or rugulose; floral tube subcylindric, 3-6 mm wide at the rim; sepals 6-10 mm long; petals dark purple; berries barely tapered at the apex, 15-25 mm long, 5-10 mm thick; Colombia and Venezuela. (8) F. nigricans 54a. Leaves and branchlets glabrous or finely puberulent, except for hairs sometimes on the adaxial leaf midvein. 55 54b. Leaves and branchlets canescent to hirtellous or densely pilose. 56 55a. Flowers red, in axillary racemes or on short, lateral branches; leaves 5-9 cm wide; sec- ondary veins 14-19 on either side of the midvein; central Peru. (13) F. macrophylla 55b. Flowers orange red, generally axillary, not in axillary racemes or on short side branches; leaves 1-4 cm wide, secondary veins 6-15 on either side of the midvein; southern Peru to Bolivia. (4) F. sanctae-rosae 56a. Leaves broadly elliptic to ovate; flowers in axillary racemes or panicles; central Peru. (15) F. ovalis 56b. Leaves lanceolate to elliptic, not ovate; flowers in terminal racemes or panicles. _. 57 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 57a. Leaves 4 or more per whorl, the margin slightly denticulate; stems canescent to hirtellous; floral tube 13-20(-24) mm long, 1.5-3 mm wide at the base; Colombia. (55) F. hartwegii 57b. Leaves 4 or less per whorl, the margin markedly denticulate to serrate; stems canescent to densely pilose; floral tube 15-28 mm long, 3-4 mm wide at the base; northern Peru. (3) F. fontinalis 58a. Leaves sessile or subsessile, petioles 0.5-3 mm long. 59 58b. Leaves not sessile or subsessile, petioles more than 3 mm long. 62 59a. Leaves 8-12 mm long, 3-5 mm wide; stems pilose; northern Peru. (33) F. confertifolia 59b. Leaves 30-120 mm long, 10-50 mm wide; stems glabrous or sparsely puberulent. 60 60a. Secondary leaf veins 4-5 on either side of the midvein; central Peru. (42) F. coriacifolia 60b. Secondary leaf veins 8-26 on either side of the midvein 61 61a. Leaves 10-24 cm long, 4-8 cm wide, the margin entire; Ecuador and northern Peru. (12) F. glaberrima 61b. Leaves 3-6 cm long, 1-5 cm wide, the margin denticulate or serrate; Bolivia. (37) F. cochabambana 62a. Leaves entirely or mostly opposite. 63 62b. Leaves mostly in whorls of 3 or more. 81 63a. Floral tube 9-13 cm long; flowers in involucres with concave, sessile bracts; central Peru. (41) F. ceracea 63b. Floral tube less than 9 cm long; flowers not involucrate with sessile, concave bracts. 64 64a. Flowers in axillary racemes or on short, lateral branches. 65 64b. Flowers in terminal racemes or panicles, not in axillary racemes or short, lateral branches. 66 65a. Floral tube 19-25 mm long; sepals 8-9 mm long, green-tipped; berry subglobose, 10-13 mm long, 8-9 mm thick. (13) F. macrophylla 65b. Floral tube 32-45 mm long; sepals 10-13 mm long, red; berry ellipsoid, 15-17 mm long, 7-10 mm thick. (14) F. macropetala 66a. Rachis elongate, 10-60 cm long. 67 66b. Rachis compact, 0.5-10 cm long. 70 67a. Secondary leaf veins 15-22 on either side of the midvein. 68 67b. Secondary leaf veins 8-14 on either side of the midvein. 69 68a. Petioles 4-10 mm long; sepals divergent after anthesis. (21) F. abrupta 68b. Petioles 20-70 mm long; sepals strongly reflexed after anthesis. (49) F. boliviano 69a. Leaves generally strigillose, 2.5— 10(— 13) cm long, 1-5 cm wide; floral tubes generally dilated near the middle and slightly constricted at the rim; petals 6-9 mm long; Hispaniola. (59) F. triphylla 69b. Leaves soft pilose, 7-18 cm long, 2-8 cm wide; floral tubes narrowly funnelform, not strongly dilated near the middle or constricted at the apex; petals 1 1-20 mm long; northern Peru. (51) F. wurdackii 70a. Secondary leaf veins mostly 3-7 on either side of the midvein; floral tube and sepals pale pink, petals purple. (10) F. pallescens 70b. Secondary leaf veins mostly 7-22 on either side of the midvein; floral tube and sepals not pale pink, petals not purple. 71 71a. Stems and leaves glabrous, puberulent, or velutinous. 72 71b. Stems and leaves coarsely pubescent (pilose, tomentose, or hispidulous). 79 72a. Floral tube 40-70 mm long. 73 72b. Floral tube 32-40 mm long. •_ 74 73a. Leaves glossy, glabrous on the upper surface; secondary veins 8-12 on either side of the midvein; stems subglabrous; Colombia. (20) F. cuatrecasasii 73b. Leaves not glossy, usually strigillose or velutinous on the upper surface; secondary veins 13-17 on either side of the midvein; stems generally puberulent; central Peru. (50) F. corymbi 'flora 74a. Racemes (4— )5— 15 cm long, suberect to nodding; leaves strigillose to subpilose; Hispaniola. (59) F. triphylla 74b. Racemes 2-5(-10) cm long, divergent to pendant, not suberect; leaves glabrous to puberulent; South America. 75 75a. Leaves oblanceolate, 10-24 cm long, 4-8 cm wide, often purple-flushed below; petioles stout, 3-8 mm long; ovary 7-9 mm long; stipules thick, persistent; Ecuador and northern Peru. (12) F. glaberrima 75b. Leaves mostly elliptic, 4-17 cm long, 2-10 cm wide, usually not purple-flushed below; 1982] BERRY— FUCHSIA SECT. FUCHSIA 75 petioles not stout, 4-30 mm long; ovary 2.5-7 mm long; stipules not thick and persistent. 76 76a. Petals 4-6 mm wide; ovary 4-7 mm long; floral tube (28-)35-40 mm long; southern Colombia. (20) F. cuatrecasasii 76b. Petals 2.5-4 mm wide; ovary 2.5-4 mm long; floral tube 23-35(-40) mm long; northern Peru to southern Colombia. 77 77a. Leaves elliptic to obovate, 2-10 cm wide, mostly opposite; sepals lanceolate, 3-5 mm wide. 78 77b. Leaves narrowly elliptic to narrowly (ob-)lanceolate, 1-4 cm wide, whorls of 3 or 4 usually present; sepals narrowly lanceolate, 2.5-4 mm wide; southern Ecuador. __ (18) F. lehmannii 78a. Branchlets finely puberulent; secondary leaf veins 9-13 on either side of the mid- vein; petioles 6-15 mm long; floral tube 15-27 mm long; southern Colombia to central Ecuador. (17) F. putumayensis 78b. Branchlets mostly strigillose; secondary leaf veins (8-) 12-15 on either side of the midvein; petioles 6-30 mm long; floral tube 23-40 mm long; southern Ecuador and northern Peru (19) F. andrei 79a. Ovary narrowly cylindric, 10-1 1 mm long; berry narrowly cylindric-fusiform, often more or less curved or with a strongly narrowed apex, 25-32 mm long; secondary leaf veins 12-18 on either side of the midvein; soMthern Peru. (48) F. vargasiana 79b. Ovary oblong, 5-10 mm long; berry oblong to globose, 12-20 mm long, not curved or with a strongly narrowed apex; secondary leaf veins 7-14 on either side of the midvein. 80 80a. Rachis 4-18 cm long; pedicels 9-1 1(-25) mm long; petals 9-16 mm long; leaf margin subdenticulate; stipules conspicuous, 3-5 mm long; northern Peru (51) F. wurdackii 80b. Rachis 1.5-5 cm long; pedicels 15-40 mm long; petals 6-10 mm long; leaf margin denticulate or serrulate; stipules 2-3 mm long; Bolivia. (47) F.furfuracea 81a. Flowers in involucres with sessile, concave bracts. 82 81b. Flowers not in involucres with sessile, concave bracts. 83 82a. Floral tube 4-5 cm long; petals 9-13 mm long. (40) F. simplicicaulis 82b. Floral tube 9-13 cm long; petals 5-7 mm long. (41) F. ceracea 83a. Floral tubes 23-30 mm long. 84 83b. Floral tubes 30-76 mm long. 87 84a. Petioles (15-)25-50 mm long, secondary leaf veins 5-7(-9) on either side of the midvein; tube and sepals pale pink-cream, the petals much darker and purple. (10) F. pallescens 84b. Petioles 4-20(-25) mm long; secondary leaf veins 7-14 on either side of the midvein; tube and sepals not pale pink-cream, the petals not purple or much darker than the sepals. 85 85a. Young stems and leaves mostly pilose; leaf margin conspicuously gland-denticulate or serrulate; floral tube widest at the rim, where 4-6 mm wide; northern Peru. (3) F. fontinalis 85b. Young stems and leaves glabrous to strigose; leaf margin entire to finely denticulate; floral tube usually widest below the rim, 6-11 mm wide at the rim. 86 86a. Flowers densely crowded on short, divergent to pendant racemes 0.5-3 cm long; petals narrowly lance-oblong, 2.5^ mm wide; southern Ecuador. __ (18) F. lehmannii 86b. Flowers generally not densely crowded, racemes suberect to nodding and 4-15 cm long; petals elliptic-ovate, 4-6 mm wide; Hispaniola. (59) F. triphylla 87a. Leaves entirely or mostly in whorls of 4 or more. 88 87b. Leaves entirely or mostly in whorls of 3. 93 88a. Petioles 12-35 mm long. 89 88b. Petioles 3-10(-15) mm long. 91 89a. Sepals 14-18 mm long, 6-9 mm wide at the base, thick-spongy and somewhat verrucose when fresh; petals trowel-shaped, 14-19 mm long, 4-7 mm wide; southern Colombia. (57) F. canescens 89b. Sepals 11-14 mm long, 3-6 mm wide at the base, not thick-spongy or verrucose when fresh; petals narrowly lanceolate to narrowly elliptic-oblong, 11-16(-18) mm long, 3-5(-6) mm wide. 90 90a. Young branches and leaves densely canescent to strigillose; leaves rarely more than 4 per whorl; stipules deciduous, not thick-callose when fresh, 1.5-2 mm long, 0.4-0.5 mm wide; southern Colombia and Ecuador. (53) F. dependens 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 90b. Young branches and leaves sparsely strigillose to strigose; leaves often more than 4 leaves per whorl; stipules thick-callose when fresh, persistent and recurved at the older nodes, 2.5-4 mm long, ca. 3 mm wide; central Colombia. (56) F. crassistipula 91a. Flowers subracemose, not in panicles; pedicels 10-53 mm long; northern Peru. (29) F. rivularis 91b. Flowers in panicles; pedicels 4-12 mm long. 92 92a. Young stems and leaves hirtellous to strigillose; central Colombia. __ (54) F. hirtella 92b. Young stems and leaves finely puberulent to subglabrous; Ecuador and southern- most Colombia. (45) F. polyantha 93a. Secondary leaf veins 15-25 on either side of the midvein. 94 93b. Secondary leaf veins 7-15 on either side of the midvein. 95 94a. Petioles 4-10 mm long; sepals not reflexed after anthesis; pedicels divergent to ascending. (21) F. abrupta 94b. Petioles 20-70 mm long; sepals strongly reflexed after anthesis; pedicels pendant. (49) F. boliviana 95a. Petals distinctly shorter than the sepals. 96 95b. Petals nearly as long or longer than the sepals. 98 96a. Floral tube 55-76 mm long; central Peru. (43) F. sanmartina 96b. Floral tube 22-40 mm long. 97 97a. Flowers tightly grouped in a raceme 0.5-3 cm long; petals 2.5-4 mm wide; southern Ecuador. (18) F. lehmannii 97b. Flowers not tightly grouped, racemes 4-15 cm long; petals 4-6 mm wide; Hispaniola. _.. (59) F. triphylla 99 100 98a. Petioles 10-48 mm long. 98b. Petioles mostly 2-10 mm long. 99a. Pedicels generally short, 9-1 1(-25) mm long; ovary cylindrical, 7-10 mm long; leaves soft pilose, 7-18 cm long; stipules pale, 3-5 mm long and persistent; northern Peru. (51) F. wurdackii 99b. Pedicels 12-40 mm long; ovary ovoid, often constricted near the apex, 5-7 mm long; leaves membranous, subglabrous to strigillose, 3.5-12 cm long; stipules dark, 1.5-2 mm long, often deciduous; Venezuela. (30) F. gehrigeri 100a. Leaves mostly in whorls of 4, secondary veins 13-15 on either side of the margin; sepal tips usually free and spreading in bud; anthers narrowly oblong, 4-4.5 mm long; northern Peru. (29) F. rivularis 100b. Leaves mostly in whorls of 3; secondary veins 7-12 on either side of the midvein; sepal tips usually connate in bud; anthers oblong, 2-3.5 mm long. 101 101a. Young stems and leaves densely pilose, often with reddish hairs; leaves narrowly elliptic to oblanceolate, often somewhat curved to one side; not glossy; northern Peru. (52) F. mathewsii 101b. Young stems and leaves subglabrous to finely hirtellous, never with reddish hairs; leaves elliptic, symmetrical, glossy on both sides; Colombia and Venezuela. (28) F. venusta 1. Fuchsia decussate Ruiz & Pavon, Fl. Peru v. Chil. 3:88, pi. 323, fig. b. 1802. Macbr., Field Mus. Nat, Hist., Bot. Ser. 13(4):552. 1941, pro parte. Munz, Proc. Calif. Acad. Sci. IV. 25:56, pi. 8, fig. 44. 1943, pro parte, type: Peru, Dept. Huanuco, abundant at Muria, 1778-1788, Hipolito Ruiz & Jose Pavon (MA, lectotype, here designated; photograph, MO). There are five Ruiz and Pavon collections of this species at MA, four at BM, and one each at F, G, MO, and NY. Since there is no collection data on any of the sheets, it is not possible to determine which sheets may be duplicates, so no isolectotypes have been designated. Fuchsia scandens K. Krause, Repert. Spec. Nov. Regni Veg. 1: 171. 1905. type: Peru, Dept. Huanuco, mountains SW of Monzon, 3,400-3,500 m, 1904-1920, August Weberbauer 3324 (B, holotype, destroyed in World War II; photograph, F). The photograph shows a poor specimen with few leaves or flowers and branches covered with lichens. Though this entity might correspond to F.ferreyrae, its leaves are closer to those of F. decussata, which the description does not exclude. 1982] BERRY— FUCHSIA SECT. FUCHSIA 11 Fuchsia fusca K. Krause, Bot. Jahrb. Syst. 37:599. 1906. type: Peru, Dept. Cuzco, Prov. La Conven- tion, below Yanamanche, near road from Cuzco to Santa Ana, 3,300-3,400 m, 1904-1920, August Weberbauer 4975 (B, holotype, destroyed in World War II; photograph, F; G, isotype). Suberect to scandent shrubs 1-3 m tall with a well-developed system of lateral, divaricate, often horizontal branches. Secondary and tertiary branches numerous, 2-4 ferrug canescent; older stems 5-14 mm thick with sublustrous, copper red bark exfo- liating in long, uneven strips. Leaves opposite or mostly ternate, firmly membra- nous, lanceolate to elliptic-obovate, acute to attenuate at the base, acute at the apex, 15-35M5) mm long, 7-15 mm wide, usually dark green and subglabrous to strigillose above with impressed veins, paler below with strigose, usually red- dish hairs along the veins; secondary veins 4-6 on either side of the mid vein, subelevated below, higher order veins not visible without magnification, margin subentire to serrulate or denticulate. Petiole 3— 10(— 14) mm long, canescent to tomentose. Stipules filiform, 2-3 mm long, ca. 0.3 mm wide, mostly deciduous. Flowers axillary and few to numerous towards the tips of branches. Pedicels slender, strigose, at times slightly verrucose, arching-pendant, 15-25 mm long. Ovary oblong, 2-3 mm long, 1 mm lone. 3-4 mm wide at the 1 0-20( 4-6 verrucose lower V2-V3. Sepals narrowly lanceolate, 8— 10(— 18) mm long, 2-3M) mm wide, narrowly acute at the apex, not much wider than the tube in bud, spreading at anthesis. Tube red to dark red, sepals red with dull green tips in bud. Petals scarlet to orange red, linear to broadly elliptic-ovate and often variable on the same plant, 5-9(-14) mm long, 1.5-4(-8) mm wide, apex acute. Nectary unlobed to obscurely 4-lobed, 1-1.5 mm high. Filaments red, 8-10 mm and 5-8 mm long; anthers oblong, 2-2.3 mm long, 1.2-1.5 mm wide, dull white. Style glabrous, red; stigma clavate to capitate, ca. 2 mm long and 1-2 mm wide, obscurely 4-cleft, white, exserted 4-8 mm beyond the anthers. Berry subglobose, subtetragonous before maturity, 8-9 mm long, 5-6 mm thick, red; seeds tan, 1.5-1.6 mm long, 0.8-1.0 mm wide. Peru. Scattered to locally frequent shrubs Martin to Cuzco Dents., 2,900-3,400 m (Fig. 55). Representative specimens examined: Peru, ayacucho: 19 km E of Tambo, Berry 3049 (MO, USM); 40-46 km NNE of Tambo, Luteyn & Lebrdn-Luteyn 6352 (MO, NY); near Cusimachay, 25 km NE of Tambo, Madison 10374-70 (MO); Sigsiqa, between Tambo and Ayna, Tovar 6148 (USM). cuzco: Km 157 of Cuzco-Quillabamba road, Berry 2574 (CUZ, MO); Km 158 of Cuzco-Quillabamba road, Berry & Aronson 3041 (MO, USM); between Yanamanche and Quellomayo, Vargas 4512 (BH, CUZ, MO), huanuco: between Acomayo and Chinchao, Ambrose et al. 2397 (BH); 3 km E of Carpish tunnel, Berry & Aronson 3086 (MO, USM); Carpish, Ferreyra 2093 (F, MO, NY, US, USM); Carpish, above Acomayo, Hutchison & Wright 5944 (F, G, LE, MICH, MO, NY, P, RSA, UC, US, USM); Muna, Pearce 513 (K); Tumanga, Woytkowski 7984 (US), junin: 64 km above Satipo to Conception, Berry & Aronson 3075 (MO, USM). san martin: valley of Rio Apisoncho, ca. 30 km above Jucus- bamba, Hamilton & Holligan 552 (K, S, UC), 1057 (K), 1093 (K, UC). Fuchsia decussata is distinguished from other members of its species group 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 55. Distribution of the Fuchsia decussata species group. by the small leaves with few secondary veins and the strongly divaricate branch- ing pattern with short tertiary branchlets. It is closely allied to F.ferreyrae, and some intergradation between them may occur (see discussion under F.ferreyrae). It occurs sympatrically with F. abrupta, F. corymbiflora, F. denticulata, F.fer- reyrae, and F. sanmartina. Pubescence and petal size and shape vary considerably throughout the range of this species. Plants from the Carpish area in Huanuco have very narrow petals nearly as long as the sepals and a soft, reddish tomentum on the branches. Plants from Cuzco, such as the one described as F.fusca, have stiffer, more erect hairs, somewhat broader floral tubes, and wider petals. Extreme petal variation is found locally or in individual plants in Dept. Ayacucho. Luteyn & Lebron-Luteyn 6352 (MO, NY) has petals 14 mm long and 8 mm wide, while the petals of Berry 3149 1982] BERRY— FUCHSIA SECT. FUCHSIA 79 (MO, USM) from the same area vary widely, at times approaching the narrowly lanceolate petals of the Huanuco populations. Sandeman 4585 (K, OXF; Dept. Junin, above Huacapistana, ca. 2,000 m, Oct. 1943) is a possible hybrid with F. abrupta. The flowers are subracemose with narrowly cylindric ovaries as in F. abrupta, but the short tubes (ca. 25 mm long) and branching system indicate parentage of F. decussata. Both species are known from the area, and less than 5% of the pollen of the presumed hybrid is stainable. Another collection from the same locality, Sandeman 4503 (K) is more similar to F. abrupta, except that it has short floral tubes less than 25 mm long. 2. Fuchsia ferreyrae P. Berry, sp- nov. type: Peru, Dept. Junin, Prov. Satipo, 68 W '073 (MO 2720583, holotype; MO, USM, Ferreyra, Director of the Museo de Histori Peru, and long-time collector of the Peruv Fuchsia decussata sensu Munz, Proc. Calif. Acad. Sci. IV. 25:56. 1943, pro parte. Frutex erectus vel scandens 1-3 m altus, ramis arcuato-patentibus 0.5-1.5 m longis. Ramuli subcanescentes vel dense strigosi pilis interdum rubris instructi. Folia 3-4-verticillata vel raro opp- osita, membranacea, reticulata, elliptica vel obovata, basi acuta vel acuminata apice acuta, 2-8 cm longa, 1-3 cm lata, superne atroviridia subtiliter strigulosaque, subtus nervis praesertim strigosis et pagina saepe cyanescenti; nervis secundariis utroque latere 6-10, margine obscure vel valde gla- nduloso-serrulata; petiolis 4-12 mm longis; stipulis filiformibus, 2-4 mm longis, deciduis. Flores atro- rubri vel atroviolacei, axillares et numerosi ad apicem ramorum dispositi; pedicellis tenuibus depe- ndentibusque, 12-30 mm longis. Tubi florales anguste infundibuliformes, ( 15— )22— 28 mm longi, basi 3-4 mm lati et conspicue nodosi inde ca. 2 mm lati valde constricti, superne dilatati summo (6-)8-9 mm lati, extus strigulosi vel pilosi intus glabrati. Sepala anguste lanceolata, acuminata, (10-) 14-16 mm longa, ca. 3 mm lata, alabastro saepe apicibus libris. Petala rubra vel violacea, elliptica, acuta, plerumque plus minusve dimidio breviora quam sepalis, 7-9 mm longa, ca. 3 mm lata. Filamenta antisepala 9-13 mm longa, antipetala 5-9 mm longa, antheris oblongis, ca. 2 mm longis ca. 1.5 mm latis. Stylus glaber, stigmate capitato, apice leviter 4-fisso, ca. 2 mm longo latoque, obscure albido. Bacca ellipsoidea vel subglobosa, ca. 12 mm longa, ca. 8 mm lata, nitens purpureaque; seminibus 1-1.2 mm longis, ca. 0.6 mm latis. Numerus gameticus chromosomatum n = 11. Erect to climbing-scandent shrubs 1-3 m tall with horizontally divergent to arcuate branches 0.5-1.5 m long. Branchlets subcanescent to coarsely strigose with reddish hairs; older branches with finely fissured, dull tan to coppery bark. Leaves in whorls of 3 or 4, occasionally opposite or subopposite, membranous, elliptic to obovate, acute to attenuate at the base, acute at the apex, 2-8 cm long, 1-3 cm wide, velvety dark green and strigillose above, strigose below especially along veins and often flushed with a bluish tinge; secondary veins 6-10 on either side of the midvein, higher order veins reticulate and usually visible without errulate 4-12 mm lone. Stioules filiform, 2-4 Flowers axillary and numerous towards the branch tips. Pedicels slender, pen- dant, 12-30 mm long. Ovary oblong, ca. 3 mm long. Floral tube narrowly fun- 3-4 base, then ± abruptly (6-)8-9 ^ly lanceolate, (10-) 14-16 mm long, ca. 3 mm wide, acuminate, often with free tips in bud, divergent at anthesis. Tube and sepals dark red to deep violet (blue red). Petals red to violet, elliptic, usually about half 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 as long as the sepals, 7-9 mm long, ca. 3 mm wide, acute at the apex. Nectary unlobed, ca. 1.5 mm high. Filaments red to violet, 9-13 mm and 5-9 mm long; anthers oblong, ca. 2 mm long, ca. 1.5 mm wide. Style glabrous, red to violet; stigma capitate, ca. 2 mm long and wide, slightly 4-lobed, dull white. Berry ellipsoid to subglobose, ca. 12 mm long, ca. 8 mm thick, lustrous purple; seeds 1-1.2 mm long, ca. 0.6 mm wide. Gametic chromosome number /i = 11. Distribution: Central Peru. Locally frequent shrubs in openings of cloud forest from San Martin to Junfn Depts., 2,600-3,150 m (Fig. 55). Specimens examined: Peru, Huanuco: Panao, Macbride 3615 (G, GH, F, K, S, US); Tambo de Vaca, Macbride 4414 (F, GH); ca. 22 km SE of Huanuco, Macbride & Featherstone 2081 (F, G, GH, US), 2107 (F, GH, K, US), 2124 (F, GH); Pillao, Woytkowski et al. 34163 (F, MO, UC, USM). junin: 68 km W of Satipo to Conception, Berry & Aronson 3074 (MO), san martin: valley of Rio Apisoncho, ca. 30 km above Jucusbamba, Hamilton & Holligan 1428 (K, S, UC). without locality: McLean s.n. (K). This species is closely related to Fuchsia decussata but has more nodose, funnelform floral tubes, generally shorter petals, longer and more reticulate leaves, and less divaricate branching. Some plants, such as the type collection, have a very striking blue-violet coloration on the leaves and flowers (Fig. 27). This species lacks the green sepal tips of F. decussata. It occurs sympatrically with F. abrup- ta y F. decussata, F. denticulata, and F. sanmartina. Around the type locality in Dept. Junfn, F. ferreyrae is found between 2,650 and 3,100 m. At ca. 2,900 m, it is sympatric with F. decussata. Although these two species occur side by side, no intermediate plants could be found. In this area, the two species differ markedly in appearance. Fuchsia decussata has flaky, coppery bark and a tight, divaricate branching system with slender, green-tipped flowers. Plants of F. ferreyrae are much more floriferous, with broader, more nodose, and totally violet flowers and laxer, simpler branching. In many herbar- ium specimens, however, it is difficult to distinguish the two species because coloration and leaf texture characters are usually lost upon drying. Several of the specimens cited under F. ferreyrae are intermediate with F. decussata in certain characters, but are included here because of their noticeably nodose and funnel- form floral tubes, larger leaves, or indication of bluish coloration in the foliage. Additional field work is needed in Dept. Huanuco to determine if these species intergrade or if they are always clearly distinct under field conditions, as found at the type locality. 3. Fuchsia fontinalis J. F. Macbr., Candollea 8:25. 1940. type: Peru, Dept. Ama- zonas, Chachapoyas, 1830-1841, Andrew Mathews (G-DEL, lectotype, here designated; photograph, MO). Fuchsia decussata sensu Munz, Proc. Calif. Acad. Sci. IV. 25:56. 1943, pro parte. Erect to scandent shrubs 0.5^4 m tall with spreading branches. Young growth canescent to densely pilose, often becoming ferrugineous with age; branchlets pilulose to tomentose; older branches 5-12 mm thick with tan, finely fissured bark. Leaves whorled, mostly ternate or quaternate, occasionally opposite, mem- branous, elliptic to (ob-)lanceolate, acute to attenuate at the base, acute at the apex, 35-110 mm long, 10-35 mm wide, strigillose above and along the veins below, basal leaves noticeably larger than the upper ones; undersurface visibly 1982) BERRY— FUCHSIA SECT. FUCHSIA 81 reticulate-veined, secondary veins 8-14 on either side of the midvein; margin conspicuously denticulate or serrate. Petioles pubescent, 5-21 mm long, mostly reddish. Stipules filiform, 2-3 mm long, ca. 0.4 mm wide, deciduous to semiper- sistent. Flowers numerous and generally densely grouped in upper leaf axils or in racemes or many branched terminal panicles; rachis 3-14 cm long with reduced leaves 15-25 mm long subtending the flowers. Pedicels slender, pendant, pubes- cent, 6— 16(— 25) mm long. Floral tube subcylindric to narrowly funnelform, 15-28 mm long, 3-4 mm wide and somewhat bulbous at the base, then constricted to ca. 2 mm wide above the nectary and widened gradually above until 4-6 mm wide at the rim, pilose to strigillose outside, glabrous inside. Sepals lanceolate, 10-13 mm long, ca. 3 mm wide, acuminate, spreading-divergent at anthesis. Tube and sepals nitid red or pink. Petals red, usually noticeably shorter than the sepals, lanceolate, 6-9 mm long, 2-3 mm wide, acute, spreading at anthesis. Nectary 4-8-lobed, ca. 1.5 mm high. Filaments pink to red, 9-14 mm and 6-10 mm long; anthers oblong, 2-2.5 mm long, 1-1.5 mm wide, dull white. Style light red, gla- brous; stigma capitate, slightly 4-lobed, 1.5-2 mm long and wide, exserted 2-4 mm beyond the anthers. Berry subglobose, 9-10 mm long, 5-9 mm thick, red; seeds tan, 1-1 .2 mm long, ca. 0.7 mm wide. Gametic chromosome number n = 1 1 . Distribution: Northern Peru. Shrubs in thickets, banks, and streamsides; en- demic to Dept. Amazonas, in upper cloud forest on the E side of the Jalca de Calla-Calla and in the mountains to the east of the Rio Utcubamba, 2,900-3,400 m (Fig. 55). Representative specimens examined: Peru, amazonas: Cerros de Calla-Calla, 18 km SW of Lei- mebamba (Km 403), Ambrose 2375 (BH); E side of Calla-Calla, 62 km NE of Balsas, Berry & Escobar 3608 (MO, USM); Cerro Puma-Urco, 10 km SE of Chachapoyas, Berry & Escobar 3624 (MO, USM); Leimebamba-La Joya trail, Boeke 1783 (MO); Leimebamba-Lajasbamba trail, Boeke 2031 (MO); Cerros Calla-Calla, E side, 18 km above Leimebamba, Hutchison & Wright 6956 (F, NY, RSA, UC, US, USM); Almirante, Mathews 1479 (K, OXF); Chachapoyas, Mathews s.n. (G-BOIS); Cerro Tinaja, Ochoa 1653 (F, US); mountains S of Tambo de Ventilla, Pennell 15794 (PH); Almirante, S and e man 59 (F, OXF); upper slopes of Puma Urco, ESE of Chachapoyas, Wurdack 670 (F, NY, RSA, US, USM); S side of Molinopampa-Diosan pass, Wurdack 1626 (RSA, US, USM). This species is allied to F. decussata and F. ferreyrae. Its flowers are more numerous, however, and are often grouped into definite inflorescences. The leaves are mostly quaternate, unlike the other two species, and the basal leaves are usually considerably larger than the upper leaves. At its lower altitudinal limit on the eastern slopes of the Jalca de Calla-Calla, F.fontinalis is sympatric and forms local hybrids with F. mathewsii along pasture borders and stone walls (Fig. 10 and discussion under F. mathewsii). Macbr., Field M M fig. 48. 1943. type: Bolivia, Santa Rosa (Department unknown), 1,600 m, 13-20 April 1892, Otto Kuntze (NY, holotype). This specimen is in very poor shape, and no intact flowers remain, but the description and locality could only apply to this species. Fuchsia boliviana Britton, Bull. Torrey Bot. Club 17:214. 1890, horn, illeg., non Carr., 1876. type: Bolivia, Dept. La Paz, Yungas, 1885, Henry H. Rusby 1813 (NY, holotype; MICH, NY, US, isotypes). 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia weberbaueri K. Krause, Repert. Spec. Nov. Regni Veg. 1:179. 1905. type: Peru, Dept. Puno, Prov. Sandia, 2,400 m, 1906, August Weberbauer 661 (B, holotype, destroyed in World War II; photograph, F; G, isotype). Fuchsia brittonii Johnston, Contr. Gray Herb. 75:39. 1925, nom. nov. pro F. boliviano Britton, horn. illeg. Fuchsia filipes Rusby, Mem. New York Bot. Gar. 7:317. 1927. type: Bolivia, Dept. La Paz, Pulcheri, ca. 3,000 m, 15 July 1921, OrlandE. White 232 (NY, holotype; GH, K, US, isotypes; photographs ofGH isotype, NY, RSA). Erect to scandent, well branched shrubs 1-3 m tall. Branchlets usually subgla- brous, occasionally pubescent, purplish red, 2-4 mm thick; older branches 5-18 mm thick with light tan, finely fissured bark. Leaves usually ternate or quaternate, the basal ones much larger than the upper ones, membranous, oblanceolate to (narrowly) elliptic, acute to attenuate at the base, acute to acuminate at the apex, 2-14 cm long, 1^4 cm wide, dark green and glabrous above, paler below (espe- cially when dry) and glabrous except often with pilose hairs along the midrib; secondary veins 6-15 on either side of the mid vein, subelevated below; margin subentire to obscurely denticulate. Petiole subglabrous to pubescent, 5-25 mm long, green to red-purple. Stipules lance-linear to narrowly triangular, occasion- ally connate, 2-4 mm long, 0.5-1.0 mm wide, deciduous. Flowers usually nu- merous, solitary in upper leaf axils or often appearing loosely racemose or pa- niculate. Pedicels slender, glabrous, 10-30(-40) mm long, spreading-divergent to drooping. Ovary cylindric, 4-5 mm long, 1-2.5 mm thick. Floral tube subcylindric to narrowly funnelform, 15— 22(— 27) mm long, 2-3 mm wide at the base, slightly constricted above the nectary and gradually widened above until 4-6 mm wide at the rim, glabrous to sparsely pilose outside, densely pilose inside in lower Vs. Sepals lance-oblong, acute, 9-13 mm long, 2.5-4 mm wide, rather obtuse-tipped in bud and broader by 1-2 mm than the rim of the tube, spreading-divergent at anthesis. Tube and sepals nitid orange red. Petals orange red, oblong, obtuse to acute, 7-9 mm long, 2-4 mm wide, spreading. Nectary unlobed, ca. 1.5 mm high. Filaments orange red, 6-8 mm and 4-6 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide, cream. Style orange red, glabrous; stigma orange red, capitate, slightly 4-lobed, ca. 1.5 mm long and wide, exserted 3^4 mm beyond the anthers. Berry ellipsoid to subglobose, 8-12 mm long, 6-10 mm thick, dark red purple; seeds 1.3-1.6 mm long and 0.8-1.0 mm wide. Gametic chromosome number n = 11. Distribution: Southern Peru and Bolivia. Scattered to locally frequent shrubs in cloud forest from Dept. Cuzco in Peru to Dept. Cochabamba in Bolivia; 1,400-3,000 m (Fig. 55). Representative specimens examined: Peru, cuzco: Machupichu, Balls B6802 (BM, F, K, NA, UC, US); ca. 13 km below Marcapata to Quince Mil, Berry & Aronson 3019 (MO); Km 170 of Cuzco-Quillabamba road, Berry & Aronson 3046 (MO); Km 127-128 of Cuzco-Pilcopata road, Berry et al. 2596 (MO, USM); Km 160 of Cuzco-Paucartambo-Pilcopata road, Berry et al. 3002 (MO, USM); San Miguel, Urubamba valley, Cook & Gilbert 1110 (GH, US); Hacienda Santa Rita, Uru- bamba valley, Dreyfus s.n. (USM); above Machupichu on old Inca trail, Gentry et al. 19384 (MO); Amaybamba, Prov. Convention, Infantes 6033 (P); Machupichu, Mexia 8078 (F, G, GH, K, MO, UC); Pillahuata, Cerro de Cusilluyoc, Pennell 13997 (F, GH, NY, PH, US); between Calca and Hipal, Quebrada de Yanatili, Raimondi s.n. (USM); Aguas Calientes, Urubamba valley, Solomon 3118 (MO); between Pillahuata and Tambomayo, Vargas 82 (BH, CAS, CUZ, F, MO); Huaillai, Marcapata, Vargas 1343 (CUZ); Km 75-76 road to Lares, Vargas 15322 (CUZ); Manfo, Km 84, Vargas 15627 (CUZ); valley of Rio Tambomayo, West 7091 (GH, MO, UC). puno: Santo Domingo 1982] BERRY— FUCHSIA SECT. FUCHSIA 83 area, Prov. Sandia, McCarroll 14 (MICH, NY, S); 2-6 km from Oconeque, Prov. Sandia, Metcalf 30557 (A, BH, G, MO, UC, US); Ollachea, Prov. Carabaya, Vargas 6869 (CUZ, RSA); vicinity of Sandia, Vargas 15151 (CUZ); San Juan de Oro, Prov. Sandia, Vargas 20604 (CUZ); Agualani-Oco- neque, Prov. Sandia, Vargas 20635 (CUZ). Bolivia, cochabamba: Montepunco, 130 km E of Cocha- bamba, Adolfo 314 (DS); Incachaca, Brooke 6693 (F, NY), 6653 (NY); road to Yungas de Tablas, Cardenas 6259 (NY, US), la paz: Chulumani, Albert de Escobar 1303 (TEX); El Chaco, Sur Yungas, Asplund 1153 (UPS); Unduavi, Brooke 6610 (BM, F, NY, U); Hacienda Simaco, road to Tipuani, Buchtien 832 (BM, F, G, K, MO, POM), 5508 (GH, NY, S, US, Z); between Unduavi and Chirca, Eyerdam 25388 (F, UC); from Puente Villa to Chulumani, Kelley 1014 (UC); near Ananca, Macha- camarca, Prov. Larecaja, Mandon 624 (BM, F, G, GH, LE, NY, RSA, S, W): Rio Pelechuco, Williams 2494 (BM, NY); Nequejahuira, Cordillera Real, Tate 675 (NY); Okara, Cordillera Real, Tate 912 (NY). This species can be distinguished from other axillary, short-flowered species by the nearly entire leaves, usually glabrous stems and leaves (except for the pilose midvein below), and the uniformly orange red, nitid flowers. In addition, the stigma is reddish, and the basal leaves are considerably larger than the upper ones. It occurs sympatrically with F. cochabambana, F. boliviana, F. denticu- lata, F. furfuracea, F. tincta, and F. vargasiana. Fuchsia sanctae-rosae has uncommonly wide ecological and altitudinal tol- erances. It is found in open, rocky situations such as the ruins of Machupichu or in shady cloud forest thickets. In Dept. Puno, Peru, it occurs on the edges of the town of Ollachea in ditches at 2,700 m; in the neighboring Dept. Cuzco, it occurs as low as 1,400 m, where subtropical floristic elements are already present (Fig. 11). Beck 1266 (MO; Bolivia, Dept. La Paz, Prov. Murillo, Valley of Zongo at 2,170 m, 10 km from Cahua towards La Paz) is a probable hybrid with F. den- ticulata. This specimen has the stout stems and firm, elliptic leaves of F. den- ticulata, but the flowers are characteristic of F. sanctae-rosae, except for the intermediate tube length. The pollen stainability is 63% of 500 grains. Probable hybrids with F. furfuracea and F. tincta are discussed under those respective species. 5. Fuchsia loxensis Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6:106, t. 536, figs. /-5. 1823. Munz, Proc. Calif. Acad. Sci. IV. 25:28, pi. 2, fig. 14. 1943; Opera Bot., Ser. B, 3:18. 1974. type: Ecuador, Prov. Loja, near Loja, ca. 2,000 m, July-Aug. 1802, Alexander von Humboldt & Aime Bonpland (P HBK. Herb., holotype, not seen; photograph, F; microfiche, MO). Fuchsia umbrosa Bentham, PI. Hartw. 176. 1845. type: Ecuador, Prov. Pichincha, Hacienda de Pinantura, near Quito, 1841-1843, Theodor Hartw eg 983 (K Bentham Herb., holotype; BM, BR, CGE 2 sheets, G, K Hooker Herb., LE, OXF, W, isotypes). Fuchsia apiculata I. M. Johnston, Contr. Gray Herb. 75:34. 1925. type: Ecuador, Prov. Azuay and Cariar, between Cuenca and Huigra, 2,700-3,300 m, 12-13 Sept. 1923, Albert S. Hitchcock 21667 (GH, holotype; photograph, UC; NY, US, isotypes). Fuchsia hvpoleuca I. M. Johnston, Contr. Gray Herb. 75:34. 1925. type: Ecuador, Prov. Loja, between Loja'and San Lucas, 2,100-2,600 m, 6 Sept. 1923, Albert S. Hitchcock 21440 (GH, holotype; photograph, UC; NY, US, isotypes). Munz, Proc. Calif. Acad. Sci. IV. 25:57, pi. 8, fig. 45. 1943; Opera Bot., Ser. B, 3:17. 1974. Scandent to mostly erect shrubs or small trees ( 1-) 1 .5-6 m tall, usually densely branched. Young growth canescent to finely pilose; older stems 8-25(-40) mm thick, with flaky, tan bark. Leaves ternate or quaternate, membranous, (narrowly) 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 elliptic to oblanceolate, acute to narrowly cuneate or subrounded at the base, acute to acuminate at the apex, 2-9(-13) cm long, 0.8-3(-4) cm wide, medium to dark green and subglabrous to strigose above, pale green and strigose-pilose below, especially along the veins; secondary veins (5-)6-10(-13) on either side of the midvein; margin (sub-)denticulate. Petioles mostly strigose, 7— 15(— 17) mm long. Stipules narrowly lanceolate, 2^4 mm long, ca. 1 mm wide, subpersistent. Flowers few to numerous and solitary in leaf axils. Pedicels drooping, 8-23(-32) mm long. Ovary ellipsoid to cylindric, 5-8 mm long, 1.5-2.5 mm thick. Floral tube narrowly funnelform, (15-)19-30(-33) mm long, 2-3(-4) mm wide and slight- ly bulbous at the base, then narrowed to 1.5-2.5 mm above the nectary and gradually widened above until 5-8 mm wide at the rim, sparsely strigose outside, pilose in lower V6 inside. Sepals ovate-lanceolate to lanceolate, 9-16 mm long, 3-5 mm wide, acute, spreading to strongly divergent or slightly reflexed at an- thesis. Tube and sepals bright scarlet. Petals dull scarlet, oblong-elliptic to sub- rotund, 6— 10(— 1 1) mm long, (2-)3-8 mm wide, acute to rounded at the apex. Nectary mostly unlobed, ca. 1 mm high. Filaments red, 5-10 mm and 4-7 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide, white. Style red, pilose in lower X A-Vy, stigma capitate, 4-cleft at the apex, 1.5-2.5 mm long, 1-2.5 mm wide, cream to scarlet. Berry ellipsoid, 4-angled before maturity, 13-18 mm long, 7-10 mm thick, red purple when ripe; seeds 1.5-2 mm long, ca. 0.8 mm wide. Gametic chromosome number n = 1 1 . Distribution: Ecuadorian Andes. Locally common in hedgerows along fields and in thickets, mainly in the moderately dry interandean valleys of Ecuador, also on the moister eastern and western slopes, from Loja to Pichincha, with a few scattered collections farther north in Imbabura and Carchi, (2,000-)2,500- 3,500(-3,800) m (Fig. 56). Representative specimens examined: Ecuador, azuay: 13 km from Cuenca past Sayausi, Berry & Escobar 3185 (MO, QCA); along Rio Yanucay above San Joaquin, Berry & Escobar 3210 (MO); along Rio Matadero W of Cuenca, Camp E-1914 (NY, RSA, US); Sayausi, Hading 1396 (NY, S); Tam- bolama and Huasi-huaico, W of Cuenca, Lehmann 4595 (K). bolivar: Chaparro de Gualicon, Loma, Cordillera Occidental, Acosta Solis 6268 (F); Urcu-corral Chillanes, Acosta Solis 6624 (F); Pucara de Telimbela, Cordillera Occidental, Acosta Solis 6818 (F); Alto de Telimbela, Acosta Solis 7154 (F); Balsapampa to San Miguel, Hading et al. 9531 (RSA). canar: near Pimo, Camp E-4135 G, MO, NY, P, RSA, S, UC, VEN); between Cuenca and Huigra, Hitchcock 21688 (US); Ventanillas, above Shoray, E slope of Cerro Yanguang, 34 km ENE of Azogues, Fosberg & Prieto 22775 (RSA, US); between Tambo and Sucsal, Giler (Camp number) E-2756 (NY, RSA). carchi: Paramos de El Angel, Benoist 3637 (P). chimborazo: Sibambe, Hacienda La Carmela, seccion Era-Pata, Cordillera Occi- dental, Acosta Solis 5432 (F); road from Pusucucho to El Placer, Cordillera Oriental, Acosta Solis 7253 (F); road from Chambo to Sanguay, Albert de Escobar 1091 (TEX); near Quinacorral, Andre in 1876 (K); slopes of Cerro Chiguazo, Lugo 512 (RSA); Javinac, ca. 15 km from Panipe, Lugo 519 (RSA); ca. 3-4 km from Puela, Lugo 576 (RSA); Chontapamba, between Puela and Banos, Lugo 741 (RSA); Manzano, 1 km from Puela, Lugo 1303 (RSA); between Banos and Riobamba, Lugo 1844 (RSA); between Guaranda and Bodegas, Remy in 1836 (P); vicinity of Huigra, Hacienda de Licay, Rose 22477 (US); road Riobamba-Huamboya, Scolnik 1533 (RSA); Pallatanga, Spruce 5798 (BM, CGE, G, K 2 sheets, NY, OXF, TCD, W); Hacienda Joyagshi, road from Sibambe to Tambo, Wiggins 10705 (POM), cotopaxi; San Miguel, Andre 4015 (K); Pilalo, Hading 4877 (LL, NY, S); from Pilalo to Macuchi, Haught 2955 (POM, US); Pilalo, Holm-Nielsen & Jeppesen 1141 (AAU, S); Panamerican highway, ca. 6 km N of Lasso, ENE of Pastocalle, Sparre 15805 (S). imbabura: Paramo de Cotacachi, from Otavalo to Apuela, Davis 276 (MO); Mojanda, Sodiro in 1903 (P). loja: Cisne, Andre s.n. (K); Loja, Andre s.n. (K); between Saraguro and San Lucas, Asplund 17973 (NY, S); 5 km S of Saraguro, Berry & Escobar 3192 (MO); 11 km S of Saraguro, Berry & Escobar 3196 (MO); Cerro Villonaco, Berry & Escobar 3204 (MO), Camp E-238 (NY, RSA); Cerros de Acacana, ca. 30 km N of Loja, 1982] BERRY— FUCHSIA SECT. FUCHSIA 85 Espinosa E-1440 (POM, RSA); mountains of Loja, Hartweg 733 (BM, CGE, G, K, LE, NY, OXF, P, W); Km 43 on Panamerican highway N of Loja, Holm-Nielsen 4708 (AAU, MO, NY, S); San Lucas, Jameson 170 (K); Santiago, Poortman 151 (P); Cerro Villonaco, 20 km W of Loja, Sparre 16244 (S). pichincha: Gualilagua, El Corazon, Acosta Solis 7110 (F); Valle Seco del Pedregal, Acosta Solis 8442 (F); near Tambillo, Andre 3681 (K); Paramo de Paguangualli, Corazon, Andre in 1876 (K); Machachi, Asplund 6226 (S); Paramos de Aloag, Benoist 3060 (P, RSA); N of Volcan Cotopaxi on Panamerican highway, Berry & Escobar 3233 (MO, QCA); INIAP station, 14 km S of Quito, Escobar 621 (MO); ascent to Cotopaxi, Espinosa E-2342 (RSA); between Tambillo and Aloag, 28 km S of Quito, Fosberg 22549 (COL, NY, RSA, US); road Quito to Santo Domingo de los Colorados, near top of cordillera, Gentry 9470 (MO, S); vicinity of Tambillo, Penland & Summers 959 (F, GH, POM, RSA); pass at Paramo el Corazon, Prescott 922 (DS, NY); near Quito, Sodiro in 1898 (P); road to Iliniza, Sparre 15785 (S); Panamerican highway, ca. 1 km N of RR, ESE of El Chaupi, Sparre 15801 (S). tungurahua: between Leito and La Cima, Acosta Solis 9024 (F); between Huambalo and Cotalo, Acosta Solis 9717 (F); between La Cima and Riochico, Acosta Solis 10234 (F); Hacienda Yanayacu, Mocha, Balls B7188 (F, K, UC, US); near Agua de Oro, W of Ambato, Heinrichs 828 (G, NY, Z); Mocha, Laudeman 35 (BM); Runtun, ca. 4 km from Banos, Lugo 1190 (MO, RSA); Cusatagua, near Ambato, Pachano 217 (NY, US); S of Banos, Penland & Summers 93 (BH, F, POM); near Banos, Spruce 5203 (BM, CGE, G, GH, K, LE, NY, OXF, P, TCD, W). zamora- chinchipe: Km 12-14 road from Loja to Zamora, Dodson & Thien 1369 (DS, MO). Fuchsia loxensis is distinguished from other axillary, short-flowered species in the following combination of characters: leaves ternate or quaternate, elliptic to subrotund petals, 4-angled ovaries, and a generally erect, shrubby to arbores- cent habit. Although it has been placed in its own species group with two other short-flowered, but rather distantly related species, the leaves and flower shape resemble some of the long-tubed members of the F. petiolaris species group such as F. ampliata. It occurs sympatrically with F. ampliata, F. harlingii, and F. vulcanica. As treated here, F. loxensis is a wide ranging and variable species including all the short-tubed, strictly axillary-flowered collections of Fuchsia in Ecuador except for F. scabriuscula and F. steyermarkii. The shape of the leaves, petals, and floral tubes as well as pubescence are characters that vary widely in this complex. The plant habit and habitat are important characters that, unfortunately, are too little known for most collections because detailed field observations have not been made. Plants that I have seen in Pichincha and Azuay were large, erect shrubs with thick basal stems and often grew in hedgerows. Plants in northern Loja were more sprawling in thickets, as is more typical in the section. There are too few critical field observations in this group, and further study may lead to the recognition of additional taxa. Some of the different geographical variants found are: The Central Valley in Pichincha, Cotopaxi, Tungurahua, and Azuay. Collec- tions from these areas are from high elevations (over 3,200 m in the north and above 2,800 m in the south) and are generally from erect, hedgerow shrubs. The habitats in these areas are somewhat drier than most fuchsias tolerate. The type of F. umhrosa has rather long, ampliate floral tubes (31-33 mm long), sepals 16 mm long, and broad leaves, but it comes from the Central Valley southeast of Quito and may just be a robust shade plant of F. loxensis. Imbabura. The two specimens from this province grow at high elevations (ca. 3,400 m) but have unusually short floral tubes (ca. 15 mm long), oblong petals, and thin, ternate leaves with narrowly cuneate to attenuate leaf bases. This might be a new or different species, but more material is needed. Bolivar and western Cotopaxi. Collections from the western slopes of the 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Andes in these provinces grow at considerably lower elevations (2,400-2,800 m) than the previous groups, and they often have larger leaves and flowers. No living plants of these entities were seen, and there might be some intergradation with F. amp Hat a. Northern Loja. Local populations occurring between Saraguro and San Lucas have an unusual white leaf undersurface and until now have been recognized as a separate species, F. hypoleuca. As noted in the original publication of that species (Johnston, 1925) however, it is an associated fungus with a thick layer of whitish mycelia, not the plant's own pubescence, that causes the unusual leaf characteristic. In the mass of mycelia and epidermal trichomes that one finds together on the leaf undersides, stalked structures called "brown caps" by D. B. O. Savile (pers. comm.) are often observed, and anatomical sections show that these are definitely part of the fungus and not glandular processes of the plant epidermis (R. Keating, pers. comm.). Fungal specimens sent to a specialist for examination were apparently sterile and could not be identified (D. B. O. Savile, pers. comm.). Wider sampling of Fuchsia populations in Loja and further eval- uation of this fungal relationship will be necessary before the relationships of these northern Loja populations with others of F. loxensis can be confirmed, however. Tungurahua and Cahar. The set of collections labelled Spruce 5203 probably is a mixture of F. loxensis (sheets at BM, CGE, NY, and OXF), while others (G, GH, and P) have nitid, dentate leaves and floral tubes ca. 35 mm long. The lack of any specific locality information makes evaluation of these specimens difficult. Hitchcock 21667 (GH, NY, US), from the Azuay-Canar border area, was described as F. apiculata, but its buds are no more pointed than other collections throughout the range of this complex, and no other distinguishing characters are apparent. 6. Fuchsia scabriuscula Bentham, PI. Hartw. 177. 1845. Munz, Proc. Calif. Acad. Sci. IV. 25:58, pi. 8, fig. 46. 1943; Opera Bot., Ser. B, 3:22. 1974. type: Ecuador, Prov. Pichincha, W slopes of the Quito Andes, Oct. 1843, Theodor Hartweg 987 (K Bentham Herb., holotype; photograph, MO; BM, BREM, CGE, G, K Hooker Herb., OXF, isotypes; photograph of B isotype, POM). Fig. 22. Decumbent to erect shrubs 0.5-2.5 m tall. Branchlets subterete, 2-5 mm thick, pilose-hispidulous with suberect, whitish hairs; older branches with tan, exfo- liating bark. Leaves opposite, very rarely ternate, membranous, with finely re- ticulate, rugulose venation, elliptic to slightly ovate, acute to rounded at the base, acute to acuminate at the apex, 2.5-12(-14) cm long, 1.5— 5(— 6) cm wide, dull green and strigose above, pale green to flushed purple and strigose below, es- pecially along the veins; secondary veins 8-14 on either side of the midvein, subelevated below; margin subentire. Petioles densely pilose, 5-25 mm long. Stipules lance-linear, 2-3.5 mm long, thick at the base, terminating in a long, dark, filiform tip, subpersistent. Flowers few and solitary in upper leaf axils. Pedicels divergent to drooping, pilose, 10-25 mm long. Ovary oblong, 5-6 mm long, 2-3 mm thick, strigose to hispidulous, green. Floral tube narrowly funnel- 1982] BERRY— FUCHSIA SECT. FUCHSIA 87 Figure 56. Distribution of the Fuchsia loxenis and F. macrophylla species groups form, 15-27 mm long, 2-3 mm wide at the base, slightly narrowed to 1.5-2.5 mm wide above the nectary and gradually widened above until 5-7 mm wide at the rim, strigose to hispidulous outside, densely pilose inside in lower Vz. Sepals lanceolate, 9-12 mm long, 3-4 spreading at anthesis. Tube and sepals whitish pink to red. Petals pink to scarlet, elliptic-oblong, 7-10 mm long, 3-4.5 mm wide, broadly acute to rounded at the apex, spreading at anthesis. Nectary usually unlobed, green, ca. 1.5 mm high. 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Filaments red to pink, 5-7 mm and 3-5 mm long; anthers elliptic-oblong, 1.5-2 mm long, ca. 1 mm wide, bright white. Style densely villous for most its length, pink; stigma subglobose, 4-cleft at the apex, ca. 2 mm long and ca. 2 mm wide, exserted 1-4 mm beyond the anthers, white. Berry oblong before maturity, be- coming subglobose when ripe, usually deep, nitid purple; seeds tan to bright purple, 1-1.3 mm long, 0.8-1.0 mm wide. Distribution: Ecuador and southern Colombia. Low shrubs locally frequent in clearings, thickets, and moist banks of cloud forest; in Ecuador, in Napo, Pichincha, and Tungurahua Provinces, on both the east and west slopes of the Andes; in Colombia, on the eastern slopes of the Nudo de Pasto in Putumayo, in the Cordillera Central as far north as Valle, and in the Cordillera Oriental to the Cauca-Huila border; 1,400-2,750 m (Fig. 56). Representative specimens examined: Colombia, cauca: Km 31-32 from Pitalito toward Mocoa, Berry 3596 (COL, MO); Rio Villalobos, vicinity of Rio Suazita, Schultes & Villarreal 5173 (COL, F, GH). huila: Km 25 of Pitalito-Mocoa road, Berry 3593 (COL, MO); Km 28-32 of road from Pitalito to Mocoa, Luteyn et al. 7550 (COL, MO, NY); La Plata, von Sneidern 2462 (S). putumayo: Mirador, between San Francisco and Mocoa, Bristol 545 (GH); Rio Susunga, 18-20 km W of Mocoa, Fosberg 20395 (NY, RSA, UC, US); Km 90-91 between Sibundoy and Mocoa, Luteyn et al. 5041 (COL, MO); Los Monos, Km 92 between San Francisco and Mocoa, Plowman & Davis 4328 (COL), valle: El Guayabo, from Palmira to Taco, Berry & Escobar 3565 (COL, MO). Ecuador, napo: Rio Chingual, Acosta Solis 13250 (F); Cuyuja, Balslev & Mads en 10476 (AAU, MO); between Baeza and Papallacta, Mexia 7339 (GH, NA, S, UC, US); Baeza to Tena, ca. 5 km S of Cosanga, Ollgaard & Balslev 10248 (AAU, MO); above Baeza on road to Quito, Plowman et al. 3889 (COL, F, GH, K). pichincha: Guaruna, Km 38 of road to Saloya, Acosta Solis 1 1018 (F); Mindo, Andre in 1876 (K); Corazon, towards Miligalli, Andre 3710 (K, NY); Salvador, below San Juan, Asplund 16184 (NY, S); 34 km W of Chillogallo towards Chiriboga, Berry & Escobar 3243 (MO, QCA); 30 km W of Cotocallao towards Nanegal, Berry & Escobar 3178 (MO, QCA); W slope Cerro Corazon, Camp E-1663 (NY, RSA); Quito-Santo Domingo road, Haught 3224 (A, BH, US); Quito Andes, Jameson 92 (BM, G, GH, P); Quito-Nono-Puerto Quito road, 13 km NW of Nono, Luteyn et al. 6519 (MO, NY); road Nono- Tandayapi, Km 43-45 along Rio Alambi, Sparre 15989 (S). tungurahua: Banos, Benoist 4223 (P); Rio Verde, Hurling et al. 10159 (MO, RSA); valley of Rio Pastaza, between Banos and Cashurco, Hitchcock 21781 (GH, NY); Montana Woma, 11 km E of Banos, Holm-Nielsen & Jeppesen 279 (AAU, DS); Volcan Tungurahua, Lehmann 4995 (F, K). without locality: Spruce 5038 (BM, CGE, G 2 sheets, GH, F, K, LE, NY, OXF, S 2 sheets, TCD, W 3 sheets). This species is easily distinguished by its finely rugulose, opposite leaves and stiff, whitish pubescence on most parts. Just a few flowers are borne on each branch; these later produce round, dark purple berries. Fuchsia scabriuscula occurs sympatrically with F. cuatrecasasii, F. hartwegii, F. macrostigma, F. orientalis, F. pallescens, F. sessilifolia, F. sylvatica, and F. verrucosa. No hy- brids have been detected with these species, despite close examination of several populations. 7. Fuchsia steyermarkii P. Berry, sp. nov. type: Ecuador, Prov. Zamora-Chinchipe, between Rancho Achupallas and Tambo Valladolid, deep forest along the W side of Rio Valladolid to Rio Molino, 1,800-2,000 m, 12 Oct. 1943, Julian A. Steyermark 54596 (NY, holotype; photograph, MO). Frutex pilosus ramosissimus ca. 2 m altus. Folia linearia, revoluta, opposita vel 3^-verticillata sed plerumque fasciculata, firme membranacea ambis extremis acuta, 20-70 mm longa, 2-3 mm lata, superne subnitida atroviridia strigosaque, subtus pallidioria praesertim in costa media margineque dense pilosa; nervis secundariis plerumque inconspicuis, margine valde revoluta; petiolis 0.2-2.0 mm 1982] BERRY— FUCHSIA SECT. FUCHSIA 89 longis; stipulis fuscatis, lanceolato-linearibus, ca. 1.5 mm longis, subpersistentibus. Flores pauci, axillares, ad apicem ramorum dispositi; pedicellis 16-18 mm longis; ovario oblongo, 4-5 mm longo. Tubi florales roseo-rubri, anguste infundibuliformes, 33-37 mm longi, basi 2.5-3 mm lati et bulbosi inde ca. 2 mm lati constricti superne gradatim dilatati summo 5-6 mm lati, extus laxe strigoso-pilosi, intus infra medium pilosi. Sepala lanceolata, acuminata, 13-15 mm longa, 3-4 mm lata, roseo-rubra. Petala coccinea, lanceolata, acuta vel subacuminata, 9-10 mm longa, ca. 3 mm lata. Filamenta anti- sepala 8-9 mm longa antipetala 6-7 mm longa, antheris oblongis, 2.5-3 mm longis ca. 2 mm latis. Stylus laxe villosus, stigmate capitato apice leviter 4-fisso, ca. 2 mm longo, 2-3 mm lato. Bacca matura non visa. Densely pilose, many-branched shrub ca. 2 m tall. Branchlets subterete, 1.5-4 mm thick, pilose, with copper-colored bark. Leaves opposite to quaternate but usually appearing fasciculate due to the very reduced lateral shoots, firmly mem- branous, linear, acute at both ends; 20-70 mm long, 2-3 mm wide, subglossy dark green and sparsely strigose above, pale and densely pilose below along central nerve and margin; secondary veins mostly inconspicuous, mid vein strong- ly prominent below, margin strongly revolute. Petioles 0.2-2 mm long. Stipules dark, lance-linear, ca. 1.5 mm long, subpersistent. Flowers few and axillary near the branch tips. Pedicels 16-18 mm long. Ovary oblong, 4-5 mm and ca. 2 mm thick. Floral tube narrowly funnelform, 33-37 mm long, 2.5-3 mm wide and bulbous at the base, narrowed to ca. 2 mm wide above the nectary and gradually widened above until 5-6 mm wide at the rim, loosely strigose-pilose outside, pilose inside in lower l /i. Sepals lanceolate, acuminate, 13-15 mm long, 3-4 mm wide. Tube and sepals rose red. Petals scarlet, lanceolate, acute to subacuminate at the apex, 9-10 mm long, ca. 3 mm wide. Nectary unlobed, ca. 1.5 mm high. Filaments red, 8-9 mm and 6-7 mm long; anthers oblong, 2.5-3 mm long and ca. 2 mm wide, cream yellow. Style red, loosely villous for most of its length; stigma capitate, slightly 4-lobed at the apex, 2 mm long, 2-3 mm wide, exserted 1-3 mm beyond the anthers. Berry not seen. Distribution: Known only from the type locality on the eastern slopes of the Andes in southernmost Ecuador (Fig. 56). This is an unmistakable species because of its linear, revolute leaves and dense, pilose pubescence. It comes from a very poorly known area of Ecuador that is still accessible only by foot or by mule. Further collections would be desirable to determine the extent of variability in leaf shape and the probable affinities of this species. It is named in honor of Dr. Julian Steyermark, undoubt- edly the foremost neotropical plant collector of this century. 8. Fuchsia nigricans Linden ex Planchon, Fl. Serres Jard. Eur. 5: t. 481. 1849. Merida, between Mendoza Mucuti type; BM, G 2 sheets, K, LE, TCD 2 sheets, US, W 2 sheets, isotypes). This same collection number at OXF was used as the type for F. caracasensis Fielding & Gardner, Sert. PI. /. 29. 1844; that specimen is different from those listed above, however, and is a natural hybrid of F. nigricans and F. gehrigeri Most Merida locality is given by Linden in the protologue. Fig. 21. 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia atroruhra I. M. Johnston, Contr. Gray Herb. 75:31. 1925. type: Colombia, Dept. Risaralda (for- merly Caldas), Santa Elena, above Santuario, 2,000-2,300 m, 7-13 Sept. 1922, Francis W. Pennell 10313 (GH, holotype; NY, isotype). Fuchsia sylvatica sensu Munz, Proc. Calif. Acad. Sci. IV. 25:68, pi. 1 1 , fig. 58. 1943, pro parte. Fuchsia adpressipilis Steyermark, Fieldiana Bot. 28:438. 1952. type: Venezuela, Edo. Lara, between Santo Domingo and Los Quebraditos, S of Las Sabanetas, above Humocaro Bajo, 2,439-2,475 m, 8 Feb. 1944, Julian A. Steyermark 55381 (F, holotype; NY, isotype). Erect to scandent shrubs 1-3 m tall with suberect to spreading branches. Young growth densely canescent with appressed or less often suberect hairs; branchlets terete, 2-5 mm thick, dull purple to light green; older branches with light brown, splitting bark. Leaves mostly ternate, opposite or rarely quaternate, membranous, smooth to sulcate nerved, mostly elliptic or obovate, but also spat- ulate or subpanduriform, acute to attenuate or cuneate at the base and sometimes noticeably unequal, acute to rounded at the apex, (4-)7-15(-18) cm long, (2-)3-8(-9) cm wide, medium-dark green and strigillose to subglabrous above, pale green and strigillose below, especially along the veins; secondary veins 1 1— 16(— 20) on either side of the midvein, sometimes red tinged; margin remotely gland-denticulate. Petioles strigillose, 12— 35(— 48) mm long. Stipules narrowly tri- angular, dark purple, 1-2 mm long, ca. 0.8 mm wide, succulent at the base, subulate at the apex, subpersistent. Flowers few to numerous, axillary in upper leaf nodes or in terminal bracteate racemes; rachis 4-22 cm long; bracts narrowly elliptic, short petiolate, loosely spaced. Pedicels slender, 3-9 mm long, suberect in bud to divergent or drooping at anthesis. Ovary narrowly cylindric-fusiform, densely canescent or strigillose, 6-11 mm long, 2-3 mm thick. Floral tube sub- cylindric, 14-22 mm long, 1.5-3 mm wide and slightly bulbous at the base, grad- ually widened above until 3-5(-6) mm wide at the rim, canescent-strigillose out- side, densely pilose inside in lower Vi. Sepals lanceolate, acute, 6-10 mm long, 3-4 mm wide, spreading at anthesis. Tube and sepals pale pink to lavender or light red. Petals much darker, generally dark purple, narrowly elliptic-oblong, acute, 5-10 mm long, 2-3.5 mm wide, suberect to spreading at anthesis. Nectary green, shallowly 4-lobed, ca. 1.5 mm high. Filaments deep purple, 5-6 mm and 3-4 mm long; anthers oblong, 1.5-2 mm long, ca. 1 mm wide, dull white. Style glabrous, pink to purple; stigma capitate, subtetragonous, 2-3 mm long, 2-3.5 mm wide, 4-lobed in upper Vi, pink, slightly exserted beyond the anthers or in contact with the antesepalous stamens at anthesis. Berry cylindric, 15-25 mm long, 6-10 mm thick, dull green to flushed purple, strigillose, slightly verrucose before maturity; seeds tan, 1.3-1.6 mm long, ca. 0.8 mm wide. Gametic chro- mosome number /z = 11. Distribution: Venezuela and Colombia. Scattered to locally frequent shrubs in forest openings, streambanks, along roadsides, and in moist thickets in mid- elevation cloud forest; in Venezuela, from Lara to Tachira at 2,100-2,650 m; in Colombia, in all three Cordilleras: in the Cordillera Oriental known only from Norte de Santander near the Venezuelan border; in the Cordillera Central in Tolima, Quindio, Caldas, and Antioquia; in the Cordillera Occidental from An- tioquia south to Cauca; 1,700-2,700 m (Fig. 57). Representative specimens examined: Venezuela, lara: below La Sabaneta, 15 km from Los Na- ranjos above Humocaro Bajo, Berry & Azuaje 2500 (MERF, MO); trail from Humocaro to Buenos Aires, below Paramo de Las Rosas, Liesner et al. 8001 (MO), 8215 (MO, VEN). merida: La Culata 1982] BERRY— FUCHSIA SECT. FUCHSIA 91 to El Valle, Badillo 5587 (MY); between El Morro and Aricagua, Loma de la Vagabunda, Badillo 6567 (MY); Guaraque to Tovar, Benitez de Rojas 2076 (MY); La Mosquera, Mucuqui, above Pueblo Nuevo, Bernardi 232 (NY); Monte Zerpa, Bernardi 653 (NY); 10 km above La Azulita to La Car- bonera, Berry & Centeno 2513 (MER, MERF, MO); 1 km below Chachopo on road to Timotes, Berry 3131 (MO, VEN); 4 km below San Eusebio to La Azulita, Berry 3454 (MO, VEN); La Carbonera, Berry 3451 (MO); between La Mucuy and Mesa de los Pinos, Berry 3438 (MO, VEN); Paramo de la Sal, Jahn 507 (VEN); above Palmira, Jahn 509 (US, VEN); Paramo de Aricagua, Jahn 1028 (US, VEN); road to El Morro, Jahn in 1910 (VEN); El Filo, between Guaraque and Tovar, Lopez-Palacios 8 (MER); trail from La Escalera to Puente de La Escalera, Luteyn et al. 6208 (MO); road to Paramo de los Colorados, Quintero et al. 1280 (MER); El Chorotal, Quintero 1616 (MER); Tienditas del Chama to Aricagua, Quintero & Ricardi 1772 (MER); between El Chorreron and La Cueva, road to Paramo de Palmira, Dpto. Miranda, Ruiz-Terdn & Dugarte 12493 (MERF); Quebrada La Honda, E of Los Nevados, Dpto. Libertador, Ruiz-Terdn 12090 (MO); between Los Corales and Los Cuadros, Steyermark 55769 (F, US); 2 km above Las Tapias, S of Bailadores, Tillett & Hbnig 738-465 (MO, VEN); road from Manzano Alto to Paramo de los Conejos, ca. 7 km NE of Merida, Wessels-Boer 2245 (MER, MO, U). tachira: 13 km E of El Portachuelo, road to Pregonero, Berry 3288 (MO, VEN); 2 km E of Zumbador, road to Queniquea, Berry 3294 (MO, VEN); 15 km E of Zumbador to Queniquea, Berry 3304 (MO, VEN); between Zumbador and San Isidro, Bunting 2548 (GH, MY); Paramo el Pantano, Charpin et al. 13446 (G); Quebrada Agua Azul, S of El Reposo, Steyermark & Liesner 118431 (MO, VEN). trujillo: Campo Elias to Bocono, Badillo 4965 (MY); 12 km W of San Rafael, old road Bocono-Trujillo, Berry 3103 (MO, VEN), 3104 (MO, VEN); 5 km above La Mesa de Esnujaque, road to Dury, Berry 3130 (MO); 8 km W of El Batatal to Bocono, Berry 3093 (MO, VEN); 3095 (MO, VEN); Agua Obispo, Funck & Schlim 797 (BM, CGE, F, G, LE, OXF, P 2 sheets, W); Rio de Agua Negra, near Bocono, Lasser 1175 (US, VEN); Misisi, slopes of Paramo de la Cristalina, Ruiz-Terdn & Lopez-Palacios 7635 (MERF, MO); Paramo de Guaramacal, 15 km from Bocono, Ruiz-Terdn 9224 (MERF, MO). Colombia, antioquia: Santa Elena, Archer 1195 (US 2 sheets); near Narino, Barkley & Johnson 180826 (COL, RSA, US), caldas: Mt. Cardal, Tracey 344 (K). choco: N of Alban, Dugand & Jaramillo 3040 (COL, US); Quebrada El Sucio above Carmen de Atrato, Fosberg & Core 21550 (RSA, S, US), norte de santander: Ocana to Pamplona, Kal- breyerlSisale 1104 (K); Prov. of Ocana, Schlim 338 (P). quindio: Rio Santa Rita, Salento, Killip & Hazen 8966 (GH, NY, PH, US); Rio Quindio, Salento, Pennell 10099 (GH): Salento, Pennell & Hazen 10144 (GH, NY, PH, US), tolima: Alto San Juan-Quindio, Andre 2072 (K); Quindio, Andre s.n. (K); La Suiza, above Ibague, Cuatrecasas 3260 (MA); Las Juntas, Combayeque, Goudot 3 (P); Rio Toche to Machin, Killip & Hazen 9564 (GH); Rio Combeina, Juntas, Schneider 1156 (COL, S); Toche, von Sneidern 3090 (COL, S); la Mediacion, Quindio, Triana 3811 (G, K, W). valle: above Las Brisas, between El Tabor and Alto de Mira, Cuatrecasas 22425 (F, RSA, VALLE). This species is vicarious with Fuchsia sylvatica, which differs in a series of minor characters such as lighter colored petals, narrower fruits, more rugulose leaves, and denser, more cordate bracts. They are geographically separated by the Rio Patfa valley in the Cordillera Occidental of southern Colombia. The pat- tern of distribution of these two species suggests a northern origin in the Cordi- llera Oriental of Colombia and Venezuela, where F. nigricans is now most com- mon and widespread. As with at least three other species in the section, it may have spread westwards into the Cordillera Central and Cordillera Occidental; F. nigricans is not found south of 3°N Latitude or in the Nudo de Pasto area. Fuchsia sylvatica may have diverged secondarily south of the Patia barrier, where it is now restricted on the western slopes of the Cordillera Occidental in Ecuador. If a northwards migration route had been used, one would expect to find members of these species in the Nudo de Pasto or in the southern parts of the Colombian cordilleras, but this is not the case. Fuchsia nigricans and F. sylvatica are distinguished from their congeners by their fine, ashy pubescence; the short-pedicellate, elongate ovaries and fruits; the petals darker than the tube or sepals; and the subtetragonous stigma. The closest relative is F. pallescens, which occurs on the opposite side of the Ecuadorian Andes from F. sylvatica and which shares the subracemose flowers, darker pet- 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 als, and elongate ovaries on short pedicels. It differs, however, in its pale whitish floral tube, semisucculent stems, and thin membranous, long petiolate leaves. Fushia sessilifolia also has small bicolored flowers with short pedicellate, elon- gate fruits, but the flowers are borne in panicles, and the leaves are subsessile, lanceolate, and quaternate. Fuchsia nigricans is the most widespread of the Venezuelan fuchsias and occurs continuously from Lara to Tachira. It is surprisingly absent from most of the Colombian side of the Cordillera Oriental, however. In Colombia, it is only one of two species that occurs in all three Cordilleras. With this wide range it is found sympatrically with five species, F. gehrigeri, F. putumayensis , F. sessili- folia, F. venusta, and F. verrucosa. Hybrids have been found with all of these except F. verrucosa ; a detailed analysis of the different hybrids is included in the section on interspecific hybridization. 9. Fuchsia sylvatica Bentham, PI. Hartw. 176. 1845. Munz, Proc. Calif. Acad. Sci. IV. 25:68. 1943, pro parte; Opera Bot., Ser. B, 3:23. 1974, pro parte. type: Ecuador, Prov. Pichincha, woods at Guayan, W slopes of Volcan Pi- chincha, 1841-1843, Theodor Hartweg 984 (K Bentham Herb., holotype; pho- tograph, MO; BM, BREM, CGE 2 sheets, G, K Hooker Herb., LE, OXF, isotypes). Erect to scandent shrubs 1-2.5 m tall. Young growth finely canescent; older stems with tan, fissured bark. Leaves mostly ternate, opposite, or rarely quater- nate, membranous, finely reticulate- veined and sometimes rugulose, elliptic to (ob-)ovate, acute to attenuate at the base, mostly acute at the apex, 4-15 cm long, 2-8 cm wide, medium-dark green and strigillose above, pale green to tinged red and strigillose below; secondary veins 11-18 on either side of the midvein; margin remotely gland-denticulate. Petioles strigillose, 6-34 mm long. Stipules dark purple, narrowly triangular, 1-2 mm long, ca. 0.7 mm wide, thick at the base, subulate at the apex, subpersistent. Flowers generally numerous in long, straggly, terminal or subterminal racemes; rachis arching-pendant or sometimes ± twisted, 5-20 cm long; bracts ovate-cordate, subsessile, tightly grouped. Ped- icels slender, divergent to pendant, 3-10 mm long. Ovary narrowly cylindric- fusiform, often narrowed towards the apex, 6-10 mm long, 2-3 mm thick. Floral tube narrowly funnelform, 14-22 mm long, 2-3 mm wide and slightly bulbous at the base, gradually widened above until 3.5-7 mm wide at the rim, canescent- strigillose outside, pilose inside in lower Vi. Sepals lanceolate, acute, 10-13 mm long, 3-5 mm wide, spreading at anthesis. Tube and sepals red to rose red. Petals darker, crimson, narrowly elliptic-oblong, acute, 5-10 mm long, 2-4 mm wide. Nectary green, shallowly 4-lobed, ca. 1.5 mm high. Filaments red, 5-7 mm and 3-5 mm long; anthers oblong, 1.5-2 mm long, ca. 1 mm thick, dull white. Style glabrous to sparsely pilose, light red; stigma capitate, subtetragonous, 2-3 mm long, 2-4 mm wide, 4-lobed in upper Vi, pink, slightly exserted beyond the an- thers. Berry subcylindric-fusiform, gradually narrowed towards the apex, ± ver- rucose before maturity, 13-17 mm long, 5-7 mm thick, red purple to subtranslu- cent; seeds ca. 1.5 mm long, ca. 0.8 mm wide. Gametic chromosome number n = 11. 1982] BERRY— FUCHSIA SECT. FUCHSIA 93 400 Km 1 Figure 57. Distribution of the Fuchsia nigricans species group. western from Cotopaxi to Imbabura Provinces, in mid-elevation cloud forest, growing in 600-3 57). Representative specimens examined: Ecuador, cotopaxi: 5 km above Pilalo towards Latacunga, Berry & Berry 2550 (MO); 4 km E of Pilalo, Escobar 1095 (MO), imbabura: 25 km W of Laguna Cuicocha on road to Inta, Berry 3171 (MO, QCA). pichincha: Los Alpes, NW side of El Corazon, Acosta Sotis 7074 (F); 26 km from Chillogallo towards Chiriboga, Berry 2537 (MO, Q); Km 32-33 W of Chillogallo, Berry & Escobar 3237 (MO), 3238 (MO), 3239 (MO), 3240 (MO); Sigsal, above Chi- riboga, Berry & Escobar 3244 (MO); 1 1-12 km NW of Nono, Croat 38823 (MO); W side of Volcan Pichincha, Jameson 709 (BM, G, K, LE, OXF 2 sheets); valley of Lloa, Jameson s.n. (K); Km 32-38 old road Quito to Santo Domingo de los Colorados, Luteyn & Lebron-Luteyn 5608 (MO, NY); Alas- pongo, trail from Nono to Gualea, Mexia 7703 (BM, GH, K, MO, NY, POM, S, U, UC, US). This species is closely related to F. nigricans, and their differences, distri- butions, and relationships are discussed under that species. It occurs sympatri- 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 cally with F. macrostigma, F. scabriuscula, and F. sessilifolia. No flowering hybrids were found with any of these species, but it is likely that nonflowering hybrids were growing between Nono and Nanegal, where I found a dense thicket of F. scabriuscula, F. sessilifolia, and F. sylvatica growing together in direct contact in 1977. The vegetative similarity of F. scabriuscula and F. sylvatica makes confirmation of this problematical, however. On the western slopes of Prov. Carchi along the Colombian border, plants were found that agreed well with F. sylvatica in all morphological characters except for their much longer flowers and fruits. Floral tubes are 45-54 mm long and fruits ca. 23 mm long in Holm-Nielsen et al. 5752 (AAU, S; Ecuador, Prov. Carchi, Km 60 of Tulcan-Maldonado road, 2,700 m) and Berry 3149 (MO, QCA; 68 km W of Tulcan to Maldonado, 2,500 m). Pollen stainability in these collections is high, 88.8% in Holm-Nielsen et al. 5752 and 94.2% in Berry 3149 (500 grains counted). More intensive collections are needed from this and surrounding areas, because it is very rich in intermediate forms and local endemics (see F. cinerea, F. corollata, F. dependens, and F. polyantha). 10. Fuchsia pallescens Diels, Notizbl. Bot. Gart. Berlin-Dahlem 14:34. 1938. Munz, Proc. Calif. Acad. Sci. IV. 25:29. 1943; Opera Bot., Ser. B, 3:20. 1974. type: Ecuador, Prov. Tungurahua, E of Patate, woods above Leito, 2,750 m, 8 March 1935, Arnold Schultze-Rhonhof 1827 (B, holotype, destroyed in World War II). neotype: Ecuador, Prov. Tungurahua, vicinity of Patate, Hacienda Leito, 2,800 m, 5 Aug. 1939, Erik Asplund 8073 (S; photograph MO). Usually well-branched herbs to subshrubs 5-18 dm tall. Branchlets subterete, semisucculent, 2-4 mm thick, subglabrous; older stems 4-8 mm thick, with dull tan, finely fissured bark. Leaves opposite or ternate, thin membranous, elliptic- ovate, attenuate to rounded at the base, subacuminate at the apex, 3.5-7(-9.5) cm long, 1 . 5— 4(— 5) cm wide, velvety dark green and glabrous to strigillose above, paler below and subglabrous to puberulent, especially along the veins; secondary veins 5-7(-9) on either side of the midvein, lightly impressed above and subele- vated below; margin gland-denticulate. Petioles slender, (15-)25-40(-50) mm long, glabrous to puberulent, green. Stipules triangular, 1-1.5 mm long, ca. 0.5 mm wide, succulent at the base, sometimes connate and reflexed on lower nodes, subpersistent. Flowers few and pendant in upper leaf axils or subracemose at the tips of branches with reduced leaves 10-20 mm long subtending the flowers. Pedicels slender, drooping, 7-15 mm long. Ovary narrowly cylindric, 5-8 mm long, 2-3 mm thick. Floral tube narrowly funnelform, 18-25 mm long, slightly nodose at the base and 1.5-2 mm wide, gradually widened above until 4-5 mm wide at the rim, glabrous to puberulent outside, densely pilose inside in lower Vi. Sepals lanceolate, acute to long acuminate, 9-15 mm long, 3-4 mm wide, spreading at anthesis. Tube and sepals subnitid whitish pink to pale red. Petals considerably darker, maroon to dark red, broadly elliptic, acute to rounded at the apex, 5-8 mm long, ca. 4 mm wide, suberect. Nectary unlobed, 1-1.5 mm high. Filaments light pink, 5-7 mm and 3-5 mm long; anthers oblong, 1-1.5 mm long, ca. 0.8 mm wide, cream to pale yellow. Style glabrous, light pink; stigma capitate, 4-lobed in upper Vi, 1 .5-2.5 mm long, 2-3 mm wide, white. Berry oblong, 1982] BERRY— FUCHSIA SECT. FUCHSIA 95 wed apically, rugose betore maturity, tu 4-6 mm thick: seeds ca. 1 mm long, 0.6-0 13 mm number n = 11. Moist itats, restricted to the eastern slopes of the Andes in Ecuador from Azuay to Napo, and disjunct to the west slopes of the Cordillera Occidental in Cauca, Colombia: 2.550-2.900 m (Fig. 57). Specimens examined: Colombia, cauca: near ridge of Cordillera Occidental, Km 41 of Popayan- Uribe-Munchique road, Berry 3570 (COL, MO); Mount El Derrumbo, Killip 7992 (NY, PH, US), Lehmann 5956 (K). Ecuador, chimborazo: road from Pusucucho to El Placer, Acosta Soils 7255 (F), 7281 (F). imbabura: Trail to Rio San Pedro, ridge S of Rio Clavadero, E of Cayambe, Wiggins 10444 (DS, NY, POM), napo: Km 209 of Quito-Baeza road, below Papallacta, Berry & Berry 2524 (MO, 0), 2527 (MO, Q); above Cuyuja, Berry & Escobar 3248 (MO); Salcedo (San Miguel), Km 55 on Salcedo-Napo road, Boeke 901 (MO); Santa Barbara de Sucumbios, 10 km E of Santa Barbara, Harlinx 4105 (S); between Cuyuja and Papallacta, Holm-Nielsen et al. 6815 (AAU, NY, S). tungu- rahua: between Leito and La Cima, Acosta Soils 9025 (F); between La Cima and Rio Chico, Acosta Solis 10242 (F). azuay: Quebradas leading into the Rio Collay, 3-8 km N of Sevilla de Oro, Camp 4897 (GH, NY, RSA, US). This species is allied to F. nigricans and F. sylvatica in its subracemose inflorescence of short flowers with dark petals and elongate ovaries. The tube and sepals are unusually pale, however, and the plants are generally more deli- cate, with thin leaves and semisucculent young stems. Small, nearly herbaceous plants of F. pallescens are commonly found growing under large Gunner a leaves, though larger, woodier plants can sometimes be found in more open areas such as roadbanks. It occurs sympatrically with F. scabriuscula and F. vulcanica. Fuchsia pallescens occupies ranges distinct from both F. nigricans and F. sylvatica. The disjunction from western Colombia to the eastern slopes of Ecua- dor is similar to that of F. putumayensis and is possibly due to a past migration across the Nudo de Pasto. The limited number of specimens available makes an accurate assessment of the variability of this species difficult, but three variants can be mentioned: 1) Hading 4105 (S), from northern Napo near the Colombian border. The petioles of this collection are short, and the leaves have 8 or 9 secondary veins instead of the normal 5 or 6. Pollen stainability of this individual is 60% (500 grains). 2) Camp 4897 (GH, NY, RSA, US), from Azuay. The floral tubes of this collection are short (ca. 18 mm), as are the nonacuminate sepals, and the petals may not Berry berulent Ecuadorian populations. Killip 7792 (NY, PH, US) and Lehmann 5956 (K) have rather stout stems and larger leaves. The tube and sepal color cannot be deter- mined from these specimens, however, so their placement in this species should be considered tentative. 11. Fuchsia orientalis P. Berry, sp. nov. type: Ecuador, Prov. Napo, between Co- May 1935, Ynes Me 1662406. holotvpe: photograph, MO 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia sylvatica sensu Munz, Opera Bot., Ser. B, 3:23. 1974. Frutex 1-2 m altus, ramulis foliis junioribus puberulo-strigulosis. Folia opposita, raro ternata, membranacea, anguste elliptica vel obovata, basi acuta, attenuata vel inaequalia, apice acuta vel acuminata, 6-21 cm longa, 2.5-6 cm lata, supra subnitida subglabrataque subtus pallidioria nervis marginibusque plerumque strigulosis vel puberulis; nervis secundaris utroque latere 12-17 subtus plerumque prominentibus, margine integerrima; petiolis strigulosis, 8-20 mm longis; stipulis lanceo- latis, subpersistentibus, 1.5-3 mm longis, 0.5-0.8 mm latis, in nodis veternis crassisculis et recurvatis. Flores scarlatini vel aurantiaco-rubri, pauci vel numerosi in racemis abruptis terminalibus vel axi- llaribus dispositi; rhachidi 5-16 cm longa; bracteis persistentibus lanceolatis 10-25 mm longis, 3-10 mm latis, petiolis 3-5 mm longis; pedicellis 3-7(-12) mm longis; ovario oblongo, 4-5 mm longo. Tubi florales anguste infundibuliformes 15-21 mm longi, basi ca. 2 mm lati et parum bulbosi inde leviter constricti superne gradatim dilatati summo 3.5-5 mm lati, extus strigulosi intus infra medium retrorso- villosi. Sepala oblongo-lanceolata, 6-8 mm longa, 3-4 mm lata. Petala late elliptica, 5-8 mm longa, 3-4 mm lata. Filamenta antisepala 6-7 mm longa, antipetala 4-5 mm longa, antheris late oblongis, ca. 1.5 mm longis, ca. 1 mm latis, albidis. Stylus ruber plerumque glaber, stigmate capitato 1.5-2 mm longo, ca. 2.5 mm lato apice leviter 4-fisso. Bacca cylindrica, verrucosa, 9-12 mm longa, ca. 5 mm crassa, rubro-purpurea; seminibus 1-1.1 mm longis, ca. 0.7 mm latis. Numerus gameticus chromo- somatum n = 11. Shrubs 1-2 m tall. Young growth puberulent to strigillose. Leaves opposite, rarely ternate, membranous, narrowly, elliptic to obovate, acute to attenuate or unequal at the base, acute to acuminate at the apex, 6-21 cm long, 2.5-6 cm wide, subnitid and subglabrous to strigillose above, paler and mostly strigillose or puberulent along the veins and margin below; secondary veins 12-17 on either side, mostly prominent below, margin entire. Petioles strigillose, 8-20 mm long. Stipules lanceolate, 1.5-3 mm long, 0.5-0.8 mm wide, mostly persistent and be- coming thick and recurved on lower nodes. Flowers few to many in abrupt, terminal and axillary, bracteate racemes; rachis 5-16 cm long, elongating in fruit; bracts lanceolate, persistent, 10-25 mm long, 3-10 mm wide, with petioles 3-5 mm long. Pedicels strigose, 3-7(-12) mm long. Ovary cylindrical, 4-5 mm long, 1.5-2 mm thick. Floral tube narrowly funnelform, 15-21 mm long, ca. 2 mm wide and slightly bulbous at the base, gradually widened above until 3.5-5 mm wide at the rim, strigillose outside, retrorse villous in lower Vi. Sepals oblong-lanceo- late, 6-8 mm long, 3^4 mm wide. Tube and sepals scarlet to orange red. Petals red to orange red, oblong to broadly elliptic, 5-8 mm long, 3-4 mm wide. Nectary unlobed or shallowly lobed, 1-1.5 mm high. Filaments red, 6-7 mm and 4-5 mm long; anthers broadly oblong, ca. 1.5 mm long and ca. 1 mm wide, white. Style red, mostly glabrous; stigma capitate, 1.5-2 mm long, ca. 2.5 mm wide, 4-cleft at the apex. Berry cylindrical, verrucose, 9-12 mm long, ca. 5 mm thick, red purple; seeds 1-1.1 mm long, ca. 0.7 mm wide. Gametic chromosome number n = 11. Distribution: Ecuador. Scattered shrubs in low elevation cloud forest on the eastern slopes of the Andes from Napo to Zamora-Chinchipe, 1 ,200-2,600 m (Fig. 57). Specimens examined: Ecuador, azuay: 1-8 km N of Sevilla de Oro, Camp E-4251 (NY, P, RSA). loja: 14 km E of Loja towards Zamora, Berry & Escobar 3199 (MO, QCA); Km 10.5 of Loja-Zamora road, Holm-Nielsen et al. 3552 (AAU, S). morona-santiago: between Rio Soldo and La Esperanza, road to Huamboya, Acosta Solis 7343 (F); Cordillera de Cutucii, W slopes, between Logrono and Yaupi, Madison et al. 3342 (MO); trail between Mirador and Pailas, Steyermark 54280 (NY), napo: Cordillera Guacamayo, slope toward Urcusiqui, Asplund 9578 (S); Cosanga, 20 km S of Baeza, Balslev & Madsen 10340 (AAU, MO); 10 km from Baeza to Cosanga, Berry & Escobar 3246 (MO); Cerro Antisana, E of Borja, Grubb et al. 1050 (K, NY); Rio Cosanga, near Cosanga, Kirkbride & Chamba 1982] BERRY— FUCHSIA SECT. FUCHSIA 97 4012 (MO). TUNGURAHUa: Rio Mapoto, Penland & Summers 282 (E, OH, POM, RSA). zamora- chinchipe: Huaico, SE of Loja, Espinosa £-660 (RSA); Km 16 Loja to Zamora, Sparre 16507 (S). This species typically has elongate racemes with persistent, narrowly lanceo- late bracts. This character is also found in the larger leaved F. pilosa, but F. orientalis is most closely allied to F. glaberrima, which has larger flowers and more sessile leaves, but similar thick stipules and persistent bracts. In Napo, it is a low elevation species on slopes leading down into the "Oriente' 1 or Amazon lowlands of Ecuador. It occurs sympatrically there with E. scabriuscula . In southern Ecuador, it occurs between 2,000 and 2,600 m, considerably higher than in the north. M Field Mus. Nat. Hist., Bot. Ser. 13(4):555. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:66, pi. 10, fig. 55. 1943; Opera Bot., Ser. B, 3:15. 1974. type: Ecuador, Prov. Tungurahua, valley of Rio Pastaza, between Banos and Cashurco, 1,300-1,800 m, 25 Sept. 1923, Alberts. Hin MO Simple to few-branched shrubs 1-3 m tall. Branchlets subterete, 3-5 mm thick, glabrous or minutely and sparsely puberulent on youngest growth, green to red- purple. Leaves opposite or subopposite near the branch tips, firmly membranous to subcoriaceous, oblanceolate, obtuse to attenuate at the base, acute to acumin- ate at the apex, 10-24 cm long, 4-8 cm wide, subnitid and glabrous above, pale to usually flushed red-violet below and glabrous to finely puberulent along the nerves; secondary veins 8-20(-26) on either side of the midvein; margin entire. Petioles short, stout, 3-8 mm long. Stipules triangular, thick, mostly connate and 2-4 2-4 6- 0-30 12 mm long, mostly glabrous. Ovary oblong, 7-9 mm long, 2-3 mm thick. Floral tube 35(-40) 6-7 at the rim, glabrous outside, densely pubescent inside for most of length. Sepals oblong-lanceolate, 8-11 mm long, 4-5 mm wide, shortly acute-tipped, subtetra- gonous in bud, spreading at anthesis. Tube and sepals red to orange red. Petals red, broadly elliptic-obovate, 7-11 mm long, 5-6 mm wide, rounded at the apex. Nectary unlobed, ca. 1.5 mm high. Filaments red, 5-6 mm and 3^4 mm long; anthers oblong, ca. 2 mm long, ca. 1 mm wide. Style densely pilose or occasion- ally glabrous; stigma capitate, subtetragonous, ca. 2 mm long, ca. 3 mm wide, 4- lobed apically, shortly exserted beyond the anthers. Berry oblong, ca. 15 mm long, 7-9 mm thick; seeds ca. 2 mm long, ca. 1 mm wide. Distribution: Scarce shrubs of the lower cloud forest limit from Tungurahua, Ecuador, to Amazonas, Peru, 1,600-1,900 m (Fig. 57). Specimens examined: Ecuador, loja: W slopes of Cordillera del Condor N W of Nudo de Sabanilla, around Tambo Cachiyacu, SE of Yangana, Steyermark 54786 (NY), morona-santiago: Plan de Milagro, ca. 10 km NW of Indanza, Jorgensen OHJ-38 (NY); W slopes Cordillera de Cutucu, trail 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 from Logrono to Yaupi, Madison et al. 3531 (MO), tungurahua: upper Rio Pastaza, Spruce 5031 (K). zamora-chinchipe: Rio de San Francisco, Poortman 318 (P). without locality: Lobb s.n. (K); Pearce 329 (K). Peru, amazonas: 12-20 km E of La Peca, Serrania de Bagua, Barbour 2581 (MO), 2665 (MO), 2686 (MO), 2718 (MO), 2734 (MO), 4171 (MO); Gentry et al. 23031 (MO), 23023 (MO); Yambrasbamba, Mathews 803 (OXF); Taulia, Mathews 1484 (K). This species can be readily distinguished by its large, opposite, oblanceolate, and subsessile leaves that are usually violet flushed below. It also has unusually thick, persistent stipules. Fuchsia glaberrima closely resembles F. orientalis in its abrupt racemes, short pedicels, and persistent bracts, but the latter has shorter flowers and longer petiolate leaves. Both are restricted to the low, eastern slopes of the equatorial Andes, but they are not known to occur together or to intergrade. This is the only species in the section to occur both north and south of the Amotape-Huancabamba zone in southern Ecuador and northern Peru. Plants from the southern end of the range (Amazonas and Loja) have more secondary veins (ca. 20 vs. ca. 10) and less hairy styles than central Ecuadorian collections. Their floral tubes are also somewhat longer (35 mm vs. 25 mm), and the inflorescences are looser and less congested. Both vein number and style pubescence can vary locally, however; Gentry et al. 23031 (MO) from Amazonas has fewer than 20 veins and a pilose style, but the close-by Gentry et al. 23023 (MO) has over 25 secondary veins and a glabrous style. 13. Fuchsia macrophylla I. M. Johnston, Contr. Gray Herb. 75:35. 1925. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):557. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:66, pi. 10, fig. 56. 1943. type: Peru, Dept. Junin, La Merced, Hacienda Schunke, 1,200 m, Aug. -Sept. 1923, Francis Macbride 5616 (F 536655, ho- lotype; photographs, NY, POM, UC; GH, US, isotypes). Erect to scandent shrubs 1-3 m tall with spreading branches. Young growth minutely puberulent, branchlets 3-6 mm thick, usually red-purple tinged; older branches with light brown, finely fissured bark. Leaves opposite or less often ternate, firmly membranous, (narrowly) elliptic to lanceolate or (ob-)ovate, acute to cuneate at the base, acute to acuminate at the apex, basal leaves generally considerably larger than the upper ones, 16-27 cm long, 5-9 cm wide, light to dark green and subglabrous above, paler and subglabrous below; secondary veins 14-19 on either side of the midvein, reddish and prominent below, anastomosing into a distinct submarginal vein; margin subentire. Petioles subglabrous, 10-35 mm long, reddish. Stipules dark, lanceolate to triangular, sometimes connate, 1-2 mm long, 0.5-0.8 mm wide, mostly deciduous. Flowers few to numerous in short lateral racemes or on short side branchlets; rachis 2-10(-12) cm long. Pedicels slender, spreading, 10-26(-32) mm long, reddish. Ovary ellipsoid, 4-5 mm long, ca. 2 mm thick. Floral tube narrowly funnelform, 19-25 mm long, ca. 2 mm wide and slightly nodose at the base, gradually widened above until 4-6 mm wide at the rim, puberulent to subglabrous outside, pilose inside for most of length. Sepals lance-oblong, acute, 8-9 mm long, 3-4 mm wide, short-pointed in bud. Tube red, sepals red with green tips. Petals scarlet, elliptic to lanceolate, acute, 7-10 mm long, 3-4 mm wide. Nectary unlobed, ca. 1 mm high. Filaments red, 4-7 mm and 3-4 mm long; anthers oblong, 1.4-1.6 mm long, ca. 0.8 mm wide, dull white. Style pubescent for most of length, red; stigma capitate, ca. 2 mm long and wide, 1982] BERRY— FUCHSIA SECT. FUCHSIA 99 shallowly 4-lobed, cream to pink, exserted 1-2 mm beyond anthers. Berry subglo- bose to ellipsoid, 10-13 mm long, 8-9 mm thick, nitid dark purple; seed tan, 1.1-1.3 mm long, ca. 0.7 mm wide. Gametic chromosome number n = 11. Distribution: Central to southern Peru, from Loreto and Huanuco to Puno; in moist thickets in low elevation cloud forest and subtropical wet forest; 1,200-2,000 m (Fig. 56). Representative specimens examined: Peru, ayacucho: 8 km below Jano, road from Tambo to Ayna, Berry 3053 (MO, USM); between Tambo San Miguel, Ayna, and Hacienda Luisiana, Dudley 11816 (F, MO, NA 2 sheets, NY, USM); Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22353 (F, GH, K, NY, S, US), cuzco: Tunquimayo, Quince Mil, Prov. Quispicanchis, Vargas 11776 (CUZ, MO); between Tambomayo and Consuelo, along Rio Tambomayo, Prov. Paucartambo, West 7113 (GH, MO, UC). huanuco: trail from Puente Durand to Exito, Pampa Hermosa, Mexia 8142 (BH, BM, F, G, GH, K, MO, NA, NY, S, U, UC); cuesta San Toribio, quebrada Cutama, Vargas 5370 (BH, CUZ); Palo Marcado, Vargas 15336 (MO, USM). junin: 44 km W of Satipo towards Conception, Berry & Aronson 3080 (MO, USM); Cumbre Yacunay above La Merced, Hutchison 1173 (F, G, GH, MICH, MO, NY, S, UC, US); Pichis trail, Killip & Smith 25436 (F, GH, NY, US); Pangoa, Mathews 1169 (K, OXF); Utcuyacu, Woytkowski 35382 (F, MO 2 sheets, UC, USM). lor- eto: La Divisoria, Ferreyra 1654 (MO, RSA, US 2 sheets, USM); below La Divisoria, Ferreyra 4299 (MO, US, USM); vicinity of Aguaytia, Mathias & Taylor 5150 (F, RSA, US, USM). pasco: San Luis de Oxapampa, Infantes 1492 (MO, USM); Vitoc, hern 2556 (F); road to Pozuzo, Pearce 213 (K); Pozuzo, Pearce 541 (K); between La Merced and Oxapampa, Saunders 547 (BM); Quillasii, Soukup 3343 (COL); Villa Rica, Woytkowski 7325 (GH, MO), puno: trail between La Oroya and Mina Santo Domingo, Hodge 6025 (CUZ, GH, US, USM). This species is closely allied to the only other two species in the section with lateral inflorescences, F. ovalis and F. macropetala. These inflorescences emerge from leaf axils in the mid-section of the stem and not apically as in most other species (Fig. 40). Fuchsia ovalis differs in its heavily pilose pubescence and long petiolate leaves, whereas F. macropetala is extremely similar to F. macrophylla, differing mainly in its longer flowers and fruits. Both these species grow close by in the Cordillera Azul near the Loreto-Huanuco border, and despite their close similarity, no intermediate forms have been observed. Fuchsia macrophylla is isolated from other species in the section by its low Peru Much eastern slopes of the Andes, and this species was probably much more common until the recent increase in habitat destruction. Fuchsia macropetala Presl, Rel. Haenk. 2:28. 1831. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):557. 1941. type: Peru, Dept. Huanuco, in mountains of Huanuco. 1790, Thaddeus Haenke (PR 495859, holotype; photograph, MO). information 1790 trnen. Erect to scandent shrubs puberulent older branches glabrescent, with finely-splitting, tan brown bark. Leaves oppo- site, firmly membranous, elliptic, mostly acute at the base, acute to shortly acu- minate at the apex, (7-)9-18 cm long, (3-)4-8 cm wide, subglabrous to strigillose purpl 13-18 on either side of the midvein, often reddish, prominent below and anastomosing into 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 a submarginal vein; margin subentire. Petiole subglabrous, 6-16 mm long. Sti- pules lanceolate to triangular, sometimes connate, 1-2 mm long, ca. 0.7 mm wide, mostly deciduous. Flowers few to many in lateral racemes or on short side branchlets 1— 6(— 10) cm long. Pedicels subglabrous, 9-26 mm long, reddish. Ovary ellipsoid, 6-8 mm long, ca. 3 mm thick, light green, finely strigillose and ± ver- rucosa Floral tube subcylindric, 32-45 mm long, 2.5-3 mm wide at the base, slightly constricted to ca. 2 mm wide above the nectary and gradually widened above until 5-7 mm wide at the rim, finely strigillose outside, pilose inside in lower Yi. Sepals lanceolate, 10-13 mm long, 3-5 mm wide, spreading at anthesis. Tube and sepals bright red. Petals red, elliptic-oblong, 10-12 mm long, 4-6 mm wide. Nectary mostly unlobed, 1-1.5 mm high. Filaments red, 7-10 mm and 5-8 mm long; anthers oblong, ca. 2 mm long, ca. 1 mm wide. Style pubescent for most its length, red; stigma capitate, ca. 2 mm long, 2-4 mm wide, slightly 4-cleft apically, dull white to pink, exserted 4-10 mm beyond the anthers. Berry ellipsoid, 15-18 mm long, 7-10 mm thick, dull dark purple; seeds 1.5-2 mm long, dull dark purple; seeds 1.5-2 mm long, 0.9-1.2 mm wide. Distribution: Central Peru. Known only from the Cordillera Azul, which is a semi-isolated Andean spur to the east of the main Andean cordilleras of Peru and which runs along the Huanuco-Loreto border before joining the main cordillera in southern Huanuco. Clearings and roadsides, 1,600-1,800 m (Fig. 56). Specimens examined: Peru, Huanuco: 37 km E of Tingo Maria, near ridge of La Divisoria, Berry & Aronson 3089 (MO, USM), 3090 (MO, USM), 3090-B (MO); La Divisoria, Gentry et al. 18880 (MO); Cuchero, 9°35'S, 75°45'W, Poeppig 1081 (BM, W), s.n. (W 2 sheets); Km 194, La Divisoria, Ridoutt s.n. (Vargas 4654) (CUZ), USM 13004 (MO, USM). loreto: Divisoria, 59 km from Tingo Maria, Allard 21226 (RSA, US), 21297 (RSA, US); just E of crest at La Divisoria, Berry & Aronson 3091 (MO); near Divisoria, Ferreyra 991 (GH, MO, NY, RSA, US, USM); 2244 (MO, US, USM), Smith & Vera 364-H-E (RSA). without locality: Poeppig 156 (G, LE, P); Poeppig s.n. (LE). This is the only species with lateral inflorescences (Fig. 40) and floral tubes more than 32 mm long. It is closely related to F. macrophylla, which shares the same low altitude distribution, lateral inflorescences, and large, elliptic leaves. Fuchsia macrophylla has shorter flowers with green sepal tips and smaller, round- er berries, however. These are the only species of Fuchsia known from the low Cordillera Azul, but it is not known if the two grow together. Further field study will be required to determine how these closely related species are isolated or if any intermediates occur. 15. Fuchsia ovalis Ruiz & Pavon, Fl. Peru v. Chil. 3:87, pi. 324, fig. a. 1802. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):560. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:64, pi. 10, fig. 54. 1943. type: Peru, Dept. Huanuco, near Muiia, 1778-1788, Hipolito Ruiz & Jose Pavon (MA 11/87, lectotype, here designated; photograph, MO). There are also four other Ruiz and Pavon sheets of this species at MA, two at BM, and one each at F, G, MO, and NY, but these lack label information and may not be part of the same collection set. Fuchsia polyanthella I. M. Johnston, Contr. Gray Herb. 75:36. 1925. type: Peru, Dept. Huanuco, Muna, trail to Tambo de Vaca, ca. 2,700 m, 5-7 June 1923, Francis Machride 4290 (F 535372, holotype; photographs, NY, UC; GH, K, isotypes). I982| BERRY— FUCHSIA SECT. FUCHSIA 101 Erect to scandent shrubs 1-3 m tall. Young leaves and shoots mostly densely pilose; older branches pilose to glabrescent, 5-13 mm thick, with light tan, finely fissured bark. Leaves opposite to quaternate, firmly membranous, broadly elliptic to ovate, rounded to acute at the base, acute to short acuminate at the apex, mature leaves 8-16 cm long, 3-7.5 cm wide, dark green and strigose above, paler green and strigose-pilose below, especially along the veins; secondary veins 14-17 on either side of the mid vein, emerging from the midvein at ±90° and becoming gradually incurved towards the margin, anastomosing into a submarginal vein; margin subentire to obscurely denticulate. Petioles pilose, 2-7 cm long, reddish. Stipules lanceolate on young nodes, often triangular, connate, and recurved on older stems, 2-4 mm long, 2-3.5 mm wide. Flowers several to numerous in lateral racemes or panicles; rachis 5-10 cm long; bracts narrowly lanceolate, long acu- 6-12 10-20 cylindrical, 5-7 mm long, ca. 2 mm thick, loosely pilose. Floral tube narrowly funnelform, 18-20 mm long, ca. 2 mm wide and slightly bulbous at the base, gradually widened above until 3-4.5 mm wide at the rim, sparsely pilose outside, densely retrorse villous inside in lower Vi. Sepals lanceolate, acuminate, 10-13 mm long, 3-3.5 mm wide, usually noticeably longer than the petals, with tips forming a point ca. 2 mm long in bud. Tube and sepals red. Petals red, oblong, acute, 5-9 mm long, ca. 3 mm wide. Nectary shallowly 4-lobed, ca. 1.2 mm high. Filaments light red, 5-6 mm and 3-4 mm long; anthers oblong, ca. 2 mm long, ca. 1 mm wide. Style glabrous, reddish; stigma capitate, ca. 3 mm long, ca. 2 mm wide, slightly 4-cleft apically, exserted 2-3 mm beyond the anthers. Berry cylindrical or ellipsoid, 12-13 mm long, ca. 6 mm thick, subtetragonous, red purple; seeds 0.9-1.1 mm long, ca. 0.7 mm wide. Distribution: Central Peru. Restricted to Depts. Huanuco, Pasco, and Junin, rare in cloud forest thickets; 2,000-2,800 m (Fig. 56). Specimens examined: Peru, huanuco: Muria, Lobb 114 (K), Pearce 511 (K), Weberbauer 6721 (F); Chinchao, Ruiz & Pavdn s.n. (BM); Saraypampa, Woytkowski 34193 (F, UC); Huamincha, Woyt- kowski in 1946 (USM). junin: Chuquishuinca, 2 km above Huacapistana, between Tarma and San Ramon, Ferreyra 454 (MO, USM); San Juan, near Huacapistana, Ferreyra 11031 (MO); Vitoc, Mar- tinet 1585 bis (P); above Huacapistana, Sandeman in 1938 (K); without locality: Lobb s.n. (K 2 sheets); Pearce s.n. (BM). This species is easily recognized by its large, broad, long-petiolate leaves and densely pilose pubescence on the leaves and flowers. The leaves, bracts, pubes- cence, and flower size are similar to those of F. pilosa, but it is placed with F. macrophylla and F. macropetala in the same species group because of its lateral inflorescence. Fuchsia ovalis is rare, and the most recent collection is from 1955 in a now deforested area (the Chanchamayo valley). Because of this, its sympatric occur- rence with other species is unknown. 16. Fuchsia pilosa Fielding & Gardner, Sert. PI. t. 27. 1844. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):561. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:65. 1943. type: Peru, Dept. Amazonas, Taulia, S of Molinopampa, 1835-1841, Andrew Mathews 1492 (OXF, holotype; photograph, MO; BM, CGE, K 2 sheets, isotypes). 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia asperifolia K. Krause, Repert. Spec. Nov. Regni Veg. 1:169. 1905. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):547. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:65. 1943. type: Peru, Dept. Amazonas, between Tambos Bagazan and Almirante, E of Chachapoyas on path to Moyobamba, 2700 m, 1904-1921, August Weberbauer 4445 (B, holotype, destroyed in World War II; photo- graph, F). Shrub 1-2 m tall with densely hirsute young growth, the hairs usually whitish, shaggy, erect, 1.0-1.6 mm long; older branches glabrescent, with light tan, finely striated bark. Leaves opposite to quaternate, usually ternate, membranous, nar- rowly to broadly elliptic or oblanceolate, acute to attenuate at the base, acute to acuminate at the apex, 5-16 cm long, 2-7 cm wide, matte green and subglabrous to villous on upper surface, slightly paler and villous below; secondary veins 10-12 on either side of the mid vein, sulcate above and prominent below, margin subentire. Petioles hirsute, 12-60 mm long. Stipules ca. 1 mm thick, prominent, lanceolate to triangular, 3-4 mm long, ca. 2 mm wide, often connate and recurved on lower nodes, persistent. Flowers numerous and crowded at the tips of ter- minal, drooping racemes; rachis 3-20 cm long; bracts narrowly lanceolate, 5-15 mm long. Pedicels slender, 3-5 mm long, ca. 1.5 mm thick. Floral tube narrowly funnelform, 16-20 mm long, slightly bulbous and ca. 2 mm wide at the base, gradually widened above until 4-5 mm wide at the rim, sparsely to moderately hirsute outside, densely retrorse villous inside in lower V2. Sepals lanceolate, long acuminate, 6-8 mm long, 2.5-3 mm wide, with free, divergent tips 2-3 mm long in bud, divergent with recurved tips at anthesis. Tube and sepals bright orange red. Petals orange red, delicate, oblong-elliptic, 6-8 mm long, 3-4 mm wide, recurved at anthesis. Nectary unlobed, ca. 1 mm high. Filaments orange red, 4-5 mm and 2-3 mm long; anthers oblong, 1.5-2 mm long, ca. 1 mm wide, white. Style glabrous, light red; stigma capitate, subtetragonous, 4-cleft apically, 1.5-2 mm long, 2-3 mm wide, dull white, exserted 1-2 mm beyond the anthers. Berry ellipsoid-oblong, 4-sulcate before maturity, 15-19 mm long, 10-12 mm thick, sparsely hirsute, reddish; seeds tan, ca. 1.5 mm long, 0.9-1.0 mm wide. Gametic chromosome number n = 1 1 . Distribution: Northern Peru. Endemic to mid-elevation cloud forest in the mountains E of the Rio Utcubamba in Dept. Amazonas; 1,600-2,400 m (Fig. 56). Specimens examined: Peru, amazonas: 9-1 1 km E of Molinopampa on Chachapoyas-Mendoza road, Berry & Escobar 3618 (MO, USM), 3619 (MO, USM), 3620 (MO, USM); above Pomacocha, Ferreyra 15170 (MO); Tresleras, on Pomacocha- Yambrasbamba trail, Ferreyra 15233 (MO); Almirante, Math- ews 1481 (K), Sandeman 54 (K, OXF); mountains S of Tambo de Ventilla, Pennell 15794 (USM); Mendoza, Woytkowski 8202 (MO), 8325 (US). This is one of several species endemic to Dept. Amazonas in northern Peru. It is most easily recognized by its stiff, hirsute pubescence, broad leaves, and terminal, drooping inflorescence of red orange, short-pedicellate flowers. Be- cause of floral and foliar similarities to F. oralis, it is placed in the F. macrophylla species group, but in its abrupt, terminal inflorescences and thick, persistent stipules it resembles other species such as F. wurdackii and F. glaberrima. Fuchsia pilosa grows together with F. rivularis in cloud forest thickets in Dept. Amazonas. No hybrids or intermediates were found between them, which is at least partly due to differences in habit. Although their basal stems may at times be entangled, F. pilosa persists as a low, upright shrub with dangling inflo- 1982] BERRY— FUCHSIA SECT. FUCHSIA 103 rescences, while F. rivularis is a scandent-climbing species with long branches and subaxillary flowers that are usually found higher above ground than F. pilosa. 17. Fuchsia putumayensis Munz, Proc. Calif. Acad. Sci. IV. 25:62, pi. 8, fig. 52. 1943. type: Colombia, Com. Putumayo, Mocoa, 23 May 1935, Hernando Garcia-Barriga 4639 (US 1593482, holotype; photographs, NY, POM, UC; AAU, COL, isotypes). Shrubs or subshrubs 1-3 m high. Young growth minutely puberulent; branch- lets terete, 2-5 mm thick, green to dull purple; older stems pale tan gray, with finely fissured bark. Leaves opposite, firmly membranous, elliptic, rounded to acute at the base, acute to subacuminate at the apex, 6-15 cm long, 3-7 cm wide, matte green and glabrous above, pale green and subglabrous to finely puberulent below along the subelevated nerves; secondary veins 9-13 on either side of the midvein, impressed above; margin subentire. Petioles 6-15 mm long, puberulent to subglabrous. Stipules lance-deltoid, thick at the base and subulate at the apex, occasionally connate, 1-2 mm long, 0.4-0.8 mm wide, deciduous. Flowers nu- merous in mostly compact, terminal and subterminal racemes; rachis l-4(-10) cm long; bracts lance-linear, 10-25 mm long, ± recurved, often deciduous. Ped- icels slender, divergent, puberulent, 8-25 mm long. Ovary oblong, 4-5 mm long, 1.5-2 mm thick, puberulent, reddish. Floral tube narrowly funnelform, 15-27 mm long, 2-2.5 mm wide at the base, slightly constricted to 1.5-2 mm wide above 4-7 berulent i 6. Sepals lanceolate, acuminate, 8-11 mm long, 3-5 mm wide, with a 1.5-2 mm long tip in bud, divergent at anthesis. Tube and sepals nitid orange to coral red. Petals orange red, delicate, elliptic-oblong, 6-9 mm long, 3^4 mm wide, obtuse or mucronate at the apex, somewhat erose-margined, strongly spreading and recurved at anthesis. Nectary unlobed, 1-1.2 mm high, ca. 0.8 mm thick. Filaments orange red, 4-6 mm and 2.5-4 mm long; anthers elliptic-oblong, 1.5-2.3 mm long, ca. 1 mm wide, cream. Style glabrous, light red; stigma subglobose, slightly 4-lobed apically, cream to pink, exserted shortly above the anthers. Berry oblong-ellipsoid, 10-12 mm long, 5-9 mm thick, subnitid red purple; seeds tan, 1-1.2 mm long. 0.6-0.8 mm wide. Gametic chromosome number n = 1 1 . shrubs Hera Occidental of Colombia from Antioquia to Valle, and on the eastern slopes of the Cordillera Central and Cordillera Oriental in Colombia and Ecuador from Cauca south to Naoo: 1.400-2.100 m (Fie. 58). Specimens examined: Colombia, antioquia: trail Encarnacion-Parque Nacional Las Orquideas, Gentry & Renteria 24607 (MO), cauca: Rio Villalobos, vicinity of Rio Sauzita, Schuttes & Villarreal 5156 (COL, GH), 5173 (F, GH). choco: 53-55 km W of Ansermanuevo on road to San Jose del Palmar, Berry 3561 (COL, MO), 3562 (COL, MO), 3563 (COL, MO); Km 120-135 of Tutunendo-El Carmen road, alto Rio Atrato, Forero et al. 6172 (MO); Quebrada El Sucio, above Carmen de Atrato, Fosberz & Core 21551 (RSA, US); 1 1 km E of San Jose del Palmar, Luteyn et al. 7328 (COL, MO NY), putumayo: Planada de Minchoy, between Sachamates and San Francisco de Sibundoy, Cua- trecasas 11436 (COL, F, US); between Sachamates and San Antonio, Ewan 16684 (COL, F, POM, US); Km 128 from Pasto to Mocoa, Plowman & Davis 4406 (COL, PSO). risaralda: Pueblo Rico, von Sneidern 5464 (F, RSA, S, US), 5546 (COL, MICH, RSA, S, US), valle: k Tokio," above El Queremal, 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Berry 3564 (COL, MO); Quebrada Robada, Alto Bonito, drainage of Rio Alban, Cuatrecasas 22323 (F, RSA, VALLE); Quebrada of Rio San Juan, above El Queremal, Cuatrecasas 23933 (F, RSA); Calima valley, Hugh-Jones 445 (COL, K, US); Langlasse 60 bis (G 2 sheets). Ecuador, napo: Cerro Antisana, forest E of Borja, Grubb et al. 1083 (K, NY); Borja, Hurling 387! (NY, S); 16.5 km NNE of Santa Rosa, road Baeza-Lago Agrio, MacBryde 728 (MO); junction Baeza-Lago Agrio road with Rio Azuela, MacBryde 808 (MO, QCA); 28 km E of Baeza, before Salado, Plowman et al. 3951 (COL, F, GH, MO, S). This species is part of a group with Fuchsia lehmannii, F. andrei, and F. cuatrecasasii that have flowers usually tightly grouped in short terminal inflores- cences with lanceolate or deciduous bracts and ± divergent pedicels. Fuchsia putumayensis has uniformly elliptic, opposite leaves like F. cuatrecasasii, but shorter, orange flowers with somewhat erose petals as in F. lehmannii. Like F. pallescens, this species is disjunct from the western slopes of the Cordillera Occidental in Colombia to the eastern slopes of the Andes in Ecuador and Colombia. Most Ecuadorian populations have longer racemes (ca. 10 cm long) than most collections from Colombia, but occasional Colombian specimens such as Langlasse 60 bis (G), from Valle, also have loose inflorescences. Although F. putumayensis is the lowest elevation fuchsia in the northern Andes and is generally altitudinally separated from other species, a probable hybrid with F. nigricans (Robinson 138, K) was found and is discussed under that species. 18, Fuchsia lehmannii Munz, Proc. Calif. Acad. Sci. IV. 25:61, pi. 9, fig. 51. 1943; Opera Bot., Ser. B, 3: 17. 1974. type: Ecuador, Prov. Morona-Santiago, east Andes of Sigsig, 1,600-1,800 m, 1850-1903, Friedrich C. Lehmann 5498 (F 550994, holotype; photographs, MO, NY, POM, UC; GH, K, US, iso- types). Scandent to erect shrubs 1-3 m tall, mostly with numerous short lateral branches 5-12 cm long near the tips of the main stems. Leaves 2-4 per node, mostly ternate or quaternate, membranous, narrowly (ob-)lanceolate to narrowly elliptic, acute to acuminate at the base, acuminate at the apex, 4-1 1(-15) cm long, l-3.5(-4.2) cm wide, subnitid dark green and glabrous above, pale green and glabrous or pubescent along veins below; secondary veins 8 — 10(— 1 1) on either side of the midvein, impressed above, margin entire. Petioles 4— 1 2(— 20) mm long, subglabrous to strigillose, mostly reddish. Stipules lanceolate, 1.5-2 mm long, 0.4-0.8 mm wide, deciduous. Flowers numerous in compact, terminal and sub- terminal racemes; rachis 0.5-3 cm long, usually ± strigose; bracts narrowly lan- ceolate, acuminate, usually reflexed, 5-15 mm long, 1-4 mm wide. Pedicels slen- der, 8-15 mm long, ± divergent, strigose or glabrous. Ovary oblong, 3-5 mm long, 1.5-2.5 mm wide, subglabrous, mostly wine red. Floral tube narrowly funnelform, 22-35 mm long, 1.5-2.5 mm wide and slightly bulbous at the base, narrowed to 1.2-2 mm wide for the basal Vz, then dilated to 6-9 mm wide before narrowing slightly towards the rim of the tube, glabrous outside, pilose inside in basal Vs. Sepals narrowly lanceolate, 10-13 mm long, 2.5-4 mm wide, generally long acu- minate, 4-angled in bud with a tip 2-3 mm long, strongly divergent to slightly reflexed at anthesis. Tube and sepals nitid orange to coral red. Petals delicate, orange red, narrowly lance-oblong, 8-1 1 mm long, 2.5-4 mm wide, acute at the tip, undulate and spreading-reflexed at anthesis. Nectary unlobed, ca. 1 mm high. 1982] BERRY— FUCHSIA SECT. FUCHSIA 105 Figure 58. Distribution of the Fuchsia putumayensis species group 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Filaments orange red, 5-6 mm and ?>-A mm long; anthers oblong, 1.5-2 mm long, 1.1-1.3 mm wide, cream. Style orange red, mostly glabrous or loosely villous in lower Vi\ stigma capitate, shortly 4-cleft at apex, 2-2.5 mm long, ca. 2 mm wide, dull cream to orange red. Berry oblong-ellipsoid, 11-15 mm long, 7-8 mm thick, turning dark purple at maturity; seeds 0.9-1 . 1 mm long, 0.7-0.8 mm wide, brown. Gametic chromosome number n = 1 1 . Distribution: Southern Ecuador. Moist thickets, streamsides, and roadbanks on the eastern slopes of Prov. Zamora-Chinchipe and Morona-Santiago; 1,600-2,250 m (Fig. 58). ■ Representative specimens examined: Ecuador, morona-santiago: Tambo Chontal to Tambo Con- suelo, slopes above Rios Negro and Chupianza, Seville de Oro trail, Camp E-I589 (NY, RSA). zamora-chinchipe: 27 km E of Loja to Zamora, Berry & Escobar 3200 (MO, QCA); Hacienda Montecristi, ca. 40 km NE of Loja, Espinosa E-146I (US); Canillones, Rio San Francisco, Fosberg 23165 (COL, NY, RSA, US); 5 km W of Tambo, Loja-Zamora road, Marling 5865 (NY, S); Km 39 Loja-Zamora road, Holm-Nielsen et al. 4066 (AAU, COL, MO, NY, S); Km 33 Loja-Zamora road, Holm-Nielsen et al. 4134 (AAU, COL, MO, NY, S); Rio Savonilla, Lehmann 7858 (F, K, US); below El Retorno, Mathias & Taylor 5200 (F, RSA, USM); trail from Mirador to Pailas, Steyermark 54281 (NY); Tambo Valladolid, Steyermark 54654 (NY). This species is most closely allied to F. putumayensis and F. andrei, both species with tight, many-flowered inflorescences of orange red flowers and deli- cate petals. It has narrower leaves than either of these species, however, longer sepals, and floral tubes that are widest below the rim of the tube. Although it has not been found sympatrically with other fuchsias, its altitudinal and geographical ranges overlap with those of F. glaberrima and F. andrei. 19. Fuchsia andrei I. M. Johnston, Contr. Gray Herb. 75:3 1. 1925. Munz, Proc. Calif. Acad. Sci. IV. 25:61, pi. 9, fig. 50. 1943. type: Ecuador or Peru (locality uncertain), Rio de Huarunamaca (as appears on K sheet, transcribed to "Rio de Huannamaca" on the sheets at F and NY), 19 Nov. (?) 1876, Edouard Andre K.820 (F 537131, holotype; photographs, MO, NY, POM, UC; K, NY, isotypes). I have been unable to find the above locality in maps or gazeteers of either Colombia, Ecuador, or Peru, where Andre collected in 1876. Ac- cording to Johnston's protologue, the locality is from Colombia. A detailed list of Andre's itinerary, however, does not mention this river for either Colombia or Ecuador (Smith, 1965). The last part of Andre's itinerary in southern Ecuador and northern Peru was not included in Smith's list, how- ever, and that is when Andre hired H. Poortman to collect for him (L. B. Smith, pers. comm.). Since Andre had already returned to Europe in Sept. 1876, this collection must have been made by Poortman, who collected mostly in Prov. Loja of southern Ecuador. Fuchsia osgoodii J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):559. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:60, pi. 9, fig. 49. 1943. type: Peru, Dept. Libertad, Uchco, 15 June 1912, Wilfred H. Osgood & Mary P. Anderson 47 (F 346773, holotype; photographs, NY, UC). Fuchsia ovalis var. aberrans J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):560. 1941. type: Peru, Dept. Cajamarca, Prov. Cutervo, Arenales, S of San Tomas, 3,000 m, 12 Dec. 1938, Harvey E. Stork & Ovid. B. Horton 10155 (F 1052360, holotype; photograph, NY; G, NA, UC, isotypes). Shrubs 1-4 m tall. Branchlets terete, 3-8 mm thick, sparsely puberulent to strigillose; older stems 8-14 mm thick, with tan, finely striated bark. Leaves 1982] BERRY— FUCHSIA SECT. FUCHSIA 107 opposite or less often ternate, membranous, (narrowly) elliptic to obovate, acute to cuneate at the base, subacuminate at the apex, (5— )7— 17 cm long, 2— 9(— 1 1) cm wide, glabrous above, subglabrous to strigose below on mid vein and along mar- gin; secondary veins (8-) 12-15 on either side of the midvein; margin entire or subdenticulate. Petioles 6-20(-30) mm long. Stipules triangular, dark, 1.5-2 mm long, ca. 1.5 mm wide, subpersistent. Flowers numerous in short, terminal and subterminal racemes; rachis 2-6 cm long; bracts lanceolate, 6-10 mm long, 2-4 mm wide. Pedicels slender, mostly strigose, ± divergent, (7—) 1 0— 1 8(— 25) mm long. Ovary oblong, 5-6 mm long, 2-2.5 mm thick. Floral tube narrowly funnelform, 23-38(-45) mm long, 1.5-2 mm wide and subnodose at the base, narrowed to 1 4-7 glabrous outside, pilose inside in lower Vi. Sepals lanceolate, acute to subacu- minate, (6— )8— 13 mm long, 4-5 mm wide. Tube and sepals orange to coral red. Petals orange red, oblong, obtuse to acute at the apex, 7-12 mm long, 2.5^4 mm wide, nectary unlobed, ca. 1 mm high. Filaments light red, 6-8 mm and 4-6 mm long; anthers oblong, 1.5-2 mm long, 1-1.2 mm wide. Style light red, glabrous to pilose in lower Vi\ stigma capitate, 4-cleft apically, 2-2.5 mm long, ca. 2.5 mm wide. Berry oblong-ellipsoid, 10-12 mm long, 6-8 mm thick, green to red; seeds tan, 1-1.2 mm long, 0.7-0.8 mm wide. Gametic chromosome number n = 11. Distribution: Southern Ecuador and northern Peru. Cloud forest and semi- open moist scrub in Prov. Piura, Cajamarca, and Amazonas, Peru and in Prov. Zamora-Chinchipe, Ecuador (1,1 30-) 1,800-3, 000 m (Fig. 58). Specimens examined: Ecuador, zamora-chinchipe: Tambo de Savanilla, Andre K.8I8 (F, GH, NY). Peru, amazonas: Cordillera Colan SE of La Peca, Barbour 3995 (MO), 4083 (MO); 13 km E of Molinopampa on Chachapoyas-Mendoza road, Berry & Escobar 3621 (MO, USM); Km 42 of Carretera Marginal from Pedro Ruiz to Rioja, Berry & Escobar 3628 (MO, USM); Prov. Bongara, Ferreyra 15170 (MO 2 sheets), 15233 (MO); E side Cerros Calla-Calla, 5 km above Leimebamba. Hutchison & Wright 4884 (UC); Rio Almirante, Sandeman in 1938 (K); Mendoza, Woytkowski 8325 km WSW of Pomacocha, Prov. Bongara, Wurdack 850 (US), cajamarca: above Tabaco- nas, 18 km SE of Huancabamba, Prov. Jaen, Fosberg 27835 (MO); Hacienda Taulis, vicinity of the Casa Hacienda, Prov. Hualgayoc, Hutchison & Bismark 6326 (F, MICH, MO, NY, P, RSA, UC, US); San Andres, Prov. Cutervo, Lopez & Sagas tegui 5436 (MO, US), Sagas tegui in 1969 (NY), Velarde Nunez 6971 (Z); Monte Seco, Prov. Hualgayoc, Soukup 3848 (F, US), piura: Cancahaque, Stork 11428 (UC). It is difficult to adequately circumscribe this species because the specimens 2-A rn Peru When tions and field observations have been made in this area, the material treated below will probably be found to comprise several distinct taxa. They are grouped together here, however, because of their similar, compact racemes and generally short-tubed flowers with slender pedicels. In these characters, the specimens included under F. andrei are related to F. lehmannii and F. putumayensis. The type specimen and Hutchison & Bismark 6326 are characterized by their large leaves ca. 16 cm long and ca. 10 cm wide. This and the somewhat narrower floral tubes distinguish them from F. lehmannii of southern Ecuador. There are a num- ber of collections from other areas, though, that show the following differences: Collections ft arde Nunez ' Horton 10155, though, which Macbride 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 described as F. ovalis var. aberrans, has some larger, basal leaves that approach those of the type of F. andrei. Stork 11428. This collection from Cajamarca has floral tubes 42-45 mm long and is recorded as growing at 1,130 m, which is much lower than the other specimens considered here. The area where it was collected has been severely altered, however, and no further populations were found when I visited the area in 1979. Fosberg 27835. The leaves of this collection are narrowly elliptic as in F. lehmannii. The same is true for Osgood & Anderson 47, the type of F. osgoodii, but it has not been possible to identify the locality of this collection, believed by Macbride to be in Dept. Libertad. Collections from Dept. Amazonas. These are considerably isolated from the specimens to the west of the Rio Maranon, and they occur in wetter, cloud forest habitats. Wurdack 850 and Hutchison & Wright 4884 have short tubes just over 20 mm long and small, elliptic leaves with subdentate margins. Woytkowski 8325 has broad leaves ca. 7 cm wide but similarly short flowers. Ferreyra 15233 has floral tubes 45 mm long. These wide differences suggest that several different entities might be involved in these collections, but further material is needed to determine their limits. 20. Fuchsia cuatrecasasii Munz, Proc. Calif. Acad. Sci. IV. 25:51, pi. 7, fig. 40. 1943; non Munz, Opera Bot., Ser. B, 3:14. 1974. type: Colombia, Com. Caqueta, Cordillera Oriental, E slopes, Quebrada del Rio Hacha, open woods in Cajon de Pulido, 1,700 m, 26 March 1940, Jose Cuatrecasas 8738 (US 1796401, holotype; photographs, MO, NY, POM, UC; COL, isotype). Subshrubs 0.5-1.5 m high. Branchlets terete to slightly flattened, 2-6 mm thick, subglabrous, green to reddish purple; older branches with light tan-gray bark. Leaves opposite, elliptic, firmly membranous, acute at both ends, smooth and glossy above, subglossy and finely strigillose below on veins and margins; secondary veins 8-12 on either side of the midvein, ± impressed above, at times purple tinged below; margin subentire. Petioles stout, sparsely strigose, 4— 14(— 17) mm long, red purple. Stipules triangular, firm, dark purple, 1-2 mm long, ca. 1.5 mm wide, subpersistent. Flowers few to many in a compact, terminal, drooping raceme; rachis l-5(-8) cm long; bracts lanceolate, 8-18 mm long, reflexed. Ped- icels 7-13 mm long. Ovary oblong, 4-7 mm long, 1.5-2.5 mm thick, strigillose, subnitid red purple. Floral tube narrowly funnelform, (28-)35-50(-55) mm long, slightly nodose and 2-3.5 mm wide at the base, narrowed to 1.5-3 mm wide in lower l /3 and widened above until 6-8 mm wide at the rim, puberulent outside, pubescent inside in lower Vs. Sepals lanceolate, acuminate, (8-) 10-20 mm long, 4-6 mm wide, tips usually free in bud for 2-4 mm. Tube and sepals nitid red to orange red. Petals orange red, elliptic to oblanceolate, 9-15 mm long, 4-6 mm wide, acute at the apex, spreading at anthesis. Nectary unlobed, ca. 1.5 mm high. Filaments orange red, 5-9 mm and 4-5 mm long; anthers oblong, 2-3 mm long, 1-1.5 mm wide. Style sparsely villous in lower Vi, orange red; stigma capitate, tetragonous, 4-lobed in upper Vi, 2-3 mm long, 2-3 mm wide, cream, exserted 1-4 mm beyond the anthers. Berry ellipsoid, 15-20 mm long, 10-15 mm thick, lustrous red purple; seeds 1.1-1.3 mm long, 0.6-0.8 mm wide. 1982] BERRY— FUCHSIA SECT. FUCHSIA 109 Distribution: Colombia. Infrequent, low shrubs in moist, open thickets in cloud forest on eastern slopes of the Andes from Caqueta south to Putumayo, 1 ,400-2,200 m (Fig. 58). Specimens examined: Colombia, cauca: Rio Villalobos, between the junction of the Rios Villalobos and Cauchos, Schultes & Villarreal 5203 (COL, DS, F, GH, MO, UC, US), huila: Km 31 of Pitalito- Mocoa road, Berry 3594 (COL, MO); Km 28-32 of Pitalito-Mocoa road, near divide, Luteyn et al. 7538 (COL, MO, NY), putumayo: Mirador, between San Francisco and Mocoa, Bristol 542 (COL, DS); between El Silencio and La Cabana, road from Sibundoy to Urcusique, Cuatrecasas 11495 (COL, US); Cordillera del Portachuelo, road Sibundoy-Mocoa, Garcia-Barriga et al. 18635 (COL); El Mirador, San Francisco-Mocoa, Idrobo & Ospina 2364 (COL); Km 90-91 between Sibundoy and Mocoa, Luteyn et al. 5043 (COL, MO, NY); San Francisco to Mocoa, Mora 3440 (PSO), 4460 (COL), Schultes & Cabrera 18625 (RSA); Km 112 from Pasto to Mocoa, Plowman & Davis 4348 (COL, PSO); Punto Buenos Aires, Cerro Portachuelo, Soejarto 1125 (GH, PSO). The short, terminal racemes of tightly packed flowers in F. cuatrecasasii is of the same type found in F. putumayensis, F. lehmannii, and F. andrei. Its flowers are considerably longer than these species, however, and the leaves are uniformly elliptic, opposite, glossy, firm, and short-petiolate. Vegetatively, F. putumayensis is the most closely related species. Fuchsia cuatrecasasii occurs sympatrically with F. putumayensis, F. scabriuscula, F. sessilifolia, and F. ver- rucosa, but it is rare, and no apparent hybrids have been detected. M. Johnston. Contr. Grav Herb. 75:37. 1925. Macbr Mus. Nat. Hist., Bot. Ser. 13(4):545. 1941. Munz fig Macbr 4541 (F 535618 types). Fig. 20. POM Fuchsia aspiazui J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):547. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:43, pi. 6, fig. 31. 1943. type: Peru, Dept. Libertad, Prov. Pataz, valley of the Rio Mixiollo, Aug. 1914, August Weberbauer 7042 (F 629319, holotype; photographs, NY, UC; F 2 sheets, GH, US, isotypes). Scandent to erect shrubs 1-3 m tall. Young branches semisucculent, terete, 3-6 mm thick, subglabrous to densely hispid; older stems 8-15 mm thick, light tan with finely fissured bark. Leaves opposite or ternate, firmly membranous, narrowly to broadly elliptic or (ob-)ovate, acute at the base, acute to acuminate at the apex, 7- 1 4(- 1 9) cm long, 3-6(-10) cm wide, subnitid dark green and sparse- ly strigose to subglabrous above, pale and strigose to densely villous or hispid below, especially along the veins; secondary veins 15-22 on either side of the midvein, strongly impressed above, anastomosing into a distinct submarginal vein; margin subentire to obscurely denticulate. Petioles strigose, 4-10 mm long. Sti- pules conspicuous, dark, (narrowly) triangular, sharply acute-tipped, 3-6 mm long, 2-3 mm wide, often connate, persistent. Flowers few to numerous and 10-35 fruit; bracts lanceolate, divergent, 15^45 is and 10-15 mm long, divergent and asce 15-35 mm long, light red. Ovary cylindric, 7-9 mm long, ca. 2 mm thick, lustrous light red, the ovary or floral tube usually somewhat deflexed so that the distal portion of the flower is pendant. Floral tube narrowly funnelform, 35-50(-60) mm 1 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 long, 2.5-3.5 mm wide and bulbous at the base, constricted to ca. 2 mm wide for the basal 5-8 mm of the tube, gradually widened above until 6-10 mm wide at the rim, glabrous outside, densely pilose inside in basal X A. Sepals lanceolate, 13-20 mm long, 4-7 mm wide, mostly long acuminate at the apex, cuspidate and tetragonous in transection in bud, conspicuously wider than the rim of the tube, divergent at anthesis. Tube and sepals nitid, bright orange red. Petals orange red, delicate, oblong, round to acute at the apex, 13-19 mm long, 5-7 mm wide, spreading to divergent at anthesis with the apical portion recurved and the mar- gins undulate. Nectary green, shallowly 4-lobed, 2.5-3 mm high and ca. 0.7 mm thick. Filaments orange red, 8-10 mm and 6-7 mm long; anthers oblong, 2.5-3 mm long, ca. 1.5 mm wide, white. Style pale red, glabrous; stigma capitate, 2-3 mm long, 2-3 mm wide, 4-cleft apically, pale red. Berry cylindrical-ellipsoid, 18-22 mm long, ca. 10 mm thick, purplish; seeds tan, 1.1-1.4 mm long, 0.8-1.0 mm wide. Gametic chromosome number n = 11. Distribution: Central Peru. Scattered shrubs in cloud forest thickets on the eastern slopes of the Andes from Depts. Libertad to Junin, 1,500-2,700 m (Fig. 58). Representative specimens examined: Peru, huanuco: Carpish, Ferreyra 1719 (MO, RSA, US, USM); Carpish, above Acomayo, Hutchison et al. 5920 (F, NY, RSA, UC, USM); above Chinchao towards summit of Carpish, Mathias & Taylor 4033 (F, RSA, USM); Tumanga, Woytkowski 7951 (US), junin: 57-58 km above Satipo on road to Conception, Berry & Aronson 3077 (MO, USM), 3078 (MO); Km 175 Satipo-Concepcion, Seibert 2379 (MO, POM, US), pasco: Pozuzo, Pearce 537 (K). Fuchsia abrupta is distinguished by its noticeably veiny leaves (due to the numerous, impressed secondary veins), large, persistent stipules, and long, ter- minal inflorescences with divergent to ascending pedicels and cylindrical fruits. Its affinities are unclear, but despite its elongate racemes, it is placed in the F. putumayensis species group because of its divergent pedicels, orange flowers with delicate petals, and narrowly lanceolate bracts. In dried specimens, it may be confused with F. corymbiflora, but that species has finely puberulent leaves, rounder fruits, and shorter racemes and pedicels. The specimen described as F. aspiazui has non-apiculate buds and unusually large leaves, but it has the same characteristic inflorescence and vegetative traits of F. abrupta. Sandeman 4503 (K; Peru, Dept. Junin above Huacapistana, ca. 2,000 m, Oct. 1943) is like F. abrupta in all characters except its much shorter floral tubes (17 mm long). Sandeman 4585 (K, OXF) from the same locality, is morphologically intermediate and is probably a hybrid with F. decussata. It is discussed further under that species. Fuchsia abrupta also occurs sympatrically corymbiflora, F. denticulata, and F. ft 104 Munz, Proc. Calif. Acad. Sci. IV. 25:34. 1943. Fuchsia petiolaris var. typica Munz, Proc. Calif. Acad. Sci. IV. 25:35, pL 4, fig. 21. 1943. Based on F. petiolaris HBK. type: Colombia, Dept. Cundinamarca, near Bogota, ca. 2,500 m, July-Sept. Humboldt che, MO). Fuchsia quinduensis Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6:105. 1823. type: Colombia, Dept. Tolima or Quindio, in the Quindio Andes, ca. 2,500 m, Sept. -Dec. 1801, Alexander von Humboldt & Aime Bonpland (P, holotype, not seen; photograph, F; microfiche, MO). 1982) BERRY— FUCHSIA SECT. FUCHSIA 1 1 1 Fuchsia curviftora Bentham, PI. Hartw. 177. 1845. type: Colombia, Dept. Cundinamarca, in Andes near Bogota, ca. 3,000 m, 1841-1843, Theodor Hartweg (K, holotype). Fuchsia smithii Munz, Proc. Calif. Acad. Sci. IV. 25:36, pi. 4, fig. 21. 1943. type: Colombia, Dept. Santander, vicinity of Vetas, 3,200-3,250 m, 16-20 Jan. 1927, Ellsworth Killip & Albert C. Smith 17300 (GH, holotype; photograph, UC; A, F, NY, US, isotypes). Fuchsia petiolaris var. bolivarensis Munz, Proc. Calif. Acad. Sci. IV. 25:35, pi. 4, fig. 22. 1943. type: Colombia, Dept. Antioquia ("Bolivar" on Pennell's label), below Paramo de Chaquiro, Cordillera Occidental, 2,800-3,100 m, 24 Feb. 1918, Francis W. Pennell 4324 (NY, holotype; photographs, GH, POM, UC). The leaves of this collection are quite slender, but are within the range of those of F. petiolaris. Low shrubs 0.5-2 m tall or climbing-scandent in trees to 5 m above ground. Young growth puberulent to pilulose; branchlets terete, 1.5-3 mm thick, puber- ulent to ferrugineous-pilose, green to light red purple; older branches 5-12 mm thick, with reddish tan, exfoliating bark. Leaves mostly ternate, up to 6 per whorl, firmly membranous to subcoriaceous, narrowly lanceolate to elliptic or obovate, acute to narrowly cuneate at the base, mostly acute at the apex, (1.5-)3-9(-ll) cm long, (0. 5-) 1—3(^4.5) cm wide, medium to dark matte green and glabrous to puberulent above, pale green and subglabrous to puberulent or pilose below; secondary veins (4-)6- 1 0(- 1 1 ) on either side of the midvein; margin denticulate to serrulate or rarely subentire. Petioles (2-)5-20 mm long. Stipules lance-linear, 2-3 mm long, ca. 0.5 mm wide, subpersistent. Flowers few to many, axillary. Pedicels pendant, (10-)15^0(-55) mm long. Ovary ovoid, 5-8 mm long, 2-2.5 mm thick. Floral tube narrowly funnelform until the slightly constricted apex, (30-)35-50(-62) mm long, 2.5^4(-6) mm wide and somewhat bulbous at the base, narrowed to 1.5— 3(— 5) mm wide in the lower Vs of the tube, then gradually to rather abruptly widened until (5-)6-ll(-12) mm wide at the rim, strigillose to puberulent outside, pilose inside for most of length. Sepals ovate-lanceolate, con- spicuously flared outwards from the tube at the base and 1-3 mm wider in bud than the rim of the tube, acute to acuminate, 11-23 mm long, (4-)6-8(-9) mm wide, with coherent or spreading tips in bud and the lobes sometimes splitting open in the middle before anthesis, spreading-divergent at anthesis, at times the basal Vi divergent and the upper Vi erect. Tube dull to subnitid rose pink, sepals usually paler, sometimes with greenish tips. Petals bright red pink, narrowly lanceolate to lance-elliptic, (narrowly) acute to rarely obtuse at the apex, (8— )12— 16(— 20) mm long, (4-)5-7(-8) mm wide, generally unguiculate at the base, margin smooth to occasionally serrulate, mostly with finely pubescent hairs over the dorsal surface or longer villous hairs along the midvein, rarely (sub-)glabrous, suberect at anthesis. Nectary green, unlobed or 4-lobed, 2-A mm high. Filaments pink, 8-15 mm and 5-12 mm long; anthers oblong, 2.5-3 mm long, 1.5-2 mm wide, dull white. Style densely pilose in basal l /2- 3 A, pink; stigma globose, 2-4 15 mm beyond the anthers. Berry globose to ellipsoid, 8-18 mm long, 7-13 mm thick, dark red at maturity; seeds tan, 2-3 mm long, 1-1.5 mm wide. Gametic chro- mosome number n — 1 1 . Distribution: Colombia and Venezuela. Low shrubs or lianas in thickets of high elevation cloud forest or in paramo shrub islands; in the Cordillera Oriental from Huila to Norte de Santander and just into Venezuela in southern Tachira; in the Cordillera Central from northern Valle to Antioquia; and in the Cordillera Occidental in Antioquia; (2,400-) 2, 900-3 ,900 m (Fig. 59). 1 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Representative specimens examined: Venezuela, tachira: between Fatima and San Vicente de la Revancha, Fernandez 1981 (MY); headwaters of the Rio Quinimari, above "Las Copas" below the Paramo de Judio, 18-20 km S of San Vicente de la Revancha, Steyermark et al. 100795 (VEN). Colombia, antioquia: Paramo Frontino, near Llano Grande, Boeke 250 (MO); summit of Morro Pelado, Anocosca-Abriaqui road, Core 456 (RSA, US); Paramo de Sonson, Guarin 3470 (COL, US). boyaca: Quebrada de San Paulino, near Alto Ritacuva, Sierra Nevada del Cocuy , Barclay & Juajibioy 7279 (RSA); Paramos NW of Belen, headwaters of Quebrada Minas, Cleef 1854 (U); between Soata and Cocuy, Valle de la Uvita, El Hatico, Cuatrecasas & Garcia-Barriga 1 161 (COL, F, US); between Soata and Cocuy, Paramo del alto de Canultal, Cuatrecasas & Garcia-Barriga 1 183 (COL, F, US); Quebrada de Susacon, Cuatrecasas & Garcia-Barriga 9815 (COL, POM, US); above Guican, Sierra Nevada de Cocuy, Grubb et al. 70 (COL, K, MSC, US); near Laguna Seca, Sierra Nevada de Cocuy, Grubb et al. 602 (COL, K, MSC, US); Ramiriqui, towards Laguna Negra, Huertas & Camargo 6324 (COL); near Montalban, road Duitama-Charala, Langenheim 3470 (COL, UC, US); Laguna la Col- orada, Paramo de Pisba, Rangel et al. 525 (COL); La Rusia, NW of Duitama, Uribe-Uribe 1075 (COL, US), caldas: Paramo de Las Letras, Barclay & Juajibioy 6293 (COL, MO, RSA); SW slopes of Nevado de Ruiz, Termales, Cuatrecasas 9218 (COL, F, US); Paramo de Ruiz, Lehmann 3067 (BM, K, US); 35 km NE of Manizales, Madison 1335 (GH); San Felix, Tomds 1866 (BH). cundi- namarca: Subachoque to Cerro El Tablazo, Berry 3539 (COL, MO); below Paramo de Choachi, Berry 3542 (COL, MO); ridge between Fomeque and Laguna de Chingaza, Cuatro Vientos, Cuatre- casas & Idrobo 26973 (US); Paramo de Chingaza, around Laguna Grande, Cuatrecasas & Idrobo 26992 (US); Rio San Juan, above San Juan, Fosberg 20797 (RSA, US); Paramo de Sumapaz, Rio Arroz, Fosberg 20835 (US); near Bogota, Hartweg 990 (BM 2 sheets, BREM, CGE, K, OXF); Guadalupe, Bogota, Haught 5061 (POM, US), 5621 (COL, POM, US); Monserrate, Rio de San Francisco, near Bogota, Hawkes & Garcia-Barriga 69 (BM, COL, K, RSA, US); Paramo de Cruz Verde, near Bogota, Pennell 2056 (NY); Rio San Cristobal, near Bogota, Pennell 2379 (GH, MO, NY, US); Boqueron de Chipaque, Schneider 621a (COL); Montserrate, Triana 198 (K, P, US); La Calera, Paramo de Palacio, Uribe-Uribe 6886 (COL); 20 km SE of Gigante, Little 8686 (US), meta: Rio Arroz, well above confluence of Quebrada Pedregal, Fosberg 20916 (US), norte de santander: 7 km above Pamplona to Bucaramanga, Berry 3533 (MO); Chitaga, near Presidente, Cuatrecasas & Garcia-Barriga 10053 (COL, F); Pamplona, Funck & Schlim 1646 (BM, CGE, F, G, LE, OXF, P, W); 20 km S of Abrego, Las Jurisdicciones, Cerro de Oroque, border with Dept. Cesar, Garcia- Barriga & Jaramillo 19758 (COL), 19808 (COL); around Ocana, Kalbreyer426 (K); between Mutiscua and Pamplona, Killip & Smith 19734 (A, GH, NY, US); W slopes Paramo del Hatico, Toledo to Pamplona, Killip & Smith 20714 (GH, NY, US); Pica-pica valley, above Tapata, N of Toledo, Killip & Smith 21121 (GH, NY), quindio: Alaska, above Salento, Pennell 9381 (PH); Paramo del Quindio, Pennell & Hazen 10058 (NY, PH). risaralda: Campoalegre-San Ramon ridge, E of Santa Rosa de Cabal, St. John 20850 (COL, NY, RSA, US), santander: Paramo de la Cruces, Funck & Schlim 1307 (BM, G, LE, P, W); La Baja, Funck & Schlim 1305 (BM, G 2 sheets, LE, OXF, W); edge of Paramo Las Vegas, Killip & Smith 15734 (A, F, NY, US); between California and Las Vegas, Killip & Smith 17236 (A, GH, NY, US); vicinity of Vetas, Killip & Smith 17335 (A, GH, US), 17361 (A, GH, NY, US), 17386 (A, GH, NY, US); W slopes of Paramo Rico, Killip & Smith 17755 (A, F, GH, NY, US), 17771 (A, GH, NY, US); Paramo de las Puentes, above La Baja, Killip & Smith 18211 (A, GH, NY, US); Paramo de Romeral, Killip & Smith 18577 (A, GH, NY, US), tolima: 47^8 km W of Fresno to Manizales, Berry 3558 (COL, MO), 3559 (COL, MO); SE slopes Mt. Tolima, Las Mesetas, Cuatrecasas 2615 (MA 2 sheets); Quindio-La Linea, Dryander 1939 (US); Volcancitos, Holton 896 (G, GH, K, NY, UC); along divide near Quindio highway, Killip & Varela 34600 (BM, COL, POM, US); Quindio, Humboldt & Bonpland s.n. (P); Rosalito, near Paramo de Ruiz, Pennell 3121 (F, GH, MO, NY, US), valle: Paramo de Bavaya, Barragan, drainage of Rio Bugalagrande, Cuatrecasas 20042 (F, US, VALLE); Quebrada de Las Vegas paramo, headwaters of Rio Tulua, Cuatrecasas 20249 (F, VALLE). This is one of the polymorphic, long-tubed, axillary-flowered species of the Fuchsia petiolaris species group that are found at high altitudes in the northern Andes. Its main distinguishing characteristics are the slightly constricted floral tubes at the rim (in most cases), the typically ovate-lanceolate sepals that flare out conspicuously at the base from the rim of the floral tube, and the acute, elliptic to lanceolate petals that are usually puberulent or villous on the dorsal surface. The petals are always smaller than the sepals and in some populations are less than half as long. Its closest relative is probably F. corollata, which occurs south of the range of F. petiolaris in the Cordillera Central. A list of their 1982] BERRY— FUCHSIA SECT. FUCHSIA 113 Table 10. Distinguishing characters of Fuchsia petiolaris and F. corollata. F. petiolaris F. corollata Leaf texture (when dry) Smooth Slightly bullate Leaf surface (when dry) Matte Glossy Length of glandular teeth on margin 0.1-0.3 mm 0.3-0.5 mm Floral tube constricted at rim Usually No Floral tube pubescence Puberulent Strigose Petal shape Lanceolate-elliptic Rhombic-broadly elliptic Petal apex Acute Obtuse to cuneate Petal pubescence (dorsal side) Puberulent to villous, rarely glabrous Glabrous Style pubescence Densely pilose in basal Glabrous to occasionally pi- V2-Y4 lose distinguishing characters is given in Table 10. The possible affinities of F. petio- laris outside of its species group are with F. gehrigeri, which commonly has pubescent petals, and similarly large seeds and globose fruits. Characters that vary widely in F. petiolaris are leaf size and shape, floral dimensions, pubescence type on stems and petals, and the size and shape of the berries. Most of the size differences can be attributed to ecological factors. Short petiolate, small-leaved and short-flowered plants typically are found in high, exposed habitats or in trees near open paramo. Well shaded plants in lower, more protected sites are usually more robust, with longer pedicels, and larger leaves and flowers. Geographical patterns of variation exist in characters such as pubescence and leaf shape, but individuals typical of one area can always be found in the other areas as well. In view of this wide degree of local and regional polymorphism, any attempt to subdivide this group taxonomically seems unwarranted. Some of the patterns of variation are described in the following paragraphs. Most collections from the northern part of the Cordillera Oriental of Colombia (Boyaca to Norte de Santander) and just over the border into Tachira, Venezuela have short pilose, ferrugineous stem pubescence. Such plants were treated by Munz as Fuchsia smithii, and these also differ from most populations from Cun- dinamarca and the Cordillera Central in their larger, narrower, more reticulate- veined leaves, petals villous mostly along the midvein, and larger, more cylin- drical fruits. Variation within this area can also be considerable, however. Grubb et al. 602 (COL, K, US) has floral tubes 52 mm long but only 5 mm wide; Cuatrecasas & Garcia-Barriga 10053 (COL, F) has the largest flowers in the species, with tubes 62 mm long and 1 1 mm wide. Its leaves are broad and nearly entire, unlike most collections from this area. Some plants of the type described above reach south to Cundinamarca, such as Cuatrecasas & Idrobo 26973 (US). Plants from the Bogota area are especially variable, however, but for the most part have fine, canescent vegetative pubes- cence and round fruits. The degree of petal pubescence is exceedingly variable between populations around Bogota, with some puberulent over the entire adaxial (BM POM 1 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 imens of F. petiolaris and F. curviflora are both from the vicinity of Bogota. The slight curvature of the tubes in F. curviflora is not unusual, and though the sepals of the dried specimen are pale, they are by no means white as the description states. Fuchsia petiolaris is widespread in the Nevado de Ruiz Massif of the Cordi- llera Central, following the same distribution pattern as F. venusta, F. hirtella, and F. nigricans, all of which are present in both the Cordillera Oriental and the Cordillera Central. Many individuals with small leaves and flowers are found in this area; these plants are often found in trees or in exposed paramo shrub islands where stunting is a common response to the harsh climatic conditions. Fuchsia quinduensis was apparently described from one such stunted plant, and Sande- man 5672 (K, OXF) is a particularly good example of leaf reduction in an exposed plant. Sometimes large, petiolate leaves can be found on the basal portions of specimens that have nearly sessile, tightly grouped small leaves on the upper branches. Sepals on plants from the Cordillera Central are usually more flared at the base than ones from the Cordillera Oriental, and they sometimes split open in bud in the middle before the tips become separated. Petals are puberulent over the entire surface and are mostly narrowly lanceolate and much smaller than the sepals. The fruits are generally small and globose as in most Cundinamarca col- lections, but the flowers are usually puberulent, and the stems are sometimes ferrugineous as in many Santander collections. Fuchsia petiolaris has also reached the high northern peaks of the Cordillera Occidental in Antioquia. Although Boeke 256 (MO), from Paramo Frontino, has long, slender flowers (51-55 mm long, 6-8 mm wide), other characters such as petal shape and pubescence are well within the range of F. petiolaris. Since Fuchsia petiolaris is the highest altitude species in its range, it usually occurs alone, but in the Cordillera Central it is occasionally sympatric with F. hirtella and F. hartwegii. It would be of considerable interest to examine more populations of the Cordillera Central between Valle and Cauca, to see if transi- tional populations occur between the closely related F. petiolaris and F. corol- la ta . 23. Fuchsia corollata Bentham, PI. Hartw. 179. 1845. type: Colombia, Dept. Cauca, near Popayan in woods of Purace, ascent to Paramo de Guanacas, ca. 3,000 m, 1842, Theodor Hartweg 993 (K Bentham Herb., holotype; pho- tograph, MO; BM, BREM, CGE 2 sheets, F, G, K Hooker Herb., LE, OXF, P, W 2 sheets, isotypes). Fuchsia canescens sensu Munz, Proc. Calif. Acad. Sci. IV. 25:26. 1943; Opera Bot., Ser. B, 3:12. 1974. Fuchsia colombiana Munz, Caldasia 4:109, UQ.fig. 1946. type: Colombia, Dept. Cauca, Cordillera Central, W slopes, headwaters of Rio Palo, paramos between la Quebrada del Duende and Las Casitas, 3,500-3,600 m, 3 Dec. 1944, Jose Cuatrecasas 18959 (A, holotype; COL, isotype). Erect to scandent shrubs 0.5-5 m tall with ascending to divergent branches. Branchlets terete, 2-3 mm thick, strigose, generally red purple; older stems 5-12 mm thick with red tan, exfoliating bark. Leaves ternate or less often quaternate, firmly membranous to subcoriaceous, elliptic to oblanceolate, acute to narrowly cuneate at the base, acute to obtuse at the apex, 20-70 mm long, 7-30 mm 1982] BERRY— FUCHSIA SECT. FUCHSIA 115 Figure 59. Distribution of the Fuchsia petiolaris species group wide, dark glossy green and glabrous to strigose above, pale green and usually strigose or villous below; secondary veins 3-8 on either side of the midvein, often red below; margin gland-serrulate, the teeth usually conspicuous and mostly 0.3-0.5 mm long. Petioles strigose, 3— 12(— 22) mm long. Stipules lance-linear, dark, 2-4 mm long, 0.7-1 mm wide, persistent and usually divergent. Flowers axillary and pendant, usually fewer than 6 per branch. Pedicels 12-24 mm long. Ovary ovoid- ellipsoid, 6-7 mm long, 2-2.5 mm thick, subglabrous to strigose. Floral tubes narrowly funnelform, (28-)35-55(-60) mm long, 2.5-3 mm wide and bulbous at base, then narrowed to 1.5-2 mm and gradually widened above until 5-8 mm wide at rim, subglabrous to strigose outside, villous inside in lower Vi. Sepals lanceolate to ovate-lanceolate, subacuminate at apex, 10— 17(— 22) mm long, 5-6(-8) mm wide, sometimes split open in middle in bud, spreading to divergent at an- 1 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 thesis. Tube subnitid red pink to scarlet; sepals paler, often greenish towards tips. Petals scarlet, almost always longer than sepals, (broadly) elliptic to rhom- 10-18C )6-10 spreading at anthesis. Nectary unlobed or shallowly 4-lobed, 1.5-2 mm high, sometimes with a few erect hairs. Filaments red pink, 10-18 mm and 7-15 mm long; anthers oblong to subreniform, 2-3 mm long, 1.5-2.2 mm wide, cream. Style red, glabrous or occasionally pilose; stigma globose, slightly 4-cleft at apex, mm wide, cream to light red. Berry subglobose, 9-13 mm long, 2-3 mm long, 2-4 mm wide, cream to light red. Berry subglobose, 9- 8-10 mm thick, purple at maturity; seeds tan, 2-2.6(-3) mm long, (1.0-) 1.2-1. 6(-2) mm wide. Gametic chromosome number n = 11, 22. Distribution: Southern Colombia and northern Ecuador; high elevation cloud shrubs 800-3 .800 Representative specimens: Colombia, cauca: Pilimbala, N slopes Volcan Purace, Aguirre 311 (COL); 1 1 km E of Totoro to Inza, Berry 3575 (COL, MO); 4 km W of Gabriel Lopez, road Totoro-Inza, Berry 3576 (COL, MO); 17 km E of Purace, Berry 3582 (MO); headwaters of Rio Palo, Quebrada del Duende, Cuatrecasas I893I (BH, F, GH, VALLE); Finca Las Mercedes, near Silvia, Granger & Rodriguez 17 (RSA, US); Manchai, NE of Silvia, Haught 5122 (COL, POM, US); Paramo Las Delicias, Lehmann B.T.1058 (GH, K, NY); Paletara, Pennell 7004 (GH, PH); Paramo de Buena Vista, Huila group, Pittier 1 188 (US), narino: between PastoandTiiquerres, Andre 3 182 (K); "EI Paramo/ 1 Km 15 of Pasto-Ipiales road, Berry 3146 (MO); Paramo El Tabano, road Pasto-Laguna La Cocha, Berry 3249 (MO, PSO); 18 km above Pasto, road to Tangua, Ewan 15918 (BM, DS, POM, US); Laguna La Cocha, Ewan 16381 (POM, US); road Tiiquerres-Ipiales, Garcia-Barriga & Hawkes 13073 (COL, US); road Ipiales-La Victoria, Paramo de la Cortadera, Garcia-Barriga & Hawkes 13091A (COL); Volcan de Sotara, Lehmann 6196 (K); trail from Sapuyes to Paramo de Gualaratan, Mora 2901 (PSO); Pasto, base of Volcan Galeras, above Obonuco, Schultes & Villarreal 8023 (COL, RSA, US); Tangua, Cubijan trail, Uribe-Uribe 5302 (COL, PSO, US). Ecuador, carchi: Tulcan-El Car- melo road, 7 km E of Panamerican Highway, Berry 3158 (MO), 3159 (MO); between El Carmelo and Santa Barbara, Berry 3165 (MO); road Tulcan-El' Pun (= El Carmelo), Mexia 7584 (BH, K, RSA, UC, US), imbabura: Hacienda Curubi, 10 km W of Otavalo towards Mojanda, Berry 3173 (MO, QCA); 11 km W of Otavalo to Laguna de Mojanda, Berry 3175 (MO, QCA); Mojanda, ca. 10 km SSW of Otavalo, Sparre 13523 (S). Probable hybrids between Fuchsia corollata and F. caucana: Colombia, cauca: Paramo de Las Papas, near Laguna Cusiyaco, Barclay & Juajibioy 5936 (COL, RSA); Volcan de Purace, Barkley et al. I8ca.ll2 (US); 45 km from Popaydn on road to Volcan Purace, Barkley & Mullen 38C712 (COL, GH); 23-35 km E of Purace, road to La Plata, Berry 3583 (COL), 3584 (MO), 3585 (COL, MO); Paramo de Purace, S of Volcan Purace along divide, San Francisco, Cuatrecasas 14644 (BH, F, VALLE); Paramo de Juntas, Km 53 (extension of Paramo de Guanacas), Cuatrecasas & Willard 26375 (COL); W side of Paramo de Purace, Killip & Lehmann 38602 (BH, US); Paramo de Las Papas, between El Boqueron and La Hoyola, Idrobo et al. 3935 (COL); road Popayan to Purace, Parque Nacional Purace, Lozano & Ruiz 1518 (COL); Parque Nacional Purace, Plowman et al. 5335 (COL), von Sneidern 1843 (S), 1847 (S), 1848 (S), Uribe-Uribe 3845 (COL, MO), huila: 35-40 km E of Purace on road to La Plata, Berry 3586 (COL, MO), 3588 (COL, MO). This is one of several closely allied and polymorphic species in the Fuchsia petiolaris species group from high elevations in the Colombian and Ecuadorian Andes. Its most characteristic features are the elliptic-rhombic petals usually exceeding the size of the sepals and the glossy, firm, gland-denticulate leaves that often dry with a slightly bullate texture. Its closest relative is F. petiolaris, which occurs in the Cordillera Oriental and Cordillera Central just north of the range of F. corollata. Since these species are widespread and variable, clear recognition of the species limits between them can be difficult, especially if one is dealing 1982] BERRY— FUCHSIA SECT. FUCHSIA 117 only with herbarium specimens. Furthermore, we do not yet know if the two species intergrade in southern Valle Dept. or northern Cauca, where their ranges are in close proximity. Based on the existing specimens, Table 10 presents a summary of characters that can be used to distinguish F. corollata from F. pet- iolaris. It must be noted, however, that hybrids of F. corollata with other fuchsias are apparently widespread in southern Colombia, and these individuals would generally fall outside the limits designated for the above species. Munz (1943, 1974) mistakenly synonymized this species under Fuchsia ca- nescens because of an erroneously numbered Hartweg collection from Geneva Hartweg but the specimen at G does not correspond to the holotype at K. It is rather a duplicate of Hartweg 993, the type collection of F. corollata. Munz was not able to examine the holotypes or other isotypes of these species, so he concluded that F. corollata was a synonym of F. canescens. The latter species is now known with certainty to be distinct and much rarer than F. corollata ; it is found mostly on the eastern slopes of the Cordillera Central in southern Colombia and has thick-tubed, or- ange, canescent flowers and larger leaves than F. corollata. Morphologically similar populations of F. corollata are found from Cauca, Colombia to Imbabura, Ecuador. The species may even reach as far south as Pichincha, but the single collection there, Benoist 4547 (P; west side of Volcan Pichincha), has rather narrow flowers and pilose styles for this species. Different ploidy levels have been found in different parts of the range of F. corollata, however. Two collections from nearby populations in Cauca were diploid, and three counts from two distinct populations in Ecuador were tetraploid (Table 4). The only differences noted between these groups were that the tetraploid popu- lations had slightly larger seeds, more markedly rhombic petals with a narrow base, and a common pyramidal shaped growth habit, with sharply ascending branchlets. Their pollen also has variable proportions of triporate grains. Most of these morphological characters vary within the tetraploid populations, how- ever, and some of the Imbabura individuals even have petals that are shorter than the sepals. Two collections from the western slopes of the Cordillera Occidental in Car- chi, Ecuador, Holm-Nielsen et al. 5580 (AAU, S) and 5674 (S; both from Km 53 of Tulcan-Maldonado road, 3,150-3,250 m), have certain characters of F. corol- lata such as somewhat rhombic petals and green sepal tips in bud, but they have nearly entire, quaternate, membranous leaves that are not typical of this species. More cytological examination and population sampling from southern Colombia (Cauca and Narino) would help to establish the pattern of ploidy level in F. corollata and to detect if any further morphological differences can be correlated with either diploid or tetraploid populations. A substantial number of specimens from Cauca differ from typical F. co- rollata populations in one or more of the following characters: small, lanceolate leaves 15-30 mm long, few (2-4) secondary veins, entire or subentire leaf margins, petals shorter than the sepals, long floral tubes mostly 55-70 mm long, and the presence of triporate pollen grains. These collections, which are listed separately at the end of the specimens citation list, are believed to be hybrids between F. corollata and F. caucana. A series of these probable hybrids was examined in 1 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 June 1979 along the road from Popayan to La Plata, in Cauca and Huila Depts., on the north face of Volcan Purace. Ascending from Popayan from the west, F. corollata {Berry 3582 ; COL, MO) was collected with F. canescens at 3,235 m. Berry 3583 to 3586 (COL, MO) were collected farther east, at higher elevations in semi-open subparamo habitats near the divide of the Cordillera Central . Each number of this series comes from a different population along a stretch of several km, where extreme variability in leaf shape and floral dimensions was found. Starting to descend to the east, the first plant recognized as F. cauca na {Berry 3587 \ COL, MO) was found at ca. 3,100 m growing next to one of the variable forms typical of the upper, intermediate area {Berry 3588; COL, MO). Between 10% and 20% of the pollen in the suspected hybrids, Berry 3583 , 3584, 3586, and 3588, have triporate grains, and stainability varies from 20% in Berry 3583 to 80% in Berry 3584. Triporate grains in Fuchsia are sometimes indicators of polyploidy, and gametic and somatic counts of Berry 3584 indeed proved to be tetraploid. The two counts of F. corollata from Cauca and the one count of F. caucana from Narino are all diploid, however, and both Berry 3582 {F. corollata) and Berry 3588 {F. caucana) have entirely biporate and highly stainable (>95%) pollen grains. It is possible, then, that if all the variable collections along the exposed ridge of the Paramo de Purace are tetraploids, they originated as allotetraploid hybrids of F. corollata and F. caucana. This is only hypothetical, but could be analyzed experimentally. Additional evidence of hybrid origin for the above series is available from Berry 3584. Normal pairing of 22 bivalents was found in meiotic preparations, but the only ripe fruit found on the plant had 35 aborted seeds and 13 ripe ones. These seeds were germinated along with other collections at the University of California Botanical Gardens at Berkeley in 1980, but only one small, weak seedling developed, from which the tetraploid root tip count was made. Another area where collections variable in leaf texture and petal size occur is the Paramo de Las Papas area of the Macizo Colombiano in southern Cauca. Hybridization may be the cause of this variability as well. The type of F. co- lombiana, Cuatrecasas 18959 (A), from northern Cauca, is distinguished by its extremely reduced leaves less than 2 cm long, but this may be due at least in part to the high elevation (3,500-3,600 m) where it was collected. Its leaves are not very different from individuals such as Berry 3585, from the probable hybrid series discussed above. Fuchsia corollata is found in close proximity to F. caucana in Narino, and it occurs sympatrically with F. canescens, F. dependens, and F. sessilifolia. 24. Fuchsia caucana P. Berry, sp. nov. type: Colombia, Dept. Cauca, 31 km E of Totoro on road to Inza, E slopes of Cordillera Central, 3,240 m, 8 June 1979, Paul E. Berry 3577 (COL 107163, holotype; MO 2 sheets, isotypes). Fig. 26. Fuchsia petiolaris sensu Munz, Proc. Calif. Acad. Sci. IV. 25:34. 1943, pro parte. Frutex (0.3-)0.5-2 m altus plerumque pauciramosus. Ramuli teretes, subglabrati vel strigosi vel puberuli, albo-virides vel rubro-purpurei; caulibus glabrescentibus cortice subnitida purpureo-bru- nnea. Folia plerumque ternata interdum opposita raro quaterna, firme membranacea, (anguste) elliptica vel lanceolato-ovata, basi acuta vel rotundata, apice plerumque acuminata, (2.5-)3.5-7(-10) cm longa, (l-)1.5-3.2(-4) cm lata, supra obscure atroviridia, subglabrata sparsim vel strigulosa, subtus pallide 1982] BERRY— FUCHSIA SECT. FUCHSIA 1 19 albo-viridia vel strigosa nervis secundariis utroque latere (3-)5-6(-7) subtus saepe rubris, paralleli- nervis; margine conspicue glanduloso-serrulata, raro subintegerrima; petiolis saepe rubro-purpureis, 2— 13(— 14) mm longis; stipulis anguste lanceolatis vel linearibus, fuscatis, 3-4 mm longis, 0.5-1 mm latis, subpersistentibus. Flores roseo-cerasinii vel (pallide) purpureo-lavandulacei, pauci in axillis supernis ramorum dispositi; pedicellis dependentibus, 12-28 mm longis; ovario ovoideo-ellipsoideo, glabro vel puberulenti-strigoso, 5-6 mm longo. Tubi florales anguste infundibuliformes, 32^45(-54) mm longi, basi 2-4 mm lati parum bulbosi, plerumque constricti l A parte inferiore inde saepe ± abrupte dilatati, summo 4-11 mm lati, extus glabrati vel puberulenti, intus pilosi. Sepala lanceolata (sub)acu- minata ( 12— ) 14— 17(— 22) mm longa, 5-6 mm lata, alabastro apice interdum albido-viridi. Petala plerumque valde breviora quam sepalis, atrorubra ve! purpurea, elliptico-ovata, 9-1 1(-14) mm longa, 5_6(-9) mm lata, obtusa vel cuneata apice interdum mucronulata. Filamenta rosea, antisepala 8-10 mm longa, antipetala 5-7 mm longa; antheris oblongis 3-4 mm longis, 2-3 mm latis luteo-eburneis. Stylus usque ad summum tubi floralis dense pilosus, roseo-ruber; stigmate roseo, late capitato, tetra- gono, 2-3 mm long, 3.5-4 mm lato, apice leviter 4-fisso. Bacca ante maturitatem 4-sulcata, glabrata vel puberulento-strigosa, in maturitate subglobosa, 11-13 mm longa, 7-9 mm crassa, nitida rubro- purpurea; seminibus rubro-brunneis vel vivide rubris, 2-2.5 mm longis, 1.1-1.8 mm latis. Numerus gameticus chromosomatum /i = 11. shrub puberulent, white green to red purpl purpl opposite or quaternate, firmly membranous, (narrowly) elliptic to lance-ovate, acute to rounded at the base, mostly acuminate at the apex, (2.5-)3.5-7(-10) cm long, (1— )1.5— 3.2(— 4) cm wide, dark velvety green and subglabrous to sparsely strigose above, pale whitish green to deep flushed purple and strigose below; secondary veins (3-)5-6(-7) on either side of the midvein, subparallel and as- cending towards the tip at ca. 45° to the midvein; margins serrulate with con- spicuous glandular teeth, rarely subentire. Petioles glabrous to strigose, usually purple, 2— 13( 3-4 long, 0.5-1 mm wide, subpersistent. Flowers few in upper leaf axils. Pedicels pendant, 12-28 mm long. Ovary ovoid-ellipsoid, glabrous to puberulent or stri- gose, 5-6 mm long, 2.5-3 mm thick. Floral tube narrowly funnelform, occasion- ally dilated in the middle and constricted again near the rim, 32-45(-54) mm long, 2-4 mm wide and slightly bulbous at the base, usually narrowed to 1.5-3 mm in lower Ys, then widened to 4-11 mm wide at the rim, glabrous to strigose or puberulent outside, pilose inside for most or the entire length. Sepals lanceolate, (sub-)acuminate at the apex, ( 12— )14— 17(— 22) mm long, 5-6 mm wide, with a short point 1-2 mm long in bud, spreading at anthesis. Tube and sepals pink cerise to (light) purple lavender, young buds sometimes whitish green. Petals usually con- siderably darker than the sepals, (dark) red-purple, elliptic-ovate, 9-1 1(-14) mm long, 5-6(-9) mm wide, obtuse to cuneate or sometimes mucronate at the apex, suberect at anthesis. Nectary unlobed, ca. 2 mm high. Filaments pink, 8-10 mm and 5-7 mm long; anthers oblong, 3^ mm long, 2-3 mm wide, yellow cream. Style densely pilose from base to near the rim, red pink; stigma broadly capitate, tetragonous, slightly 4-cleft apically, 2-3 mm long, 3—1 mm wide, pink, exserted 7-16 mm beyond the anthers. Berry 4-sulcate before maturity, glabrous to strigose or puberulent, subglobose at maturity, 11-13 mm long, 7-9 mm thick, lustrous red purple; seeds reddish brown to bright red, 2-2.5 mm long, 1.1-1.8 mm wide. Gametic chromosome number n = 1 1 . Distribution: Southern Colombia. Low shrubs in open thickets in high ele- vation elfin cloud forest, eastern slopes of the Cordillera Central in Cauca, Huila, 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Narino, and Putumayo, and in the Cordillera Occidental in the Farallones de Cali, Valle, 2,700-3,600 m (Fig. 59). Representative specimens examined: Colombia, cauca: near Rio Grande, between San Andres and Alto Todos los Santos, Core 1060 (RSA); between Jardin and San Rafael, Quebrada del Rio San Marcos, Cuatrecasas 14711 (F); Alto del Buey, Escobar & Escobar 1143 (MO); Paramo de Guanacas, Lehmann 5616 (K); Parque Nacional Purace, Paletara, Luteyn & Lebron-Luteyn 6948 (NY); Purace, Moscopan, Yepes-Agredo 274 (COL), huila: between Leticia and Parque Nacional Purace, 63 km SW of La Plata, Aguirre 274 (COL); 37^0 km E of Purace, road to La Plata, Berry 3587 (COL, MO), 3590 (COL, MO), narino: Paramo de Quilinsayaco, between El Encano and Santiago, Barclay & Juajibioy 9470 (RSA); above El Encano, Laguna La Cocha, Balls B7504 (BM, K, NA, UC, US); 24 km E of Pasto on road to Mocoa, Berry 3252 (MO, PSO); Paramo de Bordoncillo, Espinal 1023 (COL, PSO); 34 km E of Pasto, Paramo de Quilinsayaco, Espinosa E3089 (NY); above Laguna La Cocha, 16 km ESE of Pasto, Fosberg 20445 (RSA, US); El Tabano, Garcia-Barriga 4560 (COL, US); Bue- saco, Garcia-Barriga et al. 13016 (COL, US); Laguna La Cocha, Ciudadela, near Paramo de Bor- doncillo, Schultes & Villarreal 7575 (COL, GH, K, NY, RSA, US), putumayo: between Santiago and Laguna La Cocha, Alston 8398 (BM, COL, K, RSA); 8 km N of Sibundoy, Bristol 823 (DS, PSO); La Cabana, headwaters of Rio Sibundoy, valley of Sibundoy, Cuatrecasas 1 1598 (COL, F, US); Paramo de Bordoncillo, Cuatrecasas 11709 (COL, US): Paramo El Capuchino, between El Encano and Sibundoy, Garcia-Barriga et al. 18581 (COL); Paramo de San Antonio, between Laguna La Cocha and Sibundoy, Schultes 3225 (DS, GH, NY); between La Maria and Paramo de San Antonio, road from Sibundoy to Pasto, Schultes & Villarreal 7827 (GH, RSA). Fuchsia caucana is allied to other long-tubed, axillary-flowered, high-elevation species such as F. corolla ta and F. petiolaris. Munz included the specimens he saw of this species under F. petiolaris, but F. caucana is clearly distinct from that species both morphologically and geographically. Part of this confusion is due to the loss of key characters upon drying, making it difficult to distinguish this species when just herbarium specimens are available. Fuchsia caucana can generally be distinguished, though, by its acuminate-tipped leaves with few sec- ondary veins that ascend toward the tip at a sharp (±45°) angle from the midvein. When fresh, the leaves are mostly a matte, velvety green above and much paler purpl rpl the redder flowers of F. petiolaris and F. corollata. The petals are often a deep purple and are always darker and considerably shorter than the sepals. In F. corollata , the petals usually exceed the sepals and are scarlet colored. Some variation in tube shape and pubescence occurs between different pop- ulations. In Huila and Cauca, most plants are glabrous or strigose with floral tubes that are usually somewhat constricted at the rim. Plants from Narino and Putumayo are commonly puberulent with narrowly funnelform tubes widest at the rim. The disjunct populations from the Cordillera Occidental in Valle are well within the variation present in the Cordillera Central. Fuchsia caucana is sympatric with F. canescens on the upper, eastern slopes of the Cordillera Central in Cauca and Narino. Probable hybrids are found around the type locality on the road from Totoro to Inza and are discussed under F. canescens. Extensive intergradation between F. caucana and F. corollata occurs along the high ridge of the Cordillera Central on slopes of Volcan Purace and in the Paramo de Las Papas area. Fuchsia caucana is only found on the eastern slopes, and F. corollata grows mostly on the western slopes; where they approach each other near the ridge of the cordillera, widely variable local populations are found with reduced leaves and floral tubes that are often longer than either of the 1982] BERRY— FUCHSIA SECT. FUCHSIA 121 supposed parental species, but with intermediate petal size. These populations are discussed in greater detail under F. corollata. 25. Fuchsia ayavacensis Humboldt, Bonpland & Kunth, Nov. Gen. PI. 6:107. 1823. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):549. 1941, pro parte. Munz, Proc. Calif. Acad. Sci. IV. 25:21. 1943, pro parte; Opera Bot., Ser. B, 3:11. 1974, pro parte, type: Peru, Dept. Piura, near Ayabaca, ca. 2,850 m, Aug. 1802, Alexander von Humboldt & Aime Bonpland (P, holotype, not seen; photograph, F; microfiche, MO). Fuchsia townsendii I. M. Johnston, Contr. Gray Herb. 75:33. 1925. type: Ecuador, Prov. Loja, Sabiango Hill, 26 Nov. 1910, Charles H. T. Townsend A93 (US, holotype; photographs, NY, UC; fragment, GH). Fuchsia asplundiii. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):548. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:37, pi. 4, fig. 24. 1943. type: Peru, Dept. Piura, Prov. Huancabamba, above Palambla, 3,000 m, April 1912, August Weberbauer 6054 (F, holotype; photographs, NY, POM, UC; G, GH, US, isotypes). Suberect to scandent shrubs 1-4 m tall with flexuous branches to several m long when scandent. Young growth densely canescent-strigose or hirtellous with whitish hairs, older portions strigose to hirtellous; older branches with brown, exfoliating bark. Leaves mostly ternate, occasionally quaternate, membranous, oblong to elliptic or narrowly elliptic-oblanceolate, acute at the base, acute to subacuminate at the apex, 6— 15(— 18) cm long, 2.2-5(-7) cm wide, matte green and strigillose above, paler and strigose below, sometimes red- tinged; secondary veins 7-12 on either side of the midvein, at times reddish below; margin subentire 6-20( 6-25( 6-8 funnelform, (35-)40-55 base, narrowed to 2-2.5 mm wide above the nectary and gradually widened above until 5-8 mm wide at the rim, subtetragonous in transection, strigose outside, pilose inside in lower Vz. Sepals lanceolate, acuminate, 10-20 mm long, 4-6 mm wide, spreading to divergent at anthesis. Tube and sepals orange red. Petals orange red, broadly elliptic-ovate, rounded to subacute at the apex, 7-11 long, 5.5-8 mm wide, mostly about half as long as the sepals. Nectary unlobed or shallowly 4-lobed, 1.5-2 mm high. Filaments light red, 8-11 mm and 5-8 mm long; anthers oblong, 2.5-3 mm long, ca. 2 mm wide, white. Style light red, pilose in lower Vi\ stigma subobconic, 4-cleft apically, ca. 3 mm long, 3-4 mm wide, light red. Berry ellipsoid, 16-20 mm long, 8-10 mm thick, strigose, reddish; seeds 1.3-1.6 mm long, 0.8-1.1 mm wide. Gametic chromosome number n = 11. northern Peru and southernmost southern Ecuador; scattered to locally frequent shrubs 900-3.200 (Fig. 59). Representative specimens examined: Peru, cajamarca: between Cascas and Contumaza, Berry & Escobar 3601 (MO, USM); Hacienda Taulis, between the Casa Hacienda and Palmito, Prov. Hual- gayoc, Hutchison & Bismark 6401 (F, G, LE, MICH, MO, NY, RSA, UC, US, USM); Socota to San Andres, Lopez & Sagdstegui 5372 (MO); Prov. San Miguel, Weberbauer 3903 (G). piura: 5 km 1 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 above Canchaque on road to Huancabamba, Berry & Escobar 3630 (MO, USM); 38 km above Can- chaque to Huancabamba, just below summit of road, Hutchison & Wright 6577 (RSA, UC, USM); Bajada de Ayabaca to El Puente, Ochoa 1782 (DS); Cienaga Larga, Huancabamba to Cuello del Indio, Sagdstegiu et at. 8526 (MO); Ayabaca, Soukup 4349 (US). This is the southernmost member of the Fuchsia petiolaris species group. Munz (1943, 1974) confused it with F. ampliata, which occurs in central Ecuador and has much larger, rounder petals, wider floral tubes, and strongly reflexed sepals. Fuchsia vulcanica can also be confused with this species, but it generally has smaller leaves with shorter petioles and larger, wider petals. Fuchsia ay a- vacensis is characterized by its large, dull leaves with whitish, strigose-hirtellous pubescence and its often scrambling habit with long, flexuous branches. It has not been found to occur sympatrically with any other species in the section, and it is the only species centered in the relatively dry Cordillera Occidental of Peru. It grows there is moist scrub thickets and in the southernmost remnants of cloud forest on the west side of the Andes. 26. Fuchsia vulcanica Andre, Rev. Hort. 60:233, 268. 1888. type: Colombia, Dept. Narino, Volcan Azufral, near Tiiquerres, 18 May 1876, Edouard Andre 3228 (K, holotype; photograph, MO). Fuchsia hitchcockii I. M. Johnston, Contr. Gray Herb. 75:33. 1925. type: Ecuador, Prov. Azuay, be- tween Ona and Cuenca, 2,700-3,300 m, 9-10 Sept. 1923, Albert S. Hitchcock 2 1603 (GH, holotype; NY, US, isotypes). Fuchsia ayavacensis sensu Munz, Proc. Calif. Acad. Sci. IV. 25:32. 1943, pro parte; Opera Bot., Ser. B, 3:11. 1974, pro parte. Fuchsia canescens sensu Munz, Proc. Calif. Acad. Sci. IV. 25:26. 1943, pro parte; Opera Bot., Ser. B, 3:12. 1974, pro parte. Erect to scandent shrubs 0.4-3.5 m tall, sometimes semi-epiphytic on mossy tree trunks. Branchlets terete, 1.5-3 mm thick, densely hirsute to hirtellous or strigillose, usually with erect, white to tan hairs; older stems 3-10 mm thick with exfoliating bark. Leaves 3-5-verticillate, mostly ternate or quaternate, firmly membranous to subcoriaceous, narrowly to broadly elliptic to obovate, acute to obtuse at both ends, 2— 5(— 10) cm long, (0.6-)l-2.5(-4) cm wide, matte to subnitid green and usually strigose above, lighter green and hirsute to strigose below; secondary veins 3— 8(— 1 1) on either side of the midvein, often red below; margin subentire to denticulate. Petioles pubescent, 2-10(-20) mm long. Stipules narrow- ly lanceolate, 1.5-2.5 mm long, 0.3-0.7 mm wide, deciduous. Flowers generally fewer than 12 per branch, axillary and pendant. Pedicels pubescent, 5-25(-40) mm long. Ovary ovoid-ellipsoid, strongly tetragonous, 6-9 mm long, 2-3.5 mm thick, hirsute to strigose, green to dull red. Floral tube narrowly funnelform, (32-)36-50(-60) mm long, 3-4 mm wide and somewhat bulbous at the base, nar- rowed to 2-3.5 mm wide above the nectary, then gradually widened or slightly ampliate near the middle until 6-10 mm wide at the rim, hirsute to strigose or subglabrous outside, pilose inside for most its length. Sepals (narrowly) lanceo- late, acute to subacuminate at the apex, (10-)13-20(-23) mm long, 3.5-5 mm wide, spreading to divergent at anthesis. Tube and sepals subnitid scarlet, pink, or orange red, sepals sometimes dull, green-tipped. Petals red, orbicular to obovate or broadly elliptic, rounded to (broadly) acute at the apex, (8-)9-15 mm long, (6-)8-13 mm wide, spreading to divergent at anthesis. Nectary unlobed or shal- 1982] BERRY— FUCHSIA SECT. FUCHSIA 123 lowly 4-lobed, 1.5-2 mm high. Filaments red, 8-12 mm and 6-9 mm long; anthers oblong, 3-4 mm long, 1.5-2 mm wide, cream. Style red, villous for most its length; stigma conic-globose, slightly 4-cleft apically, 2-3.5 mm long, 1.5^ mm wide, red. Berry tetragonous before maturity, subglobose to ellipsoid when ripe, 11-15 mm long, 7-9 mm thick, red purple; seeds tan, 1.8-2.5 mm long, 0.9-1.2 mm wide. Gametic chromosome number n = 11(?), 22. Distribution: Southern Colombia and Ecuador. Along the Cordillera Occiden- tal from Narino, Colombia to Cotopaxi, Ecuador, in elfin forest near tree line, 3,400-4,000 m; on the Cordillera Central (eastern escarpment of the Ecuadorian Andes) from Napo to Chimborazo, from tree line at 3,900 m to 2,500 m in cloud forest; and in the mountains of Azuay and Canar from tree line to upper cloud forest. 2.850-3.400 m (Fig. 59). Representative specimens examined: Colombia, narino: Volcan Azufral, Andre s.n., (K); Volcan Galera, near Pasto, Barclay 4637 (COL); La Laguna, Mun. Cumbal, Benavides 993 (PSO); NW slopes Volcan Chiles, Ewan 16093 (POM); Volcan Cumbal, Vogel 259 (U), von Sneidern in 1941 (S). Ec- uador, azuay: Paramos de Silvan, Barclay & Juajibioy 8403 (RSA); 13-18 km from Cuenca on road past Sayausi, Berry & Escobar 3184 (MO), 3187 (MO, QCA), 3188 (MO); 35 km S of Cuenca on road to Ona, Berry & Escobar 3190 (MO, QCA); along Rio Yanucay above San Joaquin, road to Laguna Soldado, Berry & Escobar 3212 (MO, QCA), 3213 (MO), 3214 (MO), 3225 (MO, QCA); Yanasacha, Parroquia Banos, Boeke 621 (MO); road Cuenca-Angas, near Angas, Boeke 654 (MO); Paramo de Tinajillas, 30-50 km S of Cuenca, Camp E-462 (NY, RSA), E-2107 (NY, RSA), E-2108 (NY, RSA); along Rio Matadero, W of Cuenca, Camp E-1988 (NY, RSA, U); N of Paute, Camp 2590 (NY, RSA, VEN); 1-8 km N of Sevilla de Oro, Camp 4257 (NY, RSA), E-4672 (COL, G, MO, NY, P, RSA, S, US, VEN); Quebradas of the Rio Collay, 3-8 km N of Sevilla de Oro, Camp 4994 (G, MO, NY, RSA, UC, US); El Pan, Hading 1228 (NY, S); 10 km S of Cumbe, Hading 5658 (NY, S); 10-12 km N of Sevilla de Oro, 0llgaard & Balslev 9366 (AAU, MO), 9374 (AAU, MO); near Laguna Sorocucho, Prescott 827 (DS, NY), Scolnik 1461 (RSA); Rio Machangara, Sparre 18561 (S), 18601 (S), 18664 (S). canar: above Biblian on road to Canar, Berry & Excobar 3227 (MO, QCA), 3228 (MO, QCA), Hading 6296 (NY, S), Hading et al. 8614 (RSA). carchi: Paramos del Angel, near Voladero, Barclay & Juajibioy 9396 (RSA), Penland & Summers 915 (F, GH, POM); base of Volcan Chiles, ca. 12 km W of Tufino, Berry 3156 (MO, QCA), 0llgaard & Balslev 8398 (AAU); near El Angel, Popenoe 1260 (NA); Paramo El Angel, Sparre 14148 (S). chimborazo: Cordillera Oriental, between Alao & Cush- nipaccha, Acosta Solis 7201 (F), 7214 (F). cotopaxi: 17 km above Pilalo, road to Latacunga, Berry & Berry 2539 (MO), 2540 (MO); timberline on Pilalo-Latacunga road, Holm-Nielsen & Jeppesen 1396 (AAU, S), Holm-Nielsen et al. 3335 (AAU, S). imbabura: Laguna de Mojanda, 0llgaard & Balslev 8779 (AAU). napo: between Oyacachi and Comenia, Acosta-Solis 11166 (F); Cuyuja, Balslev & Madsen 10469 (AAU), 10522 (AAU); ca. 5 km above Papallacta to Quito, Berry & Berry 2521 (MO); Km 215 of Quito-Baeza road just above Lago Papallacta, Berry & Berry 2523 (MO); 2-6 km below Papallacta, Berry & Berry 2525 (MO), 2526 (MO), Gentry 12401 (MO, US); just above town of Papallacta, Berry & Escobar 3245 (MO); Lago San Marcos, E of Cayambe, Cazalet & Pennington 5312 (NY, UC, US); Quebrada Chalpichio, ca. Km 204 E of Papallacta, Croat 49412 (MO); Los Corrales, near Papallacta, Grubb et al. 208A (NY 2 sheets); below Papallacta, Hading 3994 (NY, S); 4 km W of Papallacta, Holm-Nielsen et al. 6785 (AAU, MO, NY, S); between Cuyuja and Papallacta, 10 km E of Papallacta, Holm-Nielsen et al. 6834 (AAU, NY, S); Laguna de Papallacta, 01lgaard & Balslev 8036 (AAU); ca. 7 km W of Papallacta, MacBryde & Dwyer 1240 (MO, QCA); just W of Papallacta, Plowman et al. 3870 (GH, K, MO, S); Rio Papallacta, Station IV, Prescott 1276 (MSC); Dos Arroyos, ca. 5 km NW of Laguna Norte de Papallacta, Sparre 15021 (S); Rio Calpi on Baeza- Papallacta road, Sparre 15949 (S); Papallacta, Sparre 17704 (S). pichincha: E part of Haciendas Pedregal and Yanurcu, Chaparro de Sebritana, Acosta-Solis 8353 (F); NW side of Volcan Pichincha, Yanacocha trail, Luteyn et al. 5685 (MO), tungurahua: Llanganati mountains, ridge between Rio Tope and Rio Golpe, Edwards 164 (K); Paramo de Minza, Minza Chica, Penland et al. 337 (F, GH, POM). Probable hybrids Fuchsia vulcanica x F. ampliata: Ecuador, cotopaxi: above Pilalo, 2,800 m, Barclay & Juajibioy 8076-A (RSA); 13 km above Pilalo on Quevedo-Latacunga road, ca. 3,300 m, Berry & Berry 2541 (MO), 2542 (MO), 2543 (MO), 2544 (MO), 2545 (MO), 2546 (MO), 2547 (MO); 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 12 km E of Pilalo, Escobar 1100 (MO); Prov. Cotopaxi, Gilmartin 44 (MO); Pilalo- Latacunga road at timberline, 3,400 m, Holm-Nielsen & Jeppesen 1396 (DS), 1397 (AAU, DS); Quevedo-Latacunga road, above Pilalo, 3,200-3,300 m, Holm-Nielsen et al. 3268 (COL, MO, NY, S). This species is part of the high elevation Fuchsia petiolaris species group of the northern Ancles. It and Fuchsia ampliata are closely related and distinct from the other species in their generally rotund petals and coarse, usually dense pu- bescence. Fuchsia vulcanica can be distinguished from F. ampliata by its non- reflexed sepals, strongly tetragonous ovaries, smaller leaves, and shorter petioles and pedicels. Fuchsia vulcanica is treated here as a widely polymorphic species. Intensive field work and cytological examination of more populations is badly needed and may in the future lead to the recognition of additional taxa, but the extent of variation within most parts of the range is such that no consistent morphological differences can be found to separate distinct taxa at this time. The most uniform populations occur around the type locality in southern Colombia and northern Ecuador, where plants are all densely hirsute, with leaves less than 35 mm long, and pedicels less than 8 mm long. These plants occur only at or near treeline on the upper slopes of volcanic peaks. A possible diploid count of n = ca. 11 was obtained from this area from Berry 3156. Along the same mountain range, but further south in Imbabura and Pichincha, plants are found 3-6 southernmost raploid. Four tetraploid counts were obtained from two different populations (see Table 4). In addition, triporate pollen grains, which often indicate polyploidy in the genus, have been found in representatives of different localities in this area, including Hading 1228 (NY, S; El Pan), Berry & Escobar 3225 (MO, QCA; above 0llg MO: 10-12 km N of Sevilla de Oro), and Ha Most northern Provinces. Pubescence these populations from densely hirsute to hirtellous, and the petals are particularly variable, from orbicular to elliptic and acute-tipped. The type of F. hitchcockii is unusual in its long floral tubes (ca. 60 mm long). It represents the extreme of variability in this group, with petals half as long as the sepals and pedicels 3-4 cm long. Its hirsute pubescence and round petals are characteristic of F. vulcanica, however, and other collections from the same area are variable, but generally closer to the average values of morphological char- acters for this species. There are two areas where F. vulcanica intergrades extensively with F. ampliata or else shows a great deal of polymorphism over an altitudinal gradient. In the Cordillera de Zumbagua, above Pilalo in Prov. Cotopaxi, F. vulcanica is near treeline, ca. 3,450-3,500 infrequent of intermediate forms were found; these collections are listed separately at the end of the specimens citation as probable hybrids of F. vulcanica x F. ampliata. In July 1977, Berry & Berry 2541 to 2547 were collected from a rock pile that had been made by roadbuilding machinery. This population appeared to be a 1982] BERRY— FUCHSIA SECT. FUCHSIA 125 hybrid swarm between the above two species, as shown by the wide variation in the degree of sepal reflexion, leaf size and texture, and pubescence. No significant reduction in pollen stainability occurs in the intermediate forms, however, which seems to confirm the close affinity of these two species. In 1979, two years after the initial visit, this population had been destroyed by road cleaning crews. Such repeated disturbance and rapid turnover of local populations and the tendency of F. ampliata to grow in higher and drier sites than the Zumbagua cloud forest are likely major factors in the probable breakdown of isolating mechanisms be- tween F. vulcanica and F. ampliata in Cotopaxi. The second area where considerable variability involving F. vulcanica occurs is in Napo, on the road from Quito to Baeza on the eastern slopes of the Andes. Typical, small-leaved, high elevation forms of F. vulcanica occur about 5 km above the town of Papallacta near treeline at 3,700 m. As one descends first to the town of Papallacta at ca. 3,300 m, then down as low as 2,500 m in cloud forest to the east of Papallacta, considerably larger leaved plants are found along with greater variability in pedicel length and degree of sepal reflexion. Berry & Berry 2528 (MO; ca. 12 km east of Papallacta, 2,600 m) is the extreme of leaf variation, with blades 10 cm long and 4 cm wide, with 9 secondary veins on each side of the blade. MacBryde 875 (MO; W end of Lago Papallacta, 3,300 m) has some pedicels 4 cm long and floral tubes 10-13 mm wide at the rim, both beyond the normal range of variability for F. vulcanica. AH these collections share the 3-4-verticillate, short-petiolate leaves, hirtellous pubescence, and round petals of F. vulcanica, however. Asplund 18239 (S; Papallacta, 3,000 m) is a large-leaved form that apparently has reflexed sepals. This points to possible introgression with F. ampliata, but that species is not known from the eastern slopes of the Andes in Ecuador. A gametic chromosome count of n = ca. 22 was obtained from Berry & Escobar 3245 (MO, QCA; Papallacta). More intensive sampling and cytological study of plants from this area will be necessary to understand the dynamics of these populations better. Besides F. ampliata, F. vulcanica occurs sympatrically with F. loxensis in Azuay and with F. pallescens at its lower altitudinal limits in Napo. fig 439. 1877. Hook., Bot. Mag. t. 6839. 1885. type: Ecuador, Prov. Pi< slopes of Volcan Pichincha, ca. 3,000 m, 1841-1843, Theodor Hart (K Bentham Herb.. holotvDe: Dhotoeraoh. MO: BM. BR. BREM, sheets, F, G, K Hooker Herb., OXF, P, W 2 sheets, isotypes). Fig. 23. Fuchsia ayavacensis sensu Munz, Proc. Calif. Acad. Sci. IV. 25:32, pi. 3, fig. 18. 1943, pro parte; Opera Bot., Ser. B, 3:12. 1974, pro parte. Fuchsia canescens sensu Munz, Opera Bot., Ser. B, 3:12. 1974, pro parte. Erect to scandent shrubs 1-3 m tall with mostly ascending branches. Young growth conspicuously canescent to hirtellous with white hairs; branchlets 2-3 mm thick, hirtellous to densely strigose, green to dull purple; older branches dull ash gray. Leaves ternate or occasionally quaternate, often drooping, membra- nous, (narrowly) elliptic to lanceolate, acute to narrowly cuneate at the base, acute to subacuminate at the apex, 4-9(-12) cm long, 1 .5^\ cm wide, matte green 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 and strigose above, pale green and strigose- villous below with red purple veins; secondary veins 5— 9(— 1 1) on either side of the mid vein; margin subentire to den- ticulate. Petioles strigose, 7-20 mm long, usually dull purple. Stipules lance- linear, 3-5 mm long, 0.5-1 mm wide, subpersistent. Flowers axillary and pendant, generally few. Pedicels stout, 1-2 mm thick, 15-35(-50) mm long, strigose. Ovary ovoid, usually constricted near the apex, 6-8 mm long, 3-4 mm wide, canescent- hirtellous and slightly furrowed. Floral tube funnelform, usually dilated above the middle, (32-)40-50(-65) mm long, 3-4.5 mm wide and bulbous at the base, tapered to 2-3(-4) mm wide in lower V&, widened above until (6— )8— 14 mm wide at the rim, strigose outside, villous inside for lower Vi- 3 A. Sepals lanceolate, narrowly acute at the apex, 16-23 (-28) mm long, 6-8 mm wide, with a narrow, tapered tip in bud, fully reflexed soon after anthesis. Tube and sepals bright scarlet to orange red. Petals red to orange red, rotund to orbicular or very broadly ovate, (10-) 13- 18 mm long, (9-) 10-16 mm wide, rounded to obtuse at the apex, erect to suberect at anthesis. Nectary green, shallowly 4-8-lobed, ca. 2 mm high. Filaments red or orange, 10-18 mm and 8-13 mm long; anthers oblong, ca. 4 mm long, 2-3 mm wide, cream. Style red, densely villous from the base to the rim of the tube; stigma globose to subclavate, slightly 4-cleft apically, 2-A mm long, 2-4 mm wide, red. Berry ellipsoid, usually ± tetragonous before maturity, slightly verrucose, 14-16 mm long, ca. 8 mm thick, red; seeds tan, 1.8-2.1 mm long, 0.8-1.2 mm wide. Gametic chromosome number n = 11. Distribution: Ecuador, along the Cordillera Occidental from Imbabura to Bo- livar and one collection from the Narino-Putumayo border in southern Colombia; shrubs in hedgerows on dry slopes, near streams or in upper elevation cloud forest, (2,850-)3,000-3,500 m (Fig. 60). Representative specimens examined: Colombia, putumayo: S side of Laguna La Cocha, Paramo de Santa Lucia, source of Rio Alisales, Cuatrecasas 1 1884 (COL). Ecuador, bolivar: Simiatug, Hacienda Talahua, Penland et al. 623 (F, GH, POM), cotopaxi: Quevedo-Latacunga road, above Pilalo, 2850 m, Holm-Nielsen et al. 3274 (AAU, NY, S). imbabura: Bosque Andino "Rosaspamba," E side of Mojanda, Acosta-Solis 8104 (F); ca. 8 km W of Laguna Cuicocha on road to Inta, Berry 3169 (MO, QCA); ridge of Moinala, SW of Volcan Cotacachi, Owenby 2646 (COL, MO, RSA, US). pichincha: Corazon, Andre K.819 (F, K, NY); ca. 9 km W of Chillogallo to Chiriboga, Berry & Escobar 3234 (MO, QCA); Pichincha, Jameson in 1851 (BM, CGE, G, OXF, US), Humboldt & Bonpland 3115 (F, P); Volcan Pichincha, Mexia 7654 (BH, UC, US); road from Cotocollao, near Nono, Mexia 7661 (BM, GH, K, MO, NY, POM, S, U, UC, US). This species can be easily recognized by its broad, firm-tubed flowers with nearly erect, round petals and fully reflexed sepals at anthesis. The plants are generally covered by a tomentum of white, strigose- villous hairs, and the styles are densely pilose up to the rim of the tube. Fuchsia ampliata is a member of the high elevation F. petiolaris species group and was confused by Munz (1943, 1974) with F. ayavacensis from northern Peru and southernmost Ecuador. The differences between these two species are outlined under F. ayavacensis. Fuchsia ampliata largely is restricted to areas near tree line in a small extension of the Cordillera Occidental in central Ecuador, mostly on the drier eastern slopes facing the Central Valley of Ecuador. One disjunct collection is known from the eastern slopes of the Cordillera Central in southern Colombia, however. In Cotopaxi, Ecuador, above Pilalo, it occurs in moister cloud forest but is apparently quite rare. There, it hybridizes extensively with the closely related F. vulcanica, pro- 1982] BERRY— FUCHSIA SECT. FUCHSIA 127 ducing local hybrid swarms. These are discussed more fully under F. vulcanica. Fuchsia ampliata also occurs sympatrically with F. dependens and F. sessilifolia in a few localities. 28. Fuchsia venusta Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6:105. 1823. Planch., Fl. Serres Jard. Eur. 5: t. 538. 1849; Rev. Hort. III. 4:241, /. 1850. Moore & Aynes, Gard. Mag. Bot. 2:36, fig. 2. 1850. Lindl. & Paxt., Paxton's Fl. Gard. \:79, fig. 57. 1850. Anon., J. Hort. Pract. Gard. III. 49:243, t. 1904. Fuchsia venusta var. typica Munz, Proc. Calif. Acad. Sci. IV. 25:39, pi. 5, fig. 26. 1943. Based on F. venusta HBK. type: Colombia, Dept. Cundina- marca, near Guayavalito, July-Sept. 1801, Alexander von Humboldt & Aime Bonpland (P, holotype, not seen; photographs, F, GH; microfiche, MO; P, isotype). Fig. 24. Fuchsia killipii I. M. Johnston, Contr. Gray Herb. 81:94. 1928. Munz, Proc. Calif. Acad. Sci. IV. 25:52, pi. 8, fig. 41. 1943. type: Colombia, Dept. Santander, Rio Surata, above Surata, 2,000-2,300 m, 5-6 Jan. 1927, Ellsworth P. Killip & Albert C. Smith 16695 (GH, holotype; photographs, POM, UC; NY, US, isotypes). Fuchsia venusta var. huilensis Munz, Proc. Calif. Acad. Sci. IV. 25:39. 1943. type: Colombia, Dept. Huila, Cordillera Oriental, E of Neiva, 1,800-2,300 m, 1-8 Aug. 1917, Henry H. Rushy & Francis W. Pennell874 (NY, holotype; photographs, POM, UC; F, GH, MO, US, isotypes). Fuchsia meridensis Steyermark, Fieldiana Bot. 28:439. 1952. type: Venezuela, Edo. Merida, between La Azulita and La Trampa, road to Lagunillas, 1 ,280-2,285 m, 27 Aug. 1944, Julian A . Steyermark 56162 (F 1368909, holotype; NY, isotype). Erect to scandent shrubs 1-3 m tall or lianas to 10 m above ground, with suberect or long, flexuous-pendant branches to several m long. Branchlets terete to subtriangular, 2-6 mm thick, minutely puberulent to finely hirtellous, dull wine red to bluish purple; older branches 10-40 mm thick, with tan, exfoliating bark. Leaves ternate, rarely opposite or quaternate, firmly membranous to subcoria- ceous, elliptic, rounded to acute at the base, acute to subacuminate at the apex, 5- 1 1 .5 cm long, 1 .5-4.5 cm wide, glossy dark green and subglabrous above, glossy pale green and subglabrous to pubescent along the nerves below; secondary veins 7-12 on either side of the midvein, impressed above, subelevated below; margin subentire. Petioles puberulent or strigillose, 4— 1 0(— 1 2) mm long, wine red. Sti- pules lance-deltoid, 1-1.5 mm long, ca. 0.4 mm wide, subulate at the apex, de- ciduous. Flowers axillary in uppermost leaf axils or subracemose at branch tips; when subracemose, rachis 2-4 cm long, subtending leaves usually deciduous. Pedicels slender to filiform, ascending to divergent in bud, arching or drooping ■ by anthesis, (10-)15-30(-60) mm long, glabrous to pubescent. Ovary oblong to ellipsoid, 4-8 mm long, 2-3 mm thick. Floral tube narrowly funnelform, (30-)35-60 mm long, 2.5^.5 mm wide and slightly bulbous at the base, narrowed to 2-A mm wide above the nectary and gradually widened above until 5-10 mm wide at the rim, subglabrous to puberulent outside, pilose inside for most of length. Sepals lanceolate, 14-20 mm long, 4-7 mm wide, acute to acuminate at the apex, some- times pubescent within, subulate in bud for 1-2 mm, spreading to divergent at anthesis. Tube lustrous orange red, at times greenish in bud; sepals orange red, often green-tipped in bud. Petals orange to orange red, oblanceolate, 15-22 mm long, 3-6 mm wide, undulate to crispate-margined, abruptly acute at the apex, strongly spreading and recurved at anthesis, with dorsal hairs occasionally pres- 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 ent along the midvein. Nectary green, unlobed to shallowly 4-lobed, ca. 2 mm high. Filaments light red, 10-15 mm and 8-11 mm long; anthers oblong, 2.5-3.5 mm long, 1.5-2 mm wide, cream to pale yellow. Style densely villous from the base to the rim of the tube, light red; stigma subglobose, 4-cleft apically, 2-3 mm long, 2-3 mm wide, dull white to orange red, exserted 3-15 mm beyond the anthers. Berry ellipsoid-oblong to subglobose, 8-20 mm long, 6-10 mm thick, lustrous dull green to dark purple; seeds tan, 1.5-2 mm long, 0.8-1.5 mm wide. Gametic chromosome number /z = 11. Distribution: Colombia and Venezuela. Locally frequent to scattered shrubs or lianas along forest edges, streamsides, thickets or in trees of mid-elevation cloud forest; in Venezuela, from Merida to Tachira; in Colombia in the Cordillera Oriental from Santander to Huila, and in the Cordillera Central in Tolima; 1.800-2.700 m (Fie. 60). Representative specimens examined: Venezuela, merida: La Carbonera, Aristeguieta 2847 (NY, VEN); La Loma de la Vagabunda, near El Morro, between El Morro and Aricagua, Badillo 6563 (MY); Quebrada de La Mucuy, towards El Volcan, Bernardi 280 (MER, NY); La Mucuy, Bemardi 306 (MER, NY, VEN); Alto de Monte Zerpa, Bernardi 664 (NY); El Paramito, between Merida and La Carbonera, Berry & Dugarte 2507 (MER, MERF, MO), Berry 3450 (MO, VEN); Vi km below La Trampa, Berry & Centeno 25 1 1 (MER, MERF, MO); 23 km above La Azulita on road to La Trampa, Berry & Centeno 2512 (MER, MERF, MO); 17 km E of Zumbador to Queniquea, Berry 3422 (MO, VEN); La Mucuy to Mesa de los Pinos, Berry 3431 (MO, VEN), Berry in 1980 (MO), 3433 (MO, VEN); 3-5 km below San Eusebio, road to La Azulita, Berry 3542 (MO, VEN), 3455 (MO); Paramo de Aricagua, John 1029 (GH, US, VEN); Chorrera de La Gonzales-La Isla-Jaji, Lopez-Palacios 1919 (MERF, MO); El Chorotal, Quintero 1614 (MER); El Valle Grande, between San Javier and la Sierra de la Culata, Ruiz-Terdn & Lopez-Figueiras 2116 (MERF, MO); El Yagrumal, above El Maciegal, ca. 10 km W of Merida, Ruiz-Terdn et al. 12540 (MERF, MO); Bosque Experimental de San Eusebio, We ss els-Boer 1733 (MER, MO, U, VEN). tachira: 21 km E of El Portachuelo to Pregonero, Berry 3292 (MO, VEN); 13 km E of Zumbador, Berry 3302 (MO, VEN); 1 1-17 km W of Zumbador towards San Isidro, Bunting 2564 (MY); Paramo el Pantano, Charpin et al. 13459 (G); Paramo del Zumbador, Ferrari 1171 (MY); Paramo de Angaraveca, Jahn 129 (US, VEN); between Michelena and Boca de Monte, W of Zumbador, Steyermark & Rabe 96805 (NY, U, VEN); 4.5 km above El Hato, ca. 30 km above Pregonero on road to Bailadores, Tillett & Honig 758-520 (MO). Colombia, boyaca: Chiquinquira, Ariste-Joseph A842 (US); Yanaca-Mapiri, Garcia-Barriga 491 1 (COL, US); near So- cota, Sierra Nevada del Cocuy, Grubb et al. 697 (COL, K); Km 307 above Pajarito, Rancherias, Idrobo & Jaramillo 1605 (COL); near Quebrada Chorro Grande, N side Arcabuco Range, between Duitama and Charala, Langenheim 3614 (COL, UC, US); Finca La Saria, Corregimeinto de Virolin, Lozano et al. 2393 (COL); Km 88 road to Pajarito, Quebrada La Rocha, Sastre 733 (COL, MO, P); road to Barbosa, 10 km NW of Arcabuco, Steere 7048 (BH, COL, MICH); above Corinto, road Pajarito-Sogamoso, Uribe-Uribe 6534 (COL); Tota, Yepes-Agredo 3339 (COL), cundinamarca: Quebrada Honda, Andre in 1876 (K); Facatativa, Ariste-Joseph A514 (US); ca. 2.5 km W of Salto de Tequendama, road to El Colegio, Barclay et al. 3305 (COL, NA, US); just below Zipacon, Berry 3540 (COL, MO); Km 39 of Bogota-Sibate-Fusagasuga road, Berry 3548 (COL, MO); 10 km above Choachi, Breedlove 35540 (CAS); Salto de Tequendama, Cuatrecasas 48 (COL, F, US); Boca de Monte, Cuatrecasas et al. 25816 (COL); Tena, Dawe 23 (K, US); Choachi, Dawe 350 (K, US); San Miguel and Aguabonita, Duque- Jaramillo 3323 (CM, COL); Laguna Pedro Palo, 3 km N of Tena, Fernandez & Mora 1449 (COL); 5 km NNW of Cabrera, N and W of Rio Sumapaz, Fosberg 20982 (RSA, US); 12 km WSW of Junin, on Rio Blanco, drainage of Rio Sueva, Fosberg 21479 (NY, RSA, UC, US); 21482 (RSA, US); 5 km WSW of Charquito, headwaters of Rio Subia, Fosberg 22038 (NY, RSA, US); between Pacho and Rio Negro, Garcia-Barriga 10754 (COL, US); Sasaima, La Maria, Garcia-Barriga 12584 (COL, POM, US); Km 18 of road Mosquera-La Mesa, Gentry 17050 (COL, MO); 12 km NW of Gacheta, Moquentina valley, Grant 9534 (NA, RSA, US); San Fortunato, Hart- weg 995 (BM, BREM, CGE, F, G, K, LE, NY, OXF, P, W); Zipaquira-Pacho highway, H aught 5979 (COL, POM, US); Fomeque, toward the Paramo de Chingaza, Huertas & Camargo 5536 (COL); Junin, Vereda La Cumbre, toward Claraval, Huertas & Camargo 6602 (COL); above La Florida, Idrobo & Jaramillo 2250 (COL); Las Mercedes, Lourteig 3082 (MO); between Agua Larga and Facatativa, Mayor 56 (Z); Dintel, Perez-Arbelaez & Cuatrecasas 5291 (F, COL); Zipacon, Popenoe 1056 (NA); Santandercito-La Rambla, Silva & Moreno 270 (COL); Santandercito, Uribe-Uribe 1020 1982] BERRY— FUCHSIA SECT. FUCHSIA 129 Table 1 1. Distinguishing characters of Fuchsia gehrigeri and F. venusta. F. gehrigeri Twig color Leaf surface Leaf texture Leaf margin Petiole length Flower color Petal surface Fruit shape Seed length Light red to green Matte Membranous Denticulate 10-40 mm Red Smooth Ovoid-globose 2-3 mm F. venusta Wine red Glossy Firmly membranous to subcoriaceous Entire 4-10 mm Orange red Undulate Ellipsoid 1.5-2 mm (COL, U); Pacho, Urihe-Vrihe 1718 (COL), huila: Batan, on Rio Neiva, ca. 32 km SE of Neiva, Fosberg 19296 (NY, RSA); Quebrada Ariari, above Galilea, 23 km ENE of Colombia, Fosherg 19625 (RSA, US); Yucales, on Rio Fortalecillas, 32 km E of Neiva, Fosberg 19744 (RSA, US); E of Neiva, Rusby & Pennell 641 (GH, MO, NY, US); Balsillas, on Rio Balsillas, Rusby & Pennell 722 (GH, NY, US), santander: La Baja, Funck & Schlim 1306 (BM, CGE, F, G, LE, OXF, P, W); above Surata, Killip & Smith 16604 (A, US); vicinity of La Baja, Killip & Smith 16776 (A, F, GH, NY, S, US); vicinity of Charta, Killip & Smith 18854 (A, GH, NY, US), tolima: 24 km W of Fresno to Manizales, Berry 3551 (COL, MO); between Libano and Murillo, Escobar 1008 (MO); Lfbano, Pennell 3197 (GH, NY, US). Fuchsia venusta is one of few species that often climb up trees, with its long, flexuous branches (Fig. 3). It also can grow as an upright shrub, however. It is distinguished by its ternate, short-petiolate, elliptic leaves that are glossy on both surfaces and the wine red to dull bluish purple stems. The flowers are axillary to subracemose, with orange tubes, green-tipped sepals, and recurved, undulate- crispate petals. It is closely related to F. rivularis from northern Peru. That species, however, has mostly quaternate leaves, more secondary veins, and most- ly spreading sepal tips in bud. Fuchsia gehrigeri is another related species that is sympatric and can be confused with F. venusta ; a list of their distinguishing characters is given in Table 1 1 . Considerable variation in pubescence, leaf texture, and amount of green col- oration in the tube and sepals occurs between different populations of Fuchsia venusta. Along the edges of the Bogota Plateau where this species is common, localized variation in the above characters is found. Plants near the Salto de Tequendama have subcoriaceous leaves and thick stems with erect, hirtellous pubescence, but populations nearby at Zipacon and Aguabonita on the old Bo- gota-Fusagasuga road are typically subglabrous with membranous leaves. How- ever, plants from opposite ends of the range, in Huila, Colombia and Merida, Venezuela, often show little morphological differentiation. Populations from northern Santander were described by Johnston as Fuchsia killipii. Compared to other populations of F. venusta, these plants have more numerous flowers in generally tightly clustered groups and smaller, globose fruits. I could not find populations from this area during field studies in Santander, but the cited collections appear to be localized variants of F. venusta because of their shrubby to climbing habit, ternate, glossy leaves, pilose styles, and the frequent hairs on the petals and sepals. There are a few localized populations of F. venusta in Dept. Tolima on the 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 60. Distribution of the Fuchsia venusta species group. eastern slopes of the Cordillera Central at the latitude of the Bogota Plateau; plants probably spread there from the Cordillera Oriental, the same as with F. hirtella, F. nigricans, and F. petiolaris. Fuchsia venusta occurs sympatrically with F. gehrigeri, F. hirtella, F. ni- gricans, and F. verrucosa. It apparently hybridizes with F. gehrigeri in Merida and Tachira, Venezuela. Hybrids with F. hirtella were collected in Cundinamar- ca, Colombia, along the old Bogota- Fusagasuga road. In both these cases, the putative hybrids occur at the upper altitudinal limit of F. venusta and the lower limit of F. hirtella and F. gehrigeri (Fig. 6). Hybrids between F. venusta and F. nigricans are relatively common in Venezuela. Further analysis of these hy- brids is included in the discussion of F. gehrigeri, F. hirtella, and F. nigricans. 1982] BERRY— FUCHSIA SECT. FUCHSIA 131 29. Fuchsia rivularis J. F. Macbride, Candollea 8:24. 1940; Field Mus. Nat. Hist., Bot. Ser. 13(4):562. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:27. 1943. type: Peru, Dept. Amazonas, Chachapoyas, 1830-1841, Andrew Mathews (G-BOIS, lectotype, here designated; photograph, MO; G-DEL, isolectotype). Fuchsia woytkowskii i . F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):566. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:25, pi. 2, fig. 11. 1943. type: Peru, Dept. Amazonas, Almirante, 1,900 m, Felix Woytkowski 38 (F, holotype; photographs, NY, UC). Scandent shrubs or lianas to 10 m above ground, with long, flexuous-arcuate, mostly unbranched shoots to several m long. Branchlets and young growth sub- glabrous to puberulent, older branches pilose or puberulent, glabrescent with age, with subnitid, red purple bark exfoliating in wide strips. Leaves ternate or mostly quaternate, subcoriaceous, narrowly elliptic to elliptic, acute to obtuse at the base, narrowly acute to obtuse or subacuminate at the apex, 5-9 cm long, 2-5 cm wide, nitid dark green and subglabrous above, paler green and subglabrous to pilose below; secondary veins 13-15 on either side of the midvein, midrib elevated below; margin subentire to glandular-denticulate. Petioles stout, pilose, 3-6 mm long. Stipules triangular-lanceolate, 2-3 mm long, 0.8-1.3 mm wide, thick at the base, spreading, subpersistent. Flowers few and pendant in uppermost leaf axils. Pedicels drooping to subdivergent, puberulent, 10-53 mm long. Ovary cy- lindrical, 6-8 mm long, 2-3 mm thick, generally pubescent. Floral tube narrowly funnelform, 36-65 mm long, slightly bulbous at the base and 3-5 mm wide, nar- rowed to 2-4 mm wide above the nectary, then widened gradually above until (5-)6-10 mm wide at the rim, subglabrous to pilose or strigillose outside, pilose inside. Sepals narrowly lanceolate, 16-20 mm long, 3-7 mm wide, with mostly free, spreading tips in bud 1.5-2.5 mm long, divergent at anthesis. Tube and sepals bright scarlet. Petals orange red, oblong to lance-elliptic, 15-22 mm long, 4-7 mm wide, crispate-margined, often with a few, scattered hairs on the midvein dorsally, spreading-recurved at anthesis. Nectary light green, shallowly 4-lobed, 3-4 mm high, fused to the base of the tube in lower %. Filaments red, 10-15 mm and 6-1 1 mm long; anthers narrowly oblong, 4^4.5 mm long, 1.5-2 mm wide, white. Style red, usually glabrous; stigma subobconic, 3-4 mm long, 2-3 mm wide, lightly 4-cleft apically, light red. Berry ellipsoid, 14-16 mm long, 8-10 mm thick, reddish; seeds tan, 1.1-1.4 mm long, 0.8-1.0 mm wide. Gametic chromosome number n = 11. Distribution: Northern Peru. Locally frequent in thickets, frequently climbing up trees in forest openings, in mid-elevation cloud forest of Dept. Amazonas, east and northeast of Chachapoyas, 2,100-2,600 m; a few collections are also known from Dept. Cajamarca, Prov. Cutervo, 1,900-2,650 m (Fig. 60). Representative specimens examined: Peru, amazonas: 9 km E of Molinopampa on Chachapoyas- Mendoza road, Berry & Escobar 3617 (MO, USM); Km 22 of Carretera Marginal E of Pedro Ruiz towards Rioja, Berry & Escobar 3626 (MO, USM); on road to Pomacochas, 28 km above Puente Ingenio, Hutchison & Wright 6778 (RSA, UC, USM); beyond Buenos Aires, Carretera Marginal, Dtto. Yambrasbamba, Prov. Bongara, Tillett 673-213a (VEN); Mendoza, Woytkowski 8202 (MO); hills WNW of Pomacocha, Wurdack 938 (F, NY, RSA, US), cajamarca: Socata-San Andres, Lopez & Sagdstegui 5372 (MO); Achira, near Socota, Prov. Cutervo, Velarde Nunez 7065 (Z); El Suro, Prov. Cutervo, Velarde Nunez 7022 (Z). 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Northern purplish llary differs from that species in its generally quaternate leaves, less divergent pedicels, more numerous secondary veins, and spreading sepal tips in bud. Its distribution is centered in the moist cloud forests of central Dept. Amazonas, but three col- lections were seen from across the Rio Maranon valley in Dept. Cajamarca that differ in their denser, short pilose pubescence, obtuse leaf tips, pilose styles, and slender floral tubes only 5 mm wide at the rim. Further habit and habitat infor- mation on these plants is necessary, however, before we can decide if these are distinct enough to warrant separate taxonomic recognition. Fuchsia rivularis is sympatric with F. pilosa, but the two differ markedly in habit, and no hybrids have been detected. 30. Fuchsia gehrigeri Munz, Proc. Calif. Acad. Sci. IV. 25:41, pi 5, fig. 28. 1943. type: Venezuela, Edo. Merida, Mucuruba, slopes and banks of La Canada Grande, right fork of the town's stream, 2,800-3,100 m, 14 July 1930, Wilhelm Gehriger 322 (US 1515306, holotype; F, G, GH, MO, NY, PH, VEN, iso- types). Fuchsia jahnii Munz, Proc. Calif. Acad. Sci. IV. 25:40, pi. 5 y fig. 27. 1943. type: Venezuela, Edo. Merida, Paramo de la Sal, 2,800 m, 2 Nov. 1921, Alfredo John 506 (US 1186509, holotype; photographs, NY, POM; GH, VEN, isotypes). Erect to usually scandent-climbing shrubs 2-5 m high, with arching-pendant branches. Young growth sparsely puberulent to subcanescent, rarely densely pilose; branchlets subterete, subglabrous to strigose (rarely reddish pilose); older stems 8-18 mm thick, long and flexuous if climbing, with tan, flaky bark. Leaves mostly ternate, occasionally opposite or quaternate, membranous, (narrowly) el- liptic to slightly (ob-)ovate, acute to obtuse or attenuate at the base, acute to subacuminate at the apex, 3.5-12 cm long, 1.5-5 cm wide, deep matte velvety green and strigillose to subglabrous above, pale dull green to purplish and stri- gillose below, hairs denser along the veins; secondary veins 5-12 on either side of the midvein; margin usually denticulate. Petioles light red, 10-40(-48) mm long. Stipules lanceolate, 1.5-2 mm long, divergent-spreading with age, subpersistent. Flowers axillary and clustered at the branch tips or sometimes corymbose with a rachis 2-6 cm long; flowers, pedicels, and flowering branches pendant. Pedicels loosely strigose, 12-40 mm long. Ovary ovoid, 5-7 mm long, 2-4 mm thick, strigillose to pilose. Floral tube narrowly funnelform, 40-50(-55) mm long, (2.5)3-4 mm wide and bulbous at the base, narrowed to 2-3 mm for the basal 18-22 mm of the tube, then gradually or usually abruptly widened and 8-9 mm wide near the rim, subglabrous to puberulent-strigose outside (rarely pilose), (densely) pi- lose or villous inside in lower 2-4 cm. Sepals lanceolate, acute to acuminate at the apex, ( 13—) 15—21 mm long, (4-)5-6 mm wide, spreading at anthesis. Tube and sepals subnitid red. Petals scarlet, oblong to elliptic-obovate or lanceolate, 14-21 mm long, (4-)6-8 mm wide, obtuse to acute at the apex, margin and surface smooth, often with several hairs on dorsal surface, spreading to slightly recurved at anthesis. Nectary unlobed or slightly 4-lobed, 1.5-2 mm high. Filaments light red, sometimes whitish at the base, 10-14 mm and 8-12 mm long; anthers oblong, 1982] BERRY— FUCHSIA SECT. FUCHSIA 133 2.5-3 mm long, ca. 1.5 mm wide, white. Style red, glabrous to loosely villous; stigma subglobose, 2-2.5 mm long, ca. 2 mm wide, slightly 4-cleft at the apex, dull cream to pink red. Berry ovoid to subglobose, somewhat quadrangular before maturity, 13-16 mm long, 10-13 mm thick, dark red purple at maturity; seeds 2-3 mm long, 1.1-1.4 mm wide. Gametic chromosome number n = 11. Distribution: Venezuelan Andes and the Serrania de Perija along the Colom- bian-Venezuelan border; in the Merida Andes, in Trujillo, Merida, and Tachira; in the Sierra de Perija in Zulia, Venezuela and Cesar, Colombia, 2,200-2,800(-3,100) m, in cloud forest thickets and woods (Fig. 60). Representative specimens examined: Venezuela, merida: vicinity of Pinango, La Pinita, Badillo 921 (VEN); between El Valle and Paramo La Culata, Benitez de Rojas 1523 (MY); bridge of Quebrada El Valle, 7 km above Hotel Valle Grande, Berry & Ruiz-Terdn 2504 (MER, MERF, MO); Mucuruba, Quebrada El Rincon, above the town, Berry & Ruiz-Terdn 2509 (MER, MERF, MO); above Pinango, 4 km below Las Pailitas, 43 km W of Pico El Aguila, Berry 3138 (MO, VEN); 9 km above Santo Domingo on road to Apartaderos, Berry 3139 (MO, VEN), 3310 (MO), 3599 (MO); Mesa de Los Pinos, above La Mucuy, Berry 3440 (MO, VEN), 3442 (MO), 3445 (MO, VEN); Paramo de Pinango, John 402 (US, VEN); Pueblo Llano, Lopez-Palacios 53 (MER); trail from La Escalera to Puente de la Escalera, Luteyn et al. 6212 (MO); above Las Piedras, watershed of Rio Aracay, Ruiz-Terdn et al. 8238 (MERF, MO); between El Arbolito and El Alto, N of Pinango, Ruiz-Terdn & Dugarte 12356 (MERF, MO); between El Chorreron and La Cueva, road to Paramo de Palmita, Ruiz-Terdn & Dugarte 12489 (MERF, MO), 12493 (MO), trujillo: above Jajo, Aristeguieta & Medina 3397 (VEN); 12 km above and W of San Rafael on old Bocono-Trujillo road, Berry 3098 (MO, VEN); ca. 3 km S of Las Mesitas, at Visiin, Rio La Coneja, Berry 3128 (MO, VEN); 4-5 km along old Bocono-Trujillo road, 2 km S of lateral to Burbusay, Tillett 739-573 (MO), zulia: Campamento Frontera II, near international boundary, headwaters of Rio Negro, Sierra de Perija, Tillett & Honig 746-678 (MO, VEN), 746-705 (MO, VEN), 746-766 (MO, VEN): Campamento Frontera V, on international bound- ary with Colombia, headwaters of Rio Guasare, Sierra de Perija, Serrania de Valledupar, Tillett 746- 1033 (MO, VEN), 746-1086 (MO, VEN). Colombia, cesar: Sierra de Perija, E of Manaure, Sabana Rubia, Cuatrecasas & Romero Castanedas 25061 (COL, RSA, US); Sierra de Perija, E of Manaure, Quebrada de Floridablanca, Cuatrecasas & Romero Castanedas 25224 (COL, US). This species is characterized by its thin, denticulate, and rather long-petiolate leaves; its smooth, elliptic-oblong petals with frequent hairs on the back, and the ovoid-globose fruits with large seeds 2-3 mm long. It resembles Fuchsia venusta in its subracemose flowers, similar floral dimension, and frequent hairs on the petals. Since the two occur sympatrically, a list of their distinguishing characters is presented in Table 11. It may also be related to F. petiolaris from Colombia, which has similar leaves, large seeds, and pubescent petals. Considerable leaf and pubescence variation occurs within a small area in dif- ferent populations of F. gehrigeri. Characters such as differing pedicel lengths, hair length, and sepal tip length, however, which Munz used to distinguish Fuchsia jahnii from F. gehrigeri, were shown by field studies conducted near both type localities and in intermediate areas to be too variable to be of any taxonomic importance. Plants that are unusually pilose, with smaller flowers than most populations of F. gehrigeri, are found above the town of Santo Domingo on the road to Apartaderos in Edo. Merida. The hairs on these plants are often gland tipped and become reddish with age. A typical, less pubescent and longer-tubed plant of/ 7 . gehrigeri, Ruiz-Terdn et al. 8238 (MERF, MO) comes from a valley adjacent to Santo Domingo, however, indicating that the populations at Santo Domingo may be very localized variants. Somewhat verrucose floral tubes and pedicels are found in populations north- 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 west of the city of Merida, in the Valle Grande. These plants often have papillae protruding at the base of adjacent sepals. Some introgression may be occurring with F. venusta, because Berry & Ruiz-Terdn 2505 (MERF, MO; Quebrada Valle Grande, 7 km above Hotel Valle Grande) has morphological characters interme- diate between the two species, such as smooth, elliptic petals of F. gehrigeri but subnitid, nearly entire leaves as in F. venusta. It has a pollen stainability of 66.8% (400 grains), compared to 94.7% (400 grains) for Berry & Ruiz-Terdn 2504 (MERF, MO), which is morphologically closer to F. gehrigeri. This valley has long been farmed and grazed by livestock, so these repeated disturbances may have led to an increased level of hybridization. In the same area, hybrids of F. venusta and F. nigricans are also found. In contrast to El Valle Grande, populations of F. gehrigeri and F. venusta occur on virtually undisturbed slopes of primary cloud forest above La Mucuy, in the Par que Nacional Sierra Nevada, on the opposite side of the Rio Chama from Merida. There, a continual vegetation transect can be made following the trail from 2,250 m to 3,350 m at tree line (Fig. 6). Berry 3439 (MO) was collected at 2,630 m and is apparently a hybrid between the lower elevational F. venusta and the mostly higher altitude F. gehrigeri. It has a low pollen stainability (31.5% of 400 grains) and the following intermediate morphological characters: subco- riaceous leaves, matte green above, but nitid below; petioles 8-20 mm long; and orange red flowers with smooth petals. Bernardi 664 (NY; Edo. Merida, Monte Zerpa, 2,600-2,800 m, June 1953) is another probable hybrid between F. gehrigeri and F. venusta. The floral tubes of this collection reach nearly 60 mm, and pollen stainability is 69% (1,000 grains examined). An altitudinal separation similar to that of La Mucuy occurs further south in Edo. Tachira along the road from Zumbador to Queniquea. Fuchsia venusta is again the lower altitudinal species, occurring from 2,400-2,700 m on the eastern side of the ridge above Queniquea. Berry 3295 (MO), 3296 (MO), and 3297 (MO, VEN; n = 11) occur at or just below the crest of the ridge, 4-6 km E of Zum- bador, at 2,850-2,900 m, and resemble F. gehrigeri in their denticulate, long- petiolate leaves; the smooth, elliptic petals; and the ovoid-globose fruits with large seeds. The leaves tend to be broader and more ovate than the normal range of F. gehrigeri, however, and the sepals are strongly divergent, the branches ascending, and the floral tubes only gradually widened. Possible hybrids between these populations and F. venusta were collected on the east side of the ridge, 3 km E of Zumbador, at 2,760-2,775 m. Berry 3414 (MO; n = 11) and Luteyn et al. 5358 (NY) have unusually long floral tubes up to 78 mm long, elliptic, smooth petals, and subnitid, elliptic leaves with wine red stems. The floral tube length is greater than the extremes recorded in either F. venusta or F. gehrigeri, but the other characters are intermediate. Pollen stainability of Berry 3414 is 69% (300 grains examined). Fuchsia gehrigeri is also known to hybridize with F. nigricans, and the four putative hybrid collections are discussed under that species. It is also sympatric with F. verrucosa, but does not hybridize with that species. A disjunct series of populations occurs in the Serrania de Perija, along the Colombian- Venezuela border. The specimens from this area are very similar to those from the Merida Andes except for the following differences: usually nar- 1982] BERRY— FUCHSIA SECT. FUCHSIA 135 rower leaves with narrowly cuneate to attenuate bases, some petioles as long as 48 mm, more freely exfoliating bark, and generally more corymbose flowers. Petal shape varies in the few collections available from elliptic to lanceolate, but they are generally narrower, and the tips more acute, than plants from the main part of the range. None of the petals have dorsal hairs, however, which are usually found in populations from Merida and Trujillo. More complete collections and additional field data from Perija may in the future lead to the recognition of these populations as a separate taxon closely allied to F. gehrigeri. 31. Fuchsia llewelynii J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):556. 1941. Munz, Proc. Calif. Acad. IV. 25:38, pi 5, fig. 25. 1943. type: Peru. Dept. Amazonas, La Ventana, path from Chachapoyas to Moyob- amba, among shrubs on exposed rocky slopes, 2,700-3,300 m, 21 Jan. 1930, Llewelyn Williams 7594 (F 626222, holotype; photographs, NY, UC). Low shrubs with puberulent or canescent branchlets with spinescent projec- tions, usually appearing papillose or verrucose when dry. Leaves opposite or ternate, membranous to subcoriaceous, narrowly oblong to elliptic or oblanceo- late, acute at both ends, 2.5-9(-20) cm long, 1.2-4 cm wide, glabrous to strigillose on both surfaces; secondary veins 9— 13(— 25) on either side of the midvein; margin conspicuously dentate or serrulate. Petioles 3-9 mm long. Stipules narrowly lan- ceolate or filiform, 2-5 mm long, subpersistent. Flowers few and axillary at the branch tips or subracemose. Pedicels slender, puberulent, verrucose or with spi- nescent protuberances, 2-6 cm long. Ovary narrowly oblong, 5-6 mm long, ca. (40-)45 3-4 4-8 nectary, then widened above until 5-8 mm wide at the rim, subglabrous to pu- berulent outside, pilose inside in lower %. Sepals narrowly lanceolate, acuminate, 13-17 mm long, 4-8 mm wide, with a tip ca. 2 mm long in bud. Tube and sepals dull pink. Petals pink, narrowly lanceolate to elliptic, acute to acuminate, 15-20 mm wide. Nectary unlobed, ca. 2 mm high. Filaments 12-13 mm and 10-1 1 mm long; anthers oblong, 2.5-3 mm long, 1-1.5 mm wide. Style densely pilose for most its length; stigma subobconic, ca. 2 mm long, 1.5-2 mm wide. Berry ellipsoid, 12-15 mm long, ca. 8 mm thick. Distribution: Northern Peru. Known from the mountains near La Ventana and Almirante, east of Chachapoyas and in the Serrania de Bagua, Dept. Amazonas, 2,600-3,300 m (Fig. 60). Specimens examined: Peru, amazonas: Cordillera Colan, SE of La Peca, Prov. Bagua, Barbour 3828 (MO); Almirante, Mathews 1480 (K, OXF); path between Chachapoyas and Moyobamba, We- berbauer 4437 (G). This species is apparently very rare and localized, as evidenced by the few specimens collected. The leaves are mostly narrowly oblanceolate and conspic- uously toothed, but the most characteristic features are the long pedicels and unusual stems, which have numerous papillose or spinescent projections. The flowers are borne subracemosely, and the petals are slender and more or less recurved, characters that ally it to F. venusta and F. rivularis. 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 32. Fuchsia scherffiana Andre, Rev. Hort. 60:233, 268. 1888. type: Ecuador, Prov. Zamora-Chinchipe, Alto Cruz Grande (4°32'S, 79°02'W), 19 Nov. (?) 1876, Edouard Andre (K, holotype). This is probably an H. Poortman col- lection, since he collected for Andre in southern Ecuador after Andre re- turned to Europe in Sept. 1876 (L. B. Smith, pers. comm.). Erect to scandent shrubs 0.5-3 m tall with spreading branches. Branchlets canescent-strigillose, terete, 1-2.5 mm thick, purplish; older stems 3-10 mm thick, dull purple. Leaves opposite or occasionally ternate, firmly membranous, (very) narrowly elliptic to narrowly lanceolate, acute at the base, acute to acuminate at the apex, 2.5-10 cm long, 0.8-2.3 cm wide, (very) dark nitid green and strigose above, shiny green to metallic purple and strigose with long villous hairs along the mid vein below; secondary veins 6-10 on either side of the mid vein; margin subentire to slightly revolute. Petioles strigose, purplish, 4-15 mm long. Stipules lance-linear, dark, 1-2 mm long, ca. 0.3 mm wide, subpersistent. Flowers few and axillary, pendant. Pedicels strigillose, 15-20 mm long. Ovary oblong, strongly tetragonous, 7-8 mm long, 1.5-2 mm thick, greenish. Floral tube narrowly fun- nelform, 44-50 mm long, 2.5-4 mm wide and slightly bulbous at the base, nar- rowed to 2-2.5 mm wide above the nectary, gradually widened above until 7-8 mm wide at the rim, sparsely strigose outside, villous inside in lower Vi. Sepals lanceolate, acuminate, 12-14 mm long, 3.5-5 mm wide, spreading at anthesis. Tube and sepals orange red. Petals red, narrowly elliptic-lanceolate, (narrowly) acute at the apex, 9-11 mm long, 3-5 mm wide. Nectary green, unlobed, ca. 1 mm high. Filaments red, 8-10 mm and 6-8 mm long; anthers oblong, 3-3.5 mm long, ca. 1.5 mm wide, cream. Style red, villous for ca. 3 A its length from the base; stigma globose, ca. 2 mm wide, 1.5-1.8 mm wide, reddish. Berry oblong, tetragonous, green before maturity, ca. 15 mm long, ca. 6 mm thick; mature fruits not seen. Distribution: Southern Ecuador. Known only from the Nudo de Sabanilla area near the border of Loja and Zamora-Chinchipe Provinces, ca. 2,800 m (Fig. 60). Specimens examined: Ecuador, loja: road from Yangana to Nudo de Sabanilla, S of Yangana, 1 km below top of ridge, Escobar 1547 (MO), 1547- A (MO). This is one of several localized, endemic species in southern Ecuador, in- cluding Fuchsia steyermarkii and F. lehmannii. It is characterized by its dark, metallic-colored, narrowly elliptic-lanceolate leaves, purplish stems, and long, axillary flowers with narrow petals. These characters place it closest to the F. venusta species group, and it is probably closely related to F. llewelynii of north- ern Peru. Sandeman s.n. (K; Aug. 1938) from Almirante, Dept. Amazonas in northern Peru possibly belongs in this species. It is somewhat less pubescent than the Ecuadorian plants, with less markedly angled ovaries and shorter pedicels, but agrees well in other floral and foliar characters with this species. Nearly mature fruits are present in this collection; the berries are ellipsoid, 13 mm long, and 7 mm thick. The seeds measure ca. 1.5 mm long and ca. 1 mm wide. 33. Fuchsia confertifolia Fielding & Gardner, Sert. PI. pi. 28. 1844. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):550. 1941. Munz, Proc. Calif. Acad. Sci. 1982] BERRY— FUCHSIA SECT. FUCHSIA 137 IV. 25:42, pi. 6, fig. 30. 1943. type: Peru, Dept. Amazonas, Bagasan (on path from Molinopampa to Moyobamba), 1830-1841, Andrew Mathews 1478 (OXF, holotype; photograph, MO; K, isotypes). Fuchsia dolichantha K. Krause, Repert, Spec. Nov. Regni Veg. 1:172. 1905. type: Peru, Dept. Ama- zonas, Tambo Ventilla, E of Chachapoyas, 2,400-2,600 m, 1904-1921, August Weberbauer 4390 (B, holotype, destroyed in World War II; photograph, F). Densely-branched shrubs 1-2.5 m tall. Branchlets terete, 1.5-3 mm thick, usually less than 15 cm long, conspicuously ferrugineous-hirsute; older branches with red brown bark exfoliating in slender strips. Leaves densely crowded on branchlets and upper portions of older stems with internodes mostly 2-5 mm long, 2-4 leaves per node, usually ternate, subcoriaceous, elliptic-ovate, rounded to acute at the base, acute at the apex, 8-12 mm long, 3-5 mm wide, glabrous 0-1 4-5 midvein; margin remotely glandular-denticulate or slightly revolute. Petioles hir- sute-villous, 1-2 mm long. Stipules dark, filiform, 2-3.5 mm long, ca. 0.3 mm wide, persistent, often connate and divergent to reflexed. Flowers mostly 2-8, drooping, grouped in uppermost axils at the branch tips. Pedicels hirsute, 9-15 4-5 2-4 at the base, narrowed to 1.5-3 mm wide above the nectary, gradually widened above until 7-10 mm wide at the rim, glabrous to sparsely pilose outside, pilose inside for most its length. Sepals narrowly lanceolate, acuminate, 13-18 mm long, 4-5 mm wide, with tips connate into a tip ca. 2 mm long in bud. Tube and sepals red. Petals red, narrowly triangular with long, tapered tips, 9-14 mm long, 2.5-3 mm wide, X A- V A shorter than the sepals. Nectary unlobed, ca. 1.5 mm high. Filaments red, 6-10 mm and 4-7 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide. Style red, densely pilose for most its length; stigma subobconic, 4-cleft at the apex, 2-3 mm long, ca. 2 mm wide, exserted 4-5 mm beyond the anthers. Berry ellipsoid-ovoid, ca. 10 mm long, 5-7 mm thick; seeds tan, 1.8-2.1 mm long, 1-1.2 mm wide. Distribution: Northern Peru. Endemic to cloud forest near high ridges of the mountains east of the Rio Utcubamba in Dept. Amazonas, 2,600-3,200 m (Fig. 60). Specimens examined: Peru, amazonas: between Jumbilla and San Carlos, Prov. Bongara, Weber- bauer 7153 (F, GH, POM); S side of Molinopampa-Diosan Pass, N of Molinopampa, Wurdack 1605 (F, NY, RSA, UC, US, USM). This species is very distinctive in its densely crowded, small, leathery leaves and rust-colored hirsute stems. Its affinities are unclear, but it is placed in the F. venusta species group because of the coriaceous leaves, long, subracemose flow- ers, and somewhat ridged, delicate petals. It occupies the same restricted area as F. pilosa, F. llewelynii, and F. wurdackii, yet it is probably not sympatric with any of these species since it occurs at higher elevations. fig Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):553. 1941. Munz 1 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Acad. Sci. IV. 25:23, pi. 2, fig. 9. 1943. type: Peru, fci Cenchin, in cultis," probably Dept. Lima, 1778-1788, Hipolito Ruiz & Jose Pavon (MA N° 11/ 86, lectotype, here designated; photograph, MO). In the protologue, the lo- cality is given as "Huassa-huassi et Cheuchin. ,, The first site is in Junin, and the second apparently corresponds to the "Cenchin" on the lectotype label. Figs. 19, 42. Fuchsia serratifolia Ruiz & Pavon, Fl. Peruv. Chil. 3:86, pi. 325, fig. b. 1802. Hook., Bot. Mag. /. 4174. 1845. Planch., Fl. Serres Jard. Eur. 5: /. 447. 1849. Sweet, Ornam. Fl. Gard. 2: /. 88. 1854. Essig, Nat. Hort. Mag. 13:12, photo. 1934. type: Peru, Dept. Huanuco, Muna, 1778-1788, Hi- polito Ruiz & Jose Pavon (MA, lectotype, here designated; photograph, MO). Fuchsia leptopoda K. Krause, Repert. Spec. Nov. Regni Veg. 1:171. 1905. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):555. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:24, pi. 2, fig. 10. 1943. type: Peru, Dept. Junin, Prov. Tarma, between Palca and Huacapistana, 2200-2600 m, 1904-1921, August Weherhauer 1772 (B, holotype, destroyed in World War II; photographs, F, POM; G, isotype). Fuchsia siphonata K. Krause, Repert. Spec. Nov. Regni Veg. 1:171. 1905. type: Peru, Dept. Junin, Prov. Tarma, mountains E of Huacapistana, 1904-1921, August Weherhauer 2178 (B, holotype, destroyed in World War II; photograph, F). Fuchsia tacsoniiflora K. Krause, Repert. Spec. Nov. Regni Veg. 1:172. 1905. type: Peru, Dept. Lima, near railroad from Lima to La Oroya, along streams above San Mateo, 3200 m, 1904-192 1 , August Weherhauer 252 (B, holotype, destroyed in World War II; photograph, F). Erect to scandent shrubs 1.5-4 m tall or climbing in trees to 10 m above ground. Young growth subcanescent or occasionally pilosulous; branchlets terete, subdivaricate, green to wine red; older branches 5-20 mm thick with tan, exfo- liating bark. Leaves 3-5-verticillate, mostly ternate or quaternate, rarely oppo- site, firmly membranous, (narrowly) elliptic to oblanceolate, acute to narrowly cuneate at the base, acute to subacuminate at the apex, 4-17 cm long, 1.5-6.5 cm wide, dull to nitid dark green and glabrous above, pale green and subglabrous to strigose mostly along veins and margins below; secondary veins 7-17 on either side of the midvein, often reddish below; margin denticulate. Petioles glabrous to loosely strigose, (5-)8-20(-25) mm long. Stipules triangular, firm, often con- nate, 2-2.5 mm long, ca. 1.5 mm wide, deciduous. Flowers few to numerous, axillary and pendant, usually grouped toward branch tips. Pedicels stout, smooth, 1-1.5 mm thick, 18-45 mm long, suberect in bud, drooping at anthesis, generally green. Ovary narrowly oblong, terete, 10-13 mm long, 3-4.5 mm thick, generally glabrous, green. Floral tube subcylindric, firm, walls 1-1.5 mm thick, smooth, (28-)36-47 mm long, (3-)4— 8 mm wide at the base, sometimes very slightly nar- rowed above the nectary, straight or slightly widened above until (5-)6-12 mm wide at the rim, glabrous to puberulent outside, densely villous inside above the nectary for 7-10 mm, glabrous above. Sepals lanceolate, acuminate, 17-26 mm long, 4-7 mm wide, forming a tapered point in bud, spreading to subdivergent at anthesis. Tube waxy light pink, lavender, or light red; sepals pink to light red with light green to whitish tips or margins, at times entirely whitish green. Petals orange to scarlet, usually drying purple streaked, lance-oblong to oblanceolate, obtuse to broadly acute at the apex, slightly undulate, 14-18 mm long, 4-6(-7) mm wide, suberect at anthesis. Nectary green, unlobed, 2.5-3 mm high, ca. 1.5 mm thick. Filaments pink to light red, 14-23 mm and 8-18 mm long; anthers oblong, 4-6 mm long, 2-3 mm wide, white. Style stout, pink to light red, glabrous; stigma subclavate, subentire (very slightly 4-cleft at the apex), 3-4 mm long, 2-3.5 mm wide, dull white, exserted 2-12 mm beyond the anthers. Berry ellipsoid, 1982] BERRY— FUCHSIA SECT. FUCHSIA 139 20-26 mm long, 10-12 mm thick, nitid green to red purpl seeds tan. 1.8-2.2 mm lone. ca. 1 mm wide. Gametic chromosc 11. Peru the Pacific slopes of the Cordillera Occidental of Peru in Lima and Ancash, near springs and in moist canyons, 2,800-3,500 m; on the eastern slopes of the Peruvian Andes » from Huanuco to Cuzco, in cloud forest and moist upland shrub vegetation, 2,500-3,400 m; and on the northeastern slopes of the Bolivian Andes in Depts. La Paz and Cochabamba, in cloud forest, 2,200-3,100 m (Fig. 61). Representative specimens examined: Peru, ancash: Racran, near Chiquian, Ferreyra 6208 (MO, US); Chiquian, Ferreyra 7435 (F, MO, RSA, US); Km 23, Caraz, Prov. Huaylas, Lopez-Guillen 1904 (RSA); Caraz to Laguna de Paron, Lopez et al. 8348 (MO); above Monterrey, 3 km below Huaraz, Pennell 15305 (PH). ayacucho: 41-47 km E of Tambo on road to Ayna, Berry 3050 (MO, USM), 3052 (MO, USM); above Yuraccyacu on Capprichio-Puncu trail, ca. 40 km NE of Tambo, Madison 10319-70 (MO, US, USM); above Jano, from Tambo to Ayna, Plowman & Davis 4678 (GH, K); Yanamonte, between Tambo and Rio Apurimac, Weberbauer 5641 (F, GH, US); Choimacota valley, Weberbauer 7587 (F, G, GH, S), 7587a (F). cuzco: Km 156-165 of Cuzco-Quillabamba road, Prov. La Convencion, Berry 2573 (CUZ, MO), Berry & Aronson 3042 (MO, USM), 3043 (MO), 3048 (MO); Lucumayo valley, Cook & Gilbert 1356 (US); Yuncaipata, Santa Rita, Prov. Convencion, Vargas 2656 (CUZ, MO); heights of Pintobamba, Vargas 3551 (CUZ); Quellomayo-Lucumayo, Vargas 4480 (BN, CUZ); Chawares, Prov. Convencion, Vargas 20237 (CUZ); above Alfamayo, Vargas 22778 (MO), huancavelica: Salcabamba, Prov. Tayacaja, Stork & Horton 10261 (F, UC); Montepungo, 5 km E of Surcubamba, Prov. Tayacaja, Stork & Horton 10367 (F, G, NA, UC). huanuco: Carpish, Ferreyra 1218 (MO, USM), 1730 (USM), 1832 (USM), 2088 (F, MO, NY, US, USM), 2387 (MO, US, USM); Mitobamba, above Mito, Ferreyra 6683 (MO, US, USM); Carpish, above Acomayo, Hutchison et al. 5921 (F, MO, NY, RSA, UC, US, USM); between Huanuco and Pampayacu, Kanehira 236 (GH); Km 29-35 NW of Hacienda Mitobamba, Huanuco-La Union road, Luteyn & Lebron-Luteyn 5501 (MO, NY); Muiia, trail to Tambo de Vaca, Macbride 4284 (F, GH, US); Mito, Macbride & Featherstone 1403 (F, G, GH, US); ca. 23 km SE of Huanuco, Macbride & Featherstone 2082 (F, GH, US); Mima, Pearce 141 (K); Pillao, Ruiz & Pavon 11/85 (F); Pampayacu, Sawada P.8 (F); Pillao, Woytkowski 34163a (F, UC); Saraypampa, Woytkowski 34194 (F, MO, UC). juni'n: 4 km W of Comas, road Concepcion-Satipo, Berry & Aronson 3065 (MO, USM); Comas, Berry & Aronson 3066 (MO, USM); 69 km above and W of Satipo on road to Concepcion, Berry & Aronson 3072 (MO, USM); 62 km W of Satipo to Concepcion, Berry & Aronson 3076 (MO, USM); Palca, near Tarma, Ferreyra 18738 (USM); Chilifruta, Maguire & Maguire 61646 (NY); Pangoa, Mathews 1168 (K); above Huacapistana, Sandeman 4508 (K, OXF); Chaca, 5 km from Comas, Vargas 22045 (CUZ). lima: Mani, near Huascoy, Acleto 30 (MO, USM); between Infiernillo and Rio Blanco, Asplund 10846 (S, US); Llacshishi, Prov. Huarochiri, Ferreyra 682 (MO, USM); near Palacala, above Surco, Ferreyra 3415 (MICH, MO, US); between Huascoy and Cormo, Ferreyra 18392 (MO); Infiernillo, Goodspeed et al 11549 (G, GH, NA, UC); Quebrada de San Mateo, Isern 2546 (F); San Mateo, 1 km E of city, Hutchison 669 (GH, UC, US, USM); Rio Blanco, Macbride & Featherstone 723 (F, GH); Huamatanga, Mathews 541 (CGE, K, OXF); San Buenaventura, Nee s.n. (MA 183589); Cerros de Matucana, Raimondi 188 (USM). pasco: Vitoc, Martinet 1409 (P). without locality: Ruiz & Pavon s.n. (BM, CGE, F, G, MA, MO). Bolivia, cochabamba: ca. 42 km from Cochabamba to Chapare, Berry 2584 (MO), 2585 (MO), 2586 (MO); near Chulumani on way to Yungas de Tablas, Cardenas 4087 (F, GH, RSA); Plumerito, to Rio Tablas, Cardenas 6282 (US), la paz: road to Chulumani, Albert de Escobar 1305 (TEX); Yungas, Bang 731 (F, GH, NY, PH, US); near Zongo, 32 km from crest of ridge, Beck 2172a (MO); Unduavi, Brooke 6588 (BM, F, G, NY, U), 6850 (BM); Buchtein 143 (F, G, GH, NY); Zongo valley, Holliday 2111 (K); San Felipe, Sur Yungas, Holway & Holway 632 (US); Km 50 La Paz-Chulumani, Kelley 1064 (BM, UC); near Combaya, Prov. Larecaja, Mandon 623 (GH); Rio Aceramarca, Cordillera Real, Tate 718 (NY). This is the most common and widespread member of the Fuchsia denticulata species group, an alliance of thick-tubed, axillary-flowered species centered in the Central Andes. It is characterized by its firm, subcylindrical flowers with waxy pink or light red floral tubes and narrow, green-tipped sepals, and also by the terete, ellipsoid fruits, stout pedicels, purple drying petals, large anthers 4-6 mm long, and nearly unlobed, clavate stigma. 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 61. Distribution of the Fuchsia denticulata species group Fuchsia denticulata is the only member of the genus present on the dry Pacific slopes of the Cordillera Central in central Peru. It occurs there locally in isolated humid vegetation pockets, usually growing along streams, springs, or waterfalls. Plants from Ancash and Lima, such as the one used to describe F. tacsoniiflora, 1982] BERRY— FUCHSIA SECT. FUCHSIA 141 Table 12. Comparison of diagnostic morphological characters of Fuchsia denticulata, F. aus lontana. and intermediate Paucartambo collections. 1 Leaf length Petiole length Floral tube texture Floral tube color Difference between widest and narrowest part of tube Sepal angle at anthesis Stigma color Fruit transection F. denticulata 4-17 cm (5-)8-20(-25) mm Smooth Pink to light red 2-3 mm Spreading Cream Terete 1 See text for list of specimens. Paucartambo Collections 1 5-13 cm 7-12 mm Tuberculate Dark red 5-6 mm F. austromontana 2.5-8 cm 3-12 mm Subtuberculate Dark red 4-6 mm Spreading-recurved Spreading-divergent Red pink Terete (?) Red Quadrangular generally appear more robust, with thicker stems than plants of this species grow- ing on the eastern slopes of the Andes. This is probably due, however, to their upright, shrubby habit in the sparse vegetation of the Pacific slopes, compared to the typically scandent habit of F. denticulata in the dense thickets of cloud forests on the eastern slopes. This east- west disjunction of F. denticulata in the Peruvian Andes is discussed in greater detail on p. 26. Plants with narrow leaves and floral tubes like the type of F. leptopoda appear sporadically throughout Junin and Ayacucho, but they intergrade completely with more typical populations and maintain characteristic traits of F. denticulata such as green sepal tips and purple-streaked petals when dry. In cloud forests along the eastern slopes of the Andes, Fuchsia denticulata can be found as an erect shrub, a scandent shrub, or a high climbing vine. This wide variability in habit type accounts for most of the variability found in leaf size, pubescence, and floral dimensions, which Krause and Ruiz and Pavon, used to describe several of the taxa included in the above synonymy. There is a gap of some 500 km between the nearest collections from Peru and Bolivia. The related F. austromontana occurs in the intervening areas in south- ernmost Peru, at higher elevations than would be expected for F. denticulata. rm 500 Peru Berry et al. 2595 (MO, USM), 2598 (MO, USM (MO, USM) 14047 (PH): Vareas 10 (F): Weberbauer We Hemsl meters lower than nearby populations of F. austromontana (Fig. 11). A com- parison of these plants with F. denticulata and F. austromontana is presented in Table 12, showing their combination of distinctive and intermediate traits. Munz (1943) included some of the above specimens in F. austromontana', con- rph southern Dept. Puno) is needed to deten denticulata occurs in that area. 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia denticulata occurs sympatrically with F. abrupta, F. corymbiflora, F. decussata, F. ferreyrae, and F. sanctae-rosae. A Bolivian collection, Beck 1266 (MO), is a probable hybrid with F. sanctae-rosae and is discussed under that species. 35. Fuchsia austromontana I. M. Johnston, J. Arnold Arbor. 20:242. 1939. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):548. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:22, pi. 1 ,fig. 8. 1943. type: Peru, Dept. Cuzco, Prov. Paucartambo, between Pillahuata and Acanacu, 2,800 m, 26 July 1936, James West 7083 (UC, holotype; photographs, NY, POM; GH, isotype). Erect to occasionally climbing shrub 2-4 m tall. Young growth subcanescent to whitish pilose; branchlets subglabrous to puberulent, purplish, older branches with tan-brown, splitting bark. Leaves 3-5-verticillate, firmly membranous, acute to obtuse at the base, mostly acuminate at the apex, 2.5-8 cm long, 1-3 cm wide, dark green and subglabrous above, pale green and pubescent below, hairs usually denser along the midvein; secondary veins 5-9 on either side of the midvein; margin subentire to denticulate and sometimes slightly revolute. Petioles 3-12 mm long, reddish, densely pilose to lightly pubescent. Stipules linear-lanceolate, 2-4 mm long, 0.5-1 mm wide, subpersistent. Flowers generally few and pendant from upper leaf axils. Pedicels usually tuberculate, red, 14— 35(— 55) mm long. Ovary tetragonous, 6-8 mm long, 1.5-2 mm thick. Floral tube narrowly funnel- form, walls ca. 1 mm thick, (25-)34-50 mm long, 4-6 mm wide at the base, then narrowed to 2-5 mm wide for lower Vs, widened above until 10-12 mm wide at the rim, often ± verrucose, glabrous to densely pilose outside, (densely) villous inside in lower l A- l A. Sepals lanceolate, acute to acuminate, thick-spongy, ca. 1.5 mm thick when fresh, often verrucose, 15-20 mm long, 5-7 mm wide, spread- ing to divergent at anthesis. Tube and sepals scarlet, the tips sometimes dull purple green. Petals deep red, usually drying a uniform purple, elliptic-obovate to suborbicular, 12-17 mm long, (7— )8— 13 mm wide, rounded to broadly acute at the apex, spreading at anthesis. Nectary deeply 4-lobed to unlobed, ca. 2 mm high. Filaments red, 10-14 mm and 7-10 mm long; anthers oblong, 3.5-4.5 mm long, 2-3 mm wide, white. Style red, glabrous or with a few hairs; stigma sub- globose, 4-cleft at the apex, 2-2.5 mm long, ca. 3 mm wide, red pink, exserted 1-10 mm beyond the anthers. Berry ellipsoid, reddish, tetragonous before ma- turity, 16-18 mm long, 8-11 mm thick; seeds tan, 1.4-1.7 mm long, 0.8-1.1 mm wide. Gametic chromosome number n = 11. Distribution: From southern Cuzco Dept. in Peru to just beyond the Bolivian border in Dept. La Paz, Bolivia; upper cloud forest shrubs, 2,600-3,500 m (Fig. 61). Representative specimens examined: Peru, cuzco: Acanaco, Balls B6708 (BM, K, UC, US); Km 145 of Cuzco-Quillabamba road, Berry 2568 (CUZ, MO); 3035 (MO, USM); 8 km E of Acanaco Pass, Km 101 of Cuzco-Pilcopata road, Berry et at. 2594 (MO, USM); between town of Marcapata and the banos termales, Berry & Aronson 3025 (MO, USM); Km 143-152 of Ollantaytambo-Alfamayo road, 58-67 km NW of Ollantaytambo, Luteyn & Lebron-Luteyn 6451 (MO); region of Acanacu and Cor- dillera de Tres Cruces, Luteyn & Lebron-Luteyn 6372 (NY); Pillahuata, Cerro de Cusilluyoc, Pennell 14110 (F, GH, PH, US); Km 144 from Ollantaytambo to Chaullay, Plowman & Davis 4749 (GH, K); Huaillai, Marcapata, Vargas 1351 (BH, MO); between Achirani and Medias-Mayu, Prov. Paucartam- 1982] BERRY— FUCHSIA SECT. FUCHSIA 143 bo, Dtto. Marcachea, Vargas II 121 (F, UC). Bolivia, la paz: Italaque, Prov. Camacho, Cardenas 3846 (LIL, POM). This species is closely allied to Fuchsia denticulata, in a group of species with large, axillary flowers and thick floral tubes. Fuchsia austromontana grows at higher elevations than F. denticulata and has rounder, wider petals, tetrago- nous ovaries, and darker, somewhat verrucose flowers. Fuchsia austromontana has a restricted range in southern Peru and just over the border into Bolivia. Populations that seem to intergrade with F. denticulata at intermediate elevations occur in Prov. Paucartambo, Dept. Cuzco, and are discussed under F. denticulata (Table 12 and Fig. 11). Another variant is Metcalf 30548 (A, BH, F, G, MO, UC, US; Peru, Dept. Puno, Prov. Sandia, 9 km from Limbano at Chamacani, 2,700 m); it resembles F. austromontana except for its more acute, narrower petals (15 mm long, 6-7 mm wide), strongly spreading sepal tips, elongate ovaries, and dense pubescence. More collections of the F. denticulata species group are needed from southern Peru to better evaluate the pattern of variability in the species that occur there. 36. Fuchsia harlingii Munz, Aliso 7:409. 1972; Opera Bot., Ser. B, 3:16. 1974. type: Ecuador, Prov. Loja, 14 km S of Saraguro, 3,000 m, 1-3 Aug. 1959, Gunnar Hading 6192 (S, holotype; photograph, MO). Fig. 25. Fuchsia fosbergii Munz, Aliso 7:409. 1972; Opera Bot., Ser. B, 3:15. 1974. type: Ecuador, Prov. Loja, Cruz Loma, Cerro Villonaco, 6 km W of Loja, 2,980 m, 14 Feb. 1945, Francis R. Fosberg & M. A. Giler 23036 (RSA 83773, holotype; US, isotype). Erect to scandent shrubs 1-3 m tall. Branchlets subterete, 2-4 mm thick, glabrous or rarely strigose- villous; older branches with lustrous, purple brown bark, exfoliating in wide strips with age. Leaves opposite or ternate, firmly mem- branous to subcoriaceous, narrowly elliptic to elliptic-ovate, acute to rounded at the base, acute to acuminate at the apex, 3-7 mm long, 1.5-2.5 cm wide, glabrous and subnitid dark green above, pale green and glabrous to strigose- villous below; secondary veins 4-6(-7) on either side of the mid vein, sometimes reddish below; margin glandular-serrulate. Petioles 3-6(-10) mm long. Stipules lance-deltoid, thick at the base, sometimes connate or recurved, 1-2 mm long, ca. 1.5 mm wide, subpersistent. Flowers few, pendant, and solitary in the upper leaf axils. Pedicels stout, 1-2 mm thick, 10-15 mm long. Ovary tetragonous, 6-7 mm long, 3-6 mm thick. Floral tube cylindric to narrowly funnelform, the base much wider than the ovary, (35-)44-53 mm long, very firm, 2-3 mm thick when fresh, (3-)4-9 mm wide at the base, 7-12 mm wide at the rim, gradually widened from the base to the rim or more often with subparallel sides, glabrous to villous outside, villous inside in lower Vi. Sepals lance-oblong, 13— 17(— 21) mm long, 5-8 mm wide, 2-2.5 mm thick when fresh, spreading at anthesis, the tip blunt or less often acute in bud. Tube and sepals pale red to orange. Petals red, darker than the sepals, broadly elliptic-ovate, obtuse at the apex, 10-15 mm long, 7-11 mm wide, sub- erect at anthesis. Nectary shallowly 4-8-lobed, 1-2 mm high. Filaments red, 11-14 mm and 7-10 mm long; anthers oblong, 3-5 mm long, ca. 2 mm wide, cream. Style densely villous from the base to near the rim of the tube; stigma capitate, 4-cleft apically, 3-5 mm long, 3-5 mm wide, red. Mature berry not seen. 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Distribution: Southern Ecuador, rare in cloud forest of Loja and Azuay Provinces, 2,600-3,300 m (Fig. 61). Specimens examined: Ecuador, azuay: Paramo and subparamo N and NW of Paramo de Castillo, 6-8 km NNE of Sevilla de Oro, Camp E-5184 (NY), loja: 10-12 km S of Saraguro, Berry & Escobar 3193 (MO, QCA), 3195 (MO), 3206 (MO), 3207 (MO); Nudo de Guagrauma, S of Saraguro, Correll E4I5 (LL); road from Loja to Saraguro, Dodson & Thien 587 (DS); Cerro Arcana, Espinosa & Williams 2490 (RSA); Km 51 on Panamerican Highway N of Loja, Holm-Nielsen et al. 4701 (AAU, S); Saraguro, Humboldt & Bonpland s.n. (P); 12 km S of Saraguro, King & Atmeda 7840 (MO); Chuquirbamba, Poortman 445 (P). santiago-morona or zamora-chinchipe: between Loma de Galapagos and headwaters of Rio Tintas, Steyermark 53462 (NY), without locality: Lobb s.n. (K). This is a member of the thick-tubed, axillary-flowered Fuchsia denticulata species group and is distinctive in its firm, short petiolate, few- veined leaves, the short pedicels, and the smooth, purplish stems. Collections from the type locality near Saraguro, Loja, are usually glabrous with cylindric flowers that are broad at the base and blunt-tipped. Plants from other localities, however, have flowers narrower at the base with narrowly funnelform floral tubes and acute tips. This difference in tube shape was used by Munz (1972) to describe F. fosbergii; at that time Munz had seen only one other sheet of F. harlingii. Berry & Escobar 3194 (MO), from near Saraguro, is densely hirsute, yet was collected from a population of totally glabrous plants. Camp E-5184 (NY), the northernmost col- lection from Azuay, is also unusual in its strigose- villous pubescence. Fuchsia harlingii is sympatric with the more widespread F. loxensis, but no hybrids between the two were detected. 37. Fuchsia cochabambana P. Berry, sp. nov. type: Bolivia, Dept. Cochabamba, Prov. Chapare, Km 104 of Cochabamba- Villa Tunari road, 3,100 m, 18 Dec. 1966, Roy F. Steinbach 632 (US 2533496, holotype; F, LAM, MO, NY, U, isotypes). Frutex 0.5-1.5 m altus, ramulis foliisque glabratis vel puberulentis. Folia ternata vel interdum quaterna, subsessilia, elliptica vel anguste elliptico-lanceolata, basi acuta vel obtusa, apice acuminata, 3-12 cm longa, 1-5 cm lata, nervis secundariis utroque latere 11-14, margine valde denticulata vel serrulata; petiolis 1-3 mm longis; stipulis subpersistentibus, triangularibus, 2-3 mm longis, ca. 2 mm latis. Flores aurantiacorubri vel vivide coccinei, dependentes vel nutantes, axillares vel subracemosi ad apicem ramorum congesti; pedicellis 7-30 mm longis; ovario oblongo, 5-7 mm longo. Tubi florales anguste infundibuliformes, 45-58 mm longi, basi ca. 3 mm lati, superne sensimdilatati summo 7-10 mm lati. Sepala lanceolata acuminata 16-18 mm longa, ca. 5 mm lata. Petala rubra, in sicco saepe pur- purascentia, elliptica, 10-13 mm longa, 5-8 mm lata, apice obtusa vel late acuta. Filamenta antisepala ca. 10 mm longa, antipetala 6-7 mm longa; antheris oblongis, ca. 3 mm longis, ca. 2 mm latis. Stylus ruber glaberque, stigmate rubro globoso apice leviter 4-fisso, ca. 3 mm longo, ca. 2 mm lato. Bacca matura non visa. Shrubs 0.5-1.5 m tall. Young growth subglabrous to puberulent. Leaves ter- nate or less often quaternate, subsessile, firmly membranous, mostly elliptic or narrowly elliptic-lanceolate, acute to obtuse at the base, (sub-)acuminate at the apex, 3-12 cm long, 1-5 cm wide, subglabrous above, subglabrous to usually puberulent along the veins below and usually purple-flushed; secondary veins 11-14 on either side of the midvein; margin conspicuously dentate or serrate. Petioles 1-3 mm long. Stipules triangular, membranous when young to thick and recurved when old, 2-3 mm long, ca. 2 mm wide, subpersistent. Flowers pendant, few to numerous, axillary or subracemose, but always tightly grouped at the 1982) BERRY— FUCHSIA SECT. FUCHSIA 145 branch tips. Pedicels 7-30 mm long. Ovary oblong, 5-7 mm long, ca. 2 mm thick. Floral tubes narrowly funnelform, 45-58 mm long, ca. 3 mm wide and slightly bulbous at the base, narrowed to ca. 2 mm wide above the nectary and gradually widened above until 7-10 mm wide at the rim, subglabrous to pubescent outside, pilose inside in lower Vi. Sepals lanceolate, acuminate, 16-18 mm long, ca. 5 mm wide, with a narrow tip 2.5-3.5 mm long in bud. Tube and sepals orange red to bright crimson. Petals red, often drying purplish, elliptic, 10-13 mm long, 5-8 mm wide, obtuse to broadly acute at the apex. Nectary unlobed, ca. 2 mm high. Filaments light red, ca. 10 mm and 6-7 mm long; anthers oblong, ca. 3 mm long, ca. 2 mm wide. Style red, glabrous; stigma globose, 4-cleft apically, 2-3 mm long, ca. 2 mm wide, reddish. Mature berry not seen. Distribution: Bolivia, scattered to locally frequent cloud forest shrubs in Depts. Cochabamba and Santa Cruz, 2,500-3,100 m (Fig. 61). Specimens examined: Bolivia, cochabamba: Cochabamba-Villa Tunari road, Badcock 711 (K); Corani, Cardenas 5754 (K, US); Siberia, Cardenas 5764 (US), santa cruz: 24 km W of Comarapa on Carretera Fundamental 4, Davidson 3851 (MO); Fortaleza, between Siberia and Comarapa, Vogel 499 (U). This species is included in the Fuchsia denticulata species group because of its long, firm floral tubes and purple drying petals. It differs from other members of this group in its funnelform floral tubes, nearly sessile, serrulate leaves, and flowers tightly grouped at the branch tips. This is one of only two species of sect. Fuchsia endemic to Bolivia, where it probably occurs sympatrically with F. sanc- tae-rosae. 38. Fuchsia macrostigma Bentham, PI. Hartw. 129. 1844. Fuchsia macrostigma var. typica Munz, Proc. Calif. Acad. Sci. IV. 25:31, pi. 3, fig. 17. 1943. Based on F. macrostigma Benth. Fuchsia macrostigma var. macrostigma, Munz, Opera Bot., Ser. B, 3:19. 1974. type: Ecuador, Prov. El Oro, in mountains of Paccha, 1841-1843, Theodor Hartweg (K, holotype; photograph, MO). Fuchsia longifiora Bentham, PI. Hartw. 177. 1845. Fuchsia macrostigma var. longiflora (Benth.) Munz, Proc. Calif. Acad. Sci. IV. 25:31. 1943; Opera Bot., Ser. B, 3:20. 1974. type: Ecuador, Prov. Pichincha, W side of Volcan Pichincha, between Quito and Nanegal, 1841-1843, Theodor Hartweg (K, holotype). Fuchsia spectahilis Hooker ex Lindley, Gard. Chron. 1848:319, /. May 13, 1848. Hook., Bot. Mag. 74: t. 4375. June 1, 1848. Lem., Fl. Serres Jard. Eur. 4:360, /. 1848. Paxt., Paxton's Mag. Bot. 16:255, /. 1849. Ysabeau, Journ. Hort. Prat. Belg. 7:225, t. 1849. Fancourt, Gard. Chron. 1850:71, fig. 1850. type: cultivated in London, England, by the Veitch nurseries, 1848, from seeds col- lected by William Lobb in the mountains of Cuenca, Prov. Azuay, Ecuador, William J. Hooker (K, holotype). Fuchsia macrostigma var. puhens I. M. Johnston, Contr. Gray Herb. 75:34. 1925. type: Ecuador, Prov. Chimborazo, vicinity of Huigra, mostly on the Hacienda de Licay, 3 Sept. 1918, Joseph N. Rose & George M. Rose 22479 (US 1022130, holotype; photographs, POM, UC; GH, NY, isotypes). Erect shrubs 0.5-1.5 m tall. Branchlets stout, subsucculent, 2-7 mm thick, terete or angled, green to dull purple; older branches 8-15 mm thick, dull tan, with finely fissured bark. Leaves opposite or less often ternate, firmly membra- nous, narrowly to broadly elliptic to (ob-)ovate, acute at the base, acute to acu- minate at the apex, 6-27 cm long, 3-12 cm wide, velvety dark green and sub- glabrous to puberulent above, pale green to wine purple below with strigose to 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 villous pubescence mostly along the veins; secondary veins 9-15(-22) on either side of the mid vein; margin remotely denticulate with small, glandular teeth. Petioles stout, 2-3 mm thick, 15— 30(— 35) mm long, strigose, green to purple. Stipules dark, triangular, sometimes connate, 2-2.5 mm long, ca. 2 mm wide, deciduous. Flowers few and axillary in upper nodes. Pedicels stout, 2-2.5 mm thick, 8-20(-30) mm long, pubescent, spreading to ascending. Ovary cylindrical, 8-12 mm long, 3-5 mm thick, green, verrucose, 4-8 sulcate. Floral tubes narrowly funnelform, 50-80 mm long, 2-5 mm wide and bulbous at the base, gradually widened above until 4-10 mm wide at the rim, subglabrous to villous outside, sparsely pilose inside in lower Vi\ tube firm-spongy, 1-1.5 mm thick when fresh, often curved downward in distal Vi. Sepals lance-oblong, 14-23 mm long, 7-8 mm wide, thick- spongy, 1-2 mm thick when fresh, apiculate, divergent, tips free and spreading in bud. Tube pale to dark red; sepals red with dull green tips. Petals bright red, shorter than the sepals, orbicular to broadly obovate, 12-18 mm long, 10-19 mm wide, undulate, rounded at the apex, strongly spreading. Nectary green, unlobed, ca. 3 mm high and ca. 1.5 mm thick. Filaments red, 8-12 mm and 5-8 mm long; anthers oblong, 2.5-3.5 mm long, ca. 1.5 mm wide. Style pink, sparsely pubescent to subglabrous; stigma massive, tetragonous, 3-5 mm long, 4-6 mm wide, with 4 mound-like, sticky lobes, cream to pink. Berry ellipsoid, 8-sulcate until fully ripe, verrucose, 20-22 mm long, 8-12 mm thick; seeds tan brown, 2-2.5 mm long, 1.2-1.6 mm wide. Distribution: Cordillera Occidental of Colombia and Ecuador, infrequent in moist cloud forest thickets on Pacific slopes, 1,000-2,500 m (Fig. 61). Representative specimens examined: Colombia, cauca: Km 42-47 NE of Uribe, Luteyn & Lebron- Luteyn 7418 (COL, NY), narino: between Rio Miraflores and Rio San Martin, Volcan Cumbal region, Ewan 16165 (POM, US); trail from Mayasquer to Tambo, Vogel 284 (U). valle: above Queremal, Las Colonias, Cuatrecasas 23901 (F). Ecuador, azuay: W of Patul, 3 km between Huahualcay and Rio Patul, below Pasas de Pinglion, Steyermark 52613 (NY), bolivar: Alto de Telimbela, Acosta Solis 7156 (F); Simiatug, Hacienda Talahua, Penland & Summers 614 (POM), 657 (F, POM), canar: N rim of valley of Rio Canar, Camp E-2895 (NY, RSA). carchi: Km 50 Tulcan-Maldonado road, Boeke 861 (MO); Km 71 on Tulcan Maldonado road, Holm-Nielsen et al. 6011 (AAU); Los Olivos, Mexia 7462 (F, NA, UC, US), chimborazo: Sibambe, Hacienda La Carmela, Acosta Solis 5367 (F); 5 km from Huigra, Canon del Rio Chanchan, Camp E-3376 (GH, MO, NY, P, RSA, UC, US); foot of Volcan Chimborazo, Spruce in I860 (K). cotopaxi: 5 km above Pilalo, Berry & Berry 2549 (MO); 3 km E of Macuchi, Quevedo-Latacunga road, Dodson & Gentry 10152 (MO); Pilalo, Holm-Nielsen 6 Jeppesen 1549 (AAU, DS). el oro: between La Chorita and Portovelo, Hitchcock 21168 (GH, NY, US); along trail from Sambotambo, near highway to Portovelo, Steyermark 54221 (NY), imba- bura: Cerro de Pinam, Acosta Solis 8505 (F); trail between Irubi and Apuela, N of Volcan Cotocachi, Drew E-131 (RSA, US), pichincha: valley of Rio Pilaton, below Garretas, Asplund 9709 (S); above Rio Toachi, Lopez-Palacios 4224 (MERF, MO); Km 72-74 of old road Quito-Santo Domingo de los Colorados, Luteyn & Lebron-Luteyn 5651 (NY); 13 km NW of Nono, road to Puerto Quito, Luteyn et al. 6520 (MO, NY); San Ignacio, Km 23 of Aloag-Santo Domingo road, Sparre 14583 (S); Km 37-50 along Rio Saloya, between Volcan Atacaso and Volcan Pichincha, Steyermark 52543 (NY). The flowers of this species are rather striking; they are long and narrow, usually horizontally disposed and recurved in the lower one half, with round, spreading petals, and a large, tetragonous stigma. Although it is a very distinctive species, it is placed in the Fuchsia denticulata species group because of its thick- tubed, axillary flowers, stout pedicels, and greenish sepal tips. The round petals, sulcate ovaries, and thick tubes also ally it to F. ampliata, however. Fuchsia macrostigma is restricted to the Cordillera Occidental of the northern Andes and is the only species to be found in the southernmost remnant of cloud 1982] BERRY— FUCHSIA SECT. FUCHSIA 147 forest in El Oro Province of southern Ecuador. Differences in floral tube length and degree of pubescence were used to describe the taxa listed in synonymy, but these characters vary too widely and continuously between populations to be of any taxonomic significance. No naturally occurring hybrids of this species were found, but one of the earliest artificial hybrids in sect. Fuchsia, F. dominiana (van Houtte, 1854) was a cross between F. macrostigma and F. denticulata. Fuchsia macrostigma occurs sympatrically with F. putumayensis, F. scabriuscula, F. sessilifolia, and F. syl- vatica. 39. Fuchsia magdalenae Munz, Proc. Calif. Acad. Sci. IV. 25:25, pi. 2, fig. 5. 1943. type: Colombia, Dept. Magdalena, above San Miguel, at edge of pa- ramo, 3,000 m, July 1932, William E. Seifri Z 392 (US 1572275, holotype; pho- tographs, MO, NY, UC). Fuchsia lampadaria J. O. Wright, Bot. Jour. Linn. Soc. 77: 1 13, fig. 1978. type: Cultivated in Reading, Great Britain in 1976, from seeds collected in Colombia, Dept. Magdalena, SE slopes of the Sierra Nevada de Santa Marta, Trombachuca Valley, uppermost forests, 3,300 m, 1975, by Mi- chael Adams and George Bernard, J. O. Wright (RDG, holotype, not seen). Shrubs 2-5 m tall. Young growth glabrous to lightly strigose; branchlets pur- plish, older branches red brown, with bark exfoliating in strips. Leaves ternate or less often quaternate, firmly membranous to subcoriaceous, (narrowly) elliptic to ovate, acute to rounded at the base, acute to subacuminate at the apex, 2. 5-8. 5(- 12) cm long, l^(-6) cm wide, dark green and mostly glabrous above, pale green with purplish veins and glabrous below, except for hairs usually pres- ent along the margin; secondary veins 8-14 one either side of the midvein; margin denticulate or subentire. Petioles purplish, sparsely strigose, 4-25(-30) mm long. Stipules lanceolate to triangular, dark, ca. 1 mm long, deciduous. Flowers pen- dant and solitary in upper leaf axils. Pedicels firm, 15-60 mm long, mostly red. Ovary ovoid-ellipsoid, glabrous, 7— 10(— 1 1) mm long, 3-3.5 mm thick. Floral tube subcylindric, (35-)42-60 mm long, firm-fleshy, ca. 1 mm thick when fresh, 3.5-6 mm wide at the base, sometimes slightly narrowed above the nectary, very grad- ually widened above until 6-10 mm wide at the rim, glabrous inside and outside. Sepals lanceolate, acute to subacuminate, 13— 18(— 22) mm long, 4-5(-6) mm wide, blunt-tipped in bud, spreading at anthesis. Tube slightly purplish at the base and orange red above, sepals nitid orange red with greenish tips. Petals orange red, suborbicular to obovate or subrhomboid, sometimes with irregular margins, 1 1-19 mm long, 7— 12(— 18) mm wide, usually with a narrowly acute tip. Nectary an uneven band of lustrous tissue lining the basal 3-6 mm of the floral tube, without prominent lobes. Filaments light red, 12-16 mm and 8-13 mm long; anthers ob- long, 3.5-4 mm long, ca. 2.5 mm wide, cream. Style light red to orange, glabrous; stigma capitate, lightly 4-cleft apically, 2-3 mm long, 2-A mm wide, light red to orange, exserted 4-10 mm beyond the anthers. Berry ellipsoid, 20-24 mm long, 10-12 mm thick, usually dark purple; seeds (1.5-)2-2.5 mm long, 1-1.3 mm wide. Gametic chromosome number n = 22. Distribution: Endemic to upper cloud forest of the Sierra Nevada de Santa Marta in northeastern Colombia; (2,000-)3, 000-3, 350 m (Fig. 61). 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Representative specimens examined: Colombia, magdalena: SE slopes Sierra Nevada de Santa Marta, Dirincune Sabana, watershed of Rio Donachui, Cuatrecasas & Romero Castaheda 24514 (COL, RSA, US); San Antonio, Sierra Nevada de Santa Marta, Hanbury-Tracy 504 (K), 518 (K); San Miguel, Karsten s.n. (W); Paramo in quebrada from Rio Frio, ca. 10°55'N, 73°53'W, Kirkbride & Forero 1759 (COL, MO); from Paramo to Cebolleta, Romero Castaheda 7177 (COL); Pico Hacha- Sierra Nevada, Schlim 800 (BM, F, G, K, MPU, P); between Pueblo and San Miguel, Seifriz 537 (US). This species is placed in the Fuchsia denticulata species group because of its thick- tubed, nearly cylindric, axillary flowers, but it is highly distinct from all other species in sect. Fuchsia in its non-annular, band type nectary, which resem- bles those found in sects. Ellobium and Hemsleyella (Figs. 45 and 48). Pu- bescence is completely lacking inside the tube and on the style, and the flowers have rounded petals that often have uneven margins. Fuchsia magdalenae is also tetraploid and has biporate pollen. The above assemblage of unusual character- istics suggests that this species may have been an early offshoot of the section that was isolated on an outlying portion of the Andes. Fuchsia magdalenae is restricted to the upper cloud forest and subparamo belt that more or less continuously surrounds the high peaks of the isolated Sierra Nevada de Santa Marta range. It is the only species of Fuchsia to occur there, although Wright (1978a) recently described F. lampadaria from this same area. Most of the differences he states for that species, however, are insignificant when all the specimens from the Santa Marta area are examined. The larger leaf di- mension he describes may also be related to the controlled greenhouse conditions in which his plants were grown. 40. Fuchsia simplicicaulis Ruiz & Pavon, Fl. Peru v. Chil. 3:89. pi. 322 , fig. a. 1802. Hook., Bot. Mag. t. 5096. 1859. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):563. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:41,/?/. 5 y fig. 29. 1943. Morley & Everard, Wild Flowers of the World, pL 173, fig. CI. 1970. Travis, Fuchsia Annual 1975:21, photo. 1975. type: Peru, Dept. Huanuco, woods at Mufia, 1778-1788, Hipolito Ruiz & Jose Pavon (MA N° 11/92, ■ lectotype, here designated; photograph, MO). Scandent shrubs 2-5 m tall, sparsely verticillately branched, with ultimate branches pendulous. Branchlets subterete, puberulent, older stems with light red, exfoliating bark. Leaves ternate or quaternate, membranous, linear-lanceolate to ovate-lanceolate, acute to narrowly obtuse or truncate at the base, acuminate at the apex, 8-15 cm long, 1-3.5 cm wide, glabrous above, glabrous to scattered pilose below or along the margin; secondary veins 10-18 on either side of the mid vein; margin subentire. Petioles glabrous to puberulent, 2-6 mm long. Stipules triangular, 1-2 mm long, ca. 0.9 mm wide, subpersistent. Flowers in simple to multiple, pendant, involucrate racemes, with flowers 3-4 per whorl and inter- nodes 1.5-6 cm long; rachis 8-30 cm long; bracts thinly membranous, sessile, concave, ovate-lanceolate, broadly rounded and clasping the stem at the base, acuminate at the apex, 1-5 cm long, 0.5-1.5 cm wide. Pedicels puberulent, 5-10 mm long. Ovary oblong-ellipsoid, puberulent or velutinous, 5-6 mm long, 2-2.5 mm thick. Floral tube narrowly funnelform, 40-50 mm long, bulbous and 3-4 mm wide at the base, narrowed to 1.5-2 mm wide above the ovary, then gradually widened above until 5-7 mm wide at the rim, finely pilose outside, pilose inside. 1982] BERRY— FUCHSIA SECT. FUCHSIA 149 Sepals lanceolate, puberulent on both sides, acuminate, 16-20 mm long, ca. 4 mm wide. Tube and sepals light reddish pink. Petals red, linear-lanceolate to elliptic, acute to narrowly acuminate at the apex, 9-13 mm long, 2-3 (-5) mm wide. Nectary apparently unlobed, ca. 1.5 mm high. Filaments 10-12 mm and 7-8 mm long; anthers oblong, 3-4 mm long, ca. 2 mm wide. Style pilose from base up to the rim of the tube; stigma globose, 4-parted at the apex, 2.5-3.5 mm long, ca. 2 mm wide. Berry ellipsoid, puberulent, 1 1-13 mm long, ca. 8 mm thick; seeds tan, ca. 1.2 mm long, ca. 0.7 mm wide. Distribution: Central Peru. Rare in cloud forests in Depts. Huanuco, Junin, and Pasco, eastern slopes of the Andes; 2,200-2,500 m (Fig. 62). Specimens examined: Peru, huanuco: Muna, Lobb 119 (K), Macbride 4014 (BH, F, GH, K, S, US), Pearce 133 (K). junin: Vitoc, Martinet 1310 (P). pasco: Oxapampa, Soukup 1804 (GH, US, USM). without locality: Lobb s.n. (K). This is one of two very rare and unusual species with whorls or involucres of thin, membranous bracts and short-pedicelled flowers. The other species, Fuchsia ceracea y also occurs in Huanuco, but has much larger bracts and flowers, with shorter petals. Both species are glabrous except for the puberulent or velutinous young stems, pedicels, and flowers. Two other species from Central Peru, F. coriacifolia and F. sanmartina, seem to be derived from, or at least closely related to the above species. Their relationships are discussed further under the F. sim- plicicaulis species group (p. 54). 41. Fuchsia ceracea P. Berry, sp. nov. type: Peru, Dept. Huanuco, Prov. Huanuco, in tree along stream, 5 km W of Carpish tunnel between Huanuco and Tingo Maria, 2,500 m, 4 Aug. 1978, Paul E. Berry & James Aronson 3081 (MO 2720597, holotype; MO, USM, isotypes). Fig. 31. Frutex scandens dependensque 3-6 m altus, praeter flores juvenes pedicellosque pilosos glaber. Folia opposita vel ternata, firme membranacea, lanceolata vel anguste ovata basi obtusa vel leviter auriculata aliquantum inaequalia, apice acuta, 8-15 cm longa, 4-6 cm lata, utrinque ceracea, nervis secundariis utroque latere 7-10, margine integerrima; petiolis leviter tortis 2-20 mm longis. Flores dependentes in verticillis involucratis 2-3 floralibus ad apicem ramorum disposisti; bracteis pruinosis, tenuiter membranaceis, concavis, sessilibus, late ovatis, 4-12 cm longis, 1.5-5 cm latis; pedicellis 7-15 mm longis; ovario ellipsoideo ca. 7 mm longo. Tubi florales vivide rosei vel lavandulacei, 9-13 cm longi, basi 6-7 mm lati inde 3-5 mm lati constricti superne gradatim dilatati summo 8-9 mm lati extus glabrati vel puberuli. Sepala lanceolata acuminata 28-30 mm longa 6-7 mm lata. Petala coccinea vel atropurpurea, ovata, late acuta, 5-7 mm longa, ca. 4 mm lata. Filamenta pallide rosea, antisepala 13-14 mm longa, antipetala 9-10 mm longa, antheris oblongis 4-5 mm longis ca. 3 mm latis. Stylus puberulus, stigmate magno subtetragono, 4-lobato, 4-5 mm longo ca. 4 mm lato. Bacca matura non visa. Numerus gameticus chromosomatum n = 1 1 . Climbing shrub or dangling liana in trees to 6 m above ground, the ultimate branches pendant. Plants glabrous except for puberulent young flowers and ped- icels. Leaves opposite or ternate, firmly membranous, lanceolate to narrowly ovate, obtuse to unequal or slightly auriculate at the base, acuminate at the apex, 8-15 cm long, 4-6 cm wide, waxy green on both surfaces; secondary veins 7-10 on either side of the midvein; margin entire. Petioles reddish, slightly twisted, 2-20 mm long. Stipules narrowly triangular, 1.5-2 mm long, ca. 0.7 mm wide, subpersistent. Flowers in simple, pendant, involucrate racemes, with 2-3 flowers per whorl and internodes 2-6 cm long; bracts thin membranous, concave, sessile, 1 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 pruinose, broadly ovate, acuminate at the apex, clasping the stem at the base, 4-12 cm long, 1.5-5 cm wide. Pedicels 7-15 mm long. Ovary ellipsoid, ca. 7 mm long, ca. 3 mm thick. Floral tube subcylindric to narrowly funnelform, (9-) 10-13 cm long, bulbous and 6-7 mm wide at the base, narrowed to 3-5 mm wide above the nectary, slightly widened above until 8-9 mm wide at the rim, puberulent to subglabrous outside, pubescent inside in lower Vi. Sepals lanceolate, acuminate, 28-30 mm long, 6-7 mm wide, suberect at anthesis. Tube and sepals lavender to bright pink, tube white inside. Petals crimson to dark purple, ovate, broadly acute, 5-7 mm long, ca. 4 mm wide, suberect at anthesis. Nectary light green, unlobed, ca. 3 mm high. Filaments white pink, 13-14 mm and 9-10 mm long; anthers oblong, 4-5 mm long, ca. 3 mm wide. Style lavender pink, puberulent for most its length; stigma massive, subtetragonous, 4-lobed at the apex, 4-5 mm long, ca. 4 mm wide, light pink. Berry not seen. Gametic chromosome number n = 11. Distribution: Very rare climbers in cloud forest, known just from Panao and the Carpish area of Dept. Huanuco, Peru; 2,500-2,850 m (Fig. 62). Specimens examined: Peru, huanuco: above La Molina, near Panao, Asplund 13699 (S); Carpish, Asplund 14857 (RSA — a mixed collection with F. corymbiflora). This species and Fuchsia simplicicaulis are the only two species in the genus with involucrate inflorescences and sessile concave bracts. Fuchsia ceracea has unusual waxy leaves, pruinose bracts, and very long floral tubes. The petals are less than one quarter as long as the sepals and are the proportionately most reduced of any species in sect. Fuchsia. It is extremely rare and is known only as a liana with long, unbranched, hanging stems. Fuchsia abrupt a and F. corymbiflora both grow in the same area as F. ceracea, but they are lower shrubs and occur in more open thickets. 42, Fuchsia coracifolia P. Berry, sp. nov. type: Peru, Dept. Pasco, Prov. Oxapam- pa, Huancabamba, 1863-1865, Richard Pearce 565 (K, holotype; photograph, MO). Frutex plerumque glaber 1-2 m altus. Folia coriacea subsessilia opposita vel ternata, basi rotund- ata vel subcordata apice acuta vel acuminata 3-6 cm longa 1-2.5 cm lata superne atroviridia subtus multo pallidioria, nervis strigulosis; nervis secundariis utroque latere 4-5 versus marginem gradatim inconspicuis; margine subdenticulata; petiolis 1-2 mm longis; stipulis plerumque connatis, subincra- ssatis, 2-3 mm longis, 2-3.5 mm latis, subpersistentibus. Flores rosei, numerosi et dependentes in racemo terminali et laxo dispositi, basi oppositi vel ternati apice gradatim alterni; rhachidi 3-9 cm longa; bracteis sessilibus anguste ovatis 10-20 mm longis 5-10 mm latis; pedicellis 5-15 mm longis; ovario oblongo 5-6 mm longo. Tubi florales firmi, anguste infundibuliformes, 52-58 mm longi, basi 3-3.5 mm lati et parum bulbosi inde 2.5-3 mm lati constricti superne gradatim dilatati summo 6-10 mm lati ubique plerumque glabrati. Sepala firma, ca. 1.5 mm crassa, oblongo-ovata, acuta, 17-19 mm longa 7-9 mm lata. Petala lanceolata acuta 10-12 mm longa ca. 4 mm lata. Filamenta antisepala 8-10 mm longa antipetala 6-7 mm longa, antheris oblongo-reniformibus 3-3.5 mm longis 2-2.5 mm latis. Stylus glaber, stigmate conico ca. 2 mm longo ca. 2.5 mm lato apice leviter 4-fisso. Bacca non visa. Apparently simply branched low shrubs 1-2 m tall. Branchlets glabrous, slightly angled; older stems with whitish tan bark. Leaves opposite or ternate, firmly coriaceous, rounded to subcordate at the base, acute to acuminate at the apex, 3-6 cm long, 1-2.5 cm wide, dark green and strigillose along the veins above, much paler whitish green and strigillose along the red veins below; secondary 1982] BERRY— FUCHSIA SECT. FUCHSIA 151 Figure 62. Distribution of the Fuchsia simplicicaulis and F. tincta species groups. veins 4-5 on either side of the mid vein, becoming inconspicuous towards the margin, the midvein prominent below; margin subdenticulate. Petioles 1-2 mm long, glabrous. Stipules mostly connate, firm, 2-3 mm long, 2-3.5 mm wide, subpersistent. Flowers numerous and pendant in loose, terminal racemes 3-9 cm long; bracts narrowly ovate, sessile, 10-20 mm long, 5-10 mm wide, opposite to ternate at the base and becoming alternate towards the end of the raceme. Ped- icels 5-15 mm long, glabrous. Ovary ovoid, 5-6 mm long, ca. 3 mm thick. Floral tube narrowly funnelform, firm, 52-58 mm long, 3-3.5 mm wide and slightly bulbous at the base, narrowed to 2.5-3 mm above the nectary, then gradually widened above until 6-10 mm wide at the rim, mostly glabrous outside, glabrous inside. Sepals firm, ca. 1.5 mm thick, oblong-ovate, acute at the apex, 17-19 mm 1 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 long, 7-9 mm wide. Flowers rose. Petals lanceolate, acute at the apex, 11-12 mm long, ca. 4 mm wide. Nectary unlobed, 1.5-2 mm high. Filaments 8-10 mm and 6-7 mm long; anthers oblong-reniform, 3-3.5 mm long, 2-2.5 mm wide. Style glabrous; stigma conic, slightly 4-cleft at the apex, ca. 2 mm long, ca. 1.5 mm wide. Berry not seen. Distribution: Known only from the type specimen, from central Peru, in Dept. Pasco near the Huanuco border (Fig. 62). The ovate, sessile bracts, short pedicels, and inflorescence with opposite or ternate basal flowers place this species in the Fuchsia simplicicaulis species group. It is distinct from related species in having coriaceous, nearly sessile leaves and thick floral tubes. The inflorescence of F. coriacifolia seems to be derived from the involucrate type of F. simplicicaulis and F. ceracea and is transitional between these and the alternate-flowered, racemose inflorescences of the related F. sanmartina. 43. Fuchsia sanmartina P. Berry, sp. nov. type: Peru, Dept. San Martin, Dist. Hual- laga, Valley of Rio Apisoncho, 30 km above Jucusbamba, 7°55'S, 77°10'W, 2,800 m, 6 Aug. 1965, A. C. Hamilton & P. M. Holligan 1065 (K, holotype; photograph, MO; NY, S, UC, isotypes). Frutex scandens 1.5-10 m altus, ramulis foliisque junioribus puberulenti-velutinis. Folia pleru- mque ternata interdum opposita, firme membranacea, anguste elliptico-ovata, basi obtusa vel rotundata aliquantum inaequalia apice acuta vel acuminata, 5-15 cm longa, 1.5-6 cm lata, subtus nervis ple- rumque puberulentis; nervis secundariis utroque latere 9-11, margine subintegerrima vel denticulata; petiolis puberulentis 3-20 mm longis; stipulis anguste triangularibus, ca. 2 mm longis, deciduis. Flores numerosi et dependentes, vivide rubri, racemis terminalibus dispositi; rhachidi 7-16 cm longa; bracteis 8-50 mm longis, 5-25 mm latis, petiolis 1-8 mm longis; ovario 7-8 mm longo. Tubi florales subcyli- ndrici 55-76 mm longi basi 3-4 mm lati et parum bulbosi, inde 2-3 mm lati constricti superne gradatim dilatati summo 6-8 mm lato extus puberuli. Sepala lanceolata acuminata 20-25 mm longa 4-5 mm lata. Petala rubra anguste lanceolato-elliptica, acuta, 10-16 mm longa, 3-6 mm lata. Filamenta antisepala 14-15 mm longa antipetala 9-1 1 mm longa, antheris oblongis 3.5-4 mm longis 1.5-2 mm latis. Stylus pilosus, stigmate capitato, 3-4 mm longo, 2.5-3 mm lato apice leviter 4-fisso. Bacca matura non visa. Scandent shrubs 1.5-3 m high or lianas to 10 m above ground. Young growth finely puberulent-velutinous, branchlets mostly subtriangular in transection. Leaves mostly ternate, occasionally opposite, firmly membranous, narrowly elliptic-ovate, obtuse to rounded or unequal at the base, acute to acuminate at the apex, 5-15 cm long, 1.5-6 cm wide, subglabrous to puberulent above, puberulent below, especially along the veins; secondary veins 9-11 on either side of the midvein; margin subentire to denticulate. Petioles puberulent, 3-20 mm long. Stipules nar- rowly triangular, ca. 2 mm long, deciduous. Flowers numerous and pendant in terminal racemes; rachis puberulent, 7-16 cm long; bracts 8-50 mm long, 5-25 mm wide, with petioles 1-8 mm long. Pedicels puberulent, 13-50 mm long. Ovary oblong, 7-8 mm long, ca. 2 mm thick. Floral tube subcylindric, 55-76 mm long, bulbous and 3-4 mm wide at the base, then narrowed to 2-3 mm wide above the nectary, slightly widened above until 6-8 mm wide at the rim, puberulent outside, pilose inside. Sepals lanceolate, acuminate, 20-25 mm long, 4-5 mm wide, the tips slightly spreading in bud. Tube and sepals bright red. Petals red, narrowly lance-elliptic, acute, 10-16 mm long, 3-6 mm wide. Nectary unlobed, ca. 2 mm high. Filaments 14-15 mm and 9-11 mm long; anthers oblong, 3.5-4 mm long, 1982] BERRY— FUCHSIA SECT. FUCHSIA 153 1.5-2 mm wide. Style pilose, stigma 3^4 mm long and 2.5-3 mm wide, 4-parted at the apex. Berry not seen. Distribution: Known only from the type locality and near vicinity on the eastern slopes of Dept. San Martin, Peru; 2,800-3,600 (Fig. 62). Specimens examined: Peru, san martin: Valley of Rio Apisoncho, 30 km above Jucusbamba, 3,600 m, Hamilton & Holligan 336 (K, S, UC). This species belongs to the Fuchsia simplicicaulis species group because of its narrowly elliptic-ovate leaves, nearly sessile bracts, and terminally grouped, finely puberulent flowers with petals much shorter than the sepals. It lacks the unusual sessile, concave bracts and involucrate flowers of F. simplicicaulis and F. ceracea, but it is linked to these species by F. coriacifolia, which has an intermediate type of inflorescence, with the older flowers verticillate and the younger flowers alternate. Hamilton & Holligan 1157 (K), from the type locality, differs from the pre- viously cited specimens of this species in its longer, redder pubescence, shorter petals (6-8 mm long), and much more spreading sepal tips. There are presently too few specimens available to know if this collection falls within the normal variability of F. sanmartina. 44. Fuchsia sessilifolia Bentham, PI. Hartw. 176. 1845. Hook., Bot. Mag. /. 5907. 1871. Munz, Proc. Calif. Acad. Sci. IV. 25:67, pi. 11, fig. 57. 1943; Opera. Bot., Ser. B, 3:22. 1974. type: Ecuador, Prov. Pichincha, woods of Guayan, W slopes of Volcan Pichincha, 1841-1843, Theodor Hartweg 985 (K Bentham Herb., holotype; photograph, MO; BM, BREM, CGE, K Hooker Herb., OXF, isotypes). shrub es. Young growth finely puberulent, branchlets subtetragonous, cinereous-pu- berulent, 2-6 mm thick, wine red; older branches 5-15 mm thick, with exfoliating bark. Leaves verticillate, mostly quaternate, subsessile, membranous to subco- riaceous, narrowly elliptic to elliptic or oblanceolate, rounded to attenuate or narrowly subcordate at the base, acute to acuminate at the apex, 6-20 cm long, 2-6(-8) cm wide, lustrous dark green and subglabrous above, pale green to pur- plish and pubescent to subglabrous below; secondary veins 18-25 on either side of the midvein, impressed above, the midvein prominent below; margin serrulate and commonly undulate. Petioles 1-5 mm long, dull red. Stipules triangular, thick- ened at the base, 1-2 mm long, ca. 1 mm wide, mostly deciduous. Flowers nu- merous in terminal, drooping, verticillately-branched panicles; rachis 5-30 cm long, bracts lanceolate, 10-35 mm long, 4-10 mm wide, often reflexed. Pedicels slender, strigillose, 3-5(-8) mm long. Ovary cylindrical, 5-6 mm long, 1.5-2.5 mm thick, finely strigose. Floral tube narrowly funnelform, 1 3— 1 8(— 20) mm long, 2-3 mm wide at the base, narrowed slightly to 1-2 mm above the nectary, grad- ually widened above until 4-5 mm wide at the rim, finely strigose outside, pilose inside. Sepals lanceolate, 8-1 1 mm long, 3-4 mm wide, acute to acuminate, form- ing a short point 1-2 mm long in bud, spreading at anthesis. Tube pale cream to rose green or light red; sepals light yellow green to light pink. Petals bright scarlet, 1 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 6-9 3-4 Nectary unlobed, ca. 1 mm high. Filaments red to greenish red, 5-8 mm and 3.5-5 mm long; anthers oblong, 1.5-2 mm long, ca. 1 mm wide, pale yellow. Style glabrous to loosely pilose; stigma capitate, 4-parted at the apex, 1.5-2 mm long, 1.5-2 mm wide, cream, situated at the level of the anthers or slightly exserted. Berry cylindric-oblong, 15-19 mm long, 8 seeds reddish purple to tan, 1.5-2 mm long, number n = 11. purpl Distribution: Ecuador and Colombia. In the Cordillera Occidental of the Northern Andes from Pichincha, Ecuador to Choco and Antioquia, Colombia; in the Cordillera Central from northern Napo to Tolima; and in the Cordillera Ori- ental from Putumayo to Huila; cloud forest shrubs in thickets, streambanks, and forest openings; 2,300-3,200 m (Fig. 63). Representative specimens examined: Colombia, antioquia: without locality, Kalbreyer in 1879 (K). caqueta: Pass between Garzon and Florencia, Mason 13930 (COL, GH, UC, US), cauca: above Carpinteria, Alston 8232 (BM, COL, RSA, S, US); between Guachicono and Rio Putis, Core 936 (MICH, RSA, US); Finca La Cumbre, headwaters of Rio Dinde, Core 1086a (US); Quebrada de Santo Domingo, headwaters of Rio Palo, Cordillera Central, Cuatrecasas 19297 (A, F, NY, VALLE); Aguabonita, region of Moscopan, east slopes Cordillera Central, Cuatrecasas 23555 (F, RSA); Vi- nagrita tibia, Purace, Dryander 1633 (US); west slopes of Rio Munchique, Garcia-Barriga et al. 12928 (AAU, COL, US); W of Tambo, Cordillera Occidental, Haught 5191 (COL, P, RSA, US); Paramo de Las Papas, between El Boqueron and La Hoyola, Idrobo et al. 3919 (COL, P); Volcan de Sotara, Lehmann 6220 (K); Canaan, Purace, Pennell & Killip 6674 (GH, NY, PH, US); Paletara to Calaguala, Pennell 7111 (GH, NY, PH, US); San Jose, San Antonio, Pennell 7569 (GH, NY, PH, US); above Carpinterias, between Cerros Munchique and Altamira, Perez-Arbelaez & Cuatrecasas 6145 (COL, F, US); valle de Quintero, near Pitaio, Rio Palo basin, Pittier 1427 (NY);.road from Tambo to 20 de julio, Plowman & Vaughan 5315 (COL), choco: headwaters of Rio Carmen, 8 km N of Carmen de Atrato, Fosberg 21560 (RSA, US); Dauro, Toro 1168 (NY), huila: Km 25 of Pitalito-Mocoa road, Berry 3592 (COL, MO); between Gabinete and Andalucia, Cuatrecasas 8586 (COL, F, US); Finca Merenberg, Municipio de La Playa, Diaz Piedrahita et al. 405 (COL); Lavaderos, ridge between Rios Naranjo and Granadillo, 15 km S of San Agustin, Fosberg 20062 (RSA, UC, US); 20 km from Altamira toward Florencia, Luteyn et al. 4846 (COL, MO); between Garzon and Florencia, Mason 13921 (UC); E of Neiva, Rusby & Pennell 687 (NY, US), narino: above El Encano, Laguna La Cocha, Balls 7521 (BM, COL, K, NA, UC, US); between Pasto and Rio Bobo, Barclay 4670 (COL, MO); Gruta de la Virgen, road Pasto to Yacuanquer, Benavides 31 (PSO); Aponte, Rio Majinsanoy, Bristol 1177 (COL, DS); right bank of Rio Barrancos, Municipio de La Florida, Diaz et al. 831 (COL); Encano, N end of Laguna La Cocha, Fosberg 20431 (RSA, S, US); Laguna La Cocha, Isla La Corota and Sixce-Turibamba, Garcia-Barriga et al. 13049 (COL, US); woods near Pasto, Jameson 432 (BM, CGE, G, K, LE, OXF, TCD); road from Cuatro Esquinas to Quitasol, Mora 293 (COL); road from Balalaika to Yascual, Mora 396 (COL); near crossroads of El Pedgregal-Ipiales road, Guacuan path, Mora et al. 6056 (PSO). putumayo: 2 km SW of Sibundoy, Bristol 328 (COL, DS, GH); Km 79 from Pasto to San Francisco and Mocoa, Plowman & Davis 4315 (COL); Paramo de San Antonio, road Pasto-Sibundoy, Schultes & Cabrera 18883 (US); near Sibundoy, Schultes & Smith 3061 (BH, DS, MO, NY, UC). quindio: 12 km from Salento towards Cocora, along Rio Salento, Escobar 1014 (MO), tolima: Buenavista, Andre 2134 (F, GH, K, NY): Quindio, Triana s.n. (G 2 sheets, P); Mt. Quindio before El Gallego, Triana 6125.6 (BM, COL), valle: Barragan, watershed of Rio Bugala- grande, La Laguna, Cuatrecasas 20863 (F, RSA, VALLE); La Palma, right bank Rio Pichinde, watershed of Rio Cali, Cuatrecasas 21672 (F, RSA, VALLE); Rio Cali, Pichinde, Duque-Jaramillo 4702 (COL); near Palmira, near Quebrada El Socorro, Cordillera Central, Maas & Escobar 628 (MO, U). Ecuador, carchi: Km 57 of Tulcan-Maldonado road, Boeke 828 (MO); 5 km W of El Carmelo, Berry 3163 (MO, QCA); El Pun (= El Carmelo), Asplund 16890 (K, NY, S), Mexia 7579 (POM, RSA, UC, US), imbabura: Urcusique, W slopes of Volcan Cotacachi, Acosta Solis 8225 (F). napo: Santa Barbara de Sucumbios, along Colombia border, Harling 4130 (NY, S); Rio San Pedro below mouth of Rio Clavadero, E of Cayambe peak, Wiggins 10454 (DS). pichincha: Guarumal, Km 38 of road to Saloya, Acosta Solis 11008 (F); Saloya, W of Quito, Asplund 7302 (S); Chiriboga, Asplund 17146 1982] BERRY— FUCHSIA SECT. FUCHSIA 155 (NY, S); near Atahualpa, Asplund 20314 (NY, S); above Chaupi-Saycha, Pululagua, Bell 513 (BM); above San Jose de Minas, Benoist 3963 (F); 41 km W of Cotocollao to Nanegal, Berry & Escobar 3182 (MO, QCA); 34 km W of Chillogallo to Chiriboga, Berry & Escobar 3242 (MO, QCA); Cornejo Astorga, road Quito-Santo Domingo de los Colorados, Harling et al. 9355 (MO, RSA); below Nono, Haught 3166 (A, BH, US); above Tandapi, Holm-Nielsen et al. 7130 (AAU); Alaspongo, Holmgren & Heilborn 697 (S); Mt. Corazon and Lloa, Sodiro s.n. (W); San Ignacio, road Aloag-Santo Domingo, Sparre 14596 (S); Km 37-50 along Rio Saloya, between Volcanes Atasco and Pichincha, Steyermark 52513 (NY). This is an easily recognizable species due to its large, lanceolate, subsessile, and quaternate leaves and the terminal, drooping panicles of short, pale flowers with contrasting dark petals. It is one of the most widespread species in the genus, occurring in Ecuador on both sides of the Andes and in all three Cordilleras of the northern Andes. Its distribution pattern indicates a probable origin in the Cordillera Occidental of Ecuador with a subsequent northward extension into the southern parts of the Colombian Cordilleras. As a result of this wide distribution, it is sympatric with many taxa of Fuchsia, including F. cuatrecasasii, F. corol- lata, F. dependens, F. hartwegii, F. nigricans, F. scabriuscula, F. sylvatica, F. polyantha, and F. verrucosa. It is only known to hybridize with F. nigricans (see discussion under that species). Fuchsia sessilifolia has flowers similar to F. nigricans in the contrasting petal and tube colors, short tubes, and cylindrical ovaries, but it has a much more branched inflorescence. It is very closely allied to F. polyantha, which can only be distinguished by its longer, totally red flowers. The two species occur together or in close proximity on the western slopes of the Ecuadorian Andes, and their different floral tube lengths and flower coloration may be related to different pollinators, or to the development of mechanical factors that reduce the chance of interspecific pollination. Munz fig. 37. 1943. Munz Narino, between Mayasquer and Tambo, 2,800 m, 2 Aug. 1935, Ynes Mexia 7571 (US 1662416, holotype; photographs, NY, POM; F, UC, US, isotypes). Erect shrubs 1-2 m tall. Young growth minutely puberulent-canescent; branchlets puberulent, terete, purplish. Leaves quaternate, firmly membranous, narrowly elliptic-lanceolate, acute to obtuse at the base, acuminate at the apex, 7-12 cm long, 2.5-4 cm wide, nitid dark green and subglabrous above, light green and sparsely puberulent below, especially along the veins; secondary veins 13-16 on either side of the midvein; margin subentire to serrate. Petioles puberulent, purplish, 3-10 mm long. Stipules dark, thick at the base, triangular, 1.5-2 mm long, ca. 1 mm wide, often connate and recurved when old, persistent. Flowers numerous in terminal, nodding, verticillately branched panicles; rachis 8-15 cm long; bracts narrowly lance-ovate, recurved, 10-30 mm long, 5-10 mm wide. Pedicels puberulent, 5-10 mm long. Ovary fusiform, puberulent, 6-7 mm long, ca. 2 mm thick. Floral tube narrowly funnelform, 34-43 mm long, slightly nodose and 2.5-3.5 mm wide at the base, narrowed to ca. 2 mm above the nectary, then widened above until 5-7 mm wide at the rim, finely puberulent outside, pilose inside in lower Vi. Sepals narrowly lanceolate, acuminate, 10-15 mm long, ca. 3 mm wide, with a narrow tip 1.5-2 mm long in bud. Tube and sepals red. Petals 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 63. Distribution of the Fuchsia sessilifolia species group and part of the F. boliviana species group. 1982] BERRY— FUCHSIA SECT. FUCHSIA 157 red, a little darker than the sepals, lanceolate to narrowly elliptic, subacuminate, 8-14 mm long, 2.5—4 mm wide. Nectary unlobed, ca. 1 mm high. Filaments red, 8-11 mm and 6-7 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide. Style red, sparsely pilose in lower Vr, stigma capitate, slightly 4-parted at the apex, 2-3 mm long, ca. 2 mm wide, dull white, exserted 5-4 mm beyond the anthers. Mature berries not seen, young ones oblong, 8-10 mm long, 5-7 mm thick. Gametic chromosome number n = 11. Western 200-3.300 Specimens examined: Ecuador, carchi: 50 km W of Tufino on road to Maldonado, Berry 3152 (MO, QCA); 60 km W of Tufino to Maldonado, Berry 3154 (MO, QCA); 20 km E of Maldonado on road to Tulcan, Gentry & Shupp 26321 (MO); between Moran and Olivos, Mexia 7474 (US), pichincha: Km 32-38 of old road from Quito to Santo Domingo de los Colorados, Luteyn & Lebron-Luteyn 5620 (MO, NY). Except for its longer, entirely red flowers, this species is very similar to the more widespread and apparently sympatric Fuchsia sessilifolia. They both have distinctive subsessile, narrow, quaternate leaves with a well branched, terminal, drooping inflorescence. No plants with intermediate flower tube sizes or colors have been seen, and the two species may be reproductively isolated by different lifoli 4-4 M. Johnston, J. Arnold Arbor. 20:242. 1939. Macbr Mus. Nat. Hist., Bot. Ser. 13(4):565. 1941, pro parte. Munz Sci. IV. 25:43, pi. 6, fig. 32. 1943, pro parte, type: Peru, 1 West MO subshrubs turning reddish-brown with age; older stems 4-10 mm thick, with light brown, striated bark. Leaves opposite or sometimes subopposite near the branch tips, softly membranous, ovate-elliptic to broadly ovate, acute to rounded or unequal at the base, acute to acuminate at the apex, 3-14 cm long, 1.5-9 cm wide, velvety dark green and strigillose above, pale green or commonly purple flushed and strigose below, pubescence denser along the nerves; secondary veins 1 1-18 on either side of the midvein, sulcate above; margin glandular-denticulate to coarsely dentate. Petioles strigose, 13-50 mm long. Stipules lanceolate, dark, 1-2 mm long, sub- persistent. Flowers usually 5-10 and nodding in a generally compact, terminal, corymbose raceme; rachis 1.5-7 cm long. Pedicels slender, strigose, 18-30 mm 6-8 long, 1 16-23 mm wide and slightly bulbous at the base, narrowed to 1-1.5 mm above the nectary, then widened gradually above until 3-5 mm wide at the rim, pilose outside, retrorse villous inside in lower Vz. Sepals lanceolate, 7-9 mm long, 3^1 mm wide, acute, spreading at anthesis. Tube and sepals pink to bright red. Petals oink to red. oblong, 6-9 mm long, 3-5 mm wide, obtuse to mucronate at the 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 13. Principal morphological differences between Fuchsia tincta and F. vargasiana. Leaves: Shape Color Flowers: Tube length Sepal color Sepal and petal angle at anthesis Fruits: Shape Length when ripe F. tincta Broadly elliptic-ovate Dark green above, mostly purple flushed below 12-23 mm Red pink Spreading Ellipsoid to subglobose 13-17 mm F. vargasiana Narrowly elliptic-ovate to del toid Medium-dark green above, pale green below 35-40 mm Dull green Strongly spreading to diver- gent Narrowly cylindrical, often curved and with a constrict- ed apex 25-32 mm apex, spreading at anthesis. Nectary unlobed or shallowly 4-lobed, ca. 1.5 mm 3-4 % capitate, 1.5-2 mm long, 1.5-2 mm wide, 4-parted at the apex, white to light pink, exserted 2-3 mm beyond the anthers. Berry ellipsoid to subglobose, 13-17 mm long, 8-11 mm thick; seeds 1 some number n = 1 1 . 4-1 Peru eastern Pillahuata farther north; 1, 800-2, 400(-2, 800) shady in the Prov. Specimens examined: Peru, cuzco: Km 131-137 of Cuzco-Pilcopata road, near Buenos Aires, Berry 3004 13956 (F, GH, PH, S, US); above Pintobamba, Prov. Convention, Vargas 3551a (BH, CUZ); Pilla- huata, Vargas 5128 (BH, CUZ, MO); Ticancha, Huaisanpilla, Vargas 9922 (CUZ, MO); Pillahuata, Yanamayo, Vagas 16695 (CUZ, RSA), 16703 (CUZ, RSA), Woytkowski 2 (MO, USM); between Puente Mercurio and Buenos Aires, Vargas 23138 (MO); Suecia, Woytkowski 144 (USM). This species is closely allied to Fuchsia vargasiana and F.furfuracea, which share the compact, terminal racemes, dense pubescence, and similar ovate, long- petiolate leaves. Fuchsia tincta has shorter flowers than these two species, though, and usually has deep dark green leaves with a purple flush below. It is remarkable that F. tincta and F. varagasiana are so similar morpholog- ically, yet they are sympatric and occupy the same restricted range. Both are - M _ A _ _ - Pilcopata 100 m (Km 136-137) rphological 1982] BERRY— FUCHSIA SECT. FUCHSIA 159 Table 14. Comparison of the principal morp sanctae-rosae, and a presumed hybrid. hological diiierences bet^ ween Fuchsia tincta, F. F. tincta Hybrid (Pennell 13956, NY) F. sanctae-rosae Leaves: number/node Max. blade size (L x W) Pubescence, lower surface 2 14 x 9 cm Strigose on entire surface 2-3 15 x 6 cm Strigose mostly along veins 3-4 14 x 4 cm Glabrous except hairs on midvein Flowers: Position Tube pubescence outside Racemose Pilose Axillary Strigose-pilose Axillary Glabrous Pubescence on young growth Whitish pilose Canescent Subglabrous in herbarium specimens. An enumeration of the main differences between F. tincta and F. vargasiana is presented in Table 13. Fuchsia tincta is also sympatric with F. sanctae-rosae. Pennell 13956 (NY), from Pillahuata, is apparently a hybrid between these two species. Diagnostic characters of the putative hybrid and the parental species are given in Table 14. A pollen stainability of 9.2% (500 grains examined) for this collection further supports the hypothesis that it is of hybrid origin. 47. Munz fig 1890, Miguel Bang 674 (BH, holotype; photographs, POM MO shrubs 2.5 m tall. Young growth densely hispidulous to ferrugine 2-4 es 5-10 mm thick, loosly hirtellous, with smooth, purple tan bark. Leaves op- posite or rarely ternate, membranous, elliptic-ovate, rounded to acute at the base, acuminate and often curved at the apex, 6— 15(— 17) cm long, 3-6(-7) cm wide, matte dark green and hispidulous-strigose above, slightly paler to flushed purple and hispidulous below, especially along the nerves; secondary veins 9-14 on either side of the midvein, subelevated below; margin serrulate to denticulate. Petioles pubescent, 2-5(-7) cm long. Stipules lance-linear, acuminate, dark, 2-3 mm long, 0.2-0.4 mm wide. Flowers few and pendant in short, corymbiform, terminal racemes; rachis 1.5-5 cm long with bracts 10-25 mm long. Pedicels slender, pubescent, 15^10 mm long, lengthening considerably from bud to fruit. Ovary ellipsoid-oblong, 5-8 mm long, 2-3 mm wide, densely pubescent. Floral tube narrowly funnelform, (22-)25^18 mm long, 2-3 mm wide and slightly nodose at the base, constricted to 1.5-2 mm wide above the nectary, gradually widened 6-9 10-18 mm long, 3-5 mm wide, mostly long acuminate with free tips in bud for 1-3 mm, spreading at anthesis. Tube dull pink to lavender 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 15. Comparison of morphological characters in Fuchsia furfuracea, F. sanctae-rosae , and presumed hybrids. Hybrids F. furfuracea (6653, BM, F) and (6744, BM, F) 1 (6693, BM) 1 Hybrids (6719, BM, F) i F. sanctae-rosae (6653, NY) and (6693, F, NY) 1 Stem pubescence Leaf shape Leaf pubescence Densely pilose, reddish hairs Broadly ovate Strigillose on both sides Loose villous Loose villous Glabrous Lance-elliptic Densely villous on veins below Narrowly elliptic Narrowly elliptic Villous on both sides Leaf margin Number of secondary veins Inflorescence type Serrulate Lightly serrulate Serrulate Glabrous except hairs on mid- vein below Entire 11-18 Tight terminal ra ceme 1—4 cm long 10-16 11-12 5-10 Subracemose, 6- Subracemose, 8- Axillary along 12 cm long 16 cm long uppermost 25 cm of branches i All numbers refer to W. Brooke collections from Incachaca, Dept. Cochabamba, Bolivia, col- lected in Aug. 1950. or orange red; sepals light red but becoming pale white toward the tips. Petals bright pink red to dark red, elliptic-ovate, mostly about Vi as long as the sepals, 6—10 mm long, 3-5 mm wide, acute at the apex, suberect at anthesis. Nectary unlobed or shallowly 4-lobed, ca. 1.5 mm high. Filaments lavender, 7-8 mm and 4-5 mm long; anthers oblong, 2-3 mm long, 1-1.5 mm wide. Style villous from the base to near the rim of the tube; stigma capitate, 4-cleft at the apex, 2-3 mm long, 2-3 mm wide, white. Berry ovoid-ellipsoid, becoming subglobose when fully ripe, 12-16 mm long, 6-12 mm thick, pubescent; seeds ca. 1.5 mm long, ca. 1 mm wide. Distribution: Bolivia. Rare shrubs growing mostly in dripping wet sites such as cliff overhangs and along rivulets in cloud forest of Depts. La Paz and Co- chabamba; 2,800-3,050 m (Fig. 62). Specimens examined: Bolivia, cochabamba: Incachaca, Brooke 6744 (BM, F). la paz: San Felipe, Sur Yungas, Asplund 1743 (UPS); near Chuspipata, Km 65 of La Paz-Beni RR, Barker 220 (US); valley of Zongo, Prov. Murillo, Beck 2173 (MO); 9 km from Unduavi on road to Coroico, Berry 2577 (MO); Unduavi, Cardenas 3617 (RSA); Carretera Fundamental 3, 27 km SW of Yolosa junction, Davidson 4937 (MO); Unduavi, Rusby 2511 (NY); 20 km SW of Yolosa junction toward Unduavi, Solomon 4860 (MO), 4865 (MO); Yungas valley, Tutin 1391 (BM); 19 km from Unduavi to Coroico, Vargas 22057 (CUZ); Prov. of Yungas, Weddell 2291 (P); valleys between Tipuani and Apolobamba, Prov. Larecaja and Caupolican, Weddell 4601 (P). without locality: Bridges 283 (LE), in 1846 (BM, CGE, K); Cuming 229 (W), s.n. (W). The few-flowered, compact, terminal inflorescences and mostly opposite, pu- bescent, long petiolate, ovate leaves closely ally this species to Fuchsia tincta and F. vargasiana, both of which are endemic to southern Peru. The flowers of F. furfuracea are larger than those of/ 7 , tincta, and the fruits are shorter than those of F. vargasiana. Fuchsia furfuracea is further characterized by its elongate stipules, mostly ferrugineous hairs on older growth, and pale sepals usually about twice as long as the petals. All three species in the F. tincta species group have a very restricted range. 1982] BERRY— FUCHSIA SECT. FUCHSIA 161 Table 16. Pollen stainability of Fuchsia furfuracea, F. sanctae-rosae, and suspected hybrids. Taxon Stainability 2 Fuchsia furfuracea (Brooke 6744, BM, F) Hybrid (Brooke 6693, BM) Hybrid (Brooke 6719, BM) Hybrid (Brooke 6653, F) Fuchsia sanctae-rosae (Brooke 6653, NY) 88.4% 0% 20% 8.6% 83.8% 1 All specimens collected in Aug. 1950, around Incachaca, Dept. Cochabamba, Bolivia 2 500 grains counted for each collection. Fuchsia furfuracea is known from a few localities in the "yungas," or moist, cool valleys of the northeastern slopes of the Bolivian Andes, and it is furthermore restricted to very moist habitats such as streamsides or near waterfalls. Weddell 4061 (P), from Pro v. Larecaja near the Peruvian border, has floral tubes just 22 mm long and is distinguishable from F. tincta only by its slightly longer, browner pubescence and proportionately shorter petals. There are no other collections from the Peru-Bolivia border area, however, to indicate if more intergradation between these two species occurs there. In Dept. La Paz, Fuchsia furfuracea is found close to, but at higher elevations than F. boliviana and F. sanctae-rosae; it occurs at lower elevations than F. denticulata. In Dept. Cochabamba, a series of probable hybrids with F. sanctae- Winifred rph sumed parents in Table 15, and Table 16 shows the strongly reduced pollen stainability of the same intermediates. I returned to Incachaca in July 1979 and found plants of F. sanctae-rosae abundant in weedy thickets, but no trace of F. furfuracea. The valley al furfi Munz fig namayo and Tambomayo, 2,000 m, 25 July 1936, Cesar Vargas C. 73 (BH, MO. POM West West Fuchsia tincta sensu J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):564. 1941, pro parte, sensu Munz, Proc. Calif. Acad. Sci. IV. 25:43. 1943, pro parte. Erect shrubs 1-2 m tall. Young growth finely soft pilose with white to rusty hairs; older branches 4-10 mm thick, dull tan-brown. Leaves opposite, membra- nous, narrowly ovate-elliptic to narrowly subdeltoid, rounded to obtuse or un- equal at the base, acuminate at the apex, 6-23 cm long, 2.5-9 cm wide, subnitid dark green and sparsely strigose to subglabrous above, pale green below and finely pilose mostly along the veins; secondary veins 12-18 on either side of the midvein; margin glandular- serrulate or denticulate. Petioles pubescent, 12-60 mm long. Stipules lanceolate, dark, 1-2 mm long, ca. 0.4 mm wide, deciduous. Flow- 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 ers usually 3-10 in terminal, corymbose racemes; rachis 2-8 cm long, at times with reduced leaves subtending the flowers. Pedicels ascending when young to divergent-drooping at anthesis, 10-35 mm long. Ovary narrowly cylindrical, pu- berulent, 10-11 mm long and ca. 2 mm thick. Floral tube narrowly funnelform, 35-40 mm long, 2-3.5 mm wide and slightly nodose at the base, narrowed to 1.5-2.5 mm above the nectary, then gradually widened above until 4-8 mm wide at the rim, pilulose outside, villous inside in lower %. Sepals lanceolate, acumi- nate, 1 1-16 mm long, 4-5 mm wide, with a short tip 1-2 mm long in bud, strongly spreading to divergent at anthesis. Tube red, sepals green except for the reddish base. Petals red, oblong to broadly elliptic, 10-12 mm long, 5-8 mm wide, rounded and usually with a short point at the apex, strongly spreading at anthesis. Nectary unlobed, ca. 2 mm high. Filaments red, 6-10 mm and 4-8 mm long; anthers oblong, 2-3 mm long, 1-1.5 mm wide, white. Style red pink, villous from the base to near the rim of the tube; stigma capitate, ca. 2 mm long and wide, 4-parted at the apex, white to dull pink. Berry narrowly cylindrical-fusiform, often curved and usually constricted at the apex, 25-32 mm long, ca. 8 mm thick; seeds 1.3-1.6 mm long, 0.9-1.1 mm wide. Gametic chromosome number n = 11. Distribution: Endemic to Dept. Cuzco, Peru. Locally frequent in moist shady thickets along the Rio Tambomayo, Pillahuata area of Prov. Paucartambo; 1.700-2.300 m (Fie. 62). Specimens examined: Peru, uzco: Km 132-140 of Cuzco-Pilcopata road, below Buenos Aires, Berry et al. 3000 (MO, USM), 3003 (MO), 3005 (MO, USM); between Yanamayo and Santa Isabel, valley of Rio Cosnipata, Scolnik 851 (RSA); Tambomayo, Vargas 22986 (MO); Pillahuata, Vargas 22985 (MO); Suecia, Woytkowski 128 (USM). This species is closely related to the sympatric Fuchsia tincta and to the Bolivian F. furfuracea. It has distinctive long, fusiform, curved fruits and nar- rower leaves than either of the above species, and its sepals are green-tipped. Its sympatric occurrence and apparent lack of intermediates with F. tincta are dis- cussed under that species, and a comparison of their distinguishing characters is presented in Table 13. The label of Vargas 4524 (BH) records the collection locality as "Dept. Cuzco, Prov. Convention, 3,600 m," but this is probably in error, since it is very unlikely that this species would occur at so high an elevation. Fuchsia vargasiana is sympatric with F. sanctae-rosae, F. tincta, and populations of Fuchsia related to F. denticulata (Fig. 11). 49. Fuchsia boliviana Carriere, Rev. Hort. 48:150, pi. 1876. Hemsl., Garden 11:70, pi. 1877. Johnst., Contr. Gray Herb. 75:37. 1925. Fawc. & Rendle, Fl. Ja- maica 5:413, fig. 149. 1926. Travis, Fuchsia Annual 1975:55, photo. 1975. non Britton, 1890. Fuchsia boliviana var. typica Munz, Proc. Calif. Acad. Sci. IV. 25:53, pi. 8, fig. 42. 1943. Based on F. boliviana Carr. type: Plate in Revue Horticole 48:150. 1876, drawn from plants cultivated in France in 1876 from seeds collected in the mountains of Bolivia at "6000 m" (a probable error for 6,000 ft.) in 1873 by Benedict Roezl. Figs. 34, 43, and 64 (lectotype, here designated). Fuchsia corymbiflora auct., non Ruiz & Pavon, 1802. Lindl., Bot. Reg. 26: pi. 70. 1840. Harrison, 1982] BERRY— FUCHSIA SECT. FUCHSIA 163 Floric. Cab. & Florist's Mag. 9: /. /. 1841. Paxt., Paxton's Mag. Bot. 8:7,/?/. 1841. Hook., Bot. Mag. 69: pi. 4000. 1843; Gard. Chron. 1845:192, ./ig. 1845, Sweet, Ornam. Fl. Gard. 1: /. 29. 1854. Regel, Gartenfl. 28:241, /ig. 1879. Sieb. & Voss. in Vilm., Bulmengart. ed. 3, 1:333, fig. 18%. Watson, Garden 55:74, fig. 1899. Hegi, 111. Fl. Mittel-Eur. 5(2):801,^#. 2189, a^d. 1925. Bonstedt in Pareys Blumengart. 2:79, photo. 1932. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):551. 1941, pro parte. Fuchsia corymbiflora alba Harrison, Floric. Cab. Florist's Mag. 11:97 9 fig. May, 1849. Paxt., Paxton's Mag. Bot. 16:219. 1849. van Houtte, Fl. Serres Jard. Eur. 6:29, fig. 1850. Essig. Nat. Hort. Mag. 13:6, photo. 1934. lectotype: the figure in Harrison, Floric. Cab. Florist's Mag. 17:97. 1849, illustrated from plants grown by John Salter in 1849 in London, England from plants obtained from "Continental Europe. i i Fuchsia boliviana var. luxurians I. M. Johnston, Contr. Gray Herb. 75:37. 1925. Munz, Proc. Calif. Acad. Sci. IV. 25:54. 1943. type: Venezuela, Edo. Aragua, Colonia Tovar and vicinity, 1,700- 2,300 m, 19 Feb. 1921, Henry Pittier 9252 (GH, holotype; photograph, UC; NY, US, VEN, iso- types). Fuchsia cuspidata Fawcett & Rendle, Journ. Bot. 114:105. 1926. type: Jamaica, Blue Mountains, near Woodcutter's Gap, ca. 1,200 m, 10 Aug. 1895, William Harris 5825 (BM, holotype). Fuchsia boliviana forma puberulenta Munz, Proc. Calif. Acad. Sci. IV. 25:54. 1943. type: Bolivia, Dept. La Paz, Prov. Nor Yungas, 1,800 m, Dec. 1917, Otto Buchtein 732 (F 588700, holotype; photographs, POM, UC; F, G, GH, MO, NY, US, Z, isotypes). Erect bushy shrubs or small trees 2-4.5(-6) m tall with arching-pendulous branches. Young growth usually densely canescent with fine, erect, gray- white hairs; terminal shoots 2-4 mm thick, terete to angled; older stems woody, usually hollow, 1-6 cm thick, with tan brown, splitting bark. Leaves opposite, ternate, or sometimes alternate near the uppermost branching nodes, softly membranous, narrowly to broadly elliptic or ovate, acute to rounded at the base, acute to acuminate at the apex, 5-20(-23) cm long, 3— 12(— 15) cm wide, medium to dark matte green and pubescent to subglabrous above, pale green and ashy puberulent to densely canescent below, especially along the nerves; secondary veins (12-) 15-25 on either side of the midvein, ± impressed above, prominent and often reddish below; margin glandular-denticulate. Petioles pubescent, 2-5(-7) cm long. Stipules dark, narrowly lanceolate, 0.5-1.5 mm long, ca. 0.3 mm wide, deciduous. Flowers numerous in terminal, drooping racemes or few-branched panicles, the flowers congested near the tips; rachis 5-30 cm long in flower, 15-40(-60) cm long in fruit; bracts lanceolate, reflexed, 5-25 mm long, 2-8 mm wide. Pedicels slender, pendant, pubescent, 5— 12(— 16) mm long. Ovary cylindrical, 7-11 mm long, 2-3 mm thick, strigose to puberulent. Floral tube narrowly funnelform (25-)30-60(-70) mm long, 1.5-3 mm wide and slightly bulbous at the base, grad- ually widened above until 4-7(10) mm wide at the rim, pubescent outside, pilose inside for entire length. Sepals lanceolate, acuminate, 10-20 mm long, 4-5 mm wide, tips apiculate in bud, initially spreading at anthesis, but soon becoming totally reflexed. Tube and sepals pale pink to bright scarlet, rarely pale white. Petals scarlet, acute at the apex, ± crispate with 2-3 longitudinal ridges, oblong to elliptic, 8-16(-20) mm long, 3-7(-9) mm wide, upper Vi recurved at anthesis, drying and dehiscing before the tube does. Nectary 4-lobed, green, 2-3 mm high. Filaments red, 8-15 mm and 5-10 mm long; anthers oblong, 2-3.5 mm long, 1-1.5 mm wide, white. Style slender, pilose from the base to the rim of the tube; stigma capitate or clavate, subtetragonous, shallowly 4-lobed at the apex, 2.5-3 mm long, 3-5 mm wide, cream, situated at the level of the antesepalous anthers or barely exserted beyond them. Berry cylindrical, 10-26 mm long, 8-14 mm thick, dark purple, strigose, comestible; seeds tan, 1.3-2 mm long, 0.5-1 mm wide. Gametic chromosome number n = 1 1 . 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Figure 64. Fuchsia boliviano Carriere. Drawn from living material from El Portachuelo, D.F., Venezuela. 1982] BERRY— FUCHSIA SECT. FUCHSIA 165 Distribution: The probable native range extends from northern Argentina to southern Peru, undoubtedly naturalized elsewhere outside South America and in Colombia and Venezuela; cloud forest shrubs in moist thickets; (600-) 1 ,000-3 ,000 m (Fig. 65). Representative specimens examined (South America): Venezuela, aragua: Colonia Tovar, Agostini 65 (VEN), Allan 354 (MY, NY, US, VEN), Berry 2592 (MO, VEN), Fendler in 1854-55 (K), Jahn 439 (US), Woronow 7048 (LE); between El Lagunazo and Colonia Tovar, Aristeguieta 781 (VEN), Trujillo & Fernandez 852 (MY); La Victoria to Colonia Tovar, between Las Moras and Lagunita, Benitez de Rojas 566 (MY); Hacienda El Limon, Las Aguaitas, Delascio 2057 (VEN); between Colonia Tovar and Pico Codazzi, Ruiz-Terdn & Lopez-Palacios 101 18 (MERF, MO); La Lagunita, near Colonia Tovar, Schnee 1309 (MY), ca. 5 km from Colonia Tovar towards El Limon, Solymosy S10197150 (VEN). distrito federal: between El Junquito and Colonia Tovar, Berry 2591 (MO, VEN), Wessels-Boer et al. 2445 (MO, U); 1-2 km below junction of El Junquito-Colonia Tovar road towards Carayaca, Cardenas de Guevara 433 (MY), Davidse & Morillo 3985 (MO), Luteyn et al. 5162 (MO, NY); 4 km ENE of Colonia Tovar, Steyermark 86212 (MY, NY, VEN). Colombia, antioquia: Las Minas, Barkley 17C250 (GH); near Boqueron del Toyo, Barkley & Neira 17C235 (CAS, COL, GH); canyon 20 km E of Sonson, road to El Dorado, Barkley 38C568 (COL, GH); Alto de Minas, Medellin to La Pintada, Escobar 1000 (MO); near San Jose de San Andres, Correa & Velas- quez 68 (COL, RSA, US); between Alto San Miguel and Alto Alegrias, 15 km SSE of Caldas, Fosberg 21163 (NY, RSA, US); Alto de Minas, between Caldas and Santa Barbara, Garganta 2003 (US); El Poblado, near Medellin, Hatheway in 1955 (B); Las Minitas, S of Caldas, Pennell 10970 (GH, NY); Cerro del Pedre Amaya, Boqueron, Km 22 of Medellin-Santa Fe de Antioquia road, Rivera 808 (PSO). caldas: Termales, Dryander 2756 (BH, F, VALLE); Manizales, Sandeman 5655 (COL, K). choco: N of Alban, near Dept. Valle border, Dugand & Jaramillo 3046 (US); 2-3 km N of Carmen de Atrato, Fosberg & Core 21553 (RSA, US), cundinamarca: 2.5 km W of Salto de Tequendama, Barclay et al. 3303 (COL, NA, US), Luteyn & Lebron-Luteyn 7711 (COL, MO, NY); Bogota, Duque- Jaramillo 3049 (COL, NY), Perez-Arbelaez 4802 (NY, U, US); 17 km W of Mosquera towards La Mesa, Gentry 15139 (COL, MO); Km 26-28 SW of La Mosquera towards La Mesa, Luteyn et al. 4690 (COL, MO); Zipaquira, Pring 168 (MO); Bogota, Chapinero, Schneider 353-A (COL); 2 km from Zipaquira on road to Pacho, Schultes et al. 3 (GH, K, S, US), santander: 24 km W of Jordan on Cimitarra-Velez road, Gentry & Forero 15466 (COL, MO); 29 km W of Fresno towards Manizales, Berry 3552 (COL, MO); above Cajamarca, Giradot-Armenia road, Berry 3143 (MO); La Palmilla, Quindio, Triana 3812 (G, P); Herveo, Uribe-Uribe 756 (COL), valle: N of Las Brisas, Carrizales, Cuatrecasas 22552 (F, RSA, VALLE). Peru, apurimac: Abancay, Blood & Tremelling 206 (NA): Chincheros, Ferreyra 2810 (USM), Soukup 6248 (RSA); 8 km E of Abancay, road to Cuzco, Gentry et al. 23368 (MO); Ampuy, Stork et al. 10595 (A, G, MO, NA, UC); Chirhuay-Matara, Vargas 1971 (BH, CUZ, MO); Nacchero, Vargas 8962 (CUZ); Huancarema, West 3783 (MO, UC). ayacucho: 11 km below Jano, road Tambo-Ayna, Berry 3057 (MO, USM); Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22280 (NY, RSA, US); above Jano, Plowman & Davis 4673 (GH, K); Curahuasi, Sandeman 3552 (K, OXF); Ocros, Vargas 15912 (CUZ); Prov. La Mar, Weberbauer 5555 (F, GH, US), cuzco: Paucartambo, Balls B6715 (BM, K, NA, UC, US), Plowman & Davis 4910 (GH); Urubamba, Berry 2561 (CUZ, MO), Herrera 1058 (GH, US), Soukup 28 (F), Km 169 of Cuzco-Quillabamba road, Berry & Aronson 3044 (MO, USM); San Miguel, Chavez 3108 (MO), Cook & Gilbert 915 (US); Hacienda Santa Rita, Urubamba valley, Dreyfus in 1941 (USM); Machupichu, Ferreyra 2728 (USM), Rauh 773 (RSA), Vargas 990 (BH, CAS, CUZ, MO); Lucay, Urubamba valley, Herrera 1123 (F, US); Hacienda Churu, Prov. Paucartambo, Herrera in 1926 (US); Km 105 along RR in Urubamba valley between Ollantaytambo and Machupichu, litis & Vgent 1132 (DS, GH, K, UC, USM); Punto Real, Urubamba valley, Tutin 1308 (BM); Ollantaytambo, Tutin 1340 (BM); 5 km NE of Paucartambo on road to Pillahuata, Vgent & Vargas 4410 (DS); Huaisanpilla, Vargas 9945 (CUZ); Mantla, Km 84, Prov. Calca, Vargas 15630 (CUZ); Urubamba valley, between La Maquina and Cedrobamba, West 3783 (GH, MICH, MO, NA, RSA, UC, US, USM); Pilco, valley of Rio Paucartambo, Woytkowski 320 (MO, UC); near Chacra, Prov. Paucartambo, Woytkowski 236 (USM). huanuco: 5 km SE of Carpish, Stork & Horton 9921 (F, MICH, MO, RSA, UC, US), puno: near Sandia, Ferreyra 16652 (MO); Oconeque, E of Limbani, Hodge 6166 (US); Asalaya, Vargas 11812 (CUZ); near Sandia, Vargas 14808 (CUZ), 16414 (CUZ), 2/246 (CUZ), Weberbauer 537 (G), 557 (G). Bolivia, cochabamba: Km 205 Cochabamba-Santa Cruz, Badcock 382 (K); ca. 60 km from Cochabamba to Villa Tunari, Berry 2588-B (MO); Quime, ca. 160 km from Oruro via Eucalyptus, Quimicruz mts., Brooke 5385 (BM, F), 5418 (BM); Rosal, near Rio Limon, Brooke 5706 (BM, F, G, NY, U), 5719 (BM, F); Choro, above Rio Cocapata, ca. 160 km NW of Cochabamba, Brooke 6076 (BM, F, G, NY, S, U); Incachaca, 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 —t— 300 Figure 65. Distribution of Fuchsia boliviana and F. verrucosa. 1982] BERRY— FUCHSIA SECT. FUCHSIA 167 Brooke 6653 (BM), 6670 (BM, F, NY), 6650 (BM, F), 6719 (BM), Steinbach 4306 (K), 5024 (A, G), 576J (A, F, G, GH, K, MO, PH, US); Parangani, Ayopaya, Cardenas 4086 (GH, RSA); 1 10 km NE of Cochabamba, near Chimore, S side Rio Mateo, Eyerdam 24740 (F, UC); Pocona-Tal, Steinbach 8781 (BM, GH, K, MO, NY, S); El Limbo, Steinbach 537 (G, GH, MO, NY, U, US), la paz: Chungamayo, La Sirena, Asplund 317 (UPS); El Chaco, Sur Yungas, Asplund 1131 (UPS); Yungas, Bang 327 (BM, F, GH, K, LE, MICH, MO, NY, PH, US, W); Zongo Valley, Prov. Murillo, Beck 2172 (MO), 2739 (MO); Huancane, Sud Yungas, Beck 3066 (MO); 13 km from Unduavi to Coroico, Berry 2579 (MO); Chulumani, Brooke 6560 (BM); Sirupaya, near Yanacachi, Sur Yungas, Buchtein 102 (NY), 218 (NY, US); Sorata, Cardenas 1156 (NY); 15 km SW of Yolosa junction, Davidson 4913 (MO); Caranavi, Ellenberg 6217 (GOET); near Unduavi, Eyerdam 25395 (F, UC); El Chaco, Sur Yungas, Holway & Holway 643 (US); between Puente Villa and Chulumani, Kelley 1013 (CM, UC); Sacramento, Unduavi to Coroico, Kelley 1075 (CM, F, UC); near Sorota, Mandon 622 (BM, F, G, GH, K, LE, NY, P, S, T, US, W); Nor Yungas, Mexia 04286 (GH, MO, UC); Colaya, Sur Yungas, Mexia 04292 (G, GH, MO, UC), 7809 (F, G, K, MO, S, U, UC, US); near Yungas, Rusby 1071 (BM, F, GH, K, LE, MICH, NY, P, PH, US); Okara, Tate 908 (NY); Prov. Apopaya, Weddell4179 (P). santa cruz: Samaipata, Steinbach 3755 (GH, POM); Yungas de San Mateo, Steinbach 8392 (BM, GH, K, NY, S); Comarapa, Steinbach 8459 (BM, GH, K, MO, NY, S). without locality: Bang 1806 (BM, F, G, GH, K, LE, MICH, MO, NY, PH, US, W, Z); Bang 2005 (BM, F, GH, K, MO, NY, US, W); Bang 2833 (F, GH, K, LE, MICH, MO, NY, PH, US). Argentina, jujuy: road to Valle Grande, Dep. Ledesma, Cabrera et al. 22390 (K); San Lorenzo, Jorgensen in 1911 (POM); 10-20 km from Libertador Gral. San Martin, road to Valle Grande, Krapovickas et al. 26668 (MO); Valle Grande, ca. 2 km past Abra de Canas, Legname & Cuezzo 8601 (GH); Abra de Canas, road to Valle Grande, Vervoorst & Cuezzo 7851 (W). salta: San Andres, Lorentz & Hieronymus 265 (G, S), Pierotti 312 (A, NY, UC). tucuman: Quebrada de Las Sosas, Bocher et al. 2419 (MO, S), Killip 39530 (RSA, US), Walter & Walter 621 (B), 653 (B); Casa de Piedras, ca 25 km SSW of Tucuman, Colaris 1181 (U); along Rio Las Sosas, ca. 60 km SW of Tucuman, Haas 893 (U); Sierra de Aconquija, Humbert 20929 (P, US); Rio Cicerone, Jorgensen 21 (GH, MO); Km 20 road to Tafi del Valle, Legname 96 (CAS, MO, W); Rio Cochuna, Chicligasta, Lillo 1510 (GH), O'Donnell 10429 (BM, F, UC); Quebrada Caspinchango, Lillo 7340 (GH), Venturi 9598 (A, S); road to Tafi del Valle, Monteros, Lourteig 450 (A, U, US); Villa Nougues, above Tucuman, Munz 15469 (GH, NY, POM, US); from San Javier to Villa Nougues, Ortiz in 1945 (A, BM, UC); Quebrada de Lules, Quiroga 7215 (F), Venturi 1295 (BM, CAS, F, GH, K, MO, RSA, UC, US, W), 2232 (A, POM, US, Z); Quebrada de Lules, Arroyo Grande, Sleumer 759 (B); Estancia Las Pavas, Dept. Chichligasta, Venturi 4657 (A, F, US). Brazil: Porto Alegre, cultivated, Revicely in 1898 (W). Chile: Jardin Suizo, Las Zorras, Valparaiso, cultivated, Harshberger 1046 (P, US); Prov. Concepcion, cultivated, Junge 2975 (US). Naturalized or cultivated specimens outside of South America: Azores: Pico Island, above San Roque, Brooke 11337 (BM); Graciosa, Praia, Santana, Gongalves 3113 (BM). Costa Rica, alajuela: Zarcuo, A. Smith 2775 (F). cartago: Cartago, Donnell Smith 4804 (US), san jose: San Jose, Pittier 14104 (CM, GH, U, US). El Salvador. Volcan de San Salvador, Calderon 2345 (F, US). Guate- mala, sacatepequez: San Rafael, Donnell Smith 2176 (K, US); Antigua, Standley 62331 (F). san marcos: Volcan Tajumulco, Johnston 1231 (F), Steyermark 62331 (F). India: Ootacamund, Nilgeri Hills, Anstead 122 (A), Koelz 11048 (NA), Marcovicz in 1927 (LE); Kodaikanal, Bembower 9 (MO); Madras, Shembayanu, Sauliers 656 (A). Jamaica: Cinchona Garden, Blue Mts., Downes in 1926 (BM), Harris 7643 (G), Sporne 213 (CGE), ? 7605 (BM, F, K); vicinity of St. Helen's Gap, St. Andrew, Maxon & Killip 571 (A, F); near New Haven Gap, Philipson 914 (BM), Roever4574 (MICH); Latimer River, Seifriz in 1919 (BH). Java: Preanger, Pengalengan, Hochreutiner 1463 (G); Bandang, Rauh 13 (U). Mexico, chiapas: Piedracitos, Chamula, Breedlove 8018 (DS). distrito federal: Villa Obregon, Moore 6395 (BH); Coyoacan, Woronow 2065 (LE). puebla: Huauchinango, Baldwin 14390 (LL). veracruz: Orizaba, Botteri 929 (GH, K, LE); Jalacingo, Dodds 57 (MO); Altotonga, Foster in 1938 (BH, BM); Santa Ana Atzacana, N of Orizaba, Rosas 191 (MO); Santa Cruz, Altotonga, Ventura 1026 (CAS). Portugal: Coimbra, Rhodes 37-64-126 (BH) Reunion: Foret de Bebour, Lorence & Cadet 2423 (MO); Cilac, Reserve Matambu, Lourteig 2455 (MO), Schlieben 10966 (B, Z). Soviet Union: Leningrad Botanical Garden in 1843 (LE). Sri Lanka: Hadgala Garden, Marcovicz in 1927 (LE). St. Helena: Longs, ridge above Sandy Bay, Kerr 107 (BM); Diana's Peak, Salter 38418 (BM). Unites States, California: Univ. Calif. Botanical Garden, Berkeley, Hutchison 49.802 (MO, RSA, UC, US); Los Angeles, McClintock in 1942 (NA). Unlike most species of sect. Fuchsia, which are often scandent with weak stems and branches, Fuchsia boliviana is sometimes arborescent and almost al- characters fruits 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 reflexed sepals after anthesis, and the early deciduous, ridged petals. In addition, this species generally has large, long petiolate, softly pubescent leaves that are opposite or ternate and sometimes alternate near the branch tips. It has been widely cultivated and naturalized in many countries of the world since the mid- 1800s, but has long been confused in the literature with F. corymbiflora. The true F. corymbiflora has apparently never been cultivated and is a localized endemic in central Peru that has much finer pubescence than F. boliviana. It has strictly opposite, short-petiolate leaves, rounder ovaries, much shorter inflorescences, smooth, persistent petals, and non-reflexed sepals. Fuchsia boliviana is distinct and easily recognizable from other species of Fuchsia, but it is placed in a species group with F. corymbiflora, F. mathewsii, and F. wurdackii because of their large, pubescent, mostly opposite or ternate leaves and many-flowered racemes of long flowers with narrow petals. It also resembles F. furfuracea in the often ovate leaf shape, long petioles, similar floral dimensions, and dense pubescence. As discussed in the section on reproductive biology, many populations of F. boliviana are largely autogamous. Populations from Venezuela and green- house-raised progeny from Bolivia both produced fully fertile fruits when isolated from pollinators. The unusually high degree of autogamy in this species is due to the close proximity of the stigma to the upper staminal whorl in many populations and to the loss or very slight degree of protogyny. The pattern of the latter stages of floral maturation is unique in F. boliviana. As the sepals split open and start spreading outwards at anthesis, the upper half of the petals begin to curve out- wards. Within a day, the sepals become totally reflexed, and the petals attain their final, spreading-recurved posture. About two days after anthesis, the petals shrivel up and fall off, leaving the erect stamens and style along with the sepals reflexed back against the floral tube. The flowers remain on the plant for another day or two before the entire tube usually dehisces. In addition to autogamy, vegetative reproduction is common in F. boliviana, occurring mostly by sucker growth or rooting from broken or cut stems. These two factors account to a large degree for the widely naturalized distribution of this species. The plants are prized for their very striking, numerous, red flowers in long, drooping racemes that can each last several months, and single seeds are sufficient to start extensive local populations in suitably cool, wet areas. In ad- dition, F. boliviana has broader ecological tolerances than most species in the section and can survive short periods of drought and full sunlight. The native range of F. boliviana appears to be centered in cloud forest on the eastern slopes of the Andes in southern Peru and Bolivia, possibly extending into northern Argentina. This conclusion is based on historical information as well as an analysis of the habitats and present-day distribution of the species. The only areas where F. boliviana can be said to form an integral part of relatively undis- turbed tracts of forest is in southern Peru and Bolivia. Elsewhere, this species is found only close to towns, habitations, or heavily disturbed areas, as is typical of adventives or plants escaped from cultivation. Even in southern Peru, many collections are from cultivated or escaped plants. In Venezuela, the only popu- lations of this species are found in the Cordillera de la Costa, where no other native Fuchsia occurs. Plants there grow mostly around the Colonia Tovar, an 1982] BERRY— FUCHSIA SECT. FUCHSIA 169 old German agricultural colony that has received numerous introductions of plant material from travelers and from the numerous botanists who stayed there, such as Fendler, Moritz, and Pittier. F. boliviano is very commonly cultivated in the Sabana de Bogota in Colombia and elsewhere throughout the mountains of that The northern Woodcutter naturalized (Adams, 1972 and pers. comm. to P. H. Raven). According to Vargas (1964), F. boliviano was cultivated by the Incas and used as a motif in clay and wood artifacts. It continues to be cultivated today in the Cuzco area. The earliest cultivated plants of F. boliviano in Europe were grown Peru nurseryman the seed from garden plants there (Lindley, 1840). Progeny of these plants gave rise to many naturalized populations in other parts of the world. Seeds imported from Bolivia in 1873 were also grown in Europe and used to describe F. boliviano. Several varieties or forms of F, boliviano have been described based on dif- ferences in floral tube length and degree of pubescence. F. boliviano var. luxu- rians was described by Johnston (1925) for plants with floral tubes 5-6 cm long from Venezuela, Central America, and Jamaica. In contrast, most plants from Argentina and Bolivia have much shorter floral tubes 3-4.5 cm long and shorter pubescence. The long-flowered populations generally have a more robust growth form and larger inflorescences. The leaves of the small-flowered Bolivian and Argentinian populations may be narrower in many cases than in naturalized plants, but considerable variation is found both in leaf and in floral tube dimensions. For example, Kelley 1013 (CM, UC), from Sur Yungas, Bolivia, has some floral tubes 6 cm long and others only 4 cm long. Except for the smaller size of flowers and inflorescences in most southern populations, all of the characteristic floral and vegetative traits of these plants are identical to those of the more robust natu- ralized populations, and both kinds of plants are diploid. In Dept. Cuzco, Peru, both robust, long-tubed plants as well as shorter-tubed plants are found. Plants such as Balls B6715 (BM, K, NA, UC, US), cultivated in Paucartambo, near Cuzco, have floral tubes 5-6 cm long and closely resemble the naturalized populations in northern South America, Central America, and Jamaica. As discussed above, the earliest cultivated plants outside of Peru were from Cuzco, and illustrations in Lindley (1840) and Paxton (1841) appear to be from long-tubed plants. Since cultivation of this species in the Cuzco area dates back to the Incas, it is reasonable to suppose that robust, long-flowered plants were selected for cultivation by the local inhabitants, and these were later dis- covered by European visitors who sent seeds abroad to give rise to many of the naturalized populations known today. Due to the predominant autogamy of this species, populations with different floral tube lengths or with differences in the length and density of pubescence can easily be maintained. The only cultivated or naturalized populations with short floral tubes 4 cm long or less are from Portugal (Rhodes 37-64-126, BH) and some from India. Crosses between long- tubed plants from Colonia Tovar, Venezuela, and short-tubed plants from Dept. 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 La Paz, Bolivia, were made at Missouri Botanical Garden in 1979. These have given rise to normal, vigorous plants, with intermediate flower size. A white-tubed variant of F. boliviano was described by Harrison (1849) as "Fuchsia corymbiflora alba." It has also been collected in Colombia {Berry 3552 , COL, MO; Dept. Tolima) and is characterized by a total loss of red pig- mentation in all parts of the plant except the petals. This might be due to a single mutation and is only known in cultivated or naturalized plants. In summary, F. boliviana is a well demarcated species with populations that differ in floral tube length, robustness of the plant, and degree of pubescence. These differences can be explained, however, by a fairly wide and sometimes continuous variation within local populations, a long history of cultivation and probable selection of long-flowered forms, and a high degree of autogamy, which helps maintain small morphological differences. It is a very handsome species that is widely naturalized throughout the world in moist tropical and subtropical, montane habitats, but its probable native range extends from southern Peru (Dept. Apurimac of Cuzco) to Bolivia or northern Argentina. It is known to grow sym- patrically with F. sanctae-rosae in Peru and Bolivia, but no hybrids between these two species have been detected. 50. Fuchsia corymbiflora Ruiz & Pavon, Fl. Peruv. Chil. 3:87, pi. 25, fig. a. 1802. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):551. 1941, pro parte. Munz, Proc. Calif. Acad. Sci. IV. 25:49, pi. 7, fig. 38. 1943, pro parte, non auct. pro F. boliviana. type: Peru, Dept. Huanuco, woods of Chinchao and Muna, 1778-1788, Hipolito Ruiz and Jose Pavon (MA N°ll/78, lectotype, here designated; photograph, MO). Fuchsia velutina I. M. Johnston, Contr. Gray Herb. 75:36. 1925. type: Peru, Dept. Huanuco, Yanano, ca. 2,000 m, 13-16 May 1923, J. Francis Macbride 37/5 (F 535777, holotype: fragment, G; photo- graphs, NY, POM, UC; GH, K, isotypes). Fuchsia munzii J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):559. 1941. type: Peru, Dept. Junin, Rio Masamerich, 2,100-2,200 m, 1909-1914, August Weberbauer 6648 (F 628941, holotype; pho- tographs, NY, UC; GH, UC, isotypes). Scandent to erect shrubs 1-4 m tall. Young growth finely canescent to pu- berulent; older stems and leaves velutinous to subglabrous; branches few and spreading, branchlets terete, 2-3 mm thick, red to dull purple. Leaves opposite, very rarely ternate, firmly membranous, elliptic to oblong, acute at the base, acute to acuminate at the apex, 6— 1 2(— 1 9) cm long, 3-6(-9) cm wide, matte green above, pale green below; secondary veins 13-17 on either side of the midvein, sometimes reddish below; margin subentire to obscurely glandular-denticulate. 6-20( 2 mm long, ca. 1.5 mm wide, sometimes connate, deciduous. Flowers few to many in terminal, corymbose, arching to drooping racemes or few-branched panicles; rachis 2-8 cm long; bracts lanceolate, 5-15 mm lone. Pedicels pendant to ascending in fruit. 6-15 5-6 40-65( rowed to 1.5-2 mm wide for ca. the basal 10 mm of the tube, then gradually widened above until 5-8 mm wide at the rim, minutely pubescent under a lens 1982] BERRY— FUCHSIA SECT. FUCHSIA 171 to velutinous or strigillose outside, pubescent inside for most of length. Sepals lance-oblong, 12-15 mm long, 3.5-5 mm wide, acute at the apex, buds terete and slightly mucronate, spreading-divergent at anthesis. Tube and sepals pale pink to bright scarlet. Petals darker, red, oblong, acute to obtuse-tipped, 12-17 mm long, 4-5 mm wide, spreading and equal to or longer than the sepals at anthesis. Nec- tary shallowly 4-lobed, ca. 1.5 mm high. Filaments pink to red, 8-10 mm and 5-7 mm long; anthers oblong, 2.5-3 mm long, ca. 1.5 mm wide, white. Style pink, puberulent from the base to the rim of the tube; stigma capitate, 4-cleft at the apex, 2-2.5 mm long, ca. 2.5 mm wide, cream, exserted 3-6 mm beyond the anthers. Berry subglobose, ± tetragonous before maturity, 10-12 mm long, 8-10 mm thick, red; seeds reddish tan, 1.8-2.1 mm long, 1-1.4 mm wide. Gametic chromosome number /i = 11. Distribution: Endemic to cloud forest on the eastern slopes of the central Peruvian Andes in Depts. Huanuco, Junin, and Huancavelica; (l,500-)2, 250-2, 850 m (Fig. 63). Representative specimens examined: Peru, huancavelica: without locality, Weberbauer 6565 (F, GH). huanuco: near Panao, above La Molina, Asplund 13699 (RSA); 5 km W of Carpish tunnel. Berry & Aronson 3082 (MO, USM); Carpish, Ferreyra 1828 (MO, USM); Carpish, above Acomayo, Hutchison et al. 5919 (F, MO, NY, RSA, UC, US, USM); Huacachi, near Muna, Macbride 4180 (F, G); road from Mirador to Chinchao, Mexia 7765 (GH, K, MO, UC, US); Muna, Pearce 148 (K), 512 (BM, K). junin: 55 km above and W of Satipo on road to Concepcion, Berry & Aronson 3079 (MO, USM); San Juan to Huacapistana, Ferreyra 11293 (USM); above Huacapistana, Sandeman 95 (BM, K); Km 170 Satipo-Concepcion, Seibert 2384 (MO, POM, US), without locality: Ruiz & Pavdn (BM, F, MA, MO). rm and terminal racemes of slender flowers with short pedicels and round fruits. These characters place it in the Fuchsia boliviana species group, where it is possibly most closely related to F. wurdackii from northern Peru. For over a century its name was used for the species later described as F. boliviana, a commonly cultivated and naturalized, ornamental plant from southern Peru and Bolivia that has cylindrical fruits, flowers with reflexed sepals and much longer Munz southern ternate southern munzii. They have much paler pink flowers than populations from Huanuco, and their leaves are lighter green with wine purple veins. This appears to be due to the loss of some of the dark red pigmentation in the plant. Although the Junin plants are often nearly glabrous, the characteristic fine, erect hairs of F. cory- mbiflora can usually be seen upon careful examination. They also have the same 4-sulcate, rounded fruits and finely pilose styles as F. corymbiflora. In the Carpish leaf corymbifl and F. denticulata. Fuchsia abrupta is a long-tubed species with opposite leaves that can be confused with F. corymbiflora in dried specimens, but it can be distinguished by its cylindrical fruits, divergent pedicels, often hispid pubescence, and strongly divergent sepals and petals. 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 51. Fuchsia wurdackii Munz, Brittonia 16:229. 1964. type: Peru, Dept. Amazo- nas, Prov. Chachapoyas, Quebrada Molino, 5 km below Chachapoyas, 2,200 m, 30 May 1962, John J. Wurdack 622 (US 2404206, holotype; photograph, MO). Erect shrubs 1-1.5 m tall. Young growth finely pilose, older stems pilose and dull tan, 3-10 mm thick. Leaves opposite or ternate, softly membranous, narrow- ly elliptic-ovate, acute to subacuminate at both ends, 7-18 cm long, 2-8 cm wide, shortly soft pilose on both sides; secondary veins 7-14 on either side of the midvein; margin subdenticulate. Petioles pilose, 7-43 mm long. Stipules conspic- uous, lanceolate, usually with an obtuse apex, pale membranous, 3-5 mm long, 0.5-2 mm wide, sometimes connate, usually persistent and reflexed. Flowers few to many in terminal, corymbose, drooping racemes; rachis 4-18 cm long; bracts lance-linear, denticulate, 15-50 mm long, 3-8 mm wide. Pedicels 9—1 1(— 25) mm long. Ovary oblong, 7-10 mm long, 3-4 mm thick. Floral tube narrowly funnel- form, 31-50 mm long, 2-3 mm wide and slightly bulbous at the base, gradually widened above until 5-8 mm wide at the rim, pilose to short villous outside, pilose inside in lower %. Sepals oblong, obtuse, 9-16 mm long, 3-5 mm wide, spreading at anthesis. Tube and sepals coral red. Petals red, oblong to broadly elliptic, obtuse at the apex, 11-20 mm long, 4-8 mm wide. Nectary unlobed or shallowly 4-lobed, ca. 1.5 mm high. Filaments red, 5-9 mm and 3-6 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide, white. Style red, glabrous; stigma capitate, shortly 4-lobed at the apex, 2-3 mm long, 2-4 mm wide, pink, exserted 1-3 mm beyond the anthers. Berry cylindric to ellipsoid, 17-20 mm long, 9-11 mm thick; seeds 1.6-2 mm long, 0.9-1.2 mm wide. Gametic chromosome number n = 1 1 . Distribution: Northern Peru. Endemic to moist sites on slopes leading into the Rio Utcubamba valley in Dept. Amazonas; 2,100-2,400 m (Fig. 63). Representative specimens examined: Peru, amazonas: 9 km above Leimebamba on road to Balsas, Berry & Escobar 3616 (MO, USM): Leimebamba-La Joya trail, Boeke 1786 (MO); Cerros Calla-Calla, at San Miguel, 5 km above Leimebamba, Hutchison & Bennett 4542 (UC, USM), Hutchison & Wright 4884 (UC); vicinity of Leimebamba, upstream along creek leading into Rio Utcubamba, Hutchison & Wright 4888 (F, MICH, MO, NY, RSA, UC, US, USM); Cerro Puma Urco, SE of Chachapoyas, Soukup 4097 (US). This is a long- tubed, racemosely flowered species that is distinctive in its obtuse petals and sepals, pilose pubescence, and large, pale stipules. Its rather long inflorescences with narrow bracts are similar to those of Fuchsia pilosa, but that species has much smaller flowers and coarser pubescence. Fuch- sia wurdackii shares many features with F. boliviana including the large, pubes- cent, long-petiolate leaves and lanceolate bracts, but lacks the acute, reflexed sepals and long, cylindrical fruits of that species. Its closest relative is probably F. corymbiflora, known from central Peru, which has similar flowers and short, soft pubescence, but shorter fruits and petioles. Since F. wurdackii is the lowest- altitude species in northern Peru, it does not occur sympatrically with any other species of the section, although both F . fontinalis and F. mathewsii occur nearby (Fig. 10). 1982] BERRY— FUCHSIA SECT. FUCHSIA 173 Macbride, Candollea 8:24. 1940; Field Mus Bot. Ser. 13(4):558. 1941. Munz, Proc 1943. Peru 1830-1841, Andrew Math MO Fuchsia fischeri J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4):554. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:45. 1943. type: Peru, Dept. Cajamarca, Prov. Hualgayoc, Chugar, 2,700-2,900 m, 1906, August Weberbauer 4097 (G, holotype). Fuchsia storkii Munz, Proc. Calif. Acad. Sci. IV. 25:37, pi. 6, fig. 33. 1943. type: Peru. Dept. Cajamarca, Prov. Chota, pass S of Conchan, 2,500 m, Harvey E. Stork & Ovid B. Horton 10073 (F 1052248, holotype; photograph, NY; K, UC, isotypes). Suberect shrubs 1-3 m tall, puberulent to mostly densely pilose with white to eenerallv rustv hairs. Branchlets triangular (when leaves are ternate) or rarely purpl 14 mm thick, with pilose, reddish bark exfoliating in strips. Leaves mostly ternate, occasionally quaternate, firmly membranous, narrowly elliptic to oblanceolate or elliptic, at times slightly asymmetrical and curved to one side, narrowly obtuse at the base, acute at the apex, 4-16 cm long, 1.2-5 cm wide, dull to subnitid dark green and pilose to strigose above, lighter green and mostly rusty-pilose below, with hairs rpl 12 on either side of the midvein; margin subdenticulate. Petioles pubescent to dense- ly ferrugineous-pilose, 3-10(-25) mm long. Stipules narrowly lanceolate when young, becoming thick and triangular with age, dark, 2-3 mm long, ca. 1 mm wide, subpersistent. Flowers numerous and crowded at the tips of terminal, arch- 3-6 10-25 pilose, 12-25 mm long. Ovary ellipsoid, 5-7 mm long, 2-3.5 mm thick, green to purple. Floral tube narrowly funnelform, (32-)36-63 mm long, 2.5-3.5 mm wide and slightly bulbous at the base, narrowed to ca. 2 mm wide above the nectary, then widened gradually above until 5-8 mm wide at the rim, mostly pilose outside, pilose to villous inside in lower Vi, Sepals lanceolate, blunt tipped and broader than the tube in bud, 11-16 mm long, 4-5 mm wide, lobes often splitting open in the middle before detaching at the tips, suberect to spreading at anthesis. Tube and sepals pale pink to lavender or light red. Petals slightly darker than the tube or sepals, crimson to light purple, lanceolate to narrowly elliptic, 10-16 mm long, 3-4- mm wide, obtuse to subacuminate at the apex, spreading at anthesis. Nectary unlobed, ca. 1.5 mm high. Filaments dull white to light red, 7-9 mm and 4-6 mm long; anthers oblong, 2-2.5 mm long, ca. 1.5 mm wide, white. Style pink, villous from the base to the rim of the tube; stigma subglobose, 4-cleft at the apex, 2.5-3.5 mm long, 2-3 mm wide, pink. Berry subglobose to ellipsoid, 10-16(-20) mm long, 8— 1 1 (— 1 5) mm thick, lustrous red purple; seeds 1.8-2.1 mm long, ca. 1 mm wide. Gametic chromosome number n = 1 1 . Northern Peru shrub forest along streamsides or in semidisturbed sites such as roadbanks or thickets, shrubby 500-)2, 700-3 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol.69 Leimebamba Berry & Escobar 3603 (MO, USM); 51 km E of Balsas on road to Leimebamba, Berry & Escobar 3606 (MO); 24 km above Leimebamba on road to Balsas, Berry & Escobar 3612 (MO); Prov. Chacha- poyas, Mathews 2112 (K), s.n. (BM, K, OXF); above Colcamar, Pennell 15620 (PH). cajamarca: amarca Maranon above Balsas, 5 km below summit of road to Celendin, Hutchison & Wright 5356 (F, NY, RSA, UC, USM); E of Cordillera de Cumulloa to Celendin, Pennell 15158 (PH, USM). rusty inflorescences flowers characterize this*species. It is somewhat intermediate between the Fuch- sia boliviana and the F. dependens species groups and apparently has no close former ternate occurring above the dry Maranon valley and on the moister, eastern slopes. Considerable variation in the length and redness of pubescence exists within these populations, and the plants on the western side tend to have somewhat longer petioles than eastern Maranon in Dept. Amazonas and those on the opposite slopes in Dept. Cajamarca, and the differences between them are no greater than the variation found locally on the Amazonas side. The two species described from areas farther north in Ca- jamarca, F. fischeri and F. storkii, show no significant differentiation from the populations described above. Fuchsia storkii is supposedly distinguished by its short (30 mm long) flowers, but the type at F examined by Munz proves to have only young flowers; fully extended flowers on the isotype at K have floral tubes up to 48 mm long, well within the normal range for F. mathewsii. Fuchsia fontinalis often has rusty-pilose pubescence and paniculate inflores- cences like F. mathewsii, but the rachis is more extended and the flowers shorter. Whereas F. mathewsii occurs on both sides of the north-south running Cerros de Calla-Calla (between the Maranon and Utcubamba river valleys), F. fontinalis grows only on the eastern slopes. There, the two species are sympatric between 3,200 and 3,300 m, at the lower altitudinal limit of F. fontinalis and the upper limit of F. mathewsii (Fig. 10). They were found growing together in low woods along a pasture border, and a probable hybrid between them was collected growing out of a stone wall nearby (Berry & Escobar 3613, MO, USM: 3,260 m, 15 July 1979). This individual has intermediate floral tube lengths (22-32 mm), short, but brac- teate inflorescences, and reduced pollen stainability of 36.6% of 500 grains. Another possible hybrid or backcross to F. mathewsii is Berry & Escobar 3611 (MO), found 1 km above 3613; it is much closer to F. mathewsii, with floral tubes ca. 36 mm long, but the style is glabrous, and the inflorescence is bracteate and fi 500 Fuchsia W side of Volcan Pichincha, 1838, Willia •graph, MO). 1982] BERRY— FUCHSIA SECT. FUCHSIA 175 Fuchsia corymbiflora sensu J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):551 . 1941 , pro parte; sensu Munz, Proc. Calif. Acad. Sci. IV. 25:49. 1943, pro parte; Opera Bot., Ser. B, 3:13. 1974, pro parte. Erect to usually scandent-climbing shrubs 2-10 m high, with arching to hang- ing branches 0.5-3 m long. Branchlets canescent, dull red, 2-8 mm thick, with tan bark exfoliating in longitudinal strips. Leaves mostly quaternate, rarely in whorls of 3 or 5, membranous, narrowly elliptic to ovate, acute at both ends, 5— 14(— 15) cm long, 2-5(-6) cm wide, dull green and strigillose above, incanous dry 14(— 16) on either side of the midvein, ± impressed above and prominent below, midrib reddish; margin usually undulate and conspicuously glandular-denticulate. Petioles puberulent, reddish, X l--V?> the length of the blade, 14-35 mm long. Stipules lanceolate, 1.5-2 mm long, 0.4-0.5 mm wide, deciduous. Flowers numerous in dense, drooping, terminal panicles or series of terminal and subterminal, corymbiform racemes; rachis 2-6 cm long when young, 8-25 cm long in fruit; bracts elliptic, 5-20 mm long, 3-10 mm wide, short-petiolate, sometimes reflexed. Pedicels slender, pen- dant, incanous, 5-10 mm long. Ovary ellipsoid, 5-7 mm long, 2-3 mm thick. Floral tube narrowly funnelform or subcylindric, 40-50 mm long, 3-4 mm wide and nodose at the base, narrowed to 2.5-3 mm wide in basal !4, then gradually widened above until 5-8 mm wide at the rim, puberulent outside, densely pilose inside in the narrowed part of the tube. Sepals lance-oblong, 12-13 mm long, 4-5 mm wide, obtuse to shortly acute-tipped, spreading at anthesis. Tube orange red to scarlet, sepals same but with dull green tips. Petals orange red, narrowly lanceolate, 12-14 mm long, 3^1 mm wide, acute at the apex, spreading to ± recurved at anthesis. Nectary irregularly 4-lobed, 1-2 mm high. Filaments red, 9-11 mm and 5-7 mm long; anthers oblong, 2.5-3 mm long, 1.5-2 mm wide, cream. Style totally glabrous, light red; stigma capitate to subobconic, slightly 4- parted at the apex, 2-2.5 mm long, 1.5-2 mm wide, dull pink, exserted 3-10 mm beyond the anthers. Berry ellipsoid, puberulent, 12-16 mm long, 7-12 mm thick; seeds tan brown, 1-1.2 mm long, ca. 0.6 mm wide. Gametic chromosome number n = 11. Southern Northern Pichincha and on the east slopes of the Cordillera Central in Imbabura and Carchi; also in the drier Central Valley of Ecuador from Pichincha to Carchi and in the Nudo de Pasto area in Narino, Colombia, occurring commonly in hedgerows; 2,400-3,300 m (Fig. 66). Representative specimens examined: Colombia, narino: between pasto and Rio Bobo, Barclay 4645 (COL); 8 km S of Pasto towards Ipiales, Berry 3145 (MO); Oscurana, S slopes of Volcan Gualcala, 14 km from Piedrancha, Fosberg 21186 (RSA, US); Laguna La Cocha, Paramo el Tabano, Garcia-Barriga 7822 (COL, US); Volcan Galeras, near Pasto, Lehmann 5497 (F, GH, K, S, US); road from Cuatro Esquinas to Quitasol, Mora 290 (COL); region of Pedregal, S of Yacuanquer, Schultes & Villarreal 7898 (COL, RSA); base of Volcan Galeras, above Obonuco, Schultes & Vil- larreal 8042 (COL, GH, RSA). Ecuador, carchi: between Paja Blanca and El Cucho, Acosta Solis 10476 (F); Guaca, San Gabriel, Balls 7344 (BM, K, MO, NA, UC, US); El Angel, Berry & Berry 2560 (MO); 38 km W of Tufino on road to Maldonado, Berry 3148 (MO); ca. 4 km W of El Carmelo, Berry 3164 (MO, QCA); town of J. Andrade, on Panamerican highway, Berry 3166 (MO, QCA). 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 imbabura: Rosaspampa, E slopes of Volcan Mojanda, Acosta Solis 8089 (F); 4-5 km W of Otavalo to Laguna de Mojanda, Berry & Berry 2552 (MO), Berry 3172 (MO, QCA); Mojanda, ca. 10 km SSW of Otavalo, Sparre 13480 (S). pichincha: San Jose de Minas, Benoist 3961 (P); 19 km W of Cotocollao on road to Nanegal, Berry & Escobar 3176 (MO, QCA); 10 km E of Cayambe, Hurling 4343 (NY, S); between Calacati and Nono, Hartweg 986 (BREM, CGE, G, K, LE, NY, OXF, P, W); Hacienda Yunguilla, NW of Calacati, Haught 3170 (A, BH, U); Pululahua Crater, ca. 22 km N of Quito, Humbles 6281 (AAU, NY); between Cotocollao and Nono, Mexia 7658 (BM, GH, K, MO, NY, POM, S, U, UC, US); Volcan Pichincha, Spruce 5471 (BM, CGE, G, K, NY, OXF, TCD, W). This species is part of a group with mostly quaternate leaves and paniculate inflorescences. Both Munz (1943) and Macbride (1941) treated this species under Fuchsia corymbiflora, but that is a distinct species from central Peru with entirely opposite leaves and usually racemose inflorescences. Fuchsia dependens is closely related to F. hartwegii, with shorter flowers, and to F. hirtella, which has more entire leaves with short petioles. Both these species occur north of the range of F. dependens in Colombia. It is further distinguished by its densely cinereous pubescence and long-petiolate leaves with conspicuously denticulate and some- times undulate margins. Fuchsia dependens has wide ecological tolerances for sect. Fuchsia. On the eastern slopes of the Cordillera Central in Ecuador and on the western slopes of the Cordillera Occidental, it is found in cloud forest and occurs mostly as a climber or scrambling shrub. It also occurs in the drier Central Valley of Ecuador and in the Nudo de Pasto in southern Colombia, where it grows more compactly and often as upright shrubs in hedgerows. This is an excellent example of this upland area being used as a migration corridor for a species to spread from one structural unit of the Andes to another. Other species that occur sympatrically with F. dependens are F. ampliata, F. corollata, and F. sessilifolia, but no hybrids were detected with them. 54. Fuchsia hirtella Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6:107. 1823. Munz, Proc. Calif. Acad. Sci. IV. 25:47, pi. 7, fig. 36. 1943. type: Colombia, Dept. Cundinamarca, in mountains near Fusagasuga, ca. 1,900 m, July-Sept. Humboldt MO) Climbing-scandent to suberect shrubs 2-5 m tall. Branchlets subterete, 2-4 mm thick, green to reddish and densely hirtellous; older branches 10-25 mm thick, with freely exfoliating bark. Leaves mostly quaternate, occasionally in whorls of 3 or 5, membranous, elliptic to narrowly elliptic or obovate, acute to truncate at the base, acute at the apex, 6-13 cm long, 1.8-5 cm wide, subnitid dark green and loosely strigose above, pale green and densely strigose below; secondary veins 10-15 on either side of the midvein, prominent below; margin entire to subdenticulate. Petioles strigose, reddish, 3— 8(— 15) mm long. Stipules lance-linear, hirtellous, 1.5-2 mm long, ca. 0.5 mm wide, deciduous. Inflores- cence an arching to pendant panicle at the tips of branches; flowers numerous, subtended by reduced, sometimes reflexed leaves; rachis hirtellous, 4-20 cm long. Pedicels slender, pubescent, 4-12 mm long. Ovary cylindric, 5-7 mm long, ca. 2 mm thick. Floral tube subcylindric, 34^0 mm long, 3^4 mm wide and bulbous at the base, narrowed to 2-3 mm wide in the basal Vs, then gradually widened above until 5-9 mm wide at the rim, strigillose to pilose outside, densely villous 1982] BERRY— FUCHSIA SECT. FUCHSIA 177 ** * 500 Km i » f • / \ \ \ ■ * • *. \ I t # Fuchsia hartwegu •X* Fuchsia crassistipula i * ■ ( i O Fuchsia hirtella * Fuchsia canescens D Fuchsia dependens O Fuchsia cinerea Figure 66. Distribution of the Fuchsia dependens species group. 3-4 inside in basal Vi. Sepals lanceolate, acuminate, 12-16 mm long, long-tapered to apiculate in bud, spreading to divergent at anthesis. Tube and sepals subnitid lavender to reddish pink. Petals somewhat darker, crimson, nar- sharply 3-4 spreading at anthesis. Nectary unlobed, light green, ca. 1.5 mm high and ca. 1 mm thick. Filaments pink, 8-14 mm and 5-10 mm long; anthers oblong, 2.5-3 mm long, 1.5-2 mm wide, dull white. Style slender, glabrous, pink; stigma globose to subobconic, 4-parted apically, 2-2.5 mm long, ca. 2 mm wide, pale pink to dull red, exserted 3-9 mm beyond the anthers. Berry cylindric-ellipsoid, 14- mm long, 8-9 mm thick; seeds 1.4-1.7 mm long, ca. 0.7 mm wide. Game 18 chromosome number n 11. Distribution: Colombia. In the Cordillera Oriental in Cundinamarca, mostly on the outer slopes of the Sabana de Bogota; in the Cordillera Central from Caldas 500 300 Representative specimens examined: Colombia, cundinamarca: San Miguel, Km 35-36 S of Sibate to Fusagasuga, Barclay et al. 3403 (COL, US), Berry 3543 (COL, MO); Paramo de Sumapaz, near 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Lagunitas, ca. 5 km SSW of San Juan, Cleef845l (COL, MO, U, US); Boca de Monte, Cuatrecasas et al. 25856 (COL); San Miguel, Cuatrecasas & Jaramillo 12011 (BH, COL, F, US); Fomeque, between Lago de Chingaza and Cerro Verde, Garcia-Barriga 17676 (COL); Zipaquira-Pacho highway, Haught 5977 (COL, NY, POM, US); La Vega, road to Facatativa, Rio Sabaneta, Schneider 1206 (COL, S); Bojaca, Schneider 1263 (S); Zipacon, below Boca de Monte, Uribe-Urihe & Villarreal 2560 (COL, US); 14 km S of Mosquera to La Mesa, Uribe-Uribe 5929 (COL, US), quindio: 12 km from Salento towards Cocora, along Rio Quindio, Escobar 1016 (MO); Amarguras, above Salento towards Romerales, Hawkes 418 (COL, K, US); Laguneta to Magana, old Quindio trail, Killip & Hazen 9418 (GH, NY, PH, US), risaralda: Guayabal, watershed of the Rio Otun, below El Bosque, Cuatrecasas 23306 (F, RSA, VALLE). tolima: Mediacion, Andre 2152 (K, NY); 3 km E of La Linea, between Giradot and Armenia, Berry 3144 (MO); between El Libano and Murillo, Garcia- Barriga 12274 (COL, US); La Lora to Cucarronera, new Quindio trail, Hazen 9679 (GH); Volcancitos, Holton 893 (K, NY, PH); Tochecito, Triana 3809 (G, US). This species is closely allied to Fuchsia hartwegii and F. dependens, which share a similar paniculate inflorescence and quaternate, pubescent leaves. It can be distinguished from these, however, by its subsessile or shortly petiolate leaves, hirtellous pubescence, and pinkish flowers. Its flowers are longer than those of F. hartwegii, which it apparently overlaps in range in the Cordillera Central. This is a relatively high elevation cloud forest species and is common along the moist, outer slopes of the Sabana de Bogota. As with Fuchsia venusta and F. petiolaris, it also grows adjacent to Cundinamarca in Tolima, in the Cordillera Central, where it probably spread secondarily. At its lower altitudinal limit, F. hirtella is sympatric with F. venusta and F. verrucosa. Probable hybrids with F. venusta were found between 2,500 and 2,600 m on the southeastern edge of the Sabana de Bogota in Cundinamarca, where both species are locally common and grow in identical habitats. Table 17 shows the interme- diate morphological characters of the two putative hybrids collected. Pollen stain- ability of these plants was reduced by 18 to 28% relative to their presumed parents (see Table 18). Normal pairing of 11 bivalents was found in meiotic preparations of Berry 3547 (COL, MO), but occasional chromosomal bridges and fragments were observed at anaphase I. 55. Fuchsia hartwegii Bentham, PI. Hartw. 179. 1845. Munz, Proc. Calif. Acad. Sci. IV. 25:63, pi. 9, fig. 53. 1943. type: Colombia, Dept. Cauca, in woods near Pitayo and Huambia, common in hedgerows at Huambfa, 1841-1843, Theodor Hartweg 994 (K Bentham Herb., holotype; photograph, MO; BM, BR, BREM, CGE, G, K Hooker Herb., LE, OXF, P, W, isotypes). Low shrubs or small trees 0.5-4 m tall or scandent-climbing in thickets or trees to 8 m above the ground. Young growth usually canescent; branchlets 2-A mm thick, subterete, reddish, hirtellous to strigose; older branches erect or arch- ing-flexuous on scandent bushes, 0.5-3 m long, 8-50 mm thick, with tan gray, flaking bark. Leaves mostly quaternate, occasionally up to 7 per whorl, firmly membranous, narrowly to rather broadly elliptic or oblanceolate, acute to atten- uate at the base, acute to acuminate at the apex, 3-15.5 cm long, 1.5-5.5 cm wide, subnitid dark green and strigose to subglabrous above, pale green and hirtellous to strigose below, especially along the veins; secondary veins 8-14 on either side of the midvein, impressed above, prominent and reddish below; margin glandular-denticulate. Petioles hirtellous to strigose, 5-32 mm long. Stipules lan- ceolate, thick at the base, filiform at the tip, 1-2 mm long, ca. 0.6 mm wide, 1982] BERRY— FUCHSIA SECT. FUCHSIA 179 Table 17. Comparison of morphological characters of Fuchsia hirtella, F. venusta, and pre- sumed hybrids. F. hirtella (Berry 3543) Stem color Stem pubescence Number of leaves/whorl Leaf surface Leaf pubescence Rachis length Pedicel length Floral tube color Sepal length Sepal angle at an- thesis (approx.) Petal length Petal surface Petal color Green to pink Hirtellous 4 Matte Strigose both sides Inflorescence type Paniculate 13 cm 9-12 mm Floral tube length 35-36 mm Pink 12-15 mm 30° 12-14 mm Smooth Pink Hybrid {Berry 3546) Hybrid (Berry 3547) F. venusta (Berry 3544-C, 3547) Wine red Canescent 3-4 Subnitid Strigose below Subpaniculate 10-11 cm 10-25 mm 35-38 mm Pink purple 15-18 mm 30° 11-14 mm Smooth Red purple Wine red Canescent 3-4 Wine red Puberulent 3 Subnitid Nitid Strigose on veins Glabrous both below Racemose 5-8 cm 8-22 mm 45 mm Pink purple 19 mm 90° sides Subracemose 3-4 cm 16-30 mm 30-37 mm Dull orange 16-20 mm 90° 22-23 mm Undulate Red purple 16-20 mm Undulate Orange red hermaphroditic commo 15 mm wide. Pedicels funnelform bracts broadly ovate, sometimes sessile, 6-20 mm long, 3- slender, arching to pendant, pubescent, 4-12 mm long. O to pilose, 3.5-5 mm long, 1.5-2 mm thick. Floral tube 13-20(-24) mm long, 1.5-3 mm wide and bulbous at the base, constricted to 1-2 mm wide above the nectary and gradually widened above until 2-5 mm wide at the rim, strigose to pilose outside, densely villous inside in the lower 14-46. Sepals lanceolate, 7 5-4 mm wide, acute at the apex, forming in bud, spreading at anthesis. Tube and sepals subnitid orange red to bright crimson. Petals red, very narrowly elliptic, 5-9 mm long, 1.5-3 mm wide, nar- rowly acute at the apex. Nectary green, unlobed, 1.5-2 mm high, ca. 0.8 mm thick. Filaments red, 5-8 mm and 3-5 mm long; fertile anthers oblong, 1.5-2 mm long, 0.7-1 mm wide, white, abortive ones ca. V2 as large and nondehiscing. Style glabrous, red; stigma subglobose, barely 4-parted at the apex, 1-1.5 mm long, 1- 1.5 mm wide, pale red, exserted 3-6 mm beyond the anthers. Berry globose when ripe, 6-9 mm long, 5-8 mm thick, nitid purple red; seeds tan to reddish brown, 1 . 1-1.3 mm long, ca. 0.7 mm wide. Gametic chromosome number n = 11. Most common in the Cordillera Central from southern Putumayo, as a scandent or hedgerow shrub 000 southern 100-2.300 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 18. Pollen stainability of Fuchsia hirtella, F. venusta, and presumed hybrids. Collection Stainable Pollen Grains (% of 500) Berry 3543 (F. hirtella) Berry 3546 (hybrid) Berry 3547 (hybrid) Berry 3544- C (F. venusta) 95.6% 78.4% 68.8% 96.2% Representative specimens examined: Colombia, antioquia: Paramo de Sonson, Guarin 3479 (COL, US), caldas: without locality, Dawe 741 (K, NY, US); 7 km N of Manizales, Escobar 1002 (MO); Manizales, Sandeman 5666 (COL, K, OXF); San Felix, near Salamina, Tomas 2406 (BH). cauca: Purace, Alston 8137 (BM, RSA, UC); between Silvia and Totoro, Berry 3573 (COL, MO); 21 km above Piendamo on road to Silvia, Berry 3572 (COL, MO); Quebrada Santo Domingo, headwaters of the Rio Palo, Cuatrecasas 19159 (A, BH, COL, VALLE); La Tolda, above Tacueyo, watershed of Rio Palo, Cuatrecasas 19421 (A, BH, VALLE); Purace, 17 km SE of Popayan, Fosberg 20303 (COL, RSA, S, US); Rio Santo Domingo, 20 km SE of Corinto, Fosberg 21306 (GH, RSA, S, US); from Purace to Alto de San Rafael, Garcia-Barriga & Hawkes 12856 (COL, US); Paispamba, Haught 5302 (COL, US); Cococnuco, Kiltip 6829 (GH, NY, PH, US); Central Andes of Popayan, Lehmann 5613 (F, K); Paramo de Guanacas, Lehmann 5614A (F, GH, K, PH, S, US); Alto de Pesares, Lehmann 5614B (F, K); Paramo de Purace, at Chiquin, Perez-Arbelaez & Cuatrecasas 5973 (COL, F, US); between Popayan and Purace, bridge at Uarlo, Perez-Arbelaez & Cuatrecasas 5880 (COL, F); head- waters of Rio Lopez, Rio Palo basin, Tierra Adentro, Pittier 1068 (US); Prov. of Popayan, Triana s.n. (BM); Silvia, Guambia, Yepes-Agredo 3088 (COL), cundinamarca: Fomeque, Dawe 352 (K, US); Rio Blanco valley, 6 km W of Gutierrez, Grant 9695 (RSA, US); between Guayabetal and Quetame, Vogel 175 (U). huila: ca. 75 km W of La Plata on road to Purace, Berry 3591 (COL, MO); headwaters of Rio Fortalecillas, just below Paso de Las Cruces, 39 km E of Neiva, Fosberg 19785 (NY, RSA, S, UC, US); Balsillas, on Rio Balsillas, Rusby & Pennell 789 (NY, US), 810 (NY), meta: Rio Grande, S of Cordillera de Las Cruces, Fosberg 20872 (US), putumayo: above Sibundoy on road to La Cocha, Berry 3255 (MO, PSO); Balsayaco, valley of Sibundoy, Bristol 642 (COL, DS, GH); near San Francisco, Cuatrecasas 11541 (COL, F, US); upper part of Rio Minchay, Rio Mocoa drainage, Fosberg 20401 (RSA); Portachuelo, valley of Sibundoy, Schultes & Villarreal 7725 (COL, RSA, US); Sibundoy, Schultes & Villarreal 7688 (COL, F, GH, NY, RSA, US), risaralda: above Santa Rosa de Cabal, Chaquiro-Paramo de Santa Rosa trail, S of Chaquiro Falls, St. John 20837 (GH, MICH, US), tolima: 47 km W of Fresno on road to Manizales, Berry 3557 (COL, MO); La Esper- anza, Fresno, Hanbury-Tracy 624 (K). valle: above El Guayabo on Palmira-Taco road, Berry & Escobar 3568 (COL, MO). Palmira, Duque-Jaramillo 4699 (COL); above La Magdalena, Buga, Es- pinal 2057 (MO). This species is closely allied to Fuchsia dependens and F. hirtella, based on their predominantly quaternate leaves and paniculate inflorescences. Fuchsia hartwegii is the only species with mostly paniculate inflorescences that has short flowers, very narrow petals, and globose fruits. The species, as treated here, comprises three main geographical groups that may upon further examination warrant recognition as distinct taxa. The first group, including the type collection, is centered around the moist, western, mid-elevation slopes of the Cordillera Central in Cauca, extending as far north as southern Valle and reaching south to Putumayo on the eastern slopes of the same Cordillera. These populations have orange red flowers and are common in roadside thickets, hedgerows, and along streams, where they sometimes occur as lianas. A second group is found farther north in the Nevado de Ruiz Massif of the Cordillera Central, in Caldas, Tolima, and Antioquia. These plants differ from the first group in their somewhat smaller, cherry red flowers, short-petiolate leaves, and higher altitudinal range. Of the seven collections known from this area, four are male-sterile, Berry 3557 \ Guarin 3479, St. John 20837, and Tomas 2406. Of 1982) BERRY— FUCHSIA SECT. FUCHSIA 181 the much more numerous collections from the first group to the south, only five male-sterile plants were found, H aught 5302, Lehmann 5613, Pittier 1068, Schultes & Villarreal 7688, and Triana s.n. A third group consists of collections from the Cordillera Oriental. Fosberg 19885 from Huila is very similar to more or less stunted plants that grow in exposed habitats in Cauca, and it is male-sterile. The Cundinamarca collections are all hermaphroditic with mostly fasciculate flowers and differ quite notably narrow of F. hirtella. Population studies of the different groups discussed above will be required to determine the frequency of male sterility and its relative significance in the breed- ing system of this species. The finding of male-sterile plants from widely separated localities does suggest that gynodioecy is functioning in at least some populations of F. hartwegii. If confirmed, this would be the first report of male sterility in any of the main Andean-Brazilian sections of the genus, which comprise some 75 of the estimated 100 species in Fuchsia. Male sterility is presently known only in sects. Encliandra, Kierschlegeria, Schufia, and Skinnera (Arroyo & Raven, 1975). 56. Fuchsia crassistipula P. Berry, sp. nov. type: Colombia, Dept. Tolima, 40 km W of Fresno on Mariquita-Manizales highway, 2,700 m, 4 June 1979, Paul E. Berry 3553 (COL 66345, holotype; MO 3-6 cillata plerumque quaterna, firme membranacea, elliptica vel late oblanceolata basi obtusa vel acuta 6-16 10-15 serrulata stipulis persistentibus, conspicuis, saepe connatis, subtriangularibus, patenti-recurvatis, 2.5-4 mm longis, ca. 3 mm latis, costa media 2-3 mm crassa (in vivo), in sicco aliquantum minore. Flores roseo- scarlatini, numerosi in racemis (sub-)terminalibus vel paniculis verticillatim racemosis dispositi; rhachidi costata, nutanti, 5-30 cm longa; bracteis 8-20 mm longis; pedicellis 3-8 mm longis; ovario cylindrico, 5-8 mm longo. Tubi florales anguste infundibuliformes, 32-44M8) mm longi, basi 3-5 mm lati et subnodosi inde 2-3 mm lati constricti superne gradatim dilatati summo 5-8 mm lati, extus strigulosi intus infra medium pilosi. Sepala lanceolata, acuta, 10-14 mm longa, 4-6 mm lata. Petala rubra vel atrorubra, anguste oblongo-elliptica, acuta, mucronata, 11-16(-18) mm longa, 3-5(-6) mm lata. Fi- lamenta rubra, antisepala 7-11 mm longa, antipetala 4-7 mm longa; antheris oblongis, 2.5-3 mm longis, ca. 1.5 mm latis, albis. Stylus roseus glaberque, stigmate globoso, apice leviter 4-fisso, 3^4 mm longo, 2-3 mm lato, valde roseo-rubro. Bacca cylindrica, in maturitate subglobosa, 13-15 mm longa, 9-13 mm crassa, viridis vel purpurascens; seminibus inaequaliter oblongo-triangularibus, ca. 1.5 mm longi, 0.8-1.1 mm latis. Numerus gameticus chromosomatum n = 11. Suberect to scandent shrubs 1-3 m tall. Young branches conspicuously 3-6- ridged, one ridge for each leaf in the whorl above, 3-9 mm thick, green, strigillose to subcanescent; older branches terete with dull gray brown, exfoliating bark. Leaves 3-6-verticillate, mostly in whorls of 4 or 5, firmly membranous, elliptic to broadly oblanceolate, acute to obtuse at the base, acute to acuminate at the 6- purpl 10-15 glandular-serrulate and usually purple tinged. Petioles stout, 2-3.5 mm thick, 12- 40 mm long, sparsely strigose, green to pink red. Stipules conspicuous, subtrian- 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 gular, divergent, margins and tips recurved on lower nodes, separate to fully connate, thick-callose, 2.5-4 mm long and ca. 3 mm wide when fresh, each stipule with a central longitudinal nerve 2-3 mm thick, considerably flattened and shrunk- en when dry, persistent. Flowers numerous in terminal or subterminal racemes or verticillately branched panicles; rachis ridged, spreading to drooping, 5-30 cm long; bracts 8-20 mm long. Pedicels short, pendant, 3-8 mm long. Ovary cylin- drical, 5-8 mm long, 2-4 mm thick, green, strigose. Floral tube narrowly fun- nelform, 32-46(-48) mm long, 3-5 mm wide and bulbous at the base, narrowed to 2-3 mm wide above the nectary, then gradually widened above until 5-8 mm wide at the rim, finely strigose outside, pilose inside in lower Vi. Sepals lanceolate, acute, 10-14 mm long, 4-6 mm wide, with a short tip 1-2 mm long in bud, spreading at anthesis. Tube subnitid scarlet pink; sepals similar but becoming dull purple towards the tip. Petals deeper red, narrowly elliptic-oblong, acute and ± mucronate at the apex, 1 1— 16(— 18) mm long, 3-5(-6) mm wide, spreading at anthesis. Nectary green, unlobed, ca. 2 mm high, 0.8-1 mm thick. Filaments 7- 1 1 mm and 4-7 mm long; anthers oblong, 2.5-3 mm long, ca. 1.5 mm wide, white. Style glabrous, pink; stigma globose, 4-parted at the apex, 5-4 mm long, 2-3 mm wide, bright red pink, exserted 3-8 mm beyond the anthers. Berry cylindric to sub- globose when fully ripe, 13-15 mm long, 9-13 mm thick, nitid green to flushed purple; seeds ca. 1.5 mm long, 0.8-1.1 mm wide. Gametic chromosome number n = 11. Distribution: Central Colombia. Infrequent shrubs in cloud forest thickets of the Nevado de Ruiz Massif in the Cordillera Central, in Depts. Tolima, Quindio, and Caldas; 2,600-3,000 m (Fig. 66). Specimens examined: Colombia, antioquia: Tamesis, vicinity of Medellin (Mislabel?), Torn 956 (NY), caldas: San Felix, near Salamina, Tomds 2051 (US), quindio: Salento to Lagunetas, Old Quindio trail, Killip & Hazen 9141 (GH, NY, PH, US); Penares, above Salento, Pennetl 9195 (GH, PH), 9324 (GH, NY, PH, US), tolima: 43 km W of Fresno on Mariquita-Manizales highway, Berry 3554 (COL); La Palmilla, Quindio, Mariquita, Triana 3812 (BM, G, K, P, W). This species is closely related to Fuchsia hirtella, which shares the often paniculate inflorescence, short-cylindrical ovaries, glabrous style, and very sim- ilar tube and petal dimensions. Fuchsia crassistipula is unique in its strongly ridged stems and thick, persistent stipules. The flowers are also more purple than those of F. hirtella. All these characters are usually lost upon drying, however, making it difficult to distinguish between the two species in dried specimens. However, F. crassistipula has longer, thicker petioles, more serrulate leaves, and often more than four leaves per whorl. Although F. crassistipula occurs in the same area as F. hirtella, the two are not known to actually grow together. 57. Fuchsia canescens Bentham, PI. Hartw. 178. 1845. type: Colombia, Dept. Hart MO: BREM Herb., isotypes). Shrubs 1.5-4 m tall. Young branches ridged, the number of ridges the same as the number of leaves in the whorl above, canescent to hirtellous, reddish, 3- 8 mm thick; older branches with tan, exfoliating bark. Leaves 3-5-verticillate, 1982] BERRY— FUCHSIA SECT. FUCHSIA 183 mostly quaternate, firmly membranous, (narrowly) elliptic to (ob-)ovate, acute to rounded at both ends, 4.5-10 cm long, 2-5 cm wide, medium to dark green and strigillose above, hirtellous to densely strigillose and paler below, with the mid- vein prominent and reddish; secondary veins 8-15 on either side of the mid vein; margin subentire to denticulate. Petioles stout, 15-25 mm long, hirtellous, dull purple. Stipules 0.8-1 mm thick when fresh, Ungulate to subterete in transection, subulate at the tip, 1.5-2 mm long, ca. 0.8 mm wide, divergent, occasionally connate, hirtellous, subpersistent. Flowers few to many in upper, leafy nodes or in terminal to subterminal, leafy racemes; rachis 2-10 cm long. Pedicels stout, subtuberculate, pendant, hirtellous-canescent, 5-13 mm long, dull green. Ovary cylindrical, subtuberculate, 10-11 mm long, 3-4 mm thick, hirtellous, light green. Floral tube narrowly funnelform, 34-50 mm long, firmly thick-spongy, 1-2 mm thick when fresh, 3.5-5.5 mm wide and slightly bulbous at the base, narrowed to 3-4.5 mm wide above the nectary, then gradually widened above until 5-9 mm wide at the rim, hirtellous and finely striated-tuberculate outside, densely villous inside in the lower Vi-Vi. Sepals similarly thick-tuberculate, triangular, 14-18 mm long, 6-9 mm wide, spreading to divergent at anthesis; buds tetragonous in tran- section, 1-2 mm wider at the base than the rim of the tube and with small pro- tuberances at the base of adjacent sepals, the tip obtuse or shortly pointed. Tube dull scarlet or orange red, sepals becoming dull white or greenish toward the tip. Petals darker red, subtrullate, 14-19 mm long, 4-7 mm wide, margin slightly undulate, spreading to slightly recurved at anthesis. Nectary light green, irregu- larly 4-lobed, 1.5-2 mm high, ca. 1 mm thick, often with a few erect, stiff hairs on the top. Filaments red, 9-12 mm and 5-8 mm long; anthers oblong, 3-4.5 mm long, 1.5-2 mm wide. Style orange red, glabrous; stigma capitate, 4-parted at the apex, 2.5-3 mm long, ca. 2 mm wide, red to light pink, exserted 4-9 mm beyond the anthers. Berry ellipsoid, 12-15 mm long, 8-10 mm thick, subtuberculate, green to flushed purple; seeds ca. 1.5 mm long, 0.6-0.8 mm wide. Gametic chro- mosome number n = 1 1 . Distribution: Southern Colombia. Infrequent shrubs in wet subparamo or up- per cloud forest in Cauca, Huila, and Narino; 2,800-3,350 m (Fig. 66). Representative specimens examined: Colombia, cauca: 31 km E of Totoro on road to Inza, Berry 3578 (COL, MO); 34 km E of Totoro to Inza, Berry 3579 (MO); 17 km E of Purace, Berry 3581 (COL, MO); Km 56 of Totoro-Inza road, Escobar 1042 (MO); Paramo de Las Papas, between El Boqueron and La Hoyola, ldrobo et al. 3918 (COL, P); Paletara to Calaguala, Pennell 7083 (GH, NY, PH, US), 7094 (GH, NY, PH, US); Canaan, Mt. Purace, Pennell & Killip 6669 (GH, NY, PH, US); Las Escaleretas, Moras valley, Rio Paez basin, Tierra Adentro, Pittier 1382 (US); Silvia, Guambia, Yepes- Agredo 3078 (COL), huila: ca. 40 km E of Purace to La Plata, Berry 3589 (COL, MO), narino: 31 km E of Pasto on road to Sibundoy, Berry 3254 (MO, PSO); Volcan de Sotara, Lehmann 6194 (COL). Because of an incorrectly numbered type collection of Fuchsia corollata at Geneva, Munz (1943) thought that F. canescens was synonymous with that species. Both Hartweg 992 and Hartweg 993, the type collection numbers of F. canescens and F. corollata, respectively, were collected in the same area, so one of the duplicates may have been easily mislabelled. The holotype of F. canescens allows no doubt that it is distinct from F. corollata. It has dull orange red flowers with thick, firm walls, somewhat tuberculate ovaries and tubes, short pedicels, trowel- shaped petals, and sepals that are tetragonous in bud. It is a very localized and 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 rather rare species, found only in high elevation cloud forest and subparamo elfin forests between Cauca and Narino, and mostly on the east-facing slopes of the Cordillera Central. Though the thick-tubed flowers are characteristic of the F. denticulata species group, the generally quaternate leaves, hirtellous pubescence, and mostly racemose flowers place it closer to the F. dependens species group, even though it has no close relative in that group. When dried, specimens of this species are likely to be confused with F. dependens, but F. canescens has fewer, more axillary flowers, ridged stems, stout pedicels, and hirtellous rather than cinereous pubescence. Berry 3577-B (MO) and Escobar 1038 (MO) were both collected on the upper eastern slopes of the Cordillera Central in Cauca, at ca. 3,250 m, ca. 31 km E of Totoro on the road to Inza. Both appear to be hybrids between F. canescens and F. caucana, species that have similar tube lengths and grow side by side at this locality. Escobar 1038 has thick flowers typical of F. canescens, but the leaves are subsessile, coriaceous, and elliptic, characters that place it closer to F. cau- cana. Its pollen stainability is less than 15% of 500 grains, and that of Berry 3577- B is less than 1%. Throughout their range in the Nudo de Pasto, from Cauca to Narino, F. canescens and F. caucana are almost always found growing together or nearby. One collection of F. canescens was made on the western slopes of the Cordillera Central above Purace in Cauca, where it was sympatric with F. corollata. 58. Fuchsia cinerea P. Berry, sp. nov. type: Ecuador, Prov. Carchi, just E of the town of Tufino, 3,210 m, 22 Oct. 1978, Paul E. Berry 3147 (MO 2737070, holotype; QCA, isotype). Frutex erectus vel scandens 2-5 m altus, ramulis foliisque junioribus dense cinereo-strigulosis. Ramuli angulati, caulibus teretibus. Folia quaterna vel interdum ternata, membranacea, strigulosa, anguste elliptica, basi acuta apice acuta vel obtusa, 2-5(-7) cm longa, 1— 2(— 3) cm lata, nervis secu- ndariis utroque latere 5-8(-9) subtus saepe rubrescentes, margine subdenticulata; petiolis 5— 12(— 20) mm longa; stipulis lanceolatis, 1.5-2 mm longis, ca. 0.5 mm latis, deciduis. Flores axillares, depe- ndentes, plerumque 3-4-verticillati; pedicellis (4-)8-20 mm longis; ovario ellipsoideo vel subpyriformi, 5-8 mm longo, dense cinereo. Tubi florales obscure aurantiaci, infundibuliformes, 42-50 mm longi, basi 3-3.5(-4) mm lati et leviter bulvosi inde 2-2.5(-3) mm lati constricti, superne gradatim dilatati summo 5-6(-8) mm lati, extus striguloso-pilosuli, intus pubescentes. Sepala anguste lanceolata, acuta, 15-19 mm longa, 3^4 mm lata, aurantiaca apice viridescentia. Petala aurantiaca vel coccinea, oblonga vel anguste elliptico-lanceolata, 12-15 mm longa, 4-4.5 mm lata. Filamenta antisepala 10-11 mm longa, antipetala 7-8 mm longa; antheris oblongis, eburneis, 2.5-3 mm longis, ca. 1.5 mm latis. Stylus strigulosus usque ad summum tubi, stigmate globoso apice leviter 4-fisso ca. 2 mm longo, 2-3 mm lato. Bacca ellipsoidea, in maturitate subglobosa, cinereo-velutina, ca. 12 mm longa, ca. 10 mm crassa; seminibus inaequaliter oblongo-triangularibus, 1.7-1.8 mm longis, ca. 1 mm crassis. Numerus ga- meticus chromosomatum n = 1 1 . Erect to scandent shrubs 2-5 m tall. Young growth densely cinereous-strigil- lose; branchlets angled, 1-2.5 mm thick; older branches terete, with tan bark. Leaves quaternate or less often ternate, membranous, narrowly elliptic, acute at the base, acute to obtuse at the apex, 2-5(-7) cm long, 1— 2(— 3) cm wide, matte light green and strigillose above, paler and strigillose below; secondary veins 5- 8(-9) on either side of the midrib, usually reddish brown; margin subdenticulate. Petioles strigillose, 5-12(-20) mm long. Stipules lanceolate, 1.5-2 mm long, ca. 0.5 mm wide, deciduous. Flowers axillary, pendant, usually in whorls of 3 or 4. Pedicels strigillose, (4-)8-20 mm long. Ovary ellipsoid to subpyriform, 5-8 mm 1982] BERRY— FUCHSIA SECT. FUCHSIA 185 narrow 5(-4) above the nectary, then gradually widened above until 5-6(-8) mm wide at the rim, strigillose to pilose outside, pubescent inside. Sepals narrowly lanceolate, 15-19 mm long, 3-4 mm wide, acute at the apex, spreading at anthesis. Tube and sepals dull orange, sepals with dull green tips. Petals orange to crimson, oblong to narrowly elliptic-lanceolate, 12-15 mm long, 4-4.5 mm wide, obtuse to acute at the apex, spreading at anthesis. Nectary 4-lobed, ca. 1.5 mm high. Filaments orange red, 10—1 1 mm and 7-8 mm long; anthers oblong, 2.5-3 mm long, ca. 1.5 mm wide, cream. Style strigose from the base to near the rim of the tube; stigma globose, slightly 4-lobed at the apex, ca. 2 mm long, 2-3 mm wide, orange red, exserted 4-10 mm beyond the anthers. Berry ellipsoid to subglobose at maturity, canescent to velutinous, ca. 12 mm long, ca. 10 mm thick; seeds 1.7-1.8 mm long, ca. 1 mm wide. Gametic chromosome number n = 11. Distribution: Known only from the base of Volcan Chiles along the Colom- bian-Ecuadorian border; 3,100-3,250 m (Fig. 66). Specimens examined: Colombia, narino: near village of Chiles, Camp E-340 (NY, RSA, US), Wiggins 10580 (DS, POM). Ecuador, without locality: Lobb 87 (K). This species is distinguished by its dense, fine, ashy pubescence, angled stems, small, quaternate leaves, and slender, axillary flowers. It is only known from the adjacent villages of Tufino and Chiles, on opposite sides of the Colombian- Ecua- dorian border, where it grows in secondary thickets at the edge of the towns. This area is not heavily forested and is drier than the opposite side of the Cor- dillera Occidental. The quaternate leaves, fine, ashy pubescence, and floral di- mensions are very similar to those of F. dependens , which grows nearby, but has flowers grouped in terminal panicles. Fuchsia vulcanica is possibly a related axillary-flowered species that grows close by on the slopes of Volcan Chiles, but at higher elevations; it has coarser pubescence and much wider petals. 59. Fuchsia triphylla Linnaeus, Sp. PI. 1193. 1753. Plumier, PI. Amer. pi. 133, 1892. Boss- fig. 1. 1757. Hook., Bot. Mag. /. 6795. 1885. Duren, Rev. Hort. gere 15:265, t. 1889. Watson, Garden (London) 41:32, pi. 839. chere, 111. Hort. 43:94, fig. 10. 1896. Sieb. & Voss. ex Vilm., Blumengart. ed. 3, 1: /. 84, fig. 336. 1896. Bailey, Cycl. Amer. Hort. 2:615, fig. 877. 1900. Barker & Dardeau, Flore d'Haiti 270. 1930. Essig, Nat. Hort. Mag. 13:1, fig., 15, photo. 1934. Munz, Proc. Calif. Acad. Sci. IV. 25:33,/?/. 4, fig. 20. 1943. Bailey, Stand. Cycl. Hort. 1303, fig. 1606. 1963. lectotype here des- ignated: Plate 14 in Plumier, Nov. PI. Amer. Gen. 1703. Linnaeus simply states the locality as "in America," but Lamarck (1788) reports that Plumier found the plant in Haiti "dans des lieux incultes, en allant du quartier de la Bande du Sud a celui qu'on nomme de grand Cul-de-Sac." This locality belongs to the present Departement de FOuest of Haiti; "la Bande du Sud" southern Massif Morne and flowing northwards, then westwards to Port-of-Prince. Plumier worked in Haiti and Martinique from 1689 to 1697, where he collected objects of natural history and made numerous illustrations of plants. Fig. 29. 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia racemosa Lamarck, Encycl. 2:565. 1788, nom. illeg. based on F. triphylla L.; Tabl. Encycl. pi. 282, fig. I, a-h. 1792. Descourt., Fl. Med. Antilles 2:161, pi. 109. 1822. non Sesse & Mocino, 1888. Subshrub to low shrub 3-20 dm tall with erect to drooping branches. Young growth canescent to short pilose; branchlets pubescent, reddish; older branches with tan, exfoliating bark. Leaves opposite or ternate, rarely quatemate, firmly membranous, narrowly lanceolate to elliptic or oblanceolate, acute to narrowly cuneate at the base, acute to acuminate at the apex, 2.5— 10(— 13) cm long, 1-4 (-5.5) cm wide, dull medium to dark green and strigillose above, pale green to commonly flushed metallic purple and strigose below; secondary veins (6— )7— 1 3 on either side of the midvein, mostly impressed above, elevated and often reddish below; margin subentire to finely denticulate. Petioles pubescent, 4— 15(— 25) mm long. Stipules lance-linear, 1.5-3 mm long, mostly deciduous. Flowers numerous in suberect to mostly nodding, terminal racemes, occasionally subracemose; rach- is 4— 12(— 15) cm long; leaves subtending the flowers mostly reduced, narrowly lanceolate, and deciduous, but similar to normal leaves if flowers subracemose. Pedicels lightly pilose, 10-30 mm long, divergent to nodding. Ovary ellipsoid, 5- 7 mm long, 1.5-2.5 mm wide. Floral tube 25-40 mm long, 2-3.5 and bulbous at the base, sometimes with a ring-like inflation of the tube at the base surrounding the nectary, constricted rather strongly to 1-2.5 mm wide in the lower (6— )10— 18 mm of the tube, then abruptly dilated to 8-1 1 mm wide before narrowing slightly towards the rim; pubescent outside, glabrous inside. Sepals lanceolate, 10-13 mm long, 3.5-4.5 mm wide, spreading at anthesis. Tube and sepals orange to coral red. Petals orange to coral red, elliptic-ovate, rounded to broadly acute at the apex, 6-9 mm long, 4-6 mm wide, suberect to spreading at anthesis. Nectary shallowly 8-lobed, ca. 1 mm high. Filaments orange red, 5-7 mm and 3-4 mm long; anthers oblong, ca. 2.5 mm long, ca. 1.5 mm wide, cream. Style orange red, glabrous; stigma capitate, slightly 4-lobed at the apex, ca. 1.5 mm long and wide, pale red. Berry subglobose to ellipsoid, strigose and ± 4-angled before maturity, 15-18 mm long, 11-13 mm thick, glossy red purple when ripe; seeds tan, 1.8-2.1 mm long, ca. 1 mm wide. Gametic chromosome number n = 22. (West Massif de la Hotte and Massif Matheaux, and (rare) in the northern range in the Montaignes Noires and Chaine de Plaisance. In the Dominican Republic, in the southern Sierra de Baoruco, in the central Sierra de Neiba, and in the northern Cordillera Central. Low shrubs on exposed slopes, moist banks, and edges of pine and mixed pine-broadleaf forest; (700-) 1,1 00-2, 000 m (Fig. 67). Specimens examined: Haiti, artibonite: Massif des Cahos, Montaignes Noires, Petite Riviere de 1' Artibonite, Perodin, ridge above Ingrand, Ekman 3445 (S). nord: Citadelle du roi Christophe, on mountain top 5 km SE of Milot, Bartlett 17380a (MICH, US), ouest: Foret-de-Pins, Massif de la Selle, Bailey 185 (BH, US); Massif de la Selle, Morne Franchant, Ekman 1170 (K, S, US); Massif de la Selle, Morne Cabaio, near Jardin Bois Pin, Ekman 162 1 (S); Massif des Matheaux, Grand-Bois, hills towards Toussaint, Ekman 5706 (S); Massif de la Hotte, eastern group, Grand Goave, Morne Calumette, Ekman 7339 (G, S); Massif des Matheaux, 1'Arcahaie, Habitation Counolle, Ekman 9302 (S); Morne des Commissaires, Holdridge 850 (MICH, US); le Grand Fond de Port-au-Prince, Jaeger 99 (LE); vicinity of Mission, Fond Verettes, Leonard 3681 (NY, US), 3701 (NY), 3701a (US), with- 1982] BERRY— FUCHSIA SECT. FUCHSIA 187 - 19 -r- 1 1 1 1 r 74»W 73» 72* 71" 70* 69 - 20'N O Fuchsia pnngsh«imn D Fuchsia triphylla A Hybrids Figure 67. Distribution of Fuchsia triphylla, F. pringsheimii, and naturally occurring hybrids on Hispaniola. out locality: Jaeger s.n. (B, K). Dominican Republic, azua: Loma Nalga de Maco, Ekman 6304 (S). baoruco: without locality, Howard 12 15 1 (A), la vega: vicinity of Constanza, Abbott s.n. (GH, US), Allard 16446 (US), Augusto 1150 (A, JBSD), von Tuerckheim 2956 (BM, F, G, GH, ISC, K, MO, S, U, US, W, Z); between Constanza and Valle Nuevo, Allard 17422 (US); Jarabacoa, Augusto 883 (JBSD); Valley Nuevo, Augusto 1513 (JBSD); El Convento, 10 km S of Constanza to Valle Nuevo, Berry et al. 3701 (MO, JBSD), 3702 (MO, JBSD); above El Convento, 1 1 km S of Constanza, Berry et al. 3703 (MO, JBSD); below La Siberia, new road from Constanza to Valle Nuevo, Berry et al. 3704 (MO, JBSD); La Siberia, Berry et al. 3706 (MO, JBSD); S of Jarabacoa, road to Constanza, Burch & Burch 2546 (CAS, MO); 12 km SE of Constanza, Davidse 2689 (MO); Loma de Barren), Ekman 2057 (G, K); near Jarabacoa, Fuertes 1619 (A, G, U, W); Rio Yaque, Fuertes 1720 (A, G, W); La Cienaga, confluence of Rio de la Izquierda and Rio Los Guanos to form Rio Yaque del Norte, Gastony et al. 149 (GH, US); 8 km S of Constanza, Gastony et al. 734 (US); Valle Nuevo, Jimenez s.n. (DS); Rio Grande, Constanza, Liogier 19453 (JBSD); between Valle Nuevo and Constanza, Skog 1596 (US); NW of La Cienaga or La Culata, Terborgh & Brockman 87 (NA), 88 (NA). peravia: La Nuez, border with Prov. La Vega, Berry et al. 3712 (MO, JBSD); Sierra de Ocoa, Bejucal, Ekman H. 12039 (A, F, GH, S, US); La Cueva, La Horma Arriba, Liogier 20999 (JBSD); Plan de Jiguey, Pena Franjul s.n. (JBSD). san juan: Arroyo del Oro, Canela s.n. (JBSD), Howard & Howard 8992 (BM, GH, MICH, S, US). Santiago: Pico de Igua, Jimenez 1278 (G, US). This species is easily recognized by its low growth habit, orange flowers, and peculiarly shaped floral tubes. These are nodose at the base, usually strongly constricted in the lower half, then abruptly dilated above the middle, and slightly narrowed again at the rim. Fuchsia triphylla can flower when barely woody and just 20-30 cm tall. Its inflorescences are unusual in being nearly erect or nodding in the upper half, although the individual flowers are divergent or drooping as in other species of sect. Fuchsia. Linnaeus based the genus on Plumier' s illustration of tfc Fuchsia triphylla, flore coccineo" (Plumier, 1703). It was a fairly crude drawing showing only four sta- mens, but the unusual floral tube shape clearly identifies the species, and a more complete habit illustration published later (Plumier, 1757) leaves no doubt that Plumier had illustrated the common Hispaniola fuchsia. Even though Linnaeus stated in the second edition of Genera Plantarum (Linnaeus, 1742) that the normal number of stamens in Fuchsia is eight, he followed Plumier' s plate and placed 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 the species in the Tetrandria Monogynia in the first edition of Species Plantarum (Linnaeus, 1753).. Philip Miller (1739) reported having Fuchsia triphylla in cultivation at his gardens in Chelsea, England. The seeds were sent to him by William Houstoun, presumably before Houstoun's death in 1733, and said to be from "Carthagena, New Spain." Since Cartagena (it is unclear whether he was referring to Cartagena, Colombia or to a Central American locality) is probably a coastal city, it is very unlikely that it was the actual source of Houstoun' s seed. He was known to have travelled in Mexico, Cuba, and Jamaica, but not in Hispaniola. A search of Hous- toun's and Miller's collections in the Sloane Herbarium at the British Museum failed to find any specimen of Fuchsia (J. Lewis, pers. comm.), so we cannot confirm the identification or date of introduction of what otherwise would have been the first introduction of a living fuchsia into Europe. Lamarck (1788) tried to change the name of Fuchsia triphylla to F. racemosa, presumably because he found the triphyllous condition very common among the other, recently discovered species of the genus. Lamarck's name was taken up by Descourtilz (1822), who provided an accurate color illustration of the species in his Flore Medicale des Antilles. Descourtilz travelled extensively in Haiti from 1799 to 1803 and worked partly in the mines of the Cibao mountains (Massif de la Selle), where he probably found F. triphylla. Surprisingly, he mentions having seen this fuchsia several times in "St.-Jago de Cuba," that is, Santiago, a city on the southeastern coast of Cuba. No other reference of herbarium specimen mentions any native fuchsia on Cuba, however, so he may just have seen culti- vated plants there. Fuchsia triphylla is listed in his work under "plantes stoma- chiques astrignentes," used as an antipyretic, decongestant, and as a medication in uterine ailments. Virtually unknown in Europe during the heyday of fuchsia cultivation and hybridization in the early to mid- 1800s, Fuchsia triphylla was finally introduced into England in 1882, from seeds collected by Thomas Hogg in Hispaniola in 1873 (Hemsley, 1882). It was then cultivated rather extensively and used in interspe- cific crosses (Wright, 1978b). It is a rather easy species to grow because it is adapted to considerably warmer temperatures than most fuchsias and flowers when only 2-3 dm tall. In their native range, plants of Fuchsia triphylla exhibit considerable mor- phological variability. Though they generally have a low growth habit, well-de- veloped and woody shrubs with branches several m long have been collected, such as Berry et al. 3706 (MO), from Prov. La Vega, Dominican Republic. Floral tubes generally maintain their unusual shape described previously, but the degree of constriction and the amount of swelling around the nectary varies substantially within local populations. Leaves are particularly variable in their size, shape, and coloration, and some of this variation is associated with geographically isolated populations on the different parallel mountain ranges that traverse Hispaniola. In Baoruco Massif nous and lanceolate, and they occupy the lower extremes of the species limits in size and number of secondary veins. In contrast, plants from the Cordillera Cen- tral in the Dominican Republic vary widely, but are generally much larger, thin- 1982] BERRY— FUCHSIA SECT. FUCHSIA 189 Table 19. Comparison of principal morphological characters of Fuchsia triphylla and F. prings- heimii. Leaf length Petiole length Leaf texture Leaf color Number of secondary veins Flower position Floral tube length Floral tube shape Flower color Petal length Petal width Nectary type F. triphylla 2.5-10(-13) cm 4-1 5<-25) mm Membranous Dull green above, pale to pur- plish below (6-)7-13 Racemose 25^0 mm Narrowed basally, dilated in the middle, narrowed at the rim Red orange 6-9 mm 4—6 mm Annular, mostly free from the tube F. pringsheimii 1.5_3.5(_4)cm 1-6 mm Subcoriaceous Dark glossy green above white green below 3-5 Axillary 19-31 mm Obconic-funnelform Bright red pink 13-24 mm 10-15 mm Irregular lobes adnate to the floral tube ner, and more elliptic than plants from the southern ranges. Burch & Burch 2546 (CAS, MO) and Fuertes 1720 (A, G, W) both have leaves reaching 13 cm long and 5.5 cm wide. Plants from this area typically have leaves that are heavily diffused with a purplish coloration, especially on the undersides, but this is by no means ubiquitous in all populations. Fuchsia pringsheimii is also endemic to Hispaniola. It is morphologically very distinct from F. triphylla, as can be seen by a comparison of several morpholog- ical characters in Table 19. Fuchsia triphylla is much more widespread than F. pringsheimii, probably because it is more weedy and occurs at lower elevations. As discussed previously on page 28, the ancestors of these species probably arrived from South America via long distance dispersal. The very marked mor- phological differences indicate either two separate immigrations or else a wide divergence since the original arrival of a common ancestor. Both species are tetraploid and have biporate pollen, a very unusual combination in the genus, and the two apparently hybridize in nature forming partly fertile hybrids. A typical altitudinal separation of Fuchsia triphylla and F. pringsheimii is shown in Figure 5. Although no actual overlap in the altitudinal limits of these species occurs between Constanza and Valle Nuevo, Prov. La Vega, they come very close together around 1,850-1,900 m. A probable hybrid from this approxi- mate altitude was detected, von Tuerckheim 3541 (BR), in woods below Valle Nuevo, 1,950 m, Aug. 1910. It is intermediate in leaf size, flower position, and flower shape, and has only 30% pollen stainability (400 grains examined). Informa- tion on the herbarium label also notes that the plant was growing at an intermediate elevation between populations of F. triphylla and F. pringsheimii. A large local population of Fuchsia pringsheimii {Berry et al. 3712; MO, JBSD) was found interspersed with plants of F. triphylla along the edge of a potato field in Dec. 1979 at 2,000 m at La Nuez, on the border of La Vega and 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Peravia Provinces in the Dominican Republic. No intermediates or apparent hy- brids could be found at that time, but only F. triphylla was in flower. Other hybrids have been detected mostly from Haiti or the Haiti-Dominican Republic border. Smith & Mejia 10192 (MO, JBSD) is intermediate both florally and vegetatively between the two species; it comes from the Dominican Republic along the Haitian border, Pro v. Pedernales, Carretera Internacional between Los Arroyos and Aguacate, in pines at 2,000 m, 10 Nov. 1979. Some of its leaves are 5 cm long with 6 secondary veins (tending toward F. triphylla), while the flowers are narrowed in the lower half, yet flare outward at the rim as in F. pringsheimii. The petals vary in size from 10 to 16 mm long and 7 to 9 mm wide, intermediate between the two species (see Table 19). The nectary is annular as in F. triphylla, and the pollen has numerous aborted, single-porate grains among the plumper bi- and triporate grains. The pollen stainability is 18.7% (500 grains examined). It is interesting that some triporate grains appear here, since both parents have bi- porate grains. Jimenez B-4697 (US), from Loma del Toro, in the same province and at the same elevation as Smith & Mejia 10192, also shows intermediate morphological characters and has a pollen stainability of under 10%. A final prob- able hybrid from the Dominican Republic, Gastony et al. 536 (US), occurs along the Haitian border, but farther north in the Sierra de Neiba at 1,700-2,000 m. It has a pollen stainability of ca. 75%. In Haiti, Ekman H.1893 (LL, S) has flowers intermediate between the axillary ones of Fuchsia pringsheimii and the racemose ones of F. triphylla. It was col- lected in the Massif de la Selle, Morne Franchant, on the ridge towards Godet, at ca. 1,900 m, Sept. 13, 1920. Except for a slight narrowing toward the rim, its flowers are typical of F. pringsheimii, as are the nectaries. The sepals are too short (13-15 mm long) and the petals too small (10-12 mm long and 7-9 mm wide) for that species, however. The leaves of this collection also exceed the normal length for F. pringsheimii and have 7 secondary veins. Its pollen stainability is 46.8% (500 grains examined). Ekman H.1620 (LL, MO, 2 sheets at S, US) is a set of collections in which some specimens appear to be hybrids. The two sheets at S and the one at MO are labelled, "Haiti, Massif de la Selle, Morne Cabaio, near Jardins Bois Pin, ca. 2,000 m, Aug., 1924," but the LL and US sheets say "Marigot" in place of "Morne Cabaio," so it is not clear if they were all collected together. The specimen at S with a typewritten label has numerous, axillary flowers, but constricted floral tubes as in F. triphylla. The nectary is an intermediate type between the ring of F. triphylla and the adnate nectary of F. pringsheimii, and the petals are obovate, but too small for F. pringsheimii. Its pollen stainability is 50.4% (500 grains examined). The second sheet at S, with a handwritten label, has flowers shaped much closer to F. pringsheimii, with nectaries characteristic of that species. Although its leaves reach dimensions typical of F. triphylla (5 cm long, with 7 secondary veins), its pollen stainability is over 90%. The LL sheet has flowers like F. triphylla but totally aborted pollen. The MO and US sheets present other combinations of intermediate traits. The wide differences between the different sheets of Ekman H.1620 may be the result of the segregation of individuals from a hybrid population. As discussed on p. 31, a single species of hummingbird, Chlorostilhon swainsonii, is probably the common pollinator of both F. triphylla and F. prings- 1982] BERRY— FUCHSIA SECT. FUCHSIA 191 heimii, and this may help explain the relatively large number of hybrid individuals detected. In addition, severe habitat disturbances have occurred in Hispaniola since the Spanish colonization so that much of the forest vegetation of the island has been eliminated, especially in Haiti (Liogier, 1978). This may have allowed previously separated populations of the two species to enter into closer contact, increasing the chances of intervisitation by Chlorostilbon. 60. Fuchsia pringsheimii Urban, Symb. Ant. 1:375. 1899. Barker & Dardeau, Flore d'Haiti 270. 1930. Munz, Proc. Calif. Acad. Sci. IV. 25:33, pi. 4, fig. 19. 1943. type: Dominican Republic, Prov. La Vega, near Valle Nuevo, in pines among ferns, 2,100 m, 29 May 1887, Baron Henrik F.A. Eggers 2159 (B, holotype, destroyed in World War II; BM, G, K, isotypes; photograph of K isotype, MO). Fig. 30. Erect to scandent shrubs 0.5-2 m tall with ascending to spreading branches. Branchlets terete, puberulent, pruinose, or rarely pilose, reddish purple; older stems with red brown, exfoliating bark. Leaves opposite or ternate, subcoria- ceous, narrowly (ob-)lanceolate to elliptic, acute to narrowly cuneate at the base, acute at the apex, 15-35(-40) mm long, 6-15 mm wide, dark rich green and glossy above, subglossy whitish-green to purplish below with red purple veins; upper surface glabrous to loosely strigose, strigose below, especially along the midvein and margin; secondary veins 3-5 on either side of the midvein; margin subentire to glandular-denticulate or dentate, sometimes only toothed in the upper half. Petioles sparsely strigose, 1-6 mm long, red purple. Stipules lanceolate, dark, 1- 1.5 mm long, deciduous. Flowers pendant and axillary near the branch tips. Pedicels slender, subglabrous to loosely strigose, 14^0 mm long. Ovary ellipsoid, 6-9 mm long, 2-3.5 mm thick, subglabrous, green to reddish. Floral tube obconic- funnelform, 19-31 mm long, 3-4 mm wide and slightly bulbous at the base, then widened continuously until 9-16 mm wide at the rim, subglabrous outside, gla- brous within. Sepals lanceolate, acute to acuminate at the apex, 18-25 mm long, 6-8 mm wide, suberect to spreading at anthesis. Tube and sepals bright red to pink red. Petals red to pink red, broadly obovate, emarginate to broadly truncate at the apex, 13-24 mm long, 10-15 mm wide, slightly spreading and convolute at the base at anthesis. Nectary a yellow-green band lining the basal 2-3 mm of the floral tube, irregularly-lobed and protruding slightly inwards from the tube. Filaments red, 15-18 mm and 12-13 mm long; anthers oblong, 2.5-3 mm long, 1.5-2 mm wide, white to yellow cream. Style red, glabrous; stigma clavate, 4-lobed at the apex, 3-3.5 mm long, 2-3 mm wide, pink to red. Berry oblong, ca. 15 mm long, ca. 8 mm thick; seeds ca. 2 mm thick, ca. 1 mm wide. Gametic chromosome number n = 22. Distribution: Endemic to the island of Hispaniola (West Indies). In Haiti, in the southern Massif de la Hotte and Massif de la Selle ranges and in the central Chaine des Matheaux, 1,400-2,500 m. In the Dominican Republic, in the southern Sierra de Baoruco, probably in the Sierra de Neiba, and in the Cordillera Central, (1,300-) 1,500-2,600 m; low shrubs mostly in pine forests (Fig. 67). Specimens examined: Haiti, ouest: Massif de la Selle, Morne La Visite, along path to Saltrou, Ekman 1450 (US), 1450a (S), 1450b (F, G, GH, S); Massif de la Selle, Morne Cabaio road, Ekman 1613 (S); Massif des Matheaux, Grand-Bois, Morne Moitie-Duportee, Ekman 5737 (S); Trou Bon 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Dieu, Morne La Selle, Holdridge 1055 (MICH, US), sud: Morne de la Hotte to mont. Ma Blanche, Ekman 604 (S). Dominican Republic, azua: Culo de Maco, Fuertes 1971 (A), barahona: Cana Brava, Monteada Nueva, Liogier & Liogier 25126 (JBSD). la vega: Valle Nuevo, Augusto 1495 (A, JBSD), 1507 (A, JBSD), Berry et al. 3707 (MO), 3709 (MO, JBSD), 3710 (MO, JBSD), Bueno (Jimenez Herb. #1792) (A, US), Jimenez & Withgow 1382 (US); 16 km S of Valle Nuevo, between La Piramide and La Nevera, Berry et al. 3711 (MO, JBSD); Valle Nuevo, Loma Atravesada, Ekman 13851 (S); trail from Los Tablones (2 km W of La Cienaga) to La Lagunita, Gastony et al. 287 (GH); 13 km from Valle Nuevo on road to San Jose de Ocoa, near the pyramid, Gastony et al. 716 (GH); Alto de Bandera, Liogier 25224 (JBSD); near Constanza, von Tuerckheim 3151 (BM, BR, F, G, GH, K, MO, S, U, US, W, Z). peravia: La Nuez, border with Prov. La Vega, Berry et al. 3713 (MO); San Jose de Ocoa, Cuchilla del Pino Atravesado, Ekman 1 1710 (K, S); La Cueva, La Horma Arriba, Liogier 20956 (JBSD). san juan: Sabana Nueva, Piedra del Aguacate to Rio del Oro, Howard & Howard 9014 (B, BM, GH, LE, S, US). Fuchsia pringsheimii is characterized by its axillary flowers with wide floral tubes and very large, retuse petals, and by its small, dark green, few- veined leaves. It can be distinguished from F. triphylla, the other endemic species on Hispaniola, by the morphological features outlined in Table 19. Both species are tetraploid and apparently hybridize in several different localities, even though F. pringsheimii is rather infrequent and occurs at higher elevations than F. triphylla. The putative hybrids and their pollinator and ecological relationships are further discussed under F. triphylla. Following Munz (1943), Fuchsia pringsheimii is included in sect. Fuchsia, but with considerable reservation. Its affinities with this section are doubtful because it has a unique combination of features not found in the main body of sect. Fuchsia or in any other groups in the genus. These include the large, con- volute, and retuse petals, the sepals nearly as long as the floral tubes, the partic- ular non-annular nectary, the reduced leaves, and tetraploidy with biporate pol- len. These characters suggest that F. pringsheimii may represent an early offshoot of sect. Fuchsia that arrived at Hispaniola via long-distance dispersal from South America, or possibly even an earlier line, that was related to the branch of the genus that may have reached Central America before the close of the Paleo- gene. The other species of Fuchsia on Hispaniola, F. triphylla, is also tetraploid with biporate pollen, and the two species hybridize in nature, but F. triphylla is morphologically much more similar to the Andean members of sect. Fuchsia than is F. pringsheimii. Therefore, it is possible that two separate colonizations of the genus occurred in Hispaniola. Some notable variation in leaf size and pubescence occurs between popula- tions from the Cordillera Central of the Dominican Republic and the southern Sierra de Baoruco and Massif de la Selle. Plants from the Cordillera Central have leaves mostly less than 25 mm long and less than 10 mm wide, with rather short, sparse pubescence. In the southern range, most plants have larger and broader leaves 25-40 mm long and 10-15 mm wide. Ekman 1613 (S), from Haiti, is heavily hirsute-pilose on both stems and leaves. On the Dominican side of the border in Prov. Barahona, F. pringsheimii reaches its lowest altitudinal limit at 1,300 m, according to the label of Liogier & Liogier 25126. 61. Fuchsia verrucosa Hartweg ex Bentham, PI. Hartw. 178. 1845. Munz, Proc. Calif. Acad. Sci. IV. 25:58, pi. 8, fig. 47. 1943. Rodriguez, Pittiera 6:10, fig. 1 , a-e. 1974. type: Colombia, Dept. Cundinamarca, on the road between the 1982] BERRY— FUCHSIA SECT. FUCHSIA 193 Hart entham Herb., holotype; BM, BREM, CGE, W, isotypes; photograph of B isotype, POM) 47. Fuchsia perbrevis I. M. Johnston, Contr. Gray Herb. 75:30. 1925. type: Colombia, Dept. Cundinamarca, above Sibate, 3 Jan. 1854, Isaac F. Holton 892 (NY, holotype; K, isotype). Fuchsia verrucosa var. tamaensis Steyermark, Fieldiana, Bot. 28:440. 1952. type: Venezuela, Edo. Tachira, base of Paramo de Tama, between Betania and Tama, near the Colombian border, 2,430 1944 subshrubs terete, 2-4 mm thick, verrucose, green to dull purple; older stems terete, with light gray tan, finely fissured bark. Leaves opposite, firmly membranous, broadly elliptic to obovate, acute to attenuate at the base, acute to acuminate at the apex, 5-16 cm long, 2-8 cm wide, dark matte green above, pale or flushed purple below, often with strigose hairs along the veins; mid vein prominent below, secondary 10-20 3 mm rm Petioles stout, 5-22 mm long. Stipules firm to semisucculent, (narrowly) trian- gular, 2-4 mm long, 1-2 mm wide, separate or connate, subpersistent. Flowers 4-4 solitary in upper leaf axils. Pedicels firm, spreading to drooping, 10-40 mn Ovary oblong-cylindric, tetragonous, 10-12 mm long, 2.5-3 mm wide, lu light green. Floral tube obconic, 3-6 mm long, 2.5-3 mm wide at the base, mm wide at the rim, subglabrous inside and outside. Sepals lance-oblong, acute at the apex, 7-11 mm long, 2-4.5 mm wide at the base, spreading to recurved at anthesis. Tube and sepals bright orange or scarlet. Petals red orange, oblong- obovate, 8-9 mm long, 5-6 mm wide, obtuse to subacute at the apex, margin undulate, central nerve ± thickened, spreading and convolute at anthesis. Nec- tary green and composed of 4 distinct antesepalous, ridged lobes ca. 1.5 mm high, ca. 0.5 mm thick, adnate to the base of the floral tube. Filaments light red, 2-3 mm and 1-1.5 mm long; anthers broadly reniform, 1.5-2.5 mm long, 1.5-2 mm wide, white. Style glabrous, stout, red, 6-8 5^ exserted beyond the anthers. Berry tetragonous until maturity, ellipsoid or broad- ly cylindric when ripe, 20-25 mm long, ca. 10 mm thick; seeds 1.4-1.6 mm long, ca. 0.7 mm wide. Gametic chromosome number n = 22. Distribution: Venezuela and Colombia. In the Cordillera Oriental of the North- ern Andes from southern Merida, Venezuela to Huila, Colombia and then on the Putumayo; infrequent shrubs 800-3 Representative specimens examined: Venezuela, Merida: road to Mesa de Bustamante, 5 km S of La Playita, between Bailadores and Paramo La Negra, Tillett & Honig 738-397 (MO), tachira: 6 km E of Zumbador on road to Queniquea, Berry 34/7 (MO); 13 km E of El Portachuelo on road from Bailadores to Pregonero, Berry 3287 (MO, VEN); 7 km E of El Portachuelo, road to Pregonero, Berry 3281 (MO), 3662 (MO); El Hato, road to Pregonero, Ldpez-Palacios 1969 (MERF, MO); headwaters of Rio Quinimari, Las Copas, 15 km S of San Vicente de la Revancha, Steyermark et al. 100577 (MO, VEN); Quebrada Agua Azul, S of El Reposo, Steyermark & Liesner 118460 (MO, VEN). Colombia, caqueta: 57 km from Florencia to Altamira, Luteyn et al. 4954 (COL, MO, NY, US); E of summit between Garzon and Florencia, Mason 13953 (COL, F, GH, UC, US), cundinamarca: Barroblanco, 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 Andre 1336 (K, NY); Km 34-35 of Bogota-Sibate-Fusagasuga road, Berry 3545 (COL, MO); between Pacho and Rio Negro, Garcia-Barriga 10753 (COL, US); Toquiza, Gazaunta valley, 15 km NW of Medina, Grant 10260 (COL, NA, RSA); Fusagasuga, Linden 811 (BM, BR, F, G, K, LE, OXF, W), Triana 3816 (G, K, P). huila: Km 31 of road S of Pitalito towards Mocoa, Berry 3595 (COL, MO); Gabinete, border with Caqueta, Cuatrecasas 8465 (CM, COL, F, RSA, US); headwaters of Rio Fortalecillas, below Paso de Las Cruces, 39 km E of Neiva, Fosberg 19787 (RSA, US), meta: Paramo de Sumapaz, S of Cordillera de Las Cruces, Fosberg 20868 (US), norte de santander: divide between Rio La Teja and Rio Mesme, between Pamplona and Toledo, Killip & Smith 19831 (A, F, GH, NY, US), 19976 (A, GH, NY, US); prov. of Ocana, Schlim 705 (G, K, P). putumayo: road from San Francisco to Mocoa, Mora 4374 (COL); Km 92 from Pasto to Mocoa, Plowman & Davis 4327 (COL, PSO). santander: mountains above Bucaramanga, Barkley & Araque I8S097 (COL, US, VALLE). This is probably the only species in the genus in which the ovary is clearly longer than the floral tube, and both are quadrangular in transection. It is further characterized by the strictly opposite leaves, verrucose stems, and axillary po- sition of the orange flowers. Vegetatively and geographically, Fuchsia verrucosa is consistent with its placement in sect. Fuchsia, but its extremely short floral tubes, antesepalous nectary lobes, smooth viscin threads, and tetraploidy with biporate pollen are anomalous in this section. The nectary and small flower size of F. verrucosa are most like those of the Central American F. jimenezii (sect. Jimenezia), but that species has smaller, terete ovaries, paniculate inflorescences, and a reflexed whorl of stamens. Fuchsia verrucosa occurs sympatrically with seven other species of sect. Fuchsia (F. cuatrecasasii, F. gehrigeri, F. hirtella, F. nigricans, F. scabriuscula, F. sessilifolia, and F. venusta), and despite searches in the field and in herbarium specimens, no apparent hybrid plants were found, further indicating a lack of close relatives in sect. Fuchsia. Experimental crosses with species such as F. jimenezii, F. pringsheimii, and other species of sect. Fuchsia might help elucidate the affinities of this species. Uncertain Species Fuchsia miniata Planchon & Linden, Fl. Serres Jard. Eur. 8:7, pi. 1852. lec- totype: the illustration in the preceding citation, cultivated in Belgium (?) in 1852 by Jean- Jules Linden, from seeds collected in Colombia by M. Schlim. The illustration and short description provide insufficient detail to characterize this species adequately. Although the illustration closely resembles F. crassi- stipula, the conspicuous stipules of that species are not present in the plate, and the description specifically mentions the stems as terete, whereas in F. crassi- stipula they are conspicuously ridged. A copy of the published illustration is in the herbarium at Montpellier, France (MPU), where Planchon did much of his work. The plate is accompanied in the same folder by several fragments of what appears to be F. gehrigeri, but these fragments are not attached to the herbarium sheet and do not seem to correspond to the illustration. Fuchsia platypetala I. M. Johnston, J. Arnold Arbor. 20:241. 1939. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4):561. 1941. Munz, Proc. Calif. Acad. Sci. IV. 25:30, pi. 3, fig. 16. 1943. type: Peru, Dept. Apurimac, Chincheros, along lanes in town, semi-cultivated but reported wild nearby, 2,930 m, 1 Nov. 1935, 1982) BERRY— FUCHSIA SECT. FUCHSIA 195 James West 3705 (UC, holotype, not seen; GH, MO, isotypes; photograph of GH isotype, NY). This entity is known only from cultivated material from Depts. Apurimac and Cuzco of southern Peru, and no collections more recent than 1946 have been seen. The plants are axillary-flowered with large, obovate petals that are some- what mottled when dry. They also have reflexed sepals, which strongly suggests a relationship to Fuchsia boliviana, a species extensively cultivated nearby, but with long, racemose inflorescences. Other plants referrable to F. platypetala are Vargas 6267 (CUZ), from the town of Pumamarca, Dept. Cuzco, 3,400 m, and Herrera 1514 (GH), from the city of Cuzco. Most likely these represent hybrids of F. boliviana with some other axillary flowered species of the region such as F. denticulata or F. austromontana, both of which have petals that dry streaked or mottled as in the specimens cited above. Whereas the pollen stainability of the type collection is high, the pollen of Herrera 1514 is totally aborted. Excluded Species Fuchsia involucrata Sw., Prodr. 62. 1788. = Urceolaria involucrata (Sw.) Standi., N. Am. Flora 32(2): 132. 1921. (Rubiaceae). Literature Cited Adams, C. D. 1972. Flowering Plants of Jamaica. Univ. of West Indies. Mona, Jamaica. 848 p. Alexander, M. P. 1969. Differential staining of aborted and nonaborted pollen. Stain Tech. 44:117-122. Almeda, F., Jr. 1978. Systematics of the genus Monochaetum (Melastomataceae) in Mexico and Central America. Univ. Calif. Publ. Bot. 75:1-134. Andre, E. 1888. Le centenaire des Fuchsias. Rev. Hort. 60:230-233. Arroyo, M. T. K. & P. H. Raven. 1975. The evolution of subdioecy in morphologically gynodi- oecious species of Fuchsia sect. Encliandra (Onagraceae). Evolution 29:500-511. Atsatt, P. R. & P. W. Rundel. 1982. Pollinator maintenance vs. fruit production: Partitioned reproductive effort in subdioecious Fuchsia lycioides. Ann. Missouri Bot. Gard. 68:199-208. Baillon, H. 1877. Histoire des plantes. 6:466-467. Bentham, G. 1839-1857. Plantae Hartwegianae. London. 393 p. & J. D. Hooker. 1867. Genera plantarum 1:790-791. Beuzenberg, E. J. & J. D. Hair. 1959. Contributions to a chromosome atlas of the New Zealand flora. 3. Miscellaneous families. New Zealand J. Sci. Tech. 2(4):53 1-539. Breedlove, D. E. 1969. The systematics of Fuchsia sect. Encliandra (Onagraceae). Univ. Calif. Publ. Bot. 53:1-69. , P. E. Berry & P. H. Raven. 1982. The Mexican and Central American species of Fuchsia (Onagraceae) except sect. Encliandra. Ann. Missouri Bot. Gard. 69:209-234. Brown, C. A. 1967. Pollen morphology of the Onagraceae. Rev. Palaebotan. Palynol. 3:163-180. Candolle, A. P. de. 1828. Prodr. 3:36-39. Carlquist, S. 1967. The biota of long-distance dispersal. V. Plant dispersal to Pacific Islands. Bull. Torrey Bot. Club 94:704-722. . 1975. Wood anatomy of Onagraceae, with notes on alternative modes of photosynthate movement in dicotyledon wood. Ann. Missouri Bot. Gard. 62:386-424. Chaudhuri, S. K. 1956. Cytogenetic studies in the genus Fuchsia. Ph.D. thesis, Univ. of Man- chester, England. 324 p. Cleef, A. 1979. The phytogeographical position of the neotropical vascular paramo flora with special reference to the Colombian Cordillera Oriental. Pp. 175-184, in K. Larsen & L. D. Holm- Nielsen (eds.), Tropical Botany. Academic Press, London. Couper, R. A. 1960. New Zealand Mesozoic and Cainozoic plant microfossils. New Zealand Geol. Sur. Palaeont. Bull. 32:1-87, pi. 1-12. 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Cuatrecasas, J. 1958. Aspectos de la vegetacion natural en Colombia. Revista Acad. Colombiana Ci. Exact. Fis. Nat. 10:221-264. — . 1979. Comoaracion fitoeeo Pp Salgado-Labouriau (ed.), El Medio Ambiente Paramo. Editorial Arte, Caracas, Venezuela. Descourtilz, M. E. 1822. Flore Medicale des Antilles. Vol. 2. Dietrich, W. 1977. The South American species of Oenothera sect. Oenothera (Raimannia, Ren- neria; Onagraceae). Ann. Missouri Bot. Gard. 64:425-626. Duellman, W. E. 1979. The herpetofauna of the Andes: patterns of distribution, origin, differ- entiation, and present communities. Pp. 371-459, in W. E. Duellman (ed.), The South American Herpetofauna: Its Origin, Evolution, and Dispersal. Monogr. Mus. Nat. Hist., Univ. Kansas, no. 7. Dymond, J. 1975. K-Ar ages of Tahiti and Moorea, Society Islands, and implications for the hot- spot model. Geology 3:236-240. Endlicher, S. 1840. Genera plantarum. Wien (Fr. Beck) 1836-1840. Ewel, J. J., A. Madriz & J. A. Tosi. 1976. Zonas de vida de Venezuela. Memoria explicativa sobre el mapa ecologico. Ed. 2. Editorial Sucre, Caracas, Venezuela. Eyde, R. H. & J. T. Morgan. 1973. Floral structures and evolution in Lopezieae (Onagraceae). Amer. J. Bot. 60:771-787. Fielding, H. B. & G. Gardner. 1844. Fuchsia caracasensis. Sert. PI., /. 29. Flenley, J. R. 1979. The Equatorial Rain Forest: A Geological History. Butterworths, London. 151 p. 64-68 Gentry, A. H. 1974. Flowering phenology and diversity in tropical Bignoniaceae. Biotropica t Gerth, H. 1941. Die Tertiarfloren des siidlichen Sudamerika und die ungebliche Verlagerung des Siidpols wahrend dieser Periode. Geol. Rundschau 32:321-336. Grant, K. A. & V. Grant. 1968. Hummingbirds and Their Flowers. Columbia Univ. Press, New York. Gregory, D. P. 1964. Hawkmoth pollination in the genus Oenothera. Aliso 5:357-419. Haffer, J. 1974. Avian speciation in tropical South America. With a systematic study of the Toucans (Ramphastidae) and Jacamars (Galbulidae). Publications of the Nuttall Ornithological Club, No. 14. 390 p. Hammen, T. van der. 1974. The Pleistocene changes of vegetation and climate in tropical South America. J. Biogeography 1:3-26. 1979. Changes in life conditions on earth during the past one million years. Kongel. Danske Vidensk. Selsk. Biol. Skr. 22:1-32. Haque, A. 1952. Chromosome counts of species and varieties of garden plants. Ann. Rept. John Innes Hort. Inst. 41:47-50. Harrison, J. 1849. Fuchsia corymhiflora alba. Floric. Cab. Florist's Mag. 17:97. Heinrich, B. & P. H. Raven. 1972. Energetics and pollination ecology. Science 176:597-602. Hemsley, W. B. 1876. The various garden races of Fuchsia. Garden (London) 9:284-286. . 1877. The species of Fuchsia. Garden (London) 11:70-75. . 1882. New garden plants. Gard. Chron. II. 18:263-264. Hickey, L. J. 1973. Classification of the architecture of dicotyledonous leaves. Amer. J. Bot. 60:17-33. 1980. Leaf architecture of Onagraceae. P. 69 in Abstracts of the Second International Congress of Systematics and Evolutionary Biology, Vancouver, Canada, July 17-24, 1980. Houtte, L. van. 1854. Nouvelle variete de Fuchsia. Fuchsia dominiana. Fl. Serres Jard. Eur. 10:95-96, pi. 1004. Humboldt, A. von, A. Bonpland & C. S. Kunth. 1823. Nova genera et species plantarum, vol. 6:1-240. Huynh, K. L. 1965. Contribution a Tetude caryologigue et embryologigue des Phanerogames du Perou. Schweiz. Naturforsch. Ges., Mem. Soc. Helvetique Sci., Nat. 85:1-178. Jeannel, R. 1967. Biogeographie de TAmerique australe. Pp. 401-460, in C. Delamere Deboutte- ville & E. Rapoport (eds.), Biologie de TAmerique australe, 3. C.N.R.S. Groupe Francais Argiles C. R. Reun. Etud., Paris, 834 p. Johnston, I. M. 1925. Some undescribed American spermatophytes. Contr. Gray Herb. 75:27-40. . 1939. New fuchsias from southern Peru. J. Arnold Arbor. 20:241-244. Keating, R. C. 1980. Vegetative anatomy of Onagraceae. P. 68 in Abstracts of the Second Inter- national Congress of Systematics and Evolutionary Biology. Vancouver, B.C., Canada. July 17-24, 1980. Kuhnel, J. 1960. Thaddeus Haenke. Leben und Wirken eines Forschers. Prag. 276 p. Kurabayashi, M., H. Lewis & P. H. Raven. 1962. A comparative study of mitosis in the Ona- graceae. Amer. J. Bot. 49(9): 1003-1026. 1982] BERRY— FUCHSIA SECT. FUCHSIA 197 Lack, D. 1976. Island Biology (Illustrated by the Land Birds of Jamaica). Univ. of Calif. Press, Berkeley. Lamarck, J. P. 1788. Encyclopedic methodique 2:564-566. Lauer, W. 1979. La position de los paramos en la estructura del paisaje de los Andes tropicales. Pp. 29-43, in M. L. Salgado-Labouriau (ed.), El Medio Ambiente Paramo. Editorial Arte, Ca- racas, Venezuela. Lindley, J. 1840. Fuchsia corymbiflora. Cluster-flowered Fuchsia. Bot. Reg. 26, pi. 70. Linnaeus, C. 1737. Genera plantarum. ed. 1. . 1742. Genera plantarum. ed. 2. . 1753. Species plantarum. Liogier, A. H. 1978. La flora de la Espanola: analisis, origen probable. Col. Conf. Acad. Ci. Repiiblica Dominicana 3: 1-32. Macbride, J. F. 1940. New or renamed spermatophytes mostly Peruvian. Candollea 8:21-33. . 1941. Flora of Peru. Onagraceae. Field Mus. Nat. Hist., Bot. Ser. 13(4):52 1-566. Mayr, E. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Mass. Mildenhall, D. C. 1980. New Zealand Late Cretaceous and Cenozoic plant biogeography: a contribution. Palaeogeogr., Palaeoclimatol., Palaeoecol. 31:197-233. Miller, P. 1739. The second volume of the gardeners dictionary. 3rd ed. Mosquin, T. 1966. A new taxonomy for Epilobium angustifolium L. (Onagraceae). Brittonia 18 Munz, 25 167-188. P. A. 1943. A revision of the genus Fuchsia (Onagraceae). Proc. Calif. Acad. Sci. IV. 1-138, pis. 1-16. 1946. Fuchsia colombiana, new species. Caldasia 4:109-111. 1972. Three South American species of Fuchsia. Aliso 7:409-41 1. 1974. Onagraceae, Flora of Ecuador. Opera Bot., Ser. B, 3:3—46. Paxton, J. 1841. Fuchsia corymbiflora. Paxtons Mag. Bot. 7-8. Percival, M. 1969. Floral Biology. Pergamon Press, Oxford, England. 243 p. Plowman, T. 1979. The genus Brunfelsia: a conspectus of the taxonomy and biogeography. Pp. 475-491, in J. G. Hawkes, R. N. Lester & A. D. Skelding (eds.), The Biology and Taxonomy of the Solanaceae. Linn. Soc. Symp. Ser. 7. Plumier, C. 1703. Nova plantarum americanarum genera 14-15, t. 14. . 1757. Plantarum americanarum, fasc. VI: 124-125, pi. 133, fig. I. Porcher, F. 1858. Le Fuchsia. Son histoire y sa culture. 3rd ed. Paris. 214 p. Raven, P. H. 1972a. Why are bird-visited flowers predominantly red? Evolution 26:674. . 1972b. Plant species disjunctions: a summary. Ann. Missouri Bot. Gard. 59:234-246. . 1973. The evolution of Mediterranean floras. Pp. 213-224, in H. H. Mooney & F. de Castri (eds.), The Convergence in Structure of Ecosystems in Mediterranean Climates. Springer, Berlin. . 1979a. A survey of reproductive biology in Onagraceae. New Zealand J. Bot. 17:575-593. . 1979b. Plate tectonics and Southern Hemisphere biogeography. Pp. 3-24, in K. Larsen & L. B. Holm-Nielsen (eds.), Tropical Botany. Academic Press, London. — . 1980. Hybridization and the nature of species in higher plants. Canadian Bot. Assoc. Bull Supplement to Vol. 13. No. 1:3-10. — & D. I. Axelrod. 1974 [1975]. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61:539-673. — & T. E. Raven. 1976. The genus Epilobium (Onagraceae) in Australasia: a systematic and evolutionary study. New Zealand Dept. Sci. Ind. Res. Bull. 216. 321 p. Reiter, V., Jr. 1941. The outdoor culture of the Fuchsia in coastal California. J. Calif. Hort. Soc. 2:1-20. Ruiz, H. & J. Pa von. 1802. Flora peruviana et chilensis, vol. 3:1-95. Sauer, W. 1971. Geologie von Ecuador. Gebriider Borntraeger, Berlin. 316 p. Sillitoe, R. H. 1974. Tectonic segmentation of the Andes: implications for magmatism and met- allogeny. Nature 250:542-545. Simpson, B. B. 1973. Contrasting modes of evolution in two groups of Perezia (Mutisieae; Com- positae) of southern South America. Taxon 22:525-536. . 1975a. Glacial climates in the eastern tropical South Pacific. Nature 253:34-36. . 1975b. Pleistocene changes in the flora of the high tropical Andes. Paleobiology 1: 273-294. . 1979. Quaternary biogeography of the high montane regions of South America. Pp. 157-189, in W. E. Duellman (ed.), The South American Herpetofauna: Its Origin, Evolution, and Dis- persal. Monogr. Museum of Natural History, Univ. of Kansas, no. 7. Skutch, A. F. 1973. The Life of the Hummingbirds. Crown Publishers, New York. 95 p. Skvarla, J. J., P. H. Raven, W. F. Chissoe & M. Sharp. 1978. An ultrastructural study of viscin threads in Onagraceae pollen. Pollen et Spores 20:5-143. 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 62:6-35. - & J. Praglowski. 1975. The evolution of pollen tetrads in Onagraceae. Amer. J. Bot. , & . 1976. Ultrastructural survey of Onagraceae pollen. Pp. 447-479, in I. K. Ferguson & J. Muller (eds.), The Evolutionary Significance of the Exine. Academic Press, London. Smith, L. B. 1962. Origins of the flora of southern Brazil. Contr. U.S. Nat. Herb. 35:215-249. . 1965. Itinerary of Edouard Francois Andre in his expedition to the northern Andes 1875-1876. Phytologia 12(7):401-413. Solbrig, O. T. 1976. The origin and floristic affinities of the South American temperate desert and semitemperate regions. Pp. 7-49, in D. Goodall (ed.), Evolution of Desert Biota. Univ. Texas Press, Austin. Spach, E. 1835. Les Onagraires — Onagrarieae. Hist. Nat. Veg. 4:335-416. Sussman, R. W. & P. H. Raven. 1978. Pollination by lemurs and marsupials: an archaic revo- lutionary system. Science 200:731-736. Vargas Calderon, C. 1964. Las especies del genero Fuchsia del Departamento Cuzco. Biota 5(37): 1-5. Vuilleumier, B. S. 1969 [1970]. The systematics and evolution of Perezia sect. Perezia (Com- positae). Contr. Gray Herb. 199:1-163. . 1971. Pleistocene changes in the flora and fauna of South America. Science 173:771-780. Walpers, G. G. 1843. Repert. Bot. Syst. 2:94-95. Warth, G. 1925. Zytologische, histologische und stammesgeschichtliche Fragen aus der Gattung Fuchsia. Z. Indukt. Abstamm. Vererbungslehre 38(3):20C^257. Wright, J. O. 1978a. A new species of Fuchsia L. Bot. Jour. Linn. Soc. 77:113-115. . 1978b. Fuchsia, a garden history. The Plantsman 1:181-186. POLLINATOR MAINTENANCE VS. FRUIT PRODUCTION PARTITIONED REPRODUCTIVE EFFORT IN SUBDIOECIOUS FUCHSIA LYCIOIDES Peter R. Atsatt and Philip W. Rundel 1 Abstract Populations of Fuchsia lycioides Andrews are composed of small-flowered female plants and an equal number of larger-flowered hermaphrodites, which may be female fertile, or morphologically or physiologically female sterile. A major selective force driving the evolution of separate sexes in F. lycioides is suggested to be the partitioning of limited resources associated with pollinator maintenance and fruit production in a semi-arid mediterranean climate. The hummingbird Rhodopsis vesper at a- camensis is the only known pollinator, and appears to be energetically dependent upon F. lycioides. Hermaphrodites are facultative in their fruit production, produce as much as six times more nectar than females, and feed birds both prior to flower opening and during anthesis. Females bear abundant fruit and produce only a relatively small amount of nectar during anthesis. Nectar production is extremely variable in both sexes but is unpredictable in hermaphrodites and apparently predictable in females. The unpredictability of hermaphrodite nectar production may be a key factor permitting the evolution of resource partitioning into large-flowered bird-feeding pollen plants and small-flowered reproductive individuals. Subdioecy, a state of dioecy in which populations regularly contain imper- fectly differentiated individuals in addition to strictly unisexual individuals, has evolved by at least five evolutionary pathways (Ross, 1982). The genus Fuchsia is characterized by the hermaphroditism-gynodioecy-subdioecy pathway, where- in the male sterile mutants among the hermaphrodites produce the gynodioecious state, followed by a gradual reduction of seed fertility of hermaphrodites so that these come to function largely or completely as males (Godley, 1955; Breedlove, 1969; Arroyo & Raven, 1975). The selective forces apparently responsible for the evolution of dioecy have recently been reviewed by Ross (1982), who emphasized that many of the pro- posed selection models (fertility variation, sexual selection, overdominance, and resource allocation) are scarcely or not at all distinct from each other. Cases involving pollinator influence, fruit dispersal, and predation (Bawa, 1980; Givnish, 1980) are also likely to involve differential resource allocation. If so, perhaps the most generalized model for the evolution of dioecy is simply an energetic argu- ment: that female reproductive effort is largely limited by the availability of phys- iological resources, and that separation of the sexes increases fitness by allowing more efficient use or allocation of limiting factors. The resource allocation model need not exclude the traditionally considered hypothesis of outbreeding advan- tage (inbreeding depression). Ross (1982) concludes that outbreeding advantage is not always a factor in the evolution of dioecy, and even where it occurs, it is probably accompanied by other selective forces. Freeman et al. (1980a), and Bawa (1980) have expressed similar views. In this paper we report observations on the floral biology of a facultatively 1 Department of Ecology and Evolutionary Biology, University of California, Irvine, California 92717. Ann. Missouri Bot. Gard. 69: 199-208. 1982. 0026-6493/82/0 1 99-0208$0 1.15/0 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 30 31 c a 32 35 34 x Cuesta Buenas Aires Arraydn 'l_o Serena T Punto Oscuro Pichidangui Los Molles Papudo Vina del Mar Valparaiso '•"":•:■ 600 l_j I km 40 80 120 J km tiiiy Santiago 71 73 Figure I. The geographic range of Fuchsia lycioides and the location of study sites along the coast of central Chile. subdioecious shrub, Fuchsia lycioides Andrews, and examine evidence consistent with the hypothesis that differential resource allocation (related to pollinator maintenance) may be an important selective force favoring the evolution of sep- arate sexes in this species. Fuchsia lycioides, forming the monotypic section Kirschlegeria of the genus, is restricted to a narrow coastal belt in the Mediterranean-climate zone of central Chile (Fig. 1) in a relatively severe environmental regime with only moderate winter precipitation and long dry summer conditions. At its southern limit south of Valparaiso, mean annual precipitation is 460 mm/yr while at its northern limit north of La Serena there is only slightly more than 150 mm/yr. Mean growing season temperatures vary little over this latitudinal range. Ecologically, F. lycioides is characteristic of bluff communities along the im- mediate coast where it occurs in mixed stands of deciduous and evergreen shrub 1982] ATSATT & RUNDEL— REPRODUCTIVE EFFORT IN FUCHSIA LYCIOIDES 201 species. On typical sites such as those at Pichidangui (Coquimbo Province) it occurs with other woody deciduous shrubs (Adesmia arborea and Proustia pun- gens), a mixed assemblage of deciduous semiwoody shrubs, and the evergreen Lithraea caustica. On exposed rocky headlands, as at Los Molles (Aconcagua Province) just south of Pichidangui, however, F. lycioides occurs in a predomi- nately evergreen community dominated by Lucuma valparadisea. Further north Punto hrubby varied community types, F. lycioides remains an important (dominant) species. Methods Field observations and data were collected over the entire range of the species in Chile, at Papudo, Los Molles, Pichidangui, Punto Oscuro, Arrayan, and Cuesta Buenos Aires, in the years 1974, 1975, 1977, and 1978. Differences between her- maphrodite and female flowers were characterized by measuring flower tube length and style length in one southern (Pichidangui) and one northern (Arrayan) pop- ulation. Data were also collected on style abortion in hermaphrodites, and the ratio of female to hermaphrodite individuals was determined at four sites. The standing nectar crop was measured by sampling flowers with 10 lamda micropipets. In the field, buds were sampled in late afternoon by puncturing the base of the closed flower tube with a micropipet. Open, non-pollinated flowers (lacking white pollen on the red stigma) were sampled between 800 and 1000 hours. Nectar was measured for a series of marked flowers in the late afternoon, and the flowers were then bagged and remeasured the following morning to de- termine overnight nectar production. Nectar productivity was also measured from three plants each of female and hermaphrodite individuals raised from seed in the greenhouse. Flowers were tagged and measured at 1700 hours on the first, second, and third day after opening. Results FLORAL BIOLOGY The rose-colored, dimorphic flowers of F. lycioides have four sepals, four small petals, and a cylindric floral tube. The open cup-shaped floral tube of the female flower is usually 1.5-3 mm long, with a 6-10 mm style at our primary study site at Pichidangui. Eight reduced anthers are present but do not contain pollen. The larger hermaphrodite flowers are 2.5-5 mm long, with style length ranging from 14-22 mm. Although tube length of the two flower types may oc- casionally overlap, there is a sharp gap of 3.5-4 mm between maximum style length in the small female flowers and minimum style length in the larger her- maphrodite flowers (Fig. 2). Flowers of the Arrayan population show the same pattern but are smaller in size, particularly with regard to maximum style length, reflecting the considerably drier conditions under which these plants were grow- ing. A variable proportion of the flowers on hermaphrodite plants lack styles and are therefore functionally male. The percentage of male flowers on hermaphrodite plants was quantified in the northern Arrayan and Cuesta Buenos Aires popula- 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 5 4 3 E E 2 o UJ 4 3 2 4 PICHIDANGUI FEMALES HERMAPHRODITES 1 1 1 J I l l l l l arrayAn 1 1 1 1 1 uj 5|- | m ' FEMALES 1 I ) HERMAPHRODITES O O O <£> o O O O 050); hermaphroditic. 8a. Sepals shorter than the tube; petals green, 6-12 mm long. 1. F. splendens (sect. Ellobium) 8b. Sepals mostly longer than the tube; petals showy, 10-20 mm long, never green. Cultivated, (sect. Quelusia). 9a. Tube 5-10 mm long, sepals 15-25 mm long, petals purple. F. magellanica Lam. 5 9b. Tube 10-20 mm or more long, sepals generally 25-30 mm long; variable garden hybrids. F. hybrida Hort Fuchsia sect. Ellobium (Lilja) Breedlove, Berry, & Raven, comb. nov. Ellobium Lilja, Linnaea 15:262. 1841. type: Fuchsia fulgens De Candolle; based on Spachia Lilja; non Ellobum Blume, 1826, nom. rejic. Spachia Lilja, Tidning Tradgardsskotsel allman Wextkultur 8:62. 1840, horn, illeg., non Spachea Juss. 1838. Hermaphroditic. Soft wooded, terrestrial, or epiphytic shrubs, some species tuberous. Leaves opposite or ternate, membranous, elliptic-ovate to cordate. Flowers axillary, racemose, or paniculate. Floral tube longer than sepals. Petals usually Vi length of the sepals or less. Nectary unlobed, a smooth band 0.3-0.5 mm thick lining the base of the floral tube. Stamens biseriate, erect, shorter than the sepals or exserted less than 5 mm beyond them, the antisepalous stamens longer than the antipetalous ones. Stigma green. Berry ellipsoid to narrowly cy- lindric; seeds ca. 50-ca. 200, laterally compressed, oblong to irregularly triangular in outline, 1-2 mm long, 0.5-1.5 mm wide. Gametic chromosome number n = 11. Distribution: Evergreen cloud forests and moist oak-pine forests from north- ern Costa Rica to Jalisco, Mexico, at altitudes of 1,450-3,400 m (Figs. 1 and 2). Munz (1943) placed Fuchsia splendens (including F. cordifolia) and F. ful- gens in the large, generalized sect. Fuchsia. Fuchsia decidua was included in sect. Hemsleyella, a smaller South American group with tubers, apetalous flow- ers, and often an epiphytic, dry season flowering habit. Geographically and mor- phologically, however, the three species of sect. Ellobium form a well defined group that does approach the above mentioned sections in several characters, but can be distinguished by the characters shown in Table 1. 5 See appendix for lists of specimens examined from Mexico and Central America. 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 213 20° N Figure 1. Distribution of Fuchsia splendens (sect. Ellobium). The species in sect. Ellobium follow a clear progression of specialization from F . splendens, the most generalized, to F. decidua, the most specialized. This progression consists of a reduction in petal size, change in flower position from axillary to racemose to paniculate, and the development of tubers and an epi- phytic habit in F. fulgens and F. decidua, accompanied by a marked seasonality in the phenology of flowers and leaves. Furthermore, F. splendens is the most widespread species, and F. decidua is rare. 1. Fuchsia splendens Zuccarini, Flora 15, ii, Beibl., 102. 1832. type: Los Molinos, Oaxaca, Mexico, 1827-1832, Wilhelm Karwinsky (M, holotype; photographs, F, GH, MICH, MO, POM, US; BR, G, isotypes). Lindl., Bot. Reg. 28:/?/. 27. 1842. Curtis, Bot. Mag. 70:p/. 4082. 1844. Planch., Fl. Serres Jard. Eur. 5:/. 458. 1849. Watson, Garden 55:74, fig. 1899. Hemsl., Gard. Chron. 3(45):338, t. 1909. Standi., Contr. U.S. Natl. Herb. 23:1078. 1924. Munz, Proc. Calif. Acad. Sci. IV. 25:20, pi. I, fig. 6. 1943. Standi, and Williams, Fieldiana, Bot. 24(7):534,^#. 85. 1963. Munz, N. Am. Flora II. 5:4. 1965. Chickering, Flowers of Guatemala 64, pi. 18. 1973. Fuchsia cordifolia Bentham, PI. Hartw. 74. 1841. type: Near summit of Volcan Santa Maria ( tk Xe- tuch"), ca. 3,000 m, Quezaltenango, Guatemala, Nov. 1839, Theodor Hartweg 528 (K Bentham Herb., holotype; photograph, MO; BM, CGE, G, K Hooker Herb., LE, OXF, P, U, W, isotypes). Harrison, Floric. Cab. and Florist's Mag. 9:241, pi. 1841. Lindl., Bot. Reg. 27, pi. 70. 1841. Hook., Icon. PI. 5:/. 450. 1842. Essig, Nat. Hort. Mag. 13:6, photo. 1934. Munz, Proc. Calif. 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia decidua Figure 2. Distribution of Fuchsia decidua, F. fulgens (sect. Ellobium) and F . jimenezii (sect. Jimenezia). Acad. Sci. IV. 25:21. 1943. Standi, and Williams, Fieldiana, Bot. 24(7):530. 1963. Munz, N. Am. Flora II. 5:5. 1965. Fuchsia intermedia Hemsley, Diag. PI. Nov. Mex. Centr. Amer. 1:14. 1878. type: Summit of Mt. Totontepec, 3,000 m, Oaxaca, Mexico, April 1839, Theodor Hartweg 460 (K, holotype; photo- graph, MO; BM, CGE, LE, OXF, P, W, isotypes). Standi., Contr. U.S. Natl. Herb. 23:1078. 1924. Few- to many-branched, soft- wooded shrub 0.5-2.5 m tall, terrestrial or oc- casionally epiphytic on trees. Branchlets 0.5-3 dm long, 1-3 mm thick, triangular to quadrangular, subglabrous to densely short-pilose or villous; older branches with smooth reddish bark, exfoliating freely with age, 0.5-3 m long and 8-40 mm thick. Leaves opposite or occasionally ternate, membranous, ovate to cordate, rounded to cordate at base, acute to acuminate at apex, 35-130 mm long, 20-75 mm wide, pale to dark green and subglabrous to pilose or villous above, lighter below or red tinged and similarly pubescent, especially along nerves; secondary veins 5-9 on either side of the midvein, margin dentate to serrate; petioles red- dish, pubescent, 12-80 mm long; stipules lance-filiform, 1-2 mm long, ca. 0.3 mm wide, deciduous. Flowers axillary and solitary in upper leafy nodes of the year's branches; pedicels spreading to drooping, slender, subglabrous to strigose, 35- 75 mm long; ovary narrowly cylindrical; floral tube narrowly to broadly cylindric, 20-46(-64) mm long, 4-9 mm wide and usually ventricose and laterally com- pressed at the base around the nectary, ± dilated in upper half, 5-15 mm wide at the rim, pubescent or with glandular hairs outside, subglabrous or loosely strieose inside in lower Vi: seoals lanceolate. 8-20 mm lone. 5-8 mm wide, spread- 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 215 Table 1. Comparison of sects. Ellob Hum, Fuchsia, and He >msleyella. Section Presence of Presence Petals of Tubers Type of Nectary 1 Epiphy- tism Distribution Ellobium Fuchsia Hemsleyella Yes 2 of 3 spp. Yes None No Most species Band Ring Band Yes No Yes Mexico, Central America Tropical Andes and Hispaniola Tropical Andes 1 Band nectary is smooth and fully adnate to the floral tube; ring nectary is annular and mostly free from the tube. Three of the approximately 60 species in sect. Fuchsia have nectaries adnate to the tube, but they are variously lobed. ing at anthesis; tube rose to bright red, sepals green with reddish base; petals olive green, ovate, 6-12 mm long, 4-8 mm wide, rounded to cordate at the base, subacuminate at the apex, erect; nectary a lustrous yellow band ca. 0.4 mm thick 4-8 10-20 6-14 mm long and inserted on the tube ca. 1 mm below the insertion of the petals; anthers oblong, yellow, 2-3 mm long, 1-2 mm wide; style glabrous, pale green, 40-76 mm long; stigma green, subconic, 2-3 mm long, 1-2 mm wide, 4-parted at the apex. Berry elongate, 20-40 mm long, 5-8 mm thick, verrucose, green to dark purple when ripe; seeds 1.5-2 mm long, ca. 0.5 mm thick. Gametic chro- mosome number, n = 1 1 . Distribution: Mexico to Costa Rica. Scattered in a few moist forest localities in Guerrero and Oaxaca, more common in cloud forest and moist oak-pine forest from Chiapas to Costa Rica, 2,000-3,400 m (Fig. 1). Flowering throughout the year. Representative specimens examined: Mexico, chiapas: San Cristobal las Casas, NE side of Zonte- huitz, Breedlove 7799 (BM, DS, GH); ca. 2 km W of Nabenchauck, municipio Zinacantan, Breedlove 7493 (DS, GH); N & W slopes of Cerro Mozotal, road from Huixtla to Siltepec, Breedlove 25858 (MO); Cerro Huitepec, Ghiesbreght 698 (GH, K, LE, MO); Chamula, Linden 675 (G, K, LE); La Cola del Diablo, ca. 20 km N of San Cristobal Las Casas, Luteyn 3566 (DUKE); Mt. Tacana, Matuda 2322 (GH, NA, K); Motozintla, Pinabeto, Matuda 15466 (F, IPN, LL). Guerrero: Pie de la Cuesta-Toro Muerto, Hinton et al. 11222 (GH, K, NY, US); Cerro Teotepec, Rzedowski & McVaugh 184 (DS, ENCB, MICH, TEX); Tlacotepec, Paray 2012 (ENCB); Piedra Ancha, Ga- leana district, Hinton et al. 14223 (GH, MICH, NY, US). Guatemala, chimaltenango: Chichoy Pass, Hunnewell 14675 (GH); Volcan Acatenango, Hunnewell 14766 (GH); Cerro de Tecpam, Standley 61000 (A), el progreso: between Finca Piamonte and Volcan Santa Luisa, Stey- ermark 43567 (BH). huehuetenango: Cerro Pixpix, above San Idelfonso Ixtahuacan, Steyermark 50642 (BH); Cruz de Limon, between San Mateo Ixtatan and Nuca, Steyermark 49814 (BH); San Juan Ixcoy, Cordillera de los Cuchumatanes, Steyermark 50046 (BH). jalapa: Montana Miramundo, Steyermark 32833 (F). jutiapa: Asuncion, Mita, Volcan Suchitan, Steyermark 31944 (F). quezalte- nango: Cerro Quemado, Kellerman 5935 (LL, US); Cuesta de El Caracol, 5-8 km N of San Juan Ostuncalco, Williams et al. 22777 (G, S, W); Volcan Santa Maria, Skutch 867 (A, BM, G), Skinner s.n. (K); Volcan Zunil, Steyermark 34686 (F, US), quiche: Chiul, Heyde & Lux 1890 (GH, MO); above Nebaj toward Acatenango, Hunnewell 14776 (GH). sacatepequez: Volcan de Agua, Donnell Smith 2174 (F, GH, M, NY, PH). san marcos: San Luis, ca. 6 km W of Ixchiguan, Beaman 3243 (DUKE); San Sebastian, Volcan Tajulmulco, Steyermark 35844 (F, NY). solalA: Volcan San Pedro, N slopes, Steyermark 47260 (BH); Volcan Atitlan, Steyermark 47526 (F, G); ca. 18 km SE of Totonica- pan, Webster et al. 11760 (MO); Cerro Metia Tecum, 5-10 km S of Los Encuentros, Williams et al. 41729 (BM, MICH), totonicapan: Between Los Encuentros and Totonicapan, Bunting 1302 (F); ca. 13 km S of Totonicapan, Williams et al. 22909 (G). El Salvador, chalatenango: summit of 2 1 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN | Vol. 69 Los Esemiles, Tucker 1116 (BH, G, LL, MICH, NY, PH). santa ana: Cerro Montecristo, Allen 7154 (F, GH, LL, NY, US). Costa Rica, cartago: near Ojo de Agua camp, Dayton 3029 (F, MO); Villa Mills, Holm & litis 516 (A, BM, F, G, MO, NY), Raven 22051 (DS); Cerro Sakira at La Asuncion, Wilbur & Stone 10036 (DUKE), san jose: Cerro de la Muerte, Burch 4716 (DUKE, MO); trail from Canaan to Chirripo via Los Angeles, N of Rio Talari, Burger & Liesner 7396 (F, S); Cordillera de Talamanca, Pacific slopes of Chirripo massif, Davidse & Pohl 1619 (MO); Cerro de la Vueltas, Pittier 10501 (BR, G, US); 5 km SE of La Asuncion, before La Georgina, Wilbur 14386 (DUKE, MO); 9 km NW of La Asuncion, Wilbur 14407 (DUKE). Cultivated specimens: Austria, Vienna, 1903 (W). Ecuador, pinchincha: Hacienda Cataguango, W of Amaguana, 1976, Ortiz s.n. (QCA). Germany, Berlin, 1905, Bothe s.n. (B). Guatemala, Guatemala City, 1947, Brenckle 47-431 (NY). Ireland, Dublin Botanical Garden, 1844 (K). Soviet Union, Leningrad, 1845 (LE). United States, California: Berkeley, Hutchison 54.522. (UC). This species is unique in its elongate ovary and fruits, green petals, and ven- tricose, laterally compressed base of the floral tube. The peculiar compression of the floral tube has been well illustrated in horticultural journals; e.g., by Planchon (1849) and Hemsley (1909). Fuchsia splendens has the widest range in sect. Ellobium, from Costa Rica to Guerrero, Mexico. Variation in leaf shape is considerable, but it is mostly related to the degree of development of the leaf. On the same individual, the young upper leaves are often ovate while the lower ones are cordate. Pubescence varies to a much greater extent, with densely villous hairs on Linden 657 (G, K, LE; from Chiapas); glandular hairs on Breedlove 9347 (BM, GH; also from Chia- pas); finely pubescent hairs or even subglabrous plants in most Costa Rican col- lections. The most variable character of all is floral tube dimensions. Differences in floral tube lengths were used to describe F. cordifolia, longer-tubed than F. splendens, and F. intermedia, of intermediate length. Field studies and analysis of herbarium specimens have shown no geographic or ecological pattern to this variation, however. The type of F. cordifolia has floral tubes ca. 40 mm long; the type of F. splendens has floral tubes ca. 25 mm long. This does not nearly cover the variation found, however, since Skutch 867 (A, BM, G), from Volcan Santa Maria in Guatemala, has floral tubes to 64 mm long. Skinner s.n. (K) is from the same mountain at a similar elevation, yet it has floral tubes only 20 mm long. In view of this variation and the presence, throughout the range, of the common assemblage of unusual characters mentioned above, we are including both F. cordifolia and F. intermedia as synonyms of F. splendens. Diploid gametic chromosome counts of n = 1 1 were obtained from Breedlove 7796 (DS) and Raven 22051 (DS). Both Bentham (1841, p. 74) and Standley & Williams (1963) noted that inhabitants of the Volcan Santa Maria area called this species "Melocotoncito" (little peach). Hunters in that area would commonly rely on the conspicuous, epiphytic bushes of F. splendens for their sour, juicy fruits when water was not available. 2. Fuchsia fulgens De Candolle, Prodr. 3:39. 1828. type: Plate 362 of Sesse & Mocino's Flora Mexicana Icones, ined. (Copy at G). The collection of illus- trations made in Mexico by the Exoedicion Real de Botanica was lent bv Mocino to A. De Candolle in iolle in 1813. These were copied by De Candolle in requested their return in 1817 on very short notice (Sprague, 1926). De Candolle later went on to describe some 270 new species Mocino 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 217 from these illustrations, including F. fulgens. Plate 362 undoubtedly corre- sponds to the specimens collected by Sesse et al. and used to describe F. racemosa Sesse & Mocino (= F. fulgens); the type locality for F. fulgens is therefore the same as for F. racemosa (in the mountains of Patzcuaro, Mi- choacan, Mexico). Lindl., Bot. Reg. 24:/. /. 1838. Curtis, Bot. Mag. 67:f. 3801. 1840. DC, Caiques Dessins Mocino Sesse \\pl. 362. 1874. Raimann in Engl. & Prantl, Nat. Pfl. 3(7): 200,^. 1893. Watson, Garden 55:1 5, fig. 1899. Standi., Contr. U.S. Natl. Herb. 23:1078. 1924. Essig, Nat. Hort. Mag. 13:9, photo. 1934. Munz, Proc. Calif. Acad. Sci. IV. 25:55, pi. 8, fig. 43. 1943. Chittenden, Diet. Gard. 2:846,^. 1951. Munz, N. Am. Flora II. 5:6. 1965. Morley & Everard, Wild Flowers of the World,/?/. 173 , fig. Fl. 1970. Spachia fulgens (DC.) Lilja, Tidning Tradgardsskotsel allman Wextkultur 8:62. 1840. Ellobium fulgens (DC.) Lilja, Linnaea 15:262. 1841. Fuchsia fulgens pumila Carriere, Rev. Hort. 53:150, 1881. type: Rev. Hort. 53: 150, plate, 1881. No specimens of this entity were seen. The plant illustrated is from garden material of unspecified origin, cultivated in France ca. 1881. Fuchsia racemosa Sesse & Mocino, PI. Nov. Hisp. 58. 1888; non Lamarck, 1788. type: On boulders in the mountains of Patzcuaro, Michoacan, Mexico, Sept. 1790, Martin Sesse, Jose Mocino, Juan Diego del Castillo & Jose Maldonado 521 1 (MA, lectotype here designated. A second sheet of F. fulgens at MA has a Ruiz & Pavon label, but it is probably a duplicate of the type material). Fl. Mex. 101. 1893. Soft- wooded shrubs 0.5-3 m tall, with thickened, tuberous underground parts, often epiphytic in trees or on rocks. Branchlets 4-25 cm long, 1-5 cm thick, semi- succulent, quadrangular, subglabrous to strigose, reddish; older branches and stems freely exfoliating, 1-6 cm thick. Leaves opposite, soft-membranous, ovate to cordate, rounded to cordate at base, acute at apex, (5— )9— 15 cm long, (3-)5- 12 cm wide, pale green above, lighter below and red-tinged; young leaves canes- cent to tomentose, the older ones short villous; secondary veins 8-10 on either side of the midvein, margin denticulate to serrulate with red glandular teeth; petiole stout, strigose, 3-8 cm long; stipules subtuberculate, lance-deltoid, 1.2-2 mm long, 0.2-0.5 mm wide, deciduous. Flowers several to many in terminal, drooping racemes, the rachis 3-20 cm long; bracts reflexed, ovate to elliptic, 0.5- 2.5 cm long; pedicels slender, strigose, 10-20 mm long; ovary cylindrical, 12-15 mm long, 3-4.5 mm thick, strigose; floral tube narrowly funnelform, 50-65 mm long, 2-4 mm wide at the base, gradually widened above until 7-10 mm wide at the rim, loosely villous outside and inside; sepals lanceolate, subacuminate, 12- 17 mm long, 4—6 mm wide, spreading or suberect at anthesis; tube pink to dull red, sepals pale red and yellow green towards the apex; petals bright red, elliptic to ovate, 6-9 mm long, 4—6 mm wide, rounded at base, acute at apex, erect; nectary a smooth yellow green band 0.3-0.4 mm thick lining the basal 3-5 mm of the tube; filaments light red, the antisepalous ones 9-10 mm long, the antipe- talous ones 5-7 mm long; anthers white, 2.5-3 mm long, 1-2 mm thick; style slender, pink, pubescent in lower Vi, 60-82 mm long; stigma green, subconic, 2- 3 mm long, ca. 1.5 mm thick. Berry elongate, 20-30 mm long, 8-15 mm thick, glabrous to sparsely strigillose, dark purple; seeds 1-1.5 mm long, ca. 1 mm thick, ovate to subtriangular in outline. Gametic chromosome number, n = 11. Distribution: Mexico. Mostly epiphytic, in oaks or on rocks, especially around 2 1 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 seepages and waterfalls along the Trans-Mexican Volcanic Belt from Jalisco to the State of Mexico and in the Sierra Madre Occidental of Guerrero, 1,450-2,300 m (Fig. 2). Flowering in the wet season, June through September. Representative specimens examined: Mexico, Guerrero: Omiltemi, 60 km W of Chilpancingo, Row- ell 3047 (MICH), Tillett 637-150 (DS, GH). jalisco: Huejotitan, Diguet s.n. (S); E slopes of Volcan Colima, Goldsmith 22 (DS, F, GH, MO, NY, UC, US); Zapototan de Hildalgo, S of Guadalajara, Gregory & Eiten 230 (MICH, MO); San Juan Cosala, Villarreal de Puga 28 (IPN); 10-15 km S of Autlan," Wilbur & Wilbur 1438 (DUKE); E of Manantlan, ca. 22 km S-SE of Autlan, Wilbur & Wilbur 1816 (DUKE). Mexico: Amanalco, Valle de Bravo, Dressier 2433 (US); Tequexquipan, Temascal- tepec District, Hinton 823 (BM, K); Rincon del Carmen, Temascaltepec District, Hinton 1744 (BM, G, GH, K); Salitre-Canitas, Hinton et al. 4311 (BM, G, K); Comunidad, Temascaltepec District, Hinton 4894 (A, BM, K); Ipericones, Temascaltepec District, Hinton 3893 (K), Hinton et al. 8071 (DS, GH, K, MICH, NY, U); Nanchititla, Matuda et al. 30368 (GH, MEXU). michoacan: Km 16 of Uruapan-Los Reyes road, Beaman 2418 (GH); 26 km E of Morelia, Breedlove 7726 (DS); NW of Uruapan at San Juan Nuevo, Breedlove 15776 (BM, DS); near Morelia, Hartweg 286 (CGE, G, K, LE, OXF, W); Zitacuaro, Hinton et al. 11883 (GH, K); S of Torricillas, Coalcoman District, Hinton 13987 (H, NA, PH); S of Naranjillo, Hinton et al. 13952 (PH); Tancftaro, Leavenworth 303 (F, GH, MICH, MO, NY); Paracho, km 47-48 of Guadalajara-Uruapan road, Moore & Wood 4050 (A, BH, GM, DUKE, MICH); Patzcuaro, Pringle 4123 (BH, BM, G, GH, K, LE, MO, PH, S, W, Z); Taras- con, Pringle I1988M (BH, GH, K). Cultivated specimens: Austria, Vienna, 1849 (W). France, Nantes, 1839 (BM); Paris, 1843 (P). Germany, Munich, 1843 (BR); Heidelberg, 1923 (BH); Berlin, 1966, R196 (B). Great Britain, 1836- 1843, Saul s.n. (NA). Mexico, Veracruz: Jalapa, 1881, Kerber s.n. (W). Soviet Union, Leningrad, 1840 (LE). United States, California: Berkeley, 1965, Hutchinson 49.803.1 (UC, NA); Santa Barbara, 1910, Popenoe 618 (NA). The long-tubed, many-flowered racemes of Fuchsia fulgens resemble those of South American species such as F. boliviana and F. dependens. It is easily distinguished from these species, however, by the presence of tubers, the occa- sionally epiphytic habit, the cordate leaves, and the greenish sepals and stigma. It is sympatric with F. decidua in Jalisco and Guerrero, but F. decidua flowers when leafless in the dry season (December-May), and F. fulgens flowers with leaves in the rainy season (June-November). Both Hinton 3893 (K) and Hinton et al. II 883 (GH, K) were collected in May and are unusual in having just a few young expanding leaves at the shoot apices and flowers borne on dense racemes from the axils of older leafless branches, much like F. decidua. Diploid gametic chromosome counts of n = 1 1 were obtained from Breedlove 15776 (BM, DS) and Hutchison 49-803 (UC). 3. Fuchsia decidua Standley, Publ. Field Mus. Bot. 4:248. 1929. type: La Bufa, Real Alto, Sierra Madre Occidental, 2,500 m, Jalisco, Mexico, 30 January 1927, Ynes Mexia 1601 (F 579815, holotype; A, BM, C, CAS, DS, G, MICH, MO, NY, UC, US, Z, isotypes). Munz, Proc. Calif. Acad. Sci. IV. 25:75, pi. 12, fig. 75. 1943, N. Am. Flora II. 5:7. 1965.— Fig. 3. Shrub 0.5-2 m high, terrestrial, in rock crevices, or epiphytic in trees as high as 10 m above the ground, with fleshy tubers 2-4 cm thick and stoloniferous lateral shoots. Branchlets 5-15 cm long, 3-6 mm thick, subquadrangular, gla- brous; older branches 0.5-ca. 3 m long, 6-20 mm thick, with very loosely peeling, copper brown bark. Leaves opposite, chartaceous, elliptic to ovate-cordate, cu- neate to cordate at base, acute to acuminate at apex, 9-17 cm long, 5-8 cm wide, sparsely strigillose above, glabrous below; margin subentire to denticulate; de- ciduous, leafy in the wet season (June-November); petiole glabrous to strigillose, 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 219 d 3 cm Figure 3. Fuchsia decidua Standley. — a. Inflorescence. — b. Flower, longitudinal section, a & b from McVaugh 26157 (MICH), Jalisco, Mexico. — c. Leaves. From MacDougall H.52I (MO), Oa- xaca, Mexico. — d. Tuber. From Mexia 1061 (A), Jalisco, Mexico. 3-5 cm long; stipules triangular, 1-1.5 mm long, ca. 0.5 mm wide, deciduous. Flowers numerous in compact, lateral panicles on new shoots arising from the nodes of older stems, the rachis 2-6 cm long, drooping, flowering when leafless (December-May); pedicels slender, glabrous, 6-11 mm long, to 15 mm in fruit; 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 floral tube narrowly funnelform, 30-48 mm long, 1.5-3.5 mm wide at base, then slightly constricted above nectary and gradually widened above until 5-9 mm wide at rim, glabrous to sparsely strigose outside, glabrous within; sepals lance- oblong, 8-13 mm long, 3-6 mm wide, apex obtuse, spreading to suberect at anthesis; tube and sepals reddish pink; petals red, fleshy, oval to reniform, 1.5- 3.5 mm long, 1.5-4 mm wide, erect; nectary a smooth yellow green band 3-5 mm high lining the base of the tube; antisepalous filaments 10-17 mm long, antipetal- ous filaments 5-12 mm long; anthers 1.5-3 mm long, 1-2 mm wide; style slender, 55-66 mm long; stigma green, slender, 1-1.5 mm long, ca. 1 mm thick. Berry ellipsoid, 8-12 mm long, 4-6 mm thick, glabrous, greenish purple; seeds 1.5-2 mm long, 0.5-1.0 mm thick, ovate in outline. Gametic chromosome number, n = 11. Me Madre May) Specimens examined: Mexico, Guerrero: Omiltemi, ca. 60 km W of Chilpancingo, Breedlove 1582 1 (DS, LL), Lachica s.n. (IPN); W of Filo de Caballo, Breedlove 15829 (DS); Highway 95 ca. 6 km W of turnoff to Chichihualco, Croat 45630 (MO); Cerro Alquitran, Kruse s.n. (DS); Chichihualco, Camotla, Cerro de la Pastilla, Rzedowski 16435 (IPN, MICH); Campamento EI Gallo, Rzedowski & McVaugh 189 (DS, MICH), jalisco: Sierra de Manantlan, above Durazno, Boutin 3036 (MO); Mesa de los Gallos, Gonzalez 843 (MO); SSE of Autlan, SE of Ahuacapan, McVaugh & Koelz 918 (MICH); SE of Autlan, trail from Chante, above Rancho Manantlan, McVaugh 10295 (MICH); near Santa Monica, McVaugh 14047 (MICH); Autlan, Sierra de Manantlan, between El Chante and Cuzalpa, McVaugh 23100 (IPN, MICH); Ayutla, San Miguel de la Sierra, McVaugh 23500 (MICH); Sierra de Manantlan, 25 km S of El Chante, McVaugh 26157 (MICH), oaxaca: S of Tlaxiaco, San Andres Chichahuastla, Cerro Zarzamora, MacDougall H.52I (MO), s.n. (US). Cultivated specimens: Mexico, distrito federal: Mexico City, Kruse in 1967 (DS). oaxaca: Oa- xaca, MacDougall 468 (US). The tiny, thick petals and leafless, dry-season flowering distinguish Fuchsia decidua from the related F.fulgens and F. splendens. The habit of F. decidua is unusual and is shared only by some members of the South American sect. Hemsleyella. Fuchsia decidua grows epiphytically, mostly on the trunks of oaks, but also terrestrially on rocks. Plants with thick, fleshy, prostrate stems ca. 2 cm thick extending up to nearly 3 m long are rooted in moss and rich humus (F. Boutin, T. MacDougall, pers. comm.). This species is well adapted to the rather seasonal climate of the areas in which it occurs. The tubers store food and water during the wet season, when the plants are leafy and actively growing vegeta- tively. In the dry season, the plants are deciduous and flower on stems from the previous season's growth. The seeds are apparently spread by birds from tree to tree and from cliff to cliff. A single somatic chromosome count of 2n = 22 was obtained from Breedlove 15821 (DS). Fuchsia sect. Jimenezia, Breedlove, Berry & Raven, sect. nov. type: Fuchsia jimenezii Breedlove, Berry & Raven. Hermaphroditicus. Folia opposita ellipticaque. Flores racemosi vel interdum paniculati; necta- rium disco annulari styli basin cingens et basin tubi adnatum; stamina inaequalia, filamentis antisepalis erectis et supra tubum exsertis, antipetalis in tubum reflexis inclusisque. Bacca subglobosa, seminibus multis, compressis. 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 22 1 Hermaphroditic. Glabrous shrubs. Leaves opposite, elliptic to lance-elliptic. Flowers in terminal racemes or few-branched panicles. Floral tubes less than 5 mm long, shorter than sepals or petals. Nectary annular, irregularly 4-lobed, adnate to the tube. Stamens biseriate, the antisepalous stamens erect, the anti- petalous stamens reflexed and included in the tube. Berry subglobose; seeds many, ca. 50-ca. 100, laterally compressed, 0.9-1.2 mm long, 0.5-0.7 mm wide. Gametic chromosome number, n = 1 1 . 4. Fuchsia jimenezii Breedlove, Berry & Raven, sp. nov. — type: Monte verde, exposed crest of the Sierra de Tilaran, Puntarenas, Costa Rica, 23 June, 1967, Thomas Emmel (DS 614763, holotype; BM, CAS, ENCB, F, GH, K, LE, LL, MICH, NT, RSA, US, isotypes). This species is dedicated to Alfonso Jimenez Munoz, a long time student of the Costa Rican flora, who first called this entity to our attention as an undescribed species in 1967. Fig. 4. Fuchsia arborescens sensu Woodson & Schery, Ann. Missouri Bot. Gard. 46:328. 1959, pro parte. Frutex erectus vel scandens 0.4-2.0 m altus, omnino glaber; rami ad 3 cm crassi cortice subtilitei fissurato. Folia opposita, subcoriacea, elliptica vel lanceo-elliptica, (3-)6.5-l 1.5 cm longa, 2.5-4.8 cm lata. Flores racemosi vel interdum paniculati, racemo 5-20 cm longo; pedicelli erecti vel divergentes, tenues, 6-12 mm longi, fructu ad 18 mm longo; tubus floralis subcylindricus vel subobconicus, 2.5- 4.5 mm longus, apice 2-4.5 mm latus, basi 1.5-2.5 mm latus; sepala late lanceolata, 4-6 mm longa, basi 2.5-3.5 mm lata, divergentes; tubus sepalaque rubra vel roseo-rubra; petala rosea, suborbicularia vel ovata, 4-6 mm longa, 3-4 mm lata, basi rotundata, apice rotundata vel acuta, erecta; discus annularis lobis plerumque quattuor vel pluribus supra basin tubi 1-1.5 mm elevatus; filamenta anti- sepala 1-2 mm longa erectaque, antipetala 0.8-1.5 mm longa, in tubum reflexa inclusaque, antherae 1.5-2 mm longae, 0.9-1.2 mm crassae; stylus crassus 6-8 mm longus stigmate ca. 1 mm longo, ca. 1 mm crasso. Bacca subglobosa, 10-12 mm longa, in maturitate ca. 10 mm crassa, rosea ad atrorubra, seminibus multis 0.9-1.2 mm longis, 0.5-0.7 mm crassis, in ambitu oblongibus vel subtriangularibus. Numerus gameticus chromosomatum, n =11. Scandent subshrub 0.5-I.5(-2) m tall, glabrous throughout. Branchlets 1-5 dm long, 1-3 mm thick, quadrangular; older branches terete, with pale tan, finely fissured bark. Leaves opposite, subcoriaceous, elliptic to lance-elliptic, acute at base, acute to acuminate at apex, (3— )5— 1 1.5 cm long, (l-)2.5-5 cm wide, dark green above, flushed purple beneath; secondary veins 10-12 on either side of the midvein, margin subentire to denticulate with small, glandular teeth; petiole stout, 1-2 mm thick, red, 4-9 mm long; stipules semisucculent, deltoid, dark when dry, 0.8-1.0 mm long and wide, often connate, deciduous. Flowers numerous in ter- minal racemes, rarely axillary, or occasionally in 2-3-branched panicles; rachis 4-6 5-4 5-4 4-6 4-6 mm lone. 3-4 rounded at the base, rounded to acute at the apex; nectary an irregularly to 4-lobed ring-shaped disc adnate to the base of the floral tube, 1-1.5 mm high; filaments dull red, the antisepalous filaments erect and exserted above the rim of the tube, 1-1.5 mm long, the antipetalous filaments reflexed and included in the tube, 0.8-1.4 mm long; anthers white, 1.5-2.0 mm long, 0.9-1.2 mm thick; style 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 9 mm Figure 4. Fuchsia jimenezii Breedlove, Berry & Raven. — a. Habit; rachis and flowers are divergent to pendant in life. — b. Antipetalous anther. — c. Flower, longitudinal section. From Wilbur 14238 (MO), Monteverde, Puntarenas, Costa Rica. stout, 6-8 mm long, the stigma capitate and obscurely 4-cleft at the apex, 1-1.5 mm long, ca. 1 mm thick. Berry subglobose, 10-12 mm long, ca. 10 mm thick when ripe, light to dark red, lustrous; seeds reddish, 0.9-1.2 mm long, 0.5-0.7 mm thick. Gametic chromosome number, n = 11. Distribution: Panama and Costa Rica. Scattered in mostly secondary vege- tation of wet evergreen cloud forests from Chiriqui Province of northern Panama 1982] BREEDLOVE ET AL— MEXICAN AND CENTRAL AMERICA FUCHSIA 223 to the northernmost extension of Puntarenas Province of Costa Rica, 1,500-1,900 m (Fig. 2). Flowering throughout the year. Specimens examined: Costa Rica, cartago: Ca. 10 km S of Tapanti on road above Rio Grande de Orosi, Burger & Burger 7561 (MO); Tapanti Reserve, Rio Dos Amigos, Croat 36223 (MO); 8.5 km S of Tapanti, Lent 1372 (BM, F, MO), heredia: Rio Vueltas, E slope of Volcan Barba, Burger & Liesner 6438 (F, GH); Vara Blanca, Chrysler 5016 (F); Alto de los Robles, San Rafael, Jimenez 2533 (F); Rio Patria, SE slopes of Volcan Barba, Jimenez 2300 (F); vicinity of Cerro Chompipe between Rio la Vueltas and Quebrada Cabru, Luteyn & Wilbur 4441 (CAS, DUKE); San Isidro, Cerro de las Lajas, Standley & Valerio 51567 (US); 12 km NE of San Rafael, Wilbur et al. 15981 (DUKE); vicinity of Cerro Chompipe, between Rio Las Vueltas and Rio Nuevo, Wilbur & Luteyn 18566 (DUKE, MICH, MO), puntarenas: Monteverde, Sierra de Tilaran, Almeda et al. 2022 (DUKE, MO), Burger & Baker 9746 (U), Burger & Gentry 8759 (F), Haber 434 (MO), Lorence & Pierce 1722A (MO), Palmer 55 (CR), 75 (CR), 171 (CR), Solomon 5370 (MO), Wilbur 14238 (DUKE, MO), san jose Vicinity of Cascajal, 3-6 km beyond Las Nubes, Almeda et al. 2360 (MO); Alto La Palma, above Rio Hondura, Lent 1341 (DUKE, F, S); near Quebrada Grande, 3 km NW of Cascajal, Lent 2310 (F); La Hondura, Standley 36587 (US). Panama, bocas del toro: Robalo trail, N slopes of Cerro Horqueta, Allen 4965 (G, MO, RSA): E slope of La Zorra to Divide, Kirkbride 831 (MO). chiriquI: Bajo Mono-Robalo trail, W slopes of Cerro Horqueta, Allen 4816 (G, RSA); trail from Rio Palo Alto road, near peak of Pate Macho, Hammel 5785 (MO); Cerro Colorado, ca. 32 km from Rio San Felix, Sullivan 310 (MO), 317 (MO); near Bajo Chorro, Woodson & Schery 700 (MO). This species is remarkable in being totally glabrous. The antipetalous stamens reflexed down into the floral tube is found elsewhere in the genus only in sect. Encliandra, which differs strongly in its subdioecious to dioecious, axillary flow- ers, strongly four-lobed stigmas, and few-seeded berries. A gametic chromosome number of n Haber Fuchsia sect. Schufia (Spach) Munz, Proc. Calif. Acad. Sci. IV. 25:84. 1943. Schufia Spach, Hist. Nat. Veg. Phan. 4:41 1. 1835. type: Fuchsia arborescens Sims. Hermaphroditic, morph ives ODDOsite or 3-4 ve 10-25 and 9-20 cm wide. Floral tube, sepals, and petals nearly equal in length. Nectary annular, smooth to ±4-lobed, adnate to the base of the floral tube. Stamens biseriate, erect, the antisepalous stamens longer than the antipetalous stamens. Berry subglobose, with a waxy bloom when ripe; seeds ca. 50-ca. 100, laterally compressed, oval to irregularly triangular in outline, 1-1.5 mm long, 0.5-0.9 mm wide. Gametic chromosome number, n = 11. Fuchsia arborescens and F. paniculata have generally not been separated in the past. Fuchsia arborescens was first collected in Mexico in 1790 by Sesse and Mocino, but the name Fuchsia arbor ea Sesse & Mocino was not published until their manuscripts were rediscovered and published in 1888 (McVaugh, 1977). Sims (1825) first published a description and illustration of F. arborescens; his plate was dated December, 1825, preceding by just a month Lindley's publication of the same name (Lindley, 1826). Lemaire (1848) recognized that Central Amer- ican plants of this group had more ridged and serrate leaves than the typical F. arborescens from Mexico, and he consequently described his Guatemalan plants as a variety of that species. Subsequently, Lindley (1856) gave specific status to these serrate-leaved plants under the name F. paniculata. The male sterility and floral dimorphism prevalent in most populations of 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Fuchsia paniculata have provided another source of confusion in this group. Donnell Smith (1898) first recognized two distinct flower sizes in his Guatemalan collections and called the larger-flowered ones F. arborescens var. (?) megalan- tha (perfect-flowered plants). In their field studies of the Panamanian flora, Wood- son & Seibert (1937) realized that the different flower sizes were due to sexual differences and termed the species ^polygamodioecious/' Seibert noted that only the pistillate plants bore fruit and had smaller flowers but larger stigmas than "those plants which are apparently incapable of fructification" (Woodson & Sei- bert, 1937, p. 196). In his 1943 monograph of the genus, Munz failed to recognize the different sexual morphs and instead named three different forms of F. ar- borescens. As understood here, Fuchsia arborescens consists entirely of hermaphroditic plants and is quite uniform throughout its range. The leaves are larger than those of F. paniculata and have a smooth surface and entire margins. The flowers of F. arborescens are broader in bud than those of either floral morph of F. pani- culata, and the anthers are oblong and larger than the ovoid-reniform anthers in perfect flowers of F. paniculata (Fig. 5). Also, the stigma of F. arborescens is always conspicuously four-lobed and well exserted above the anthers. Fuchsia arborescens and F. paniculata are allopatric except for a small over- lap of ranges in northern Oaxaca and in Veracruz. Some unusually pubescent collections occur in the area of overlap in Oaxaca. In F. paniculata, Jurgensen 530 (G) and Galeotti 3035 (W) are pilose, and short, dense pubescence is found on two collections referable to F. arborescens, Smith 615 (GH) and Karwinsky: 46 (G). This last specimen has large leaves with serrulate margins, but the flowers and stamens agree much more with those of F. arborescens than with those of F. paniculata. The general appearance of these fuchsias strongly suggests that of certain Rubiaceae. In northern Panama, F. paniculata is sympatric with Psychotria an- gustifolia HBK, and the two can only be distinguished upon close examination. They probably share common pollinators. ? 5. Fuchsia arborescens Sims, Bot. Mag. pi. 2620, 1825. Lindl., Bot. Reg. pi. 943. 1826. — type: Plate 2620 in Sims, Bot. Mag. 1825, lectotype, here desig- nated, from cultivated material grown from seed sent in 1823 by George Bullock, who obtained the seeds from plants in cultivation at the Botanic Garden in Mexico City. The Mexican plants were probably sown by Martin Sesse and are likely progeny of the type specimen of Fuchsia arborea Sesse & Mocino. Schufia arborescens (Sims) Spach, Hist. Nat. Veg. Phan. 4:411. 1835. Fuchsia arborescens var. typica Munz, Proc. Calif. Acad. Sci. IV. 25: 85, pi 14, fig. 75. 1943. Fig. 5a. Fuchsia arborea Sesse & Mocino, PI. Nov. Hisp. 58. 1888. type: Uruapan, Michoacan, Mexico, Sept. 1790, Martin Sesse, Jose Mocino, Juan Diego del Castillo & Jose Maldonado 5216 (F 845520, lectotype here designated; MA, possible isolectotype; the one specimen of F. arborescens at MA has a Ruiz & Pavon label and is annotated as F. paniculata, but it is most probably mislabelled from a Sesse et al. collection. The sheets at MA and F appear to have been taken from the same plant). Hermaphroditic. Erect, woody shrubs or small trees 3-8 m tall, mostly glabrous throughout. Branchlets ascending, 1-4 dm long, 2-5 mm thick, subtriangular to 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 225 . '.' u • III > i 5 mm . c d Figure 5. a. Fuchsia arborescens Sims. Bud and flower, longitudinal section. From Feddema 2872 (MO), Guerrero, Mexico, b-d. Fuchsia paniculata Lindley. — b. Bud and perfect flower, longi- tudinal section; from a gynodioecious population. From Breedlove 42742 (MO), Chiapas, Mexico, c. Pistillate flower, longitudinal section. From Raven 20969 (DS), Cartago, Costa Rica. — d. Perfect flower, longitudinal section, from a hermaphroditic population. From Ventura 3631 (MO), Veracruz, Mexico. quadrangular; older branches and main trunk 5-25 cm thick. Leaves opposite or 3-4-verticillate, firmly membranous to subcoriaceous, smooth, elliptic to oblan- ceolate or obovate, acute to narrowly cuneate at base, acute to acuminate at apex, 10-21 cm long, 4-8 cm wide, deep lustrous green above, paler below; margin entire; petiole 12-30 mm long; stipules broadly triangular, occasionally connate, 0.8-1.2 mm long, 1.0-1.5 mm wide, deciduous. Flowers erect and nu- 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 merous in terminal, di- to trichotomously branched panicles, 10-25 cm long, 9- 20 cm wide; pedicels erect, slender, 9-18 mm long; floral tube subcylindric to obconic, 3.5-6 mm long, 1-1.5 mm wide at base, ± enlarged around nectary, 2- 4 mm wide at rim, usually glabrous inside and outside; flowers widened toward apex in bud; sepals oblong to lanceolate, acuminate, 4-1 1 mm long, 1.5-2.5 mm wide, spreading to reflexed at anthesis; tube and sepals rose-purple; petals lav- ender, lance-oblong to elliptic, 4-9 mm long, 1.5-3 mm wide, broadly acute at base, acute at apex, erect to spreading; nectary a smooth and irregularly lobed ring 1-1.6 mm high adnate to the base of the tube; antisepalous filaments 5-12 mm long, antipetalous ones 3-9 mm long, pink-purple; anthers oblong, 1.5-2.3 mm long, 1-1.5 mm thick; style 9-17 mm long, glabrous, pink, the stigma always exserted above the anthers, 1.5-3 mm wide, with 4 blunt lobes 1-2 mm long, lavender. Berry subglobose, 8-12 mm long, 7-10 mm thick, purple with a glau- cous, waxy bloom when ripe; seeds 1-1.5 mm long, 0.5-0.8 mm thick. Gametic chromosome number, n = 11. Distribution: Mexico. Occasional small trees mostly in barranca vegetation (moist ravines) of pine-oak forest in the Sierra Madre Occidental from Durango to Oaxaca and through the Trans-Mexican Volcanic Belt to Veracruz and then into northern Oaxaca, 1,750-2,500 m (Fig. 6). Flowering mostly in the dry season (December-May), but also well into the wet season. Representative specimens examined: Mexico, durango: El Palmito, Breedlove & Gregory 14250 (DS); ca. 8 km NE of Palmito, Gentry & Gilly 10629 (MICH). Guerrero: Filo del Caballo, Breedlove 15832 (DS); Cruz de Ocote, 50-60 km W of Chilpancingo, Feddema 2872 (DUKE, MICH, MO, PH); Campo Morado, Mina, Hinton et al. II 173 (GH); Galeana, Toro Muerto, Hinton 14213 (DS, F, GH, NA, NY, US); W of Cerro Azul, Mina, Hinton et al. 14952 (GH); Milpillas-Ayotac road, ca. 24 km SW of Filo de Caballo, Reveal et al. 4266 (MO); San Antonio-Buenos Aires, Montes de Oca, Hinton et al. 14013 (GH, MICH, NA, NY, U); W of Petlacala, Mina, Mexia 9052 (BH, G, GH, LL, MO, NY, S, U, UC); Omiltemi, W of Chilpancingo, Rzedowski 15992 (MICH), jalisco: W slopes of Sierra de San Sebastian, 15-30 km N of Mascota, Anderson & Anderson 5659 (DUKE, MICH); ca. 24 km SW of Chante, Sierra Manantlan, Gentry & Gentry 23514 (MICH); Sierra de Halo, McVaugh & Koelz 1236 (MICH); 15 km SE of Autlan, trail from Chante to Rancho Manantlan, McVaugh 10267 (DUKE, G, MICH, MO, TEX); Nevado de Colima, near Atenquique, McVaugh et al. 11770 (MICH, RSA). Mexico: Tequexquipan, Hinton 3520 (GH, MICH, NY, US); Rincon, Temascaltepec, Hinton et al. 5287 (GH, NY, US); Cumbre of Cimientos, Temascaltepec, Hinton et al. 8850 (GH); Cerro de Corona, Zacualpan, Matuda 30739 (MEXU). michoacan: Coalcoman, Barroloso, Hinton 15756 (DS, MICH, NY); ca. 28 km E of Morelia, King & Soderstrom 5020 (MICH, NY, TEX, UC, US); Falls of Tzararacua, S of Uruapan, hangman 3289 (N A, PH); Uruapan, Leavenworth & Hoogstraal 1257 (F); Sierra de Manantlan, McVaugh 13921 (MICH), morelos: Sierra de Morelos, Hinton et al. 17096 (MICH): Tepoxtlan, Miranda 517 (MEXU); Tlayacapan, Paray 2109 (MEXU); mountains above Cuernavaca, Pringle 6825 (BH, BM, BR, F, G, GH, K, LE, M, MICH, MO, NY, PH, POM, RSA, S, UC, VT, W, Z). oaxaca: Finca La Soledad, 184 km S of Oaxaca, Alexander 587 (MEXU, MICH, NY, UC, US); near San Juan, Arnold 3 (CAS); Coyula, Cuicatlan, Cancino 2438 (US); Highway 175, 6 km S of Suchixtepic, Croat 46026 (MO); Hacinda de San Luis, Gonzalez 426 (GH); Canada de los Molinos, valley of Oaxaca, Jurgensen 489 (BM, G); La Cumbre de los Molinos, Karwinski? 46 (G); between Juquila and Nopala, Nelson 2417 (US); Sierra de San Felipe, Pringle 6242 (A, BH, BM, BR, F, G, GH, LE, M, MO, NY, PH, POM, RSA, S, UC, US, VT, W, Z); 4 km S of Lachao, Oaxaca-Puerto Escondido, Rzedowski 19541 (MO), puebla: San Vicente, near Puebla, Nicolas in 1909 (G). Veracruz: Jalapa, Galeotti 3035 (BR, LE, NY); La Luz, near Cordoba, Kerher s.n. (G); El Esquilon, municipio Jilotepec, Ventura 4843 (MICH, MO). Cultivated specimens: France: 1826 (G). Great Britain: 1826, Lindley 943 (CGE); Edinburgh, 1964, Meyer 8211 (NA). India: Madras, 1850, Golley 389 (LE). Madeira: Santa Roque, Hilldebrand s.n. (Z). Mexico, Veracruz: Orizaba, 1885, Gray s.n. (GH). Netherlands: Baarn, 1955, Mennaga 4137 (BH, U). West Germany: Berlin, 1963, R181 (B). 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 227 Fuchsia arborescens 8,5 «W 20*N Figure 6. Distribution of Fuchsia arborescens and F. paniculata (sect. Schufia). Fuchsia arborescens is commonly found in barrancas, or deep moist ravines, mostly in the Sierra Madre Occidental of Mexico. In "La Vegetacion de Nueva Galicia," Rzedowski & McVaugh (1966) list F. arborescens as one of the low tree components of the "mesophytic mountain forest" formation, which occurs very discontinuously throughout the Sierra Madre and occupies less than 1% of the total vegetation cover for the area. This is the most moist formation of central, western Mexico, with rainfall estimated at 1,000-2,000 mm per year; the forma- tion is essentially restricted to the barrancas, where a higher humidity can be maintained than in adjacent oak-pine forest. Fuchsia fulgens and F. decidua are listed as epiphytes in the same formation. A diploid gametic chromosome count of ai = 11 was obtained from Breedlove 18713, A somatic count of In =22 was obtained by D. Breedlove from a culti- vated plant at Stanford University, California, grown from seeds collected at the University of Mexico Botanic Garden in Mexico City (no voucher). An additional diploid somatic count is cited by Kurabayashi et al. (1962) for Raven 14745 (RSA). 6. Fuchsia paniculata Lindley, Gard. Chron. 1856:301. 1856. type: Guatemala, 1855, George U. Skinner 48 (CGE lectotype, here designated; photograph, MO). — Figs. 5b, c, d. Fuchsia arborescens auct. non Sims, 1825: Essig, Nat. Hort. Mag. 13:2, photo, 3. 1934; Munz in 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Woodson & Schery, Ann. Missouri Bot. Gard. 45:218. 1959; Standi. & Williams, Fieldiana, Bot. 24(7):528. 1963. Fuchsia arborescens var. syringaeflora Lemaire, Fl. Serres Jard. Eur. 4\4\b, fig. 1848. type: Culti- vated in the garden of L. van Houtte, France, from seeds received in 1847 from Guatemala (K lectotype here designated); Sieb. & Voss. in Vilm., Blemengart, ed. 3, 1:332, /. 84. fig. 335. 1896. Fuchsia syringaeflora (Lem.) Carriere, Rev. Hort. 45:311, fig. 1873. Fuchsia arborescens var. (?) megalantha Donnell Smith, Bot. Gaz. 18:2. 1893. type: Volcan Aca- tenango, Dept. Sacatapequez, Guatemala, Mar. 1892, John Donnell Smith 4269 (US, holotype; GH, K, isotypes). Fuchsia leibmannii H. Leveille, Bull. Geogr. Bot. 22:24. 1912. type: Volcan Irazii, ca. 2,800 m, Prov. San Jose, Costa Rica, 1845-1848, A.S. Oersted 3297 (C, holotype; photograph, GH; G, isotype). Oersted, who collected in Costa Rica and Nicaragua from 1845 to 1848, and F. Liebmann, who collected in Mexico from 1840 to 1842, collaborated jointly on their collections upon their return to Europe. Leveille mistook the type for a Liebmann collection, hence the specific epithet and the locality cited in the protologue as "Mexique: mont Frasu" (= Irazii). Fuchsia arborescens forma tenuis Munz, Proc. Calif. Acad. Sci. IV. 25:86. 1943. type: Vara Blanca de Sarapiqui, between Volcan Poas and Volcan Barba, 1,740 m, Prov. Heredia, Costa Rica, Oct., 1937, Alexander Skutch 3357 (US, holotype; A, MO, NY, S, isotypes). Fuchsia arborescens forma parva Munz, Proc. Calif. Acad. Sci. IV. 25:86. 1943. type: Finca Pirineos, below Santa Maria de Jesus, Dept. Quezaltenango, Guatemala, Paul Standley 68287 (POM, holotype; F, isotype). Gynodioecious or subdioecious. Erect woody shrubs or small trees 3-8 m tall, usually glabrous. Branchlets ascending, 1-4 dm long, 2-5 m thick, subtriangular to quadrangular; older branches and main trunk 2-15 cm thick. Leaves opposite or 3^-verticillate, subcoriaceous, elliptic to oblanceolate, acute to narrowly cu- neate at base, acute to acuminate at apex, 5-15.5 cm long, 2-5.5 cm wide, deep lustrous green above, paler below; margin minutely to coarsely serrate; petiole 8-26 mm long; stipules triangular, often connate, 0.8-1.2 mm long, 0.7-1.5 mm wide, deciduous. Flowers erect and numerous in terminal, di- to trichotomously branched panicles, 10-14 cm long, 9-12 cm wide, narrowed towards the apex. Perfect flowers: Pedicels slender, 8-12 mm long; floral tube cylindric to subob- conic, 4-8 mm long, 1-2 mm wide at base, 1.5-3 mm wide at the rim, often pilose within; sepals lanceolate, 5-10 mm long, 1.1-2.5 mm wide, spreading to reflexed at anthesis; tube and sepals rose-purple; petals lavender, lanceolate to elliptic, 4-10 mm long, 1.1-3.5 mm wide, acute at both ends, erect to spreading; nectary smooth to irregularly lobed ring 1.3-2 mm high adnate to the base of the tube; antisepalous filaments 4-13 mm long, antipetalous ones 2-11 mm long, pink; anthers broadly ovoid to reniform, 0.8-1.6(-2) mm long, 0.7-1.4 mm thick: style sparsely pilose, either exserted beyond anthers with a shortly 4-lobed stigma or included below the anthers and the stigma strongly reduced, lavender. Pistillate flowers: Similar to the perfect ones except in the following characters: floral tube smaller, 3-5.5 mm long, 0.7-1.7 mm wide at base, 1.5-2.2 mm wide at rim; sepals 3.5-7 mm long, 0.7-1.3 mm wide; petals 2.3-4.5 mm long, 0.6-1.3 mm wide; nectary 0.8-1.4 mm high; antisepalous filaments 1.5-4.9 mm long, antipetalous ones 1.0-3.9 mm long; anthers abortive, 0.4-0.8 mm long, 0.2-0.6 mm thick; style stout, 7-1 1 mm long, the stigma well exserted above the stamens, 4-lobed, lobes 0.6-1.9 mm long. Berry subglobose, 4-9 mm long, 4-7 mm thick, purple with a glaucous, waxy bloom; seeds 1-1.4 mm long, 0.5-0.8 mm thick. Gametic chro- mosome number, n = 11. Distribution: Mexico to Panama. Frequent in moist oak-pine and evergreen 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 229 Mexico Puebla (Fig. 6). Flowering throughout the year. (80G-) 1,200-3 Representative specimens examined: Mexico, chiapas: Zinacantan, NW of Navenchauc, Breedlove 5971 (DS); Chamula, Breedlove 7145 (DS); Rayon, N of Pueblo Nuevo, near Puerto el Viento, Breed- love 10172 (DS, BM); Huistan, Breedlove 7343 (BM); N & W slopes of Cerro Mozotal, road Huixtla- El Porvenir, Breedlove & Smith 22793 (MO); Cerro Huitepec, W of San Cristobal las Casas, Breedlove 23007 (MO); 16 km NW of Rizo de Oro, SE of Cerro Baiil, Breedlove 24891 (MO); SE slopes of Volcan Tacana, above Talquian, Breedlove 29475 (MO); near summit of Chuchil Ton, NE of Bochil, Breedlove 34657 (MO); Laguna Pojo, near Lago Tsiskla, Breedlove 37087 (MO); logging road, Las Margaritas-Campo Alegre, Breedlove 41 114 (MO); near Niquivil, junction of ridge to Cerro Boqueron, Breedlove 42742 (MO); 4 km N of Colonia San Jose, ca. 12 km S of Layon, Hansen et al. 1688 (MICH); San Bartolo, Linden 39 (G); Mt. Ovando, Matuda 437 (F, MEXU, MICH, US); Siltepec, Matuda 812 (MEXU, MICH); Mt. Pasitar, Matuda 1695 (A, MEXU, MICH, NY); Volcan Tacana west, Matuda 2924 (A, LL, IPN, MICH); Mt. Pale, near Porvenir, Matuda 4670 (A); near Motozintla, Matuda 15447 (LL, MEXU, TEX); Cerro del Boqueron, Purpus 6970 (GH, MO, NY, UC, US); near Pueblo Nuevo Solistahuacan, Raven & Breedlove 20004 (DS, GH); Jitotal, Roe et al. 1182 (MICH). oaxaca: Cuyamecalco, Cuicatlan, Conzatti 3497 (MEXU); ca. 60 km N of Ixtlan de Juarez, road to Tuxtepec, Denton 1688 (MICH); Tonaguia, Galeotti 3035 (LE, W); Nolasco, Galeotti 3058K (F); Rt. 175 near Cerro Pelon, 34 km SW of Valle Nacional, Hill 1730 (MO); Monte Tepitongo, Sierra San Pedro Nolasco, Talea, Jurgensen 530 (G); Vista Hermosa to Comaltepec, Martinez 242 (MO); ca. km 125 of Tuxtepec-Oaxaca road, Smith & Tejeda 4497 (NA); ca. 16 km S of Villa Hermosa, Torke et al. 504 (MO), puebla: Road to Presa de Apulo, N of Oriental-Tezuitlan road, Koch & Fryxell 7716 (MO); NE of Tezuitlan, Pineda 686 (MICH); San Juan Abuacatlan, Salazar s.n. (MEXU); Colihuit, municipio Hueytamalco, Ventura 389 (MICH); Agua de Obispo, municipio Tezuitlan, Ventura 13641 (MO); ca. 8 km NE of Teziutlan, road to Tlapacoyan, Webster & Breckon 15463 (DUKE, GH). Vera- cruz: Orizaba, Botteri 58 (F, GH); Jalapa, Galeotti 3035 (G); Cerro de San Cristobal, W of Orizaba, Landman 3587 (PH); Mirador, Liebmann 3237 (BH, BR, G, GH, MO, MSC, UC, US); Huahesco, Mohr 1865 (US); Zacuapan, Tenampa, Purpus 4330 (A, BM, GH, MO, UC); Chiconquiaco, Canada del Huerfano, Rosas 580 (MO); Ahuacatitlan, municipio Jalacingo, Ventura 283 (MICH, MO); Lomas de Santa Rita, municipio Yecualta, Ventura 3631 (MO); Xico, Ventura 4519 (MICH); Tepezingo, Ventura 15051 (MO); Cuesta Grande de Jalacingo, Schiede & Deppe 526 (LE, W). Guatemala, alta verapaz: Chucaneb, Donnell Smith 182 (G, GH, PH); Tactic, von Tuerckheim 8395 (A, F, M, NY, US); Coban, von Tuerckheim 11713 (BR, C, F, FI, G, GH, LE, MO, US, W, Z). baja verapaz: Union Barrios, Contreras 11245 (MO); ca. 7 km NE of Purulha, Croat 4 1329 (MO), chimaltenango: Tres Cruces, N side of Volcan Acatenango, Beaman 3996 (GH); Volcan Acatenango, Hunnewell 14764 (GH); Chichavac, Skutch 255 (US); Las Calderas, Volcan Acatenango, Standley 61962 (F); Patzum, Williams & Molina 1 1843 (F, GH): road to Panajachel, between Los Idolos and Chochoyos, Williams 16214 (F). chiquimula: Concepcion de las Minas, Montana Norte, Steyermark 31042 (F). escuintla: S slopes of Volcan de Agua, Almeda 756 (DUKE). Guatemala: 6 km SE of Guatemala City, Harmon 1980 (MO); below San Lucas and Guatemala City, Molina et al. 16662 (F). huehue- tenango: Sierra de los Cuchumatanes, San Mateo Ixtatan, Breedlove 1 1636 (DS); Jacaltenango, Seler & Seler 2599 (GH); Yalambo, Seler & Seler 2869 (GH, US), quezaltenango: road to Fuentes Georginas, Boeke 164 (MO); 3 km S of Santa Maria Electric Plant, Roe et al. 718 (G); Volcan Zunil, Skutch 877 (A); above Mujulia, Standley 85487 (F); Montana Chicharro, lower SE slopes of Volcan Santa Maria, Steyermark 34282 (A, F). quiche: Nebaj, Contreras 5083 (LL); San Miguel Uspantan, Heyde & Lux 3001 (G, GH, M, MO, NY, POM, US); retalhuleu: Finca Helvetias, Muenscher 12430 (BH). sacatepequez: Volcan Acatenango, Donnell Smith 2469 (GH, US); Volcan de Fuego, Salvin s.n. (W); Santa Maria de Jesus, Volcan de Agua, Standley 65006 (F). san marcos: La Trinidad, above San Rafael, Croat 40881 (MO); 22 km W of San Marcos, Harmon & Fuentes 4744 (MO); Volcan Tacana, San Rafael, Steyermark 36331 (F, POM); Finca El Porvenir, White 5421 (MO); ca. 15 km W of San Marcos slopes of Cerro Tumbador, Williams et al. 23061 (F, G); near Alta Frater- nidad, between San Rafael Pie de la Cuesta and Palo Gordo, Williams et al. 27163 (F). solalA: 2 km W of Godinez, Harmon & Dwyer 2624 (GH, MO); above Lake Atitlan, W of Panajachel, Williams et al. 25347 (F). suchitepequez: S slope of Volcan Atitlan, Hatch & Wilson 361 (BH, F); Finca Moca, Skutch 1552 (A, F). zacapa Sierra de las Minas, Finca Planados, Steyermark 29996 (F); Volcan de Monos, Steyermark 42382 (BH, F). El Salvador, chalatenango: E slopes of Los Esemiles, Tucker 1006 (BG, G, LL, MICH, NY, P, PH, UC, US), santa ana: Volcan Santa Ana, Carlson 698 (A, F, MO, UC); Cerro Miramundo, Carlson 987 (F); Cerro Montecristo, 24 km NE of Metapan, Croat 42341 (MO). Honduras, cortes: Montana San Idalfonso, Molina 11462 (F). elparaIso: Mon- 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 tana de Yuscaran, Rodriguez 1956 (F); El Volcan, Williams & Molina 12187 (F, GH, MICH, MO). francisco morazan: Mt. Uyuca, near Zamorano, Carlson 2459 (GH); 6 km N of Tegucigalpa, Clewell 3016 (MO); Mt. San Juancito, Williams & Molina 12752 (F, GH); Montana La Tigra, ca. 15 km NW of Tegucigalpa, Williams et al. 23266 (BM, F, G, GH, LL, MICH, NY, UC, US, W). intibuca: near La Esperanza, Hazlett 1244 (MO); Cordillera de Opalaca, Calaveras, Molina 6434 (US), la paz: Sabanetas, Cordillera Guajiquiro, Molina & Molina 13918 (F); Montana Verde, Cordillera Guajiquiro, Molina 24392 (F). ocotepeque: 2 km E of Continental Divide on road between Nueva Ocotepeque and Santa Rosa de Copan, Harmon & Dwyer 4104 (GH, MO); Mt. Cocal, Cordillera Merendon, 20 km W of Ocotepeque, Molina 2214 (F); El Portillo, Cordillera Merendon, 20 km E of Nueva Ocote- peque, Molina et al. 31353 (MO). Nicaragua, jinotega: Cordillera Isabela, Macizos de Penas Blancas, Grant 7334 (A, F); Hwy 3, ca. 2 km NW of Aranjuez road, Stevens 9186 (MO); ca. 8 km 5 of Jinotega, Wilbur & Almeda 16510 (DUKE, MO), matagalpa: between Jinotega and Matagalpa, Bunting & Licht 983 (DUKE, F); Santa Maria de Ostuma, Williams et al. 23422 (F, G, W); road to La Fundadora, N of Santa Maria de Ostuma, Williams et al. 24955 (F, G); near Xelaju, Williams et al. 29239 (GH). Costa Rica, alajuela: Highway 120 to Volcan Poas, 1.5-3 km from the crater, Almeda 382 (DUKE); Palmira del Naranjo, Brenes 3521 (F, NY); ca. 12 km N of Carrizal, between Volcan Poas and Volcan Barba, Croat 35464 (MO); Zarcero, Palmira, Jimenez 2567 (F); Palmira, Smith 2145 (A, NA); Volcan Poas, Smith 6504 (GH, POM, US), cartago: 20 km SE of Empalme, Burger & Stolze 5237 (DUKE, F); near Volcan Irazii, Carlson 3581 (F, GH, S); Trinidad to Volcan Turrialba, Lent 722 (MO); Rio Tiribi, road from Ipis to Tierra Blanca, Lellinger & White 993 (F, MICH, MO); Volcan Turrialba, Pittier 7508 (F, MO, US); I km above El Empalme, Raven 20969 (DS); below San Jose de Chicua, road to Volcan Irazii, Raven 20974 (DS); Cerro Guardian, S slope of Irazii, Webster et al. 12143 (F); ca. 4 km NE of Pacayas towards Santa Cruz de Tenerife, Wilbur 6 Luteyn 13890 (F, GH, MO); ca. 4 km E of Rancho Redondo, SW flank of Volcan Irazii, Wilbur 14273 (DUKE, MO); between Hacienda Central and Finca Quemado, Wilbur 14339 (DUKE), guan- acaste: ca. 7 km NE of San Vicente, above Cascajal, Wilbur & Almeda 16720 (DUKE), heredia: Vara Blanca intersection, trail to Volcan Poas, Almeda et al. 2198 (DUKE); Porrosati, N of Barba, Crosby & Crosby 6611 (MO); Los Cartagos, Volcan Poas road, Raven 20980 (DS, GH); Vara Blanca de Sarapiqui, Skutch 3357 (A, G, MO); Volcan Barba Massif, Tonduz 1734 (W); San Rafael, E of Vara Blanca, Utley & Utley4169 (MO); Volcan Barba, Wilbur & Teeri 13679 (DUKE, F, GH, MICH, MO), san jose: Las Nubes, Allen 713 (A, MO); Abra, Chirripo Massif, Davidse & Pohl 1534 (MO); between Guayabillos and Cabeza de Vaca, Dodge & Thomas 4936 (GH, MO); Randho Redondo, Dodge & Thomas s.n. (MO); Cerro de la Muerte to San Isidro del General, Lewis 5047 (DUKE); 5 km N of Santa Maria de Dota, Lent 3910 (MO); ca. 6 km N of Copey, Little 6016 (A, MO); El Empalme, Stork 4545 (MICH, UC); La Palma, Tonduz 7410 (BM, F, GH, MO, POM, US); Copey, Tonduz 11666 (G, GH, W); ca. 7 km NE of San Vicente, above Cascajal, Wilbur & Almeda 16720 (DUKE); above Quebrada Varela, E of San Isidro, Wilbur & Luteyn 18177 (DUKE). Panama, chi- RiQUf: Nueva Suiza, Allen 1351 (GH, MO); Cerro Punta, Allen 3505 (BM, G, GH, MO, S, U); Finca Arco Iris, Boquete to Palo Alto, Beliz 210 (MO); 2 km towards Cerro de la Muerte, Correa 1256 (COL, DUKE, MO); between Alto de Guayabo and continental divide, Correa et al. 2821 (MO); Monte Rey, above Boquete, Croat 15772 (MO); above Guadelupe, Croat & Porter 16021 (MO); La Cumbres, near Cerro Punta, Croat & Porter 16136 (MO); Cerro Azul, E of Boquete, Croat 26838 (MO); N of San Felix, near Chiriquf-Bocas del Toro border on road to Cerro Colorado, Croat 33485 (MO); La Popa, above Boquete, DArcy & DArcy 6356 (MO); Alto Respinga, DArcy 9950 (MO); valley of Rio Viejo, N of Volcan City, Duke 8997 (MO); Cerro Horqueta, Duke et al. 13633 (DUKE, MO); Finca Collins, Ebinge 689 (MO); Cerro Punta to Las Nubes, Hammel 1371 (MO); N slope of Volcan Barii, E of Bajo Chorro, Hammel 2992 (MO); Cerro Hornito, Hammel 3052 (MO); Las Nubes, Liesner 289 (GH, MO); 8-15 km from Hato de Volcan, Luteyn 844 (DUKE); Alto Quiel to Bajo Mono, Luteyn 3708 (DUKE); Bajo Grande to Cerro Punta, Nee 9973 (GH, MO); upper Rio Chiriqui Viejo, vicinity of Monte Lirio, Seibert 246 (MO); Volcan Barii, near Cerro Punta, Stern & Chambers 86 (A, MO); El Volcan, White 4 (GH, MO); Bajo Quiel to Bajo Mono, Wilbur et al. 11992 (DUKE); S slopes of Cerro Horqueta, N of Boquete, Wilbur et al. 13443 (DUKE); Casita Alta, Volcan de Chiriqui, Woodson et al. 797 (A, MO, NY, POM). panamA: Cerro Jefe, 15-20 km beyond Goofy Lake (Lago Cerro Azul), Duke 8029 (MO). Naturalized specimens: Colombia, cundinamarca: Nemocon, between road and railroad tracks, Garcia Barriga 19387 (COL, US), boyaca: Labranzagrande, Guevara 389 (US). Sri Lanka: Nuwara Eliya to Hakgala, just beyond pass to Nuwara Eliya, Rte. A-5, Read & Desautels 2278 (MO). Tan- zania: Kiefingelo, W of Usumbara, Greenway 5896 (FHO, K). United States, Hawaii: bulldozed Kilauea Forest Reserve, Hawaii, Degener & Degener 31 130 (A, B, NA, Z); ravines over lava, Oahu, Leith et al. s.n. (MO). Cultivated specimens: Colombia, cundinamarca: Bogota, Duque-Jaramillo 2970 (COL, NY). France: Van Houtte nurseries, 1849 (K). Kenya: Nairobi, 1973, Raven 26162 (MO). New Zealand: 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 23 1 Whangarei, originally brought from Tahiti, Engelhorn s.n. (MO). Zimbabwe: Salisbury, Bie gel 4406 (NA). Sri Lanka: Hangola Garden, Marcovicz in 1927 (LE). United States, California: Santa Barbara, 1970, Hall 7736 (UC); Berkeley, Raven 49.801 (DS); Los Angeles, 1901, Braunton s.n. (BH). Massachusetts: Harvard University, Boston, 1876 (GH). Hawaii: ik 29 Miles/' Volcano, Hawaii, Degener & Degener 31131 (B, MO). Fuchsia paniculata has leaves with minutely to coarsely serrate margins and a ridged surface; its flowers are consistently more slender and generally smaller than those of/ 7 , arborescens, but they are much more variable. Apparently all populations of F. paniculata north of the Isthmus of Tehuantepec consist entirely of hermaphroditic plants; the flowers are more or less uniform, with small (less than 1 mm long), reniform anthers, and exserted, moderately four-lobed stigmas (Fig. 5d). The populations south of the Isthmus of Tehuantepec, however, are morphologically gynodioecious with perfect flower morphs of different popula- tions varying in style, length, and stigma size. The perfect flowers of these south- ern populations of F. paniculata vary from the type just described for the pop- ulations north of Tehuantepec to flowers with very small stigmas held well below the stamens on a short, slender style (Fig. 5b and c). Field observations were made on several populations of Fuchsia paniculata in Costa Rica by Arthur Weston in April, 1971. At one locality in Prov. Cartago, between El Empalme and La Trinidad, a random selection of the population yielded 11 pistillate and 15 perfect-flowered plants. All perfect-flowered plants were found to be functionally staminate, because no developing fruits could be found on them. This agrees with observations in Chiriquf Province of Panama by Woodson & Seibert (1937, p. 196). Examination of herbarium specimens of per- fect-flowered individuals from Panama and Costa Rica do show, however, a num- ber of collections with some ripe or developing fruits, for example, Hammel 3052 (MO) and Wilbur et al. 13443 (DUKE). In general, however, far fewer fruits are produced on perfect-flowered plants than on pistillate plants in the populations south of the Isthmus of Tehuantepec, and this can be seen easily both in the field and on herbarium specimens. In addition, fewer seeds apparently develop in fruits of the perfect-flowered plants. The perfect flowers are thus largely female-sterile, and the populations south of the Isthmus of Tehuantepec are consequently sub- dioecious like the three morphologically gynodioecious species of sect. Enclian- dra (Arroyo & Raven, 1975). Perfect-flowered plants with small stigmas and short styles, raised at the Mis- souri Botanical Garden from seed of Breedlove 42742 (MO) from Chiapas, Mex- ico, regularly produced moderate numbers of selfed fruits with viable seeds. Some of these seeds were raised until flowering, yielding 9 hermaphroditic and 4 pis- tillate individuals. Although the sample size was quite small, this does indicate that male sterility in F. paniculata is not controlled by a dominant gene, as seems to be the case in sect. Encliandra (Arroyo & Raven, 1975). The male sterility in sect. Schufia may therefore be more similar to that found in the unrelated sect. Skinnera, in which it is controlled by a recessive gene (Arroyo & Raven, 1975). The incomplete female sterility in the perfect-flowered plants of Fuchsia pan- iculata south of the Isthmus of Tehuantepec, the presence of only hermaphroditic populations of the same species north of Tehuantepec, and the presence of an entirely hermaphroditic, closely related species, F. arborescens, clearly repre- sents a dynamically evolving progression from hermaphroditic to dioecious plants, 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 where first male sterility, then female sterility, have been selected to promote increased outcrossing. Diploid gametic chromosome counts of n = 1 1 were obtained from the fol- lowing collections of Fuchsia paniculata : Breedlove 5971 , 7343, and Raven 20995. In addition, a count of n = 11 with normal meiotic pairing was obtained from an experimental cross between two different populations of F. paniculata, one from Oaxaca (Hill 1730, MO) and the other from Costa Rica (L. Fournier, El Empalme, Cartago; no voucher). Fuchsia sect. Fuchsia, type: Fuchsia triphylla L. Eufuchsia Baillon, Hist. PI. 6: 467. 1877, as section. Munz, Proc. Calif. Acad. Sci. IV. 25:15. 1943. Hermaphroditic. Erect, scandent, or climbing shrubs. Leaves opposite or whorled. Flowers generally pendant; axillary, racemose, or paniculate. Floral tube usually longer than sepals. Petals usually more than id the length of the sepals and generally not convolute at anthesis. Nectary annular, surrounding the style and free from the tube except for the basal Vi or less, rarely fully adnate to the floral tube. Stamens biseriate, shorter than the sepals or exserted less than 5 mm beyond them, the antisepalous stamens longer than the antipetalous stamens. Berry with ca. 50-ca. 200 seeds, these compressed laterally, irregularly triangular or obovate in outline, 1-2.5 mm long, 0.7-1.9 mm wide. Gametic chromosome numbers, n = 1 1, 22. 7. Fuchsia boliviana Carriere, Rev. Hort. 48:150, pi. 1876. type: Plate in Carr., Rev. Hort. 48:150. 1876. Lectotype here designated. The plant illustrated is from garden material cultivated in France, ca. 1876, from seeds collected in the mountains of Bolivia in 1873 by Benedict Roezl. For full synonymy see Berry (1982). Fuchsia corymbiftora auct. mult., non Ruiz & Pavon. 1802. Erect, openly-branched shrubs 1.5-4(-6) m tall. Branchlets ascending at base, nodding towards apex, 1-4 dm long, 3-7 mm thick, terete to angled, densely tomentose; older branches and trunks 1-5 cm thick. Leaves opposite, sometimes alternate or ternate, soft-membranous, elliptic to ovate, rounded to acute at base, acute to acuminate at apex, 5-20(-23) cm long, 3—1 2(— 1 5) cm wide, dark matte green above, paler below, soft pubescent on both surfaces; margin denticulate with numerous glandular teeth; petioles pubescent, 2-5(-7) cm long, stipules dark, filiform, 1-2 mm long, ca. 0.3 mm wide, deciduous. Flowers numerous in ter- minal, drooping racemes or few-branched panicles, the flowers congested toward the tip; rachis 5-40(-60) cm long; bracts reflexed, lanceolate, 5-25 mm long; pedicels slender, 5-16 mm long; floral tube narrowly funnelform, 30-60(-70) mm long, 1.5-3 mm wide at base, gradually widened above until 5-8 mm wide at rim, pubescent outside, pilose inside; sepals lanceolate, acuminate, 10-20 mm long, 4-5 mm wide, initially spreading but soon becoming fully reflexed, the tips con- nivent in bud; tube and sepals pale pink to usually bright scarlet; petals red, oblong to lanceolate, acute at apex, 8-16(-20) mm long, 3-7(-9) mm wide, ± crispate with 2-3 longitudinal ridges; petals shrivel and fall off before tube de- hisces; nectary annular, 4-lobed, 2-4 mm high, mostly free from the tube; fila- 1982] BREEDLOVE ET AL.— MEXICAN AND CENTRAL AMERICA FUCHSIA 233 ments red, the antisepalous filaments 8-15 mm long, the antipetalous filaments 5-10 mm long; anthers oblong, 2-3.5 mm long, 1-1.5 mm thick, white; style red, pubescent, the stigma capitate, subtetragonous, 2-3.5 mm long, 3-5 mm wide, 4- parted at apex, cream. Berry ellipsoid to cylindric, 10-25 mm long, 8-14 mm thick, dark purple, comestible; seeds tan, 1.5-2.0 mm long, 0.5-1.0 mm thick. Gametic chromosome number, n = 1 1 . Distribution: Escaped from cultivation and locally established on moist slopes from Puebla south to Costa Rica, always near habitations. (Native to southern Peru, Bolivia, and northern Argentina.) Flowering throughout the year. Specimens examined (both naturalized and cultivated): Mexico, chiapas: Piedracitos, Chamula, Breedlove 8018 (DS). distrito federal: Villa Obregon, Moore 6395 (BH); Coyoacan, Woronow 2065 (LE). puebla: Huauchinango, Baldwin 14390 (LL). Veracruz: Orizaba, Botteri 929 (LE); Jalacingo, Dodds 57 (MO); Altotonga, Foster in 1938 (BH, BM); Santa Ana Atzacan, N of Orizaba, Rosas 191 (MO); Santa 7 Cruz, Altotonga, Ventura 1026 (CAS). Guatemala, sacatepequez: San Rafael, Donnell Smith 2176 (K, US); Antigua, Standley 62331 (F). san marcos: Tajumulco, Volcan Tajumulco, Johnston 1231 (F), Steyermark 36510 (F). El Salvador: Volcan de San Salvador, Calde- ron 2345 (F, US). Costa Rica, alajuela: Zarcuo, A. Smith 2775 (F). cartago: Cartago, Donnell Smith 4804 (US), san jose: San Jose, Pittier 14104 (CM, GH, U, US); Guadalupe, Quiros 1289 (F). Fuchsia boliviana is perhaps the most widely naturalized species in the genus, at least in tropical and subtropical areas. Outside of its probable native range of southern Peru to northern Argentina, it is also naturalized in Colombia, Vene- zuela, Jamaica, Hawaii, Java, Reunion, India, and a number of other countries. A single introduced plant can give rise to extensive local populations, because it self-pollinates very effectively and also reproduces vegetatively by stem shoots. Its striking drooping terminal racemes stay in flower through the year, and it is tolerant of much drier and harsher conditions than most species in the genus. For these reasons, F. boliviana is a frequently cultivated and escaped shrub in many villages throughout Central America and Mexico. A diploid gametic chro- mosome count of n = 11 was obtained from Breedlove 8018 (DS). Literature Cited Arroyo, M. Kalin, & P. H. Raven. 1975. The evolution of subdioecy in morphologically gyno- dioecious species of Fuchsia section Encliandra (Onagraceae). Evolution 29( 3): 500-51 1. Berry, P. E. 1982. The systematics and evolution of Fuchsia sect. Fuchsia (Onagraceae). Ann. Missouri Bot. Gard. 69:1-198. Breedlove, D. E. 1969. The systematics of Fuchsia section Encliandra (Onagraceae). Univ. Calif. Publ. Bot. 53:1-69. Bullock, W. 1824. Six months residence and travels in Mexico. London, John Murray. 530 pp. Donnell Smith, J. 1893. Undescribed plants from Guatemala. X. Bot. Gaz. 18:1-7. Harrison, J. 1841. Embellishments. Fuchsia cordifolia. Floric. Cab. & Florist's Mag. 9:241, pi. 205. Hemsley, W. B. 1876. The various races of garden fuchsias. Garden 9:284-286. . 1909. Fuchsia splendens and the allied species. Gard. Chron. 3(45):338, t. Kurabayashi, M., H. Lewis & P. H. Raven. 1962. A comparative study of mitosis in the Ona- graceae. Amer. J. Bot. 49:1003-1026. Lemaire, C. 1848. Fuchsia arborescens var. syringae flora. Fl. Serres Jard. Eur. 4:4\6, fig. Lindley, J. 1826. Fuchsia arborescens. Tree Fuchsia. Bot. Reg., pi. 943. . 1856. New plants. Fuchsia paniculat a. Gard. Chron. 1856:301. McVaugh, R. 1977. Botanical results of the Sesse & Mocirio expedition (1787-1803) 1. Summary of excursions and travels. Contrib. Univ. Michigan Herb. 1 1(3):97— 195. Munz, P. A. 1943. A revision of the genus Fuchsia (Onagraceae). Proc. Calif. Acad. IV, 25: 1-137. Planchon, J. E. 1849. Fuchsia splendens. Fl. Serres Jard. Eur. t. 458. 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Raven, P. H. 1979. A survey of reproductive biology in Onagraceae. New Zealand J. Bot. 17:575- 593. Rzedowski, J. & R. McVaugh. 1966. La vegetacion de Nueva Galicia. Contrib. Univ. Michigan Herb. 9(1):1-123. Sims, J. 1825. Fuchsia arborescens. Laurel-leaved Fuchsia. Bot. Mag., pi. 1620. Sprague, T. A. 1926. Sesse and Mocino's Plantae Novae Hispaniae and Flora Mexicana. Kew Bull. Misc. Inf. 417-426. Standley, P. & L. O. Williams. 1963. Flora of Guatemala, Onagraceae. Fieldiana Bot. 24(7): 525-563. Woodson, R. E. & R. J. Seibert. 1937. Contributions toward a Flora of Panama. I. Collections in the Provinces of Chiriqui, Code, and Panama by R. J. Seibert during the summer of 1935. Ann. Missouri Bot. Gard. 24:175-210. Appendix Fuchsia hyhrida Hort. ex Sieb. & Voss. in Vilm, Blumengart. 3, 1:332. 1896. Specimens examined (presumed cultivated): Mexico, hidalgo: Real del Monte, Salazar s.n. (MEXU). michoacan: Hacienda Coahuayula, Emrick 204 (F). puebla: San Pedro, Nicolas s.n. (NY). Vera- cruz: Jalapa, Calzada 2086 (MO); Altotonga, Dodds 41 (MICH); La Orduna, Municipio Coatepec, Jimenez 70 (MO); Congregacion Dos Pocitos, Municipio Tonayan, Mdrquez et al. 456 (MO), 457 (MO), 459 (MO); Casa Quemada, Km 36 of Teocelo road, Cosautlan, Mdrquez et al. 965 (MO); Zoncuantla (La Pitaya), Municipio Coatepec, Murrieta 103 (MO). Guatemala, alta verapaz: Vi- cinity of Coban, Standley 91231 (F), 92468 (F). Guatemala: without locality, Aguilar 125 (F), 171 (F), 173 (F). huehuetenango: Wahshaklahung Pyramid, San Mateo Ixtatan, Breedlove 8658 (DS). sacatepequez: Antigua, Standley 63836 (F). Costa Rica, san jose: San Jose, Umana 20 (F). Fuchsia magellanica Lamarck, Encyc. 2:565. 1788. Specimens examined (presumed cultivated): Mexico, puebla: Arsene s.n. (US), san luis porosf: San Luis Potosf, Schauffeur in 1879 (A). Panama, chiriquI: Boquete, Dwyer 7022 (MO). THE DENNIS STANFIELD AWARD The Dennis Stanfield Memorial Fund has been established to assist persons of scientific merit to undertake botanical research on tropical African plants. Awards were made in 1974, 1977, and 1980, and the award in 1983 will be£250. Application forms should be ob- tained from the Executive Secretary of the Linnean Society of London, Burlington House, Piccadilly, London W1V OLQ, United Kingdom, and should be returned to the same address by 31st March 1983. The award is to be used for such items as travel, equipment, books, computing time, research expenses, and the like, in connection with any aspects of botanical research. The award is open both to amateurs and professionals. THE 1982 JESSE M. GREENMAN AWARD The 1982 Jesse M. Greenman Award has been won by Walter S. Judd for his publica- tion kfc A monograph of Lyonia (Ericaceae)" (Jour. Arnold Arbor. 62:63-209; 315-436. This monographic study is based on a Ph.D. Dissertation from the Department of Biology, Harvard University. The Greenman Award, a cash prize of $250, is presented each year by the Missouri Botanical Garden. It recognizes the paper judged best in vascular plant or bryophyte systematics based on a doctoral dissertation that was published during the previous year. Papers published during 1982 are now being considered for the 16th annual award, which will be presented in the summer of 1983. Reprints of such papers should be sent to: Greenman Award Committee, Department of Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. In order to be considered for the 1983 award, reprints must be received by 1 July 1983. INDEX Synonyms are italicized. Page numbers of main entries are in bold face, those of illustrations or maps are in italics. Aglaiocercus kingi 31 Anthracothorax dominicus 3 1 Araucaria 8 Arracacia incisa 27 Ascidiogyne 26 Berberis 8 Bombus 207 Brebissonia Spach 4, 67 microphylla Humboldt, Bonpland & Kunth 67 Cactaceae 9 Camissonia arenaria 64 Chlorostilbon swainsonii 31, 51, 190, 191 Chusquea 8 Circaea 4, 30 Clarkia 33, 65 Compositae 26, 28 Drimys 8 Ellobium Lilja 68, 212 fulgens (DC) Lilja 217 Ellobum Blume 68, 212 Encliandra Zuccarini 68 parviflora Zuccarini 68 Epilobium 48 Espletia 24 Fuchsia Sect. Breviflorae de Candolle 4 Sect. Ellobium 3, 4, 6, 7, 9-11, 55, 59, 62, 63, 65, 66, 209, 212-213, 215, 216 Sect. Encliandra 3, 4, 6, 7, 9-11, 33, 59, 66, 181, 209, 210, 231 Sect. Eufuchsia Baillon 68 Sect. Fuchsia 2-4, 5, 6, 7, 10, 11, 16, 19, 21-23, 59, 66, 209, 210, 212, 214, 232-233 Sect. Hemsleyella 1, 3, 4, 5, 7, 9-11, 16, 31, 33, 55, 59, 62, 63, 65, 66, 209, 210, 212, 214 Sect. Jimenezia 3, 4, 6, 7, 9-11, 59, 66, 194, 209, 210, 220-221 Sect. Kierschlegeria 1, 3-5, 7, 9, 10, 11, 33, 59, 65, 181, 200, 209, 210 Sect. Longiflorae de Candolle 4 Sect. Macrostemonae de Candolle 4 Sect. Quelusia 1, 3, 4, 5-7, 10, 33, 59, 65, 66, 209 Sect. Schufia 4, 6, 7, 9-11, 30, 59, 66, 181, 209, 210, 223-224, 231 Sect. Skinnera 3, 6, 9-11, 59, 63, 65, 66, 181, 209, 210, 231 abrupta I. M. Johnston 34, 47, 53, 57, 78, 79, 80, 105, 109-110, 142, 150, 171 adpressipilis Steyermark 90 ampliata Bentham 18, 34, 40, 49, 53, 61, 85, 86, 115, 122, 124, 125-127, 146, 176 andrei I. M. Johnston 34, 53, 56, 104, 105, 106-108, 109 apiculata I. M. Johnston 83, 86 arbor ea Sesse & Mocirio 211, 223, 224 arborescens Sims 209-211, 221, 223, 224-227, 228, 231 forma parva Munz 228 forma tenuis Munz 228 var. megalantha Donnell Smith 224, 228 var. syringaeftora Lemaire 228 var. typica 224 asperifolia K. Krause 102 aspiazui J. F. Macbride 109, 110 asplundii J. F. Macbride 121 atrorubra I. M. Johnston 90 austromontana I. M. Johnston 19, 27, 34, 53, 140, 141, 142-143 ayavacensis Humboldt, Bonpland & Kunth 26, 34, 53, 115, 121-122, 125, 126 boliviana Britton 81 boliviana Carriere 14-16, 27, 30-32, 34, 54-56, 58, 59, 61, 63, 83, 162-170, 171, 172, 210, 211, 218, 232, 233 brittonii I. M. Johnston 82 canescens Bentham 2, 34, 54, 56, 57, 61, 117, 118, 120, 122, 125, 177, 182-184 caracasensis Fielding & Gardner 38, 43, 89 caucana P. Berry 17, 34, 37, 40, 49, 53, 61, 62, 115, 117, 118-121, 184 ceracea P. Berry 34, 5/, 54-56, 58, 59, 110, 149-150, 151, 153 cinerea P. Berry 26, 34, 54, 94, 177, 184-185 coccinea Aiton 67 cochabambana P. Berry 53, 57, 58, 83, 140, 144-145 colombiana Munz 38, 114, 118 confertifolia Fielding & Gardner 53, 56-58, 130, 136-137 cordifolia Bentham 209, 211, 212, 216 coriacifolia P. Berry 54, 57-59, 149, 150-152, 153 corollata Bentham 17, 34, 37, 40, 53, 61, 65, 94, 112, 114-118, 120, 121, 155, 176, 183, 184 corymbiflora Ruiz & Pavon 34, 54, 58, 78, 110, 142, 150, 156, 162, 168, 170-171, 172, 175, 176, 211 alba Harrison 163, 170 forma puberulenta Munz 163 var. luxurians I. M. Johnston 163, 169 crassistipula P. Berry 16, 34, 46, 54, 56, 57, 58, 177, 181-182 cuatrecasasii Munz 24, 39, 53, 56-58, 88, 104, 105, 108-109, 155, 194 curviflora Bentham 111, 114 cuspidata Fawcett & Rendle 163 cyrtandroides Moore 9, 28, 33 decidua Standley 209, 210, 211-213, 214, 219- 220, 227 decussata Ruiz & Pavon 27, 52, 56, 58, 76-79, 80, 81, 110, 142 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 denticulata Ruiz & Pavon 14, 19, 22, 26, 27, 29, 34, 40, 46, 53, 55, 59, 61, 62, 78, 80, 83, 110, 137-142, 143-148, 162, 171, 183 dependens Hooker 14, 15, 35, 54, 58-60, 94, 118, 127, 155, 171, 174-176, 177, 178, 180, 184, 185, 218 dolichantha K. Krause 137 dominiana 40, 147 encliandra Steudel 68 excorticata (J. R. & J. G. A. Forster) Lin- naeus f. 67 ferreyrae P. Berry 35, 50, 52, 56, 59, 61, 78, 79-80, 81, 110, 142 filipes Rusby 82 fischeri J. F. Macbride 173 fontinalis J. F. Macbride 18, 35, 52, 56, 59, 78, 80-81, 172 fosbergii Munz 143, 144 fulgens de Candolle 209, 211-213, 214, 216- 218, 227 pumila Carriere 217 furfuracea I. M. Johnston 27, 54, 83, 151, 158, 159-161, 162, 168 fusca K. Krause 77, 78 gehrigeri Munz 15, 23, 25, 32, 35, 38, 39, 41-43, 53, 56, 61, 89, 92, 113, 129, 130, 132-135, 194 glaberrima 1. M. Johnston 26, 52, 55-58, 93, 97-98, 102, 106 harlingii Munz 49, 53, 56, 61, 140 f 143-144 hartwegii Bentham 16, 17, 30, 34, 35, 54-56, 88, 114, 155, 176, 177, 178-181 hirtella Humboldt, Bonpland & Kunth 24, 35, 39,54, 56, 114, 130 hitchcockii I. M. Johnston 122, 124, 176-178, 179-182, 194 hybrida 2 1 1 hypoleuca I. M. Johnston 83, 86 intermedia Hemsley 214, 216 involucrata Sw. 195 jahnii Munz 132, 133 jimenezii Breedlove, Berry & Raven 194, 209, 210, 214, 220, 221-223 juntasensis Kuntz 63 killipii I. M. Johnston 127, 129 lampadaria J. O. Wright 147, 148 lehmannii Munz 35, 53, 104-106, 107-109, 136 leihmanni H. Leveille 228 leptopoda K. Krause 138, 141 llewelynii J. F. Macbride 53, 56, 130, 135, 136, 137 longiflora Bentham 145 loxensis Humboldt, Bonpland & Kunth 14, 18, 32, 35, 52, 56, 65, 83-86, 87, 125, 144 lycioides Andrews 10, 33, 199-208 macropetala Presl 32, 40, 52, 87, 99-100, 101 macrophylla I. M. Johnston 14, 27, 32, 35, 40, 52, 58, 60, 87, 98-99, 100-102 macrostigma Bentham 18, 25, 40, 53, 58, 61, 62, 88, 94, 140, 145-147 var. longiflora (Benth.) Munz 145 var. puhens 1. M. Johnston 145 magdalenae Munz 22, 24, 28, 35, 53, 56, 61- 63, 65, 140, 147-148 magellanica Lamarck 10, 67, 211 mathewsii J. F. Macbride 18, 26, 35, 54, 56, 59, 81, 156, 168, 172, 173-174 meridensis Steyermark 127 microphylla Humboldt, Bonpland & Kunth 68, 206 miniata Planchon & Linden 194 munzii J. F. Macbride 170, 171 nigricans Linden ex Planchon 13, 15, 30, 32, 35, 36, 38, 41-44, 47, 51-53, 55, 59, 61, 89- 92, 93, 95, 104, 114, 130, 134, 155, 194 orientalis P. Berry 18, 36, 52, 57, 88, 93, 95- 97, 98 osgoodii J. F. Macbride 106, 108 ovalis Ruiz & Pavon 52, 57, 59, 87, 99, 100-101, 102, 108 var. aberrans J. F. Macbride 106, 108 pallescens Diels 18, 36, 52, 55, 58, 61, 88, 91, 93, 94-95, 104, 125 paniculata Lindley 30, 209-211, 223, 224, 225, lll-lhl perbrevis I. M. Johnston 193 petiolaris Humboldt, Bonpland & Kunth 14, 16, 24-26, 36, 52, 53, 55, 61, 85, 110-114, 7/5-1 18, 120, 122, 124, 126, 130, 133, 178 var. bolivarensis Munz 1 1 1 pilosa Fielding & Gardner 36, 52, 53, 55-57, 59, 61, 64, 87, 97, 101-103, 132, 137, 172 platypetala I. M. Johnston 38, 194 polyantha Killip ex Munz 26, 32, 36, 48, 54, 56, 58, 94, 155-157 polyanthella I. M. Johnston 100 pringsheimii 14, 28, 31, 36, 37, 39, 40, 50, 54, 58, 61-63, 65, 187, 189, 190, 191-192, 194 putumayensis Munz 18, 44, 53, 56, 58, 59, 61, 64, 65, 92, 95, 103-104, /05-1O7, 109, 110, 147 quinduensis 1 10, 1 14 racemosa Lamarck 186, 188 racemosa Sesse & Mocirio 217 regia (Vandelli) Munz 67 rivularis J. F. Macbride 36, 53, 55, 56, 61, 102, 103, 129, 130, 131-132, 135 sanctae-rosae Kuntz 19, 27, 36, 38, 52, 78, 81- 83, 142, 145, 159-162, 170 sanmartina P. Berry 54, 78, 80, 149, 151, 152- 153 scabriuscula Bentham 18, 24, 39, 47, 52, 56, 57, 62, 85, 86-88, 94, 95, 97, 109, 147, 155, 194 4 scandens K. Krause 76 scherffiana Andre 53, 130, 136 serratifolia Ruiz & Pavon 138 sessilifolia Bentham 24, 32, 36, 39, 43, 48, 54, 56-59, 61, 62, 88, 92, 94, 109, 118, 127, 147, 153-155, 156, 157, 176, 194 simplicicaulis Ruiz & Pavon 27, 54, 59, 60, 148-149, 150, 151, 153 siphonata K. Krause 138 smithii Munz 111, 1 13 1982] SUBJECT INDEX 237 spectabilis Hooker ex Lindley 145 splendens Zuccarini 209, 211, 212, 213-216 steyermarkii P. Berry 26, 52, 57, 58, 85, 87, Laurelia 8 Kirschlegeria H. G. L. Reichenbach 68 Lae stadia 28 88-89, 136 storkii Munz 173 Lepechinia 8 Loasaceae 9 sylvatica Bentham 18, 25, 36, 52, 57, 88, 90, Lopezieae 33 91,92-94,95,96, 147, 155 syringaeflora (Lem.) Carriere 227 tacsoniiflora K. Krause 138, 140 thymifolia Humboldt, Bonpland & Kunth 68, 206 tincta I. M. Johnston 19, 27, 36, 40, 54, 55, 58, 83, 141, 151, 157-159, 160-162 townsendii I. M. Johnston 121 triphylla Linnaeus 14, 28, 31, 33, 37, 39, 40, 50, 51, 54, 55, 58, 59, 65, 66, 185-191, 192 tuberosa 141 umbrosa Bentham 83, 85 vargasiana Munz ex Vargas 19, 27, 36, 40, 54, 55, 83, 141, 151, 158, 160, 161-162 velutina I. M. Johnston 170 venusta Humboldt, Bonpland & Kunth 12, 15, 16, 24, 36, 39, 41, 42, 49, 50, 51, 53, 55, Ludwigia 2-4, 55 Lyciopsis Spach 4, 68 thymifolia (Humboldt, Bonpland & Kunth) Spach 68 Mellisuga minima 31 Monochaetum 48 Myrinia Lilja 68 microphylla (Humboldt, Bonpland & Kunth) Lilja 68 Nahusia Schneevoogt 67 coccinea (Aiton) Schneevoogt 67 Nothofagus 8, 10 Nyctaginaceae 9 Ocreatus underwoodii 31 Oenothera 32, 48 Onagreae 33 Perezia 8 56, 61, 65, 92, 114, 127-130, 132-137, 178- Podocarpus 8 180, 194 var. huilensis Munz 127 verrucosa Hartweg 25, 30, 32, 37, 39, 41, 50, 54, 55, 57-59, 61-63, 65, 88, 92, 109, 130, 134, 155, 766, 178, 192-194 var. tamaensis Steyermark 193 Puya 24 Quelusia Vand. 67 regia Vand. ex Vellozo 67 Quilusia J. D. Hooker 68 Rhodopsis vesper atacamensis 201, 204, 206, Rubiaceae 57 207 vulcanica Andre 14, 18, 37, 40, 53, 65, 95, Schufia Spach 4, 68, 223 122-125, 126, 127 weberbaueri K. Krause 82 woytkowskii Munz 131 arborescens (Sims) Spach 68, 224 Skinner a J. R. & J. G. A. Forster 67 Spachea Juss. 68, 212 wurdackii Munz 18, 37, 54, 57, 59, 102, 137, Spachia Lilja 212 156, 168, 171, 172 Gongylocarpus 65 Gunnera 95 Hauya 55 Ilex 8 Kierschlegeria Spach 68 lycioides (Andrews) Spach 68 fulgens (DC) Lilja 68, 217 Thilcum Molina 67 tinctorium Molina 67 Tilco Adanson 67 Zygophyllaceae 9 The previous issue of the Annals of the Missouri Botanical Garden, Vol. 68, No. 4, pp. 505-686, was published on 29 March 1982. Systematics Symposia Published In 1953 the Missouri Botanical Garden began holding a series of annual systematics symposia each fall. Since 1969, with the sixteenth symposium, the Garden has pub- lished the proceedings of most in its quarterly series, ANNALS OF THE MISSOURI BOTANICAL GARDEN. Those published symposia which are available are listed below, and the price of all except the twenty-fourth is $10.00. The twenty-fourth, which was reprinted separately from the ANNALS, is available for $6.00. Many of these symposium issues of the ANNALS contain other papers, but each issue listed below contains the complete proceedings of the particular symposium. The proceedings of the 1981 symposium will be published during the second half of 1982, and if you wish to purchase a copy of that symposium, you may subscribe at this time. Upon publication, your copy will be sent. Sixteenth Annual Symposium, 1969— TROPICAL 56(3). 127 pp. $10.00. 1970—HY TEMATICS. 59(3). 142 pp. $10.00. Twentieth Annual Symposium, 1973— PLANT-ANIMAL COEVOLUTION. 61(3). 132 pp. $10.00. Twenty-first Annual Symposium, 1974— BIOGEOGRAPHY. 62(2). 143 pp. $10.00. Twenty-second Annual Symposium, 1975— EVOLUTION AT THE POPULATION 100 Twenty-third Annual Symposium. 1976— CHKMOSYSTEMATICS. 64(2). 98 pp. $10.00. Twenty-fourth Annual Symposium, 1977— SYSTEMATICS STUDIES IN AFRICA. 223 pp. $6.00. 978— PALYNOLOGY 248 pp. $10.00. 979— EVOLUTION 04 Twenty-seventh Annual Symposium. 1980— REPRODUCTIVE STRATEGIES IN PLANTS AND ANIMALS. 68(2). 146 pp. $10.00. Twenty-eighth Annual Symposium, 1981— BIOLOGICAL STUDIES IN CENTRAL AMERICA. About 140 pp. $10.00— Available Fall 1982. To place an order use this form or a photocopy of it. Orders should be prepaid; a $1.00 fee will be added to orders requiring invoices. No shipments are made until payment is received. Mail form with your check or money order, payable to Missouri Botanical Garden, to: Department Eleven Missouri Botanical Garden P.O. Hox 299 St. Louis. MO 63166-0299 U.S.A. Please send the SYMPOSIA checked above to: Name Address D Payment enclosed. □ Send invoice ($1.00 fee will be added to total) Postal Code Country Monographs in Systematic Botany from the Missouri Botanical Garden Volumes of the Monographs published to date are listed below. Prices are prepaid and postpaid. 1. A Provisional Checklist of Species for Flora North America (Revised). Edited by Stanwyn G. Shetler & Laurence E. Skog. xix + 199 pages. Paper. 1978. Out-of-print. 2. Missouri Wildflowers of the St. Louis Area. Erna R. Eisendrath. 390 pages. $7 Mosses. Second Printing. Marshall Magill. vii + 43 pages. Paper. 1978. [reprinted 1981]. $4.00. 4. Orchids of Panama. A facsimile reprint of the Orchidaceae, Flora of Panama. with a checklist of the orchids of Panama as known today by Robert L. Dressier, viii + 484 + XXVI pages. Paper. 1980. Price reduced to $20.00. 5. Index to Plant Chromosome Numbers, 1975-1978. Edited by Peter Goldblatt. vii + 553 pages. Paper. 1981. $15.00. 6. Les Ombelliferes. A reprint of the original, 1977, edition. Edited by A.-M. ■ '*-% ■ m « w ■ a A. & M ^ __ . rfl T S~** % • • *~* ^ .-*. *-». a **> t" 1"V C\£\ Marc & J. Carbonnier. ix + 865 + iv Daees. Paoer. 1982. $50.00 7. Musci Austro-Americani. A reprint of the original, 1869, edition in the Journal of the Linnean Society, Botany. By G. Mitten. 659 + vi pages $15.00 Name Address Postal Code □ □ invoice ($1.00 fee will & added to total). □ a leek here it you wish iu f— standing order for future voi umes. Country Your standing order #: Signature OF THE m OLUME 69 1982 NUMBER 2 ■L ^X.A% -'■; " *«*n Boton^ Garcfej /Wl/ w • ' CONTENTS Th e Systematics and Evolution of Epilobium (Onagraceae) in South Amer- lca James C. Solomon A Revision of the Southwestern Species of Amsonia (Apocynaceae) Steven P. McLaughlin A Synopsis of Moraea (Iridaceae) with New Taxa, Transfers, and Notes p eter Goldblatt C °rm Morphology in Hesperantha (Iridaceae, Ixioideae) and a Proposed ^ Infrageneric Taxonomy Peter Goldblatt Note s on Geissorhiza (Iridaceae): The Species in Madagascar Peter Goldblatt Illin <>is Solanaceae in the Missouri Botanical Garden Herbarium and Bio- graphical Sketches of Some Collectors Robert H. Mohlenbrock .... - ( VOLUME 69 1982 NUMBER 2 OF THE OTMKM, The Annals contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden. Papers origi- nating outside the Garden will also be accepted. Authors should write the editor for information concerning arrangements for pub- lishing in the Annals. Editorial Committee Nancy Morin, Editor Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden & St. Louis University Peter Goldblatt Missouri Botanical Garden Published four times a year by the Missouri Botanical Garden, St. Louis, Missouri 63110. ISSN 0026-6493 For subscription information contact the Business Office of the Annals, P.O. Box 368, 1041 New Hampshire, Lawrence, Kansas 66044. Subscription price is $45 per volume U.S., Canada, and Mexico, $50 all other countries. Four issues per volume. Second class postage paid at Lawrence, Kansas 66044 © Missouri Botanical Garden 1983 OF THE > VOLUME 69 1982 NUMBER 2 THE SYSTEMATICS AND EVOLUTION OF EPILOBIUM (ONAGRACEAE) IN SOUTH AMERICA 1 James C. Solomon 2 Abstract Recent field, cytological, experimental hybridization, and morphological studies indicate that the patterns of diversity in South American species of Epilobium can best be understood by recognizing 12 native species. These have been derived from at least five independent pre-European introductions of Epilobium into South America, with subsequent diversification in some of these lines. Two groups have ancestral affinities with North America: Epilobium ciliatum ssp. ciliatum is consubspecific with North American populations; the primarily northern Andean group, consisting of E. denticulatum and allied species, has no close relatives in the Northern Hemisphere, but has a chromosomal ar- rangement identical with that of some North American species. Two other lines are derived directly from Australasian ancestors: Epilobium hirtigcrum is conspecific with a widespread species of Aus- tralia and New Zealand, and E. conjungens, with its unusual creeping habit and consistently opposite ' 1 thank Peter Raven for his continuing interest, advice, and patience during the course of this study. Steven Seavey provided most of the cytological information reported here. Peter Hoch, T. P. Ramamoorthy, and Warren L. Wagner provided many interesting discussions that were useful in formulating the ideas presented here. I thank my wife, Andrea, for her help throughout this study. In South America, Armando Hunziker provided full logistical support during our stay at Cordoba, Argentina. In addition, Mary Arroyo, Osvaldo Boelcke, Jose Diem, Clodomiro Marticorena, Arturo Martinez, David Moore, Melica Munoz, Edmundo Pisano, Carlos Ramirez, and Otto Zollner answered many questions and provided seeds or other useful information that added greatly to the value of this study. 1 thank the directors and curators of the following herbaria for allowing me to examine the material entrusted to their care: A, AAU, ARIZ, ASU, B, BA, BAA, BAB, BAF, BH, BM, BP, BR, BREM, C, CAS, COL, CONC, CONN, CORD, CTES, DS, DUKE, E, ECON, F, G, GB, GH, GOET, H, HAL, HBR, HIP, JE, K, L, LA, LAM, LD, LIL, LP, LTR, LZ, M, MA, MER, MICH, MO, MPU, MSC, MU, MY, NA, NY, OXF, P, PH, POM, PR, PRC, RJ, RNG, RSA, S, SGO, SI, SP, U, UC, UPS, US, USM, VALD, VEN, W, WIS, WTU, WU, Z. This material is based upon research supported by the National Science Foundation under Grants No. DEB 78-20219 (Doctoral Dissertation Research Improvement, to J.S.) and DEB 78-23400 (to Peter H. Raven), and is published with the support of National Science Foundation Grant No. DEB 82-01242. The Foundation provides awards for research and education in the sciences. The awardee is wholly responsible for the conduct of such research and preparation of the results for the publi- cation. The Foundation, therefore, does not assume responsibility for such findings or their interpre- tation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author and do not necessarily reflect the views of the National Science Foundation. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, USA (reprint requests). Other correspondence to: Casilla 20206, La Paz, Bolivia. Ann. Missouri Bot. Gard. 69: 239-335. 1982. 0026-6493/82/0239-0335/$09.75/0 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 leaves, is closely allied to E. brunnescens ssp. brunnescens of New Zealand. Progenitors of these species presumably arrived by long-distance dispersal via the prevailing westerlies. The remaining species are southern South American, and show no close affinities with species found in other geo- graphical areas. Fertility data from experimental hybrids suggest a close alliance between some of the morphologically distinct species in this group. Their precise relationships, however, and the origin of possible ancestors, remains obscure, although Australasia appears to be the most likely source. Three other species have been introduced into South America in historical times, two from Europe and one from North America. Epilobium, the largest genus in the Onagraceae, comprises approximately 185 species distributed worldwide in temperate zones on every continent except Ant- arctica. In tropical regions the species are restricted to temperate montane hab- itats. Many species are extremely variable and widespread, while others are high- ly restricted. In general, the genus has been characterized as ifc taxonomically difficult." During the last 20 years, a steadily growing number of papers have been published that provide modern treatments for the species of Epilobium encoun- tered in Europe (Raven, 1968), Africa (Raven, 1967b), parts of western Asia (Chamberlain & Raven, 1972; Raven, 1964) and eastern Asia (Hara, 1965; Raven, 1962, 1967a), Australasia (Raven & Raven, 1976), and North America (Munz, 1965; Hoch & Raven, 1981b). With the exception of some large parts of con- tinental Asia, the only region without a recent treatment is South America. Prior to the present study, the only papers dealing specifically with Epilobium in South America were those by Samuelsson (1923, 1930). Carl Haussknecht (1884) produced a monograph of worldwide scope, which is still the only com- prehensive treatment available. Samuelsson discussed a species only when new specimens amplified or diverged from the limits given by Haussknecht; he de- scribed new taxa based on new interpretations of morphology or material col- lected since Haussknechfs time. In several places Samuelsson pointed out the difficulty of separating some of the taxa he recognized, including some of his new ones, and this suggested that there were a number of taxonomic problems among the South American species of this genus. Since the early 1920s a substantial quantity of new herbarium material has become available, amounting to many times more than Samuelsson had for study. During these same intervening years, there have been fundamental changes in our concept of the kinds of units that should be regarded as species. A reassessment of the South American species of Epilobium in the light of these concepts was viewed as necessary to bring the species into conformity with the approach and philosophy used by Raven & Raven (1976) and Hoch (1978). Equally important are recent chromosomal studies (Seavey, 1972; Seavey & Raven, 1977a, 1977b, 1977c, 1978; Hair, Raven & Seavey, 1977), especially a preliminary survey of South American plants (Seavey & Raven, 1977c), in which it was suggested that, based on morphology and the distribution of chromosomal end arrangements, the Epilobium species in South America may have been de- rived from ancestors from both North America and Australasia. In order to un- derstand the implications of this interesting biogeographical problem, it was first necessary to delimit taxa that were present in South America. Once the taxa were 1982| SOLOMON— EPILOBIUM IN SOUTH AMERICA 24 1 circumscribed, additional studies would provide some understanding of the in- terrelationships among them. The following questions were kept in mind as this study progressed: How many taxa would it be useful to recognize? Did populations exhibit morphological patterns that had a geographical basis or did the same patterns appear numerous times in different areas? Are the taxa sympatric; do they hybridize, and, if so, what role has hybridization played in the differentiation of natural populations? Where did the taxa originate? What is their relationship with each other, and what impact did historical/geological processes have on their evolution and cur- rent distribution? An attempt has been made to answer these questions in greater or lesser detail, depending on the data that could be brought to bear on the subject. Materials and Methods An extensive field examination was made of hundreds of populations in the central and southern Andes of Argentina and Chile, and in the altiplano of Bolivia. To supplement the field observations, nearly all of the available herbarium spec- imens representing the genus in South America were studied. Seeds collected in the field, from herbarium specimens, or provided by cor- respondents, were grown in greenhouses at the Missouri Botanical Garden for observation. All plants grown from a single seed source, which may be one or more plants from one collection, are here defined as a strain. Many strains were used in making additional experimental hybrids to expand the studies reported by Seavey & Raven (1977c). These were examined cytologically. Buds were fixed in 1:3 acetic alcohol, hydrolyzed in 1 N HC1 for 10-15 minutes at 60°C, stained in acetocarmine and squashed for chromosome observations. In addition, a mea- sure of pollen fertility in the experimental hybrids and suspected natural hybrids was obtained using the malachite green-acid fuchsin-orange G stain of Alexander (1969). Two buds (or flowers, if buds were unavailable) were selected from each plant, pollen from each bud was stained and 100 tetrads scored for aborted and nonaborted grains. The scores for filled grains from the two buds were averaged to give a percentage of stainable pollen for each plant. Generally, two or more individuals of each artificial hybrid strain were averaged to arrive at the figures presented. In some instances, if only one bud was available, the percentage for that bud alone was used, or if there was a striking disparity between the two scores, the higher one was used, since it represents a potential fertility under good conditions. Generic Relationships Epilobium is one of two genera currently recognized in the tribe Epilobieae. This tribe differs significantly from the six other tribes of the family Onagraceae in that it has minute, heteropycnotic chromosomes (Kurabayashi et al., 1962); sheds pollen, in most species, as tetrads; has gametic chromosome numbers of /; = 9, 10, 12, 13, 15, 16, 18, and 19, and multiples of some of these numbers; and occurs primarily in moist or seasonally moist habitats (Raven, 1976). 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 In addition to Epilobium, Epilobieae contains the genus Boisduvalia, which is represented in western North America by five species, one of which is shared with South America, and a sixth species endemic to South America (Raven & Moore, 1965). Although Boisduvalia is clearly closely related to Epilobium, it differs in having an annual habit (only three species of Epilobium are annual); angular-fusiform seeds without a terminal coma (Seavey et al., 1977); and chro- mosome numbers of n = 9, 10, 15, and 19. Only n = 15 is shared with five species of Epilobium, but they have been derived independently of Boisduvalia (Raven, 1 976) . Epilobium itself is clearly delimited, although two genera, Zauschneria and Chamaenerion, are often segregated from Epilobium. Including these groups, Epilobium consists of six well defined sections (Raven, 1976). Section Cordylo- phorum, with three species, and sect. Zauschneria, with two, are closely related, share a number of morphological features, and have a base chromosome number of x = 15. Polyploidy is known in some subspecies of E. canum (Greene) Raven (sect. Zauschneria). Two other sections contain the only annual species in the genus, two in sect. Crossostigma and one, E. paniculatum Nutt. ex Torr. & Gray, in sect. Xerolobium. These sections are characterized by unusual mor- phologies and the chromosome numbers, n = 13 or 16 and n - 12, respectively. The center of distribution for these four sections lies in western North America. A fifth, very distinctive, group is sect. Chamaenerion. Unlike the preceding four sections, the center of distribution for the seven species of sect. Chamaenerion is in Eurasia, with only two, E. angustifolium L. and E. latifolium L., occurring in North America. Section Chamaenerion has a base chromosome number of -v = 18, with polyploids known only in E. angustifolium and E. latifolium. The sixth section, sect. Epilobium, accounts for the bulk of the genus. The five sections enumerated above contain only 15 species out of a total of approx- imately 185, leaving about 170 in sect. Epilobium, all of which have a gametic chromosome number of n = 18. These are widely, but unevenly, distributed in moist, open habitats throughout the temperate regions of the world. For example, Eurasia has more than 70 species, but Africa has only 1 1, with seven of these shared with Eurasia. There has been a remarkable radiation of this section in New Zealand unparalleled elsewhere in the world, resulting in 37 species, of which 31 are endemic and the remainder shared with Australia. The most prim- itive member of sect. Epilobium, E. rigidum Hausskn., is found in western North America, where four of the other sections are also restricted. This pattern of distribution, along with the great morphological and chromosomal diversity ex- hibited by these sections, leads to the conclusion that Epilobium probably orig- inated in western North America (Raven, 1976). The native South American species are all members of sect. Epilobium. In the present treatment 12 native species are recognized. These are distributed along the length of the Andes from Costa Rica, northern Venezuela and Colombia to Cape Horn, except for E. hirtigerum, which is restricted to eastern Argentina, Uruguay, and southern Brazil. The greatest concentration of species is found in the central and southern Andes of Argentina and Chile. In addition, three other species have been introduced and persisted or become naturalized in historical times. Epilobium obscurum and E. tetragonum subsp. lamyi, from Europe, are also 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 243 members of sect. Epilobium. The third species, E. paniculatum (sect. Xero- lobium) has been introduced from North America. A fourth species, E. au- gust if olium, representing sect. Chamaenerion, has been collected once in Chile (Valdivia, Valdivia Prov., Chile, Calvert in 1914, BM). If this specimen is correctly labeled, then the plant was probably cultivated and has failed to persist or become naturalized. In the absence of additional information, it has been excluded from further consideration. Haussknecht (1884) grouped the species of Epilobium into assemblages based on stigma lobing, seed shape, and other conspicuous morphological features. Samuelsson (1923) maintained these groups for the South American species as outlined by Haussknecht and created two additional ones by removing the taxa allied to E. australe from the Platyphylla, placing them in his Australia, and by segregating E. nivale and E. fragile as Nivalia. Although Haussknecht's cate- gories did bring together species of similar morphologies, the arrangement of the groups and some of their component species do not reflect our current under- standing of the genus. Until a much more detailed understanding of the reticulate morphological patterns seen in sect. Epilobium is acquired, it does not seem warranted to recognize any assemblages such as those proposed by Haussknecht. Morphology In dealing with the South American species of Epilobium sect. Epilobium, a variety of morphological characters have been useful in delimiting the species, especially habit, perennating structures, pubescence, leaf shape, dentition and arrangement, and seed shape and surface features. Habit. Epilobiums in South America exhibit a diverse array of habit and growth form. Some species have erect or ascendent stems (e.g., E. ciliatum, E. glaucum); others are lax or decumbent (E. nivale), and often caespitose (E. den- sifolium, E. fragile, E. nivale). Perhaps the most unusual growth form is that of creeping stems, rooting along their length and with flowers arising from subter- minal nodes, characteristic of E. conjungens. This habit is shared only with a number of species from Australasia (Raven & Raven, 1976). All the species in sect. Epilobium are perennial herbs. They produce a variety of perennating parts for vegetative reproduction and persistence during the win- ter. All of the South American species will germinate, flower, and set seed during a single growing season, so it is potentially possible for them to be facultatively annual. This does not, however, seem to be an important factor except, perhaps, in those species that show the greatest tendency to weediness (e.g., E. ciliatum). There are very great differences in branching patterns, from simple to pro- fusely branched. This trait is so variable within species or even populations, however, and so strongly influenced by the age of the plant and the microclimatic conditions under which it grows, that it is of little taxonomic value. In the Northern Hemisphere, a wide variety of highly specialized perennating structures has evolved, including compact underground buds with fleshy, imbri- cate cataphylls (turions), leafy rosettes, rhizomes, corms at the tips of thin sto- lons, and leafy soboles or shoots. Only Epilobium ciliatum, of the native South American species, produces turions or leafy rosettes, usually toward the end of 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 69 a growing season. In general, the other species are less specialized in this respect, reproducing vegetatively by buds at or near the ground surface or by more or less elongated leafy shoots from the bases of the stems. The leafy shoots in E. barbeyanum often take the form of runners 20 cm or more long. Two species, E. australe and E. glaucum, produce short or elongate scaly rhizomes just under the ground surface, which, in the case of E. glaucum, may be up to 20 cm long. These various modes of perennation generally result in clumped groups of stems, although solitary, single-stemmed individuals can be found in many species. Epi- lobium ciliatum is usually solitary, since each plant produces only one or two turions or rosettes, which give rise to the following year's plant. As previously mentioned, E. conjungens is quite unusual among the South American species. Its stems form small mats of repent branches, a habit that has apparently evolved through the suppression of the main shoot with a concomitant proliferation of basal shoots (Raven & Raven, 1976). A conspicuous feature of two of the clumped, caespitose species, E. densi- folium and E. fragile, is the thick, woody, often elongate rootstock. This is in striking contrast to the normally fibrous root system produced by other species. Pubescence. There are four basic types of hairs found among the species under consideration. The presence or absence of each type has been very useful, either alone or in combination with other characters, in separating taxa. Two species, E. glaucum and E. nivale, are conspicuous by the absence of pubescence of any type, and the often blue-glaucous, waxy bloom in the former. Very rare glabrous individuals of E. ciliatum have been seen, but normally this species is pubescent. 1. Strigillose. Hairs of this type are sharp-pointed, appressed, or at least fal- cate, upwardly curved, and typically 0. 1-0.2 mm long. Following Raven & Raven (1976) and Hoch (1978), these hairs have been termed strigillose. This is by far the most common pubescence type found in Epilobium. The density and distri- bution of strigillose hairs varies considerably within and between species, but they are found in nearly all species except for the glabrous ones mentioned above. There is some variation in how closely appressed the strigillose hairs are on an individual plant. Often the hairs on the ovaries are closely appressed, while those elsewhere on the plant are more spreading. In E. puberulum the basic strigillose hair type is generally longer, ranging from 0. 15 to 0.4 mm, and often somewhat spreading. A second modified hair type occurs in E. conjungens. In this species the hairs are much smaller, 0.02-0.08 mm long, and often curled in dried specimens. It is not known if these hairs are straight in living plants, but because they are distinctive, this type of pubescence has been termed puberulent. 2. Glandular. These hairs are more or less appressed to erect, blunt-tipped, 0. 1-0.2 mm long. In living plants each hair usually exudes a minute droplet of liquid. When they are sufficiently abundant, as in E. barbeyanum and some pop- ulations of E. ciliatum, the entire plant may feel moist or viscid to the touch. Normally, when dried, these hairs become flat and twisted with an evident round- ed tip. Glandular hairs are absent in E. australe, E. densifolium, E. puberulum, and E. conjungens. In the other species their density and distribution vary con- 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 245 siderably, but they are generally present in the inflorescence, on the ovaries and floral tube, and along the margins of young leaves. 3. Long, erect. A distinction is made here between two types of erect hairs that are each restricted to a single species. The long, very fine, soft, erect hairs found in E. hirtigemm measure between 0.25 and 0.6 mm long and are designated as villous. A number of populations of E. denticulatum from southern Peru and western Bolivia contain plants that were described by Samuelsson (1923) as E. hirtum. These are characterized by long erect or spreading hairs, 0.2-0.4 mm long, which are here termed hirsute. Long hairs of this type intergrade continuously with the slightly shorter strigillose pubescence mentioned earlier, both in length of the hairs and in the degree to which they are appressed. 4. Appressed, blunt-tipped. These are generally small, 0.05-0.1 mm, blunt- tipped hairs that are tightly appressed to both surfaces of the leaf blade. They have been found in only E. denticulatum and E. pedicellare. The distinction between these hairs and the glandular ones discussed earlier is very fine. The appressed hairs are smaller but merge imperceptibly with the glandular ones in the region of the petiole and lower leaf blade. In living plants, the appressed hairs never produce an exudate. Older leaves are often glabrate, so the appressed hairs may be found only on the younger leaves. In plants of E. denticulatum that are hirsute, the hairs on the leaves may also be erect, and are perhaps under the same type of genetic control. Leaves. The basic leaf type found in South American species is lanceolate, thin, and acute at the apex. In a few instances, however, species deviate from this shape sufficiently for the leaf outline to be a useful taxonomic character. Ovate, thick leaves are typical in E. australe, and elliptic to orbicular blades are found in E. conjungens. The type and size of teeth on the leaf margins can also be helpful. The majority of species have few, denticulate teeth. In contrast, the margins of leaves in E. ciliatum are usually serrulate with numerous teeth. Epilobium pedicellare is quite unusual in having leaves with coarsely serrate margins, the teeth of uneven sizes. The few, narrow, sharply pointed, forward-directed teeth in E. hirtigemm are also diagnostic. These last two species are also distinctive in their mostly alternate leaves. Other species normally have opposite leaves, or leaves that are alternate only in the inflorescence. Inflorescences. Inflorescences in Epilobium are only partially useful as a diagnostic feature. They may vary from simple to paniculate and densely branched. The nature of the inflorescence depends upon the branching pattern exhibited by the individual plant. In general, inflorescences are terminal, erect, and highly variable in the total number of flowers produced during a growing season. Each branch of the inflorescence normally presents only one or a few flowers each day, which only last for that given day. As an example, E. densifolium seldom produces more than five or six flowers per stem during an entire season. A single plant, however, may bear many more flowers than this because of its densely branched caespitose habit. Flower production in other species is usually much 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 more prolific. For example, a large plant of E. ciliatum may produce several hundred or more flowers in a single season. The nodding inflorescence and flowers of E. denticulatum are very characteristic, although this feature may be lost in preparing specimens or if the plants are collected very late in the growing season. Epilobium conjungens exhibits one of the most specialized inflorescence types in Epilobium. Instead of producing a discrete, terminal, several-flowered inflo- rescence, it bears only a few flowers singly along the stem, which continues to grow and root at the tip. Flowers. Most Wh hirtigerum, E. conjungens, many populations of E. ciliatum, and occasionally in E. denticulatum. Epilobium puberulum is unique in having salmon pink petals. Flower size, color, and morphology alone are generally not sufficient to separate any species but are useful in combination with other characters. The only excep- tion to this is E. puberulum, which has a very distinct morphology as well as petal color. At anthesis the petals of E. puberulum are imbricate and each one is bent at about 90° a short distance above its insertion on the floral tube. Thus, the four petals produce a flat plane at right angles to the axis of the floral tube, with a small circular opening formed by the bends in the petals. The stigma is posi- tioned in this opening at about the same level as the bend in the petals, with the dehisced anthers of the four longer stamens pressed between the stigma and the petal bend. The four antipetalous stamens have extremely short filaments, usually less than 0.6 mm long, much shorter than in any other species. Among the other species, there is considerable variation in petal size, with certain ranges characteristic of a few species, such as E. densifolium, which has very large petals, but the magnitude of overlap and the variability of some species precludes the use of flower size by itself as a taxonomic character. Another unique floral feature is the lavender or bluish staminal filaments found in E. denticulatum and E. pedicellare. Additional notes on floral features are discussed under floral biology. Seeds. The morphology of seeds has been considered a useful feature in the taxonomy of Epilobium for a very long time (e.g., Haussknecht, 1884; Samuels- son, 1923). Within recent years several surveys of seed surface morphology using scanning electron microscopy have been conducted by Berggren (1974), Raven & Raven (1976), Hoch (1978), and Seavey et al. (1977). The last study is of particular importance because it is a worldwide survey of all the sections and includes species from all the continental areas where Epilobium occurs. Three basic types of seed surfaces are recognized by Seavey et al. (1977). Papillose: each cell has a more or less prominent, regular or irregular convex portion in the center of the cell. Ridged: this is a specialization of the papillose type in which the papillae are flattened laterally and fused or nearly fused end- to-end in longitudinal rows to form ridges. Reticulate: the surface cells show only a regular polygonal reticulum formed by the radial walls, without a papilla. Nearly all of the species of Epilobium in South America have papillose seed surfaces, which is the typical surface morphology for the majority of species in 1982] SOLOMON— EPILOB1VM IN SOUTH AMERICA 247 section Epilobium (Seavey et al., 1977). Four species have been illustrated in Seavey et al. (1977): E. denticulatum (Figs. 64-66; as E. hirtum, Figs. 73-75), E. australe (as E. sp., Figs. 70-72), E. densifolium (as E. cf. pauciflorum, Figs. 61- 63), and E. glaucum (Figs. 67-69), and a fifth, E. hirtigerum in Raven & Raven (1976, Fig. 3b). These examples represent the full range of papillose seeds found in South America. Epilobium ciliatum is the only species in South America that has nonpapillose seeds. This species produces conspicuously ridged seeds, a type known only in E. ciliatum and the relictual western North American E. oreganum Greene, both members of the E. ciliatum complex (Hoch, 1978). Seeds from South and North American populations are illustrated in Seavey et al. (1977, Figs. 169-174). Fo- veolate cells are also found on the seeds of E. ciliatum, but such seeds do not characterize any distinct taxonomic group. Some populations of E. ciliatum from southern Patagonia and Tierra del Fuego may have at least part of the upper seed surface with a reticulate pattern. The area covered by foveolate cells varies in size, but the seeds always have longitudinal ridges along the sides and at the micropylar end. This situation is somewhat analogous to that reported by Hoch (1978) for some populations of E. ciliatum subsp. glandulosum but has not proceeded to the complete elimination of the ridges. Seed length is variable, ranging from 0.7 to 2 mm, and is somewhat useful as a diagnostic feature. The majority of species have seeds between 0.8 and 1.4 mm long, with substantial variation within species. Epilobium puberulum has consis- tently small seeds, 0.7-0.9 mm long, and E. densifolium has the largest seeds, 1.4-2 mm long. Except for the largest seeds off. densifolium, seed size overlaps too broadly to be used alone to separate taxa. An additional useful character is the short, often rather broad, pellucid cha- lazal appendage found in six species, E. ciliatum, E. denticulatum, E. pedicellare, E. fragile, E. puberulum, and E. nivale (cf. Seavey et al., 1977; Hoch, 1978). Cytogenetics and Crossing Relationships Cytogenetics. All of the species of Epilobium sect. Epilobium that have been examined cytologically have a gametic chromosome number of n = 18. Num- bers for South American species, some of which are reported here for the first time (Table 1), conform to this pattern, which is based on studies of hundreds of strains from all parts of the world (e.g., Seavey & Raven, 1977a, 1977b, 1977c). Chromosomes in Epilobium are uniform in morphology. At meiotic metaphase I they are small (2-4 fim long), heteropycnotic, and exhibit no evident karyotypic markers; they are indistinguishable from one another. In 1968, Mosquin found evidence for reciprocal translocations while exam- ining chromosomal pairing in experimental hybrids between several species. These initial results have been elaborated on considerably by Seavey (1972), Seavey & Raven (1977a, 1977b, 1977c, 1978), and Hair, Raven, & Seavey (1977), so that the number and distribution pattern of reciprocal translocations in section Epilobium is now known in some detail. Three major species groups, which account for nearly all species in sect. Epilobium, have been detected on the basis of reciprocal translocation differ- 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol . 69 Table I. Chromosome numbers in South American species of Epilobium. All strains formed 18 bivalents in meiosis. Vouchers are deposited at the Missouri Botanical Garden (MO) unless oth- erwise indicated. Full localities listed under respective species. E. australe. Chile, Marticorena et al. 475, 834. E. barbeyanum. Chile, Solomon 4285, 4296. E. ciliatum subsp. cilia turn. Argentina, Solomon 4712. Chile, Eyerdam 10296 (NY); Marticorena et al. 852, 970 ; Moore 412 (LA). E. denticulatum . Bolivia, Solomon 4938. Ecuador, Berry 2522 ; Weydahl 153 (S). E. glaucum, Chile, Marticorena et al. 441, 991 . E. hirtitferum. Argentina, Krapovickas & Cristobal 14675. E. nivale. Argentina, Gentili in 1975. E. obscurum. Chile, F. Schlegel 6882. E. pedicellare. Bolivia, Solomon 5147. E. puberidum. Chile, Marticorena et al. 101 5 ; Moore 296 (LA). ences. The chromosomal arrangements that characterize each group have been given the arbitrary designations of AA, BB, and CC (Seavey, 1972; Raven, 1972). Hybrids within chromosomal groups produce 18 pairs of chromosomes at meiotic metaphase I, but those between groups produce various combinations of pairs and rings and/or chains. The BB arrangement is considered to be the ancestral condition in sect. Epi- lobium for the following reasons. First, one of the two most primitive species in section Epilobium, E. obcordatum A. Gray from western North America (Raven, 1976), has this arrangement. Second, the other two arrangements, A A and CC, each differ from the BB by a single reciprocal translocation. AA and CC differ from each other by two overlapping reciprocal translocations. Thus, hybrids be- tween plants with the A A and CC arrangements have 15 pairs and a ring or chain of 6 chromosomes at meiotic metaphase I, while hybrids between plants with the AA or CC arrangements and those with the BB arrangement produce 16 pairs and a ring or chain of 4. The simplest hypothesis for the derivation of these chromosomal groups is that the AA and CC arrangements were each derived independently from the BB arrangement. Third, the BB chromosome arrangement characterizes the largest number of species in sect. Epilobium and is the most widespread of the three. It is the common arrangement in Eurasia and Africa and the only one found in Australasia. A number of South American species have this arrangement, and it distinguishes a large group of species in North America. The A A arrangement is primarily restricted to the New World, where it is known from the E. ciliatum complex, consisting of five species centered in west- ern North America, several species in South America, and three apparently un- related species in Europe, E. alpestre (Jacq.) Krock., E. alsinifolium Vill., and E. atlanticum Litard. & Maire, the latter also found in the Atlas Mountains of Morocco. The third arrangement, CC, is found in a series of closely related, circum- boreal species, including E. hornemannii Reichenb., E. anagallidifolium Lam., and E. clavatum Trel., among others. Also found to have the CC arrangement is the distinctive western North American species E. luteum Pursh, which has four- lobed stigmas and cream-colored flowers. Besides these three arrangements, two additional unique ones have been dis- 1982] SOLOMON— EP1LOBIUM IN SOUTH AMERICA 249 Table 2. Strains of Epilobium used in artificial hybridization experiments. Vouchers deposited at MO; full collection information given under respective species. Acronyms and strain letters are those used in Tables 3 and 4, and Figures 1 and 2. If the name given in Seavey and Raven (1977c) differs from the nomenclature used here, that name has been placed in parentheses after the corre- sponding strain. E. australe. (AUST) (a) Argentina, Moore 1686 (E. sp.). (b) Chile, Marticorena et al. 834. (c) Argentina, Rubulis in 1976. E. barbeyanum. (BARB) Chile, Solomon 4296. E. ciliatum subsp. cilia turn. (CIL) Chile, Zollner 7868 {E. chilense Hausskn.). E. ciliatum Raf. subsp. watsonii (Barbey) Hoch & Raven. (WAT) U.S.A., California, Sharp in 1967 (E. watsonii Barbey). E. densifolium. (DENS) (a) Chile, Zollner 6245 (E. cf. pauciflorum F. Phil.). (b) Argentina, Diem 3600. E. denticulatum. (DENT) (a) Argentina, Hunziker & Ariza 20424. (b) Peru, Conrad 2715. (c) Peru, Averett 1004 (E. hirtum Samuelsson). E. xlaucum. (GLAU) (a) Chile, Marticorena et al. 7 . (b) Chile, Marticorena et al. 951 . E. hirtixerum. (HIRTIG) (a) New Zealand, Raven 25148. (b) Argentina, Krapovickas & Cristobal 14675. (c) Argentina, Troncoso et al. 2462. (d) Argentina, Solomon 4131. E. nivale. (NIV) Argentina, Gentili in 1975. E. obscurum. (OBS) (a) Spain, Raven 26069. (b) Chile, Ramirez in 1975. E. puberulum. (PUBER) Chile, Marticorena et al. 1015. covered. The first, termed DD, is found in Epilobium duriaei Gay ex Godron from Europe (Seavey & Raven, 1977a); the other, termed EE, is found in E. platy stigmata sum from eastern Asia (Seavey & Raven, 1978). Each of these arrangements differs from the BB by a single reciprocal translocation, and each is different from the A A and CC arrangements. Like the latter two arrangements, DD and EE were probably independently derived from the BB type. The chromosomal arrangements and some of the crossing relationships within the South American species have now been elucidated. Most of the strains used by Seavey & Raven (1977c), plus a number of additional ones, including several previously uncultivated species, were grown and experimentally hybridized. The hybrid progeny were then examined for chromosomal arrangement and pollen stainability. The strains used in the crossing experiments are given in Table 2, and the results from the hybrids are shown in Tables 3 and 4. As in previous studies of chromosomal arrangements in Epilobium, beginning with those of Seavey & Raven (1977a) and continuing to the present, all of the strains were standardized by crossing them with a single strain of E. ciliatum 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 3. Experimental hybrids between species of Epilobium. Acronyms and lettered strains refer to Table 2. Female parents are listed first, in alphabetical order, followed by the male parent. Data for voucher numbers below M600 are taken from Seavey and Raven (1977c). All vouchers are deposited at MO; full collection information given under respective species. Pollen Cross Chromosomal Configuration Stainability % Greenhouse Voucher AUSTa x CIL 16„ + ch4 BB X AA 32 M465 DENTb 16,, + r4 BB X AA 45 M435 DENTc I6„ + ch4 BB X AA 39 M467 GLAUa 18„ BB X BB 65 M437 WAT I6„ + r4 BB X AA 26 M469 AUSTb x DENTc BB X AA 8 M 1 542 AUSTc x DENTc BB X AA II Ml 557 PUBER 16,, + r4 BB X AA 38 M 1 562 BARB x DENTc 16„ + r4 BB X AA 12 M1948 GLAUa 18„ BB X BB 69 M 1 953 CIL x AUSTa 16„ + r4 AA X BB 27 M466 BARB 16„ + r4 AA X BB 33 M 1 954 DENTa I8„ AA X AA 37 M455 DENTb 18„ AA X AA 51 M 1 540 DENTc 18„ AA X AA 56 M1538 GLAUa I6„ + ch4 AA X BB 23 M443 WAT 18„ AA X AA 86 M 1 5 39 DENSa x GLAUa 18„ BB X BB 88 M44I OBSa 18m BB X BB M575 DENSb x GLAUa 18„ BB X BB 92 M 1 552 DENTb x AUSTa 16„ + r4 AA X BB 45 M436 CIL 18„ AA X AA 47 M456 DENSa 16„ + r4 AA X BB 15 M452 GLAUa I6„ + ch4 AA X BB 41 M439 OBSa 16„ + r4 AA X BB 33 M459 WAT 18„ AA X AA 24 M457 DENTc x AUSTa I6„ + ch4 AA X BB 34 M468 AUSTb AA X BB 7 M154I AUSTc 16„ + r4 AA X BB 25 M 1 556 CIL 18„ AA X AA 64, 61 M1537, M1964 BARB AA X BB 21 Ml 949 DENSb 16,, + r4 AA X BB 27 Ml 550 GLAUa I6„ + ch4 AA X BB 29 M444 OBSb AA X BB 31, 18 M1561, MI967 GLAUa x AUSTb AUSTc CIL DENSb DENTb DENTc PUBER GLAUb x AUSTc NIV x DENTc GLAUa OBSa x DENSa DENTb GLAUa PUBER x BARB GLAUa WAT x AUSTa DENTb 18„ I6„ + r4 n I6„ + r4 BB x BB BB x BB BB x AA BB x BB BB x AA I6„ + r4 BB x AA BB x AA 18„ BB x BB 16„ + r4 BB x AA 18„ BB x BB I8„ BB x BB I6„ + ch4 BB x AA I8„ BB x BB 16„ + r4 AA x BB I6„ + r4 AA x BB I6„ + r4 AA x BB 18„ AA x AA 84 97 22 87 46 24 44 83,68 13 23, 12 36 21 56 47 24 37 M 1 543 MI558 M442 M 1 55 1 M440 M445 M 1 546 MI559, Ml 966 M 1 554 MI555, M1965 M576 M460 M448 M 1 952 M 1 547 M470 M458 1982] SOLOMON— EP/LOBIUM IN SOUTH AMERICA 25 1 Table 4. Experimental hybrids involving Epilobium hirtigerum. Acronyms and lettered strains refer to Table 2. Female parents are listed first, in alphabetical order, followed by the male parent. Data for voucher numbers below M600 are taken from Seavey and Raven (1977c). All vouchers are deposited at MO; full collection information given under respective species. Pollen Greenhouse Cross Chromosomal Configuration Stainability % Voucher DENTa x HIRTIGb 14„ + r4 + ch4 AA x B.B, 9 M462 GLAUa x HIRTIGb I6„ + r4 BB x B,B, 9 M451 HIRTIGa x HIRTIGb 16„ + r4 BB x B,B, HIRTIGb x DENTa I4„ + 2 r4 B,B, x AA 14 M463 GLAUa 16„ + r4 B,B, x BB 7 M450 HIRTIGc 16„ + r4 B,B, x BB 38 M1958 HIRTIGd 16„ + r4 B,B, x BB 52 M1955 HIRTIGc x OBSa 18„ BB x BB 64 M1959 HIRTIGd x OBSa 18„ BB x BB 60 M1960 PUBER x HIRTIGc AA x BB 1 M196I subsp. watsonii (WAT; AA), E. obscurum (OBSa; BB), or with another strain that had previously been standardized to the first two. Only the AA and BB chromosomal arrangements are present in South Amer- ica. Their distribution among the species is summarized in Table 5. The rationale for the suspected arrangement of the three unknown species is discussed in the section on biogeographical relationships. Epilobium puberulum is presented as having the A A genome. When this species was crossed with E. glaucum, a known BB, the resulting hybrids produced 16 pairs and a ring of four chromosomes at meiotic metaphase I. There is no evidence to suggest that E. puberulum has an unusual chromosome arrangement, so it will probably be found to have the AA arrangement. The definitive crosses to known A As have been made but not yet analyzed. Epilobium hirtigerum initially presented a perplexing chromosomal situation. The original strain that was tested (HIRTIGb) was found to differ from the BB arrangement by one reciprocal translocation and from the AA by two. The ques- tion was posed as to whether this unusual arrangement characterized all popu- lations of E. hirtigerum in South America (Seavey & Raven, 1977c). Two addi- tional populations, one (HIRTIGd) from the same general area, and a second from elsewhere in the range (HIRTIGc), were tested against the original strain and a known BB. The results of these crosses are given in Table 4. It now appears likely that the BB arrangement is the normal one for E. hirtigerum in South America, as it is in Australasia, and that the original strain tested is atypical. Plants with anomalous chromosomal arrangements, such as that illustrated by E. hirtigerum, are now known in four species. In E. microphyllum A. Rich, and E. palustre L., single individuals have been found that differed from the BB arrangement by one reciprocal translocation. Other plants of these species from the same or nearby populations, however, had the BB arrangement (Raven, 1972; Seavey & Raven, 1977a). In a similar fashion, a single individual of E. glaberri- mum Barbey had an arrangement that differed from AA by one reciprocal trans- location, but plants from the same and a number of other populations were all 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Voi . 69 Table 5. Distribution of chromost )m; x\ arrangements in native South American species ot / ipi- labium. See text for further explanation of species marked with asterisk (*). Species Arrangement E. deniicu latum AA E. puberulum AA* E. ciliatum AA E. nivale BB E. barbeyanum BB E. densifolium BB E. australe BB E. glaucum BB E. hirtigerum BB* E. pedicellare Not known, probably AA E. fragile Not known, probably AA E. conjungens Not known, probably BB A A (Seavey & Raven, 1977a). It seems clear that these chromosomally differ- entiated plants probably arose spontaneously within populations. Events of this type provide an example of how the major chromosomal lines might have arisen and how differentiation between species could have proceeded. Crossing relationships. It has been well established that there are essentially no barriers to the production of viable seed between members of Epilobium sect. Epilobium, even among geographically and morphologically separated species or between chromosomal groups (e.g., Brockie, 1970; Seavey & Raven, 1977a), although various types of genetic or cytoplasmic incompatibilities may seriously disrupt development or fertility after germination (Thakur, 1965). The failure to set seed in interspecific crosses may occur due to poor pollen tube growth, or failure of fertilization, but these are generally not important factors (Raven & Raven, 1976). The results of this study support this broad conclusion. Seed set may be reduced in artificial crosses, but this is often due to technical difficulties, such as damage to small flowers when they are emasculated, transfer of an in- adequate quantity of pollen, or use of pollen from a short-styled species to cross with a long-styled one. In all cases, at least some seed was produced in inter- specific crosses. The hybrids that were made with the South American species, with few ex- ceptions, showed no developmental abnormalities or reciprocal differences such as those that have been reported by Thakur (1965), Michaelis (1954, 1965), and Raven & Raven (1976). Hybrids between E. obscurum, a European species, and E. densifolium, E. glaucum, and E. australe were all initially dwarf. Only one plant of the hybrid combination E. obscurum x E. glaucum eventually flowered vigorously. The other combinations either died or flowered only sporadically (Seavey & Raven, 1977c). Similarily, hybrids between E. denticulatum and E. obscurum were also initially dwarf, and only two individuals eventually produced normal branches and flowered. These hybrids, which involved both the AA and BB chromosomal groups (Table 5), would strongly suggest some type of genetic or cytoplasmic incompatibility specifically with E. obscurum, although its precise nature is not known. The hybrid between E. nivale and E. glaucum may also 1982) SOLOMON— EPILOB/UM IN SOUTH AMERICA 253 have been disturbed developmentally. Although the plants were vigorous vege- tatively, they produced very few flowers. Alternatively, this could be the result of the relatively low number of flowers that E. nivale normally produces under greenhouse conditions. In the absence of obvious developmental barriers to hybridization, one must utilize other data to assess the degree of relationship between species. Besides the chromosomal arrangements that divide the South American species into two groups, hybrid fertility, as indexed by the percentage of stainable pollen (Table 6), can also be used as a measure of relationship. This data must be used cau- tiously, however, because pollen stainability can vary considerably from plant to plant and even between flowers on the same plant, and it is often greatly affected by the age of the plant and the cultural conditions under which the plant has been grown. The pollen fertility of most non-hybrid plants used in the experimental hy- bridizations was greater than 85%, with two exceptions, Epilobium denticulatum (DENTc), which had 75%, and E. barbeyanum (BARB), which had 54%. Even crosses made between strains of the same species from widely separated geo- graphical areas had high fertility (e.g., E. denticulatum: Averett 1004, Peru x Billings 161 , Venezuela, 94%; E. ciliatum subsp. ciliatum, Zollner 7868, Chile x E. ciliatum subsp. watsonii, Sharp in 1967, California, 86%). The much reduced pollen fertility seen in Table 6 between species with dif- ferent chromosomal arrangements is due primarily to the high frequency of ad- jacent disjunction of the chromosomes involved in the ring of four during meiosis. The consequent duplications and deletions resulted in a substantial number of aborted gametes (Seavey, 1977). Hybrids between species with the same chro- mosomal arrangement that show reduced fertility must have other factors in- volved. Epilobium ciliatum and E. denticulatum, both with the A A arrangement, have differentiated genetically and produce hybrids with substantially reduced fertility. Within the group of species with the BB arrangement, E. glaucum when crossed with E. australe or E. densifolium produces hybrids with variable, but sometimes very high, fertility, up to 97% and 92% respectively (Table 6). These values are similar to those reported for hybrids between many closely related species in Australasia (Raven & Raven, 1976). This suggests that these three species are fairly closely related to one another, although they are morphologically distinct. The relationship of E. barbeyanum to these species is still obscure. Epilobium nivale, on the other hand, is quite strongly differentiated from E. glaucum, either genetically or cytoplasmically, as indicated by the very low hybrid pollen stain- ability and the disturbed development mentioned earlier. Its relationship with the other species remains unknown. BlOGEOGRAPHICAL RELATIONSHIPS The morphological and cytological evidence presented previously suggests that there have been a minimum of five pre-European introductions of Epilobium into South America with subsequent diversification in some of these lines. The probable source of progenitors for each group and the relationships of the cur- 254 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 X) o E c £ < in X) rt c E x c o a. V ed c <3 C/5 c cu UJ -J CQ < Q < a z < H GO OQ < < < C/3 QQ o X < a < Z a CQ < CQ > z J u UJ CQ cu Z uj Q rt rj 00 oo'^ sO 3 1— • m ri Tf (N ^^» on r- m ^ ^ on on ^r CN sC 00 r\ — 00 ^o r^ oc ON (N r4 ^r Z UJ Q < < 8 3 CI OC sO rn »0 rn r i ON 5 m r i rn oo' ON sO sO UJ cu U > Z CQ 00 H P s Z CO < < UJ D J CQ a < a c* X QQ O CQ CQ C/3 a < z < 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 255 Table 7. Distribution and probable sources of progenitors for related groups of native species of Epilobium in South America. AA and BB refer to chromosomal arrangements; see text for further discussion. Source of Group/Species Distribution Progenitor Group I (AA) North America E. denticulatum Andes, Venezuela to northern Argentina and Chile; Costa Rica E. fragile Altiplano of Peru and Bolivia E. pedicellare Altiplano of Peru and Bolivia E. puberulum Central Chile Group II (AA) E. ciliatum Southern Andes of Chile and Argentina, Falkland Islands; widespread in North America Group III (BB) E. australe E. barbeyanum Southern Andes of Chile and Argentina E. densifolium E. glaucum E. nivale Group IV (BB) E. hirtigerum Eastern Argentina, Uruguay and southern Brazil; North America Australasia Australasia widespread in Australasia Group V (BB) Australasia E. conjungens Tierra del Fuego and adjacent islands rently recognized species are summarized in Table 7 and discussed in detail be- low. The two groups with the AA chromosomal arrangement almost certainly have ancestral affinities with North America. These are the only species with the AA arrangement in the Southern Hemisphere. The other species with this arrange- ment are found primarily in North America, with only three isolated species in western Europe. Epilobium denticulatum is the most widespread and variable species of the first group in Table 7, and highly characteristic of the paramos and puna through- out the northern and central Andes. Although E. pedicellare and E. fragile are unknown chromosomally, they appear to be closely allied to E. denticulatum on morphological grounds, and will probably be found to have the A A chromosomal arrangement. Epilobium pedicellare shares with E. denticulatum the feature of appressed, blunt-tipped hairs on the leaf surfaces, a similar habit, and completely overlapping geographical ranges. The relationships of E. fragile are somewhat more problematical because of its reduced stature and restriction to extremely high elevations, but it is probably closely related to and may be directly derived from E. denticulatum. The geographical range of E. fragile is also completely contained in that of E. denticulatum, and some small specimens of E. denticu- fi frag three species are the only ones found in the northern and central Andes; all of 256 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 the others are restricted to areas mostly south of 30°S latitude. Epilobium pu- berulum, although isolated geographically in central Chile from the species dis- cussed above, is morphologically similar to some populations of E. denticulatum and has often been confused with that species. The precise relationships between these four species is not clear, and none of them resemble any extant North American species, but North America is the only reasonable source for progenitors, which probably arrived in South America by long-distance dispersal. The ancestor of these species was probably an early offshoot of the AA chromosomal line that gave rise to the members of the E. cilia turn complex in North America (Hoch, 1978). The second AA chromosomal group consists only of Epilobium ciliatum subsp. cilia turn. The South American populations, which are found in areas south of 30°S, have gone under various names, most often E. chilense, E. valdiviense, or E. magellanicum. Only recently has the question been raised as to whether pop- ulations from South America were closely related to ones from North America (Seavey et al., 1977; Seavey & Raven, 1977c), although some South American plants had been identified as North American species and vice versa many years ago (Haussknecht, 1879, 1884). In 1977, Seavey et al. discovered that the seed surface morphology of plants identified as Epilobium chilense were identical with those found in E. ciliatum in North America. As was discussed earlier, this seed type is restricted to two species in North America. In addition, these South American plants produced turions or leafy rosettes, overwintering structures that are unique in South Amer- ica but are common in a number of species in North America. An analysis of the E. ciliatum complex in North America by Hoch (1978) provided additional mor- phological features as a basis for comparison. In this study he recognized five species, including E. glaberrimum Barbey with two subspecies and E. ciliatum with three subspecies. A careful comparison of plants from many widely sepa- rated South American populations with others from North America indicated that there were few differences between them and that all the South American pop- ulations could easily be accommodated in E. ciliatum subsp. ciliatum as circum- scribed by Hoch. In fact, the variability seen in South America is only a part of that exhibited by this subspecies in North America, where it is extremely diverse and widespread. Epilobium ciliatum in South America has been derived from North America and may have originated from a single introduction. There is an unlikely possibility that Epilobium ciliatum arrived in South Amer- ica as an unintentional introduction by man. Beginning in the 1760s, with the founding of a number of missions in Alta California, increased commerce began between the western coast of North America and Peru and Chile. Prior to this time, only rare and casual visits had been made to North American areas where E. ciliatum occurs. Thus, the most likely time of introduction would have been after this date. The first South American collections of E. ciliatum were made about 1793 in central Chile, and by 1833 in Tierra del Fuego, a period of only 40 years. Considering the extensive variation and presence of E. ciliatum in many areas that were remote from human habitation over a geographic distance of 2,700 km in South America, it seems unlikely that £. ciliatum has been introduced within the past 200 to 300 years. Rather, its occurrence in South America is 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 257 probably the result of a pre-European long distance dispersal event (cf. Raven, 1963). The species with the BB chromosomal arrangement, however, probably have their affinities with Australasian species. They have been divided on the basis of morphology into three groups, one with five species, the other two with one each, representing three independent introductions (Table 7). Epilobium australe, E. barbeyanum, E. densifolium, and E. glaucum are fairly closely related, with E. nivale being somewhat anomalous, but also probably allied. These species are morphologically unlike any North American species with the BB chromosomal arrangement, but they also show no obvious affinities with any extant species in Australasia. The habit, simple leafy shoots as perennating structures, and the rather broadly obovoid seeds (cf. Raven & Raven, 1976) suggest an Australasian relationship. Epilobium hirtigerum is conspecific with a widespread species of Australia and New Zealand. For many years the South American populations were called E. brasiliense. However, during the intensive study that led to their monograph of Epilobium in Australasia, Raven & Raven (1976) discovered that the South American plants were indistinguishable from those found in many populations in Australasia, especially New Zealand. The examination of a large sample of spec- imens from South America for the present study confirms this conclusion. Phil- ibert Commerson made collections of this species in Uruguay two years prior to the first known European landings in eastern Australia or New Zealand by Cap- tain James Cook in 1769. Unless some unknown, fortuitous historical event aided introduction of this species, it seems highly probable that E. hirtigerum arrived in South America by long-distance dispersal. It also seems likely that the present populations were derived from a single introduction from Australasia. The third group contains only Epilobium conjungens. This species has what can be considered the most unusual morphology of any South American species. The stems are prostrate, growing and rooting beyond the flowers, producing small mats, with the flowers scattered singly, often widely, along the stem, and with consistently opposite leaves. This combination of features is shared with 13 species restricted to Australasia and no others. In describing E. conjungens, Skottsberg (1906) drew attention to its apparent close relationship to the creeping species of Australasia and placed it nearest to E. nummulariifolium A. Cunn. and E. brun- nescens (Cockayne) Raven & Engelhorn (as E. pedunculare A. Cunn.), based on Haussknechfs Monographic (1884). He considered it "ein weiteres Bindeglied," a wide (distant) connecting link between the floras of Tierra del Fuego, where E. conjungens is endemic, and New Zealand. The epithet "conjungens, " chosen by Skottsberg, reflects this relationship, meaning connected or united, and referring to its floristic affinities. More recently, the distinctive habit of Epilobium conjungens was interpreted to be the result of convergent evolution, since it was believed that E. conjungens had a few flowered, terminal inflorescence (Raven & Raven, 1976). An exami- nation of material unavailable to them, however, shows clearly that E. conjungens possesses all the features that are unique to the creeping Australasian species. In addition, the flowers of E. conjungens are white as they are in all other creeping species. It seems certain that E. conjungens was derived from an Australasian 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 progenitor. Although the chromosomal arrangement of E. conjungens is not known, the presence of only the BB type in Australasia suggests that E. conjungens will be found to have that arrangement also. Ancestors of these three groups probably arrived by long-distance dispersal via the prevailing westerlies. These winds blow continuously, between 30° and 60°S latitude, and are four times as strong as their Northern Hemisphere coun- terparts (Lamb, 1959). They have undoubtedly played an important role in the dispersal of organisms between southern land masses (Raven, 1973a). Epilobium seeds are highly dispersable by wind. Each seed has a coma of long silky hairs at the chalazal end. Once the capsule has dehisced, the seeds can be moved many meters by even light breezes. A striking representation of the capacity of the Southern Hemisphere westerly winds to affect movement between Australasia and southern South America is illustrated by a recent balloon experiment (Mason, 1971). A radio controlled balloon was released from Christchurch, New Zealand, held at about 12,000 m, and followed by satellite. During the 102 days of tracking, it made eight circuits of the globe, and repeatedly crossed south Australia, New Zealand, southern South America, and southern Africa. On most circuits it required only four to six days to traverse the distance between New Zealand and South America. Of special interest in connection with this pattern of dispersal is the presence of 13 species of Epilobium in the Chatham Islands, 800 km east of New Zealand. Equally suggestive of an initial origin in Australasia for the three groups of species with the BB chromosomal arrangement now found in South America is that all of them are restricted to areas south of 30°S latitude, precisely in the path of the prevailing winds. With their high fecundity and persistent autogamy, the intro- duction of a single Epilobium seed in a suitable habitat would provide the basis for a founding population. The foregoing discussion clearly shows the diverse origins of Epilobium in South America and provides examples of two major types of temperate disjuncts found in the New World, amphitropical and circumaustral (Moore, 1972; Raven, 1963, 1973b), both at the individual species level and at higher species groupings. The data available for Epilobium reinforces the conclusions of the papers just cited, that most disjuncts in common between North and South America have been derived by dispersal from north to south, and those between Australasia and South America from west to east. Evolutionary History The time of arrival and the impact of geological events on the various groups of Epilobium in South America is important to an understanding of their evolu- tionary history. In the absence of any fossil data on Epilobium in South America, except in a few apparently recent (late Pleistocene) but poorly dated pollen samples from Patagonia and Tierra del Fuego (Auer, 1958), the evidence for the time of appearance of Epilobium in South America is dependent on information about the availability of suitable habitats and on the presence of Epilobium in the presumed source areas. 1982] SOLOMON— EP1LOBIUM IN SOUTH AMERICA 259 As was discussed earlier, the genus Epilobium, based on its current distri- bution, possibly evolved in western North America and spread from there throughout the Northern Hemisphere, entering the three Southern Hemisphere continents only secondarily. In Australasia, Epilobium apparently did not arrive from Asia until the late Pliocene (Raven, 1973a; Raven & Raven, 1976). Prior to this time, from the Cretaceous to the mid-Miocene, Australasia was much further south than it is at the present and was separated initially by a gap of at least 3,000 km from points in southeast Asia (Raven, 1979b). During this period, Australasia was dominated by temperate forests (Raven, 1973a), while southern Asia was tropical, with low relief (Raven, 1979b), and hence, an area unlikely to have had Epilobium at that time. It was only during the late Pliocene, after Australasia had moved sufficiently far northward to encounter the Asian Plate, that mountains with cool-temperate climates were formed in Malaysia, Sumatra, Luzon, and New Guinea that could serve as intermediate points for the long-distance migration of temperate Asian species into Australasia. The earliest records of Epilobium in New Zealand are from the late Pliocene (Raven & Raven, 1976). Thus, Australasia could not have served as a source area for Epilobium in South America until at least the late Pliocene, and probably later. North and South America were widely separated until the Eocene, at which time South America was more or less equidistant from Africa and North America. From that point on, they moved closer to one another and gradually underwent orogenies that eventually resulted in a land connection about 3.1 million years ago (Keigwin, 1978; Marshall et al., 1979). Suitable temperate habitats for Epi- lobium, however, did not occur in South America throughout that time, except perhaps in the far South where Nothofagus forests similar to those in Australasia existed (Simpson, 1973; Vuilleumier, 1969). A review of palynological and geological studies suggests that the Andes are a very recently evolved mountain range that did not attain high elevations until the late Pliocene or even the early Pleistocene (Flenley, 1979; Hammen, 1974, 1979; Simpson, 1975, 1979). The present-day Andes are composed of a rather large number of structural-tectonic units that have undergone separate orogenies involving uplift, folding, faulting, and vulcanism, but resulting in a more or less concurrent elevation of the entire range. At the end of the Cretaceous most of the land that now forms the Andes had been elevated above sea level and gradual uplifting continued until the Miocene. During this time, elevations in excess of 1,000 m were probably infrequent, at least in the north (Hammen, 1979). Throughout most of this period the Andean region was covered with tropical forests that extended much farther south than at present and merged gradually with subtropical and temperate forest elements in the far south (Cei, 1979). Beginning in the Miocene and continuing into the Pliocene increased vulca- nism and crustal movements uplifted the mountains even more, but it is doubtful if there were significant areas of elevation over 2,000 m (Simpson, 1979). During this same period, the initial phases of climatic deterioration in southern South America began, which ultimately culminated in the glacial events of the Pleistocene. The major trend was toward a cooler, drier climate, with increasing 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 69 rain shadow effects from the rising Andes, which resulted in the breakup of Patagonian forest lands and the migration of more mesic tropical and subtropical elements northward (Baez & Scillato, 1979). The middle and late Pliocene saw a major increase in orogenic activity that continued into the Pleistocene and resulted in an average uplift of 1,000 to 3,000 m throughout the Andes. Thus, all of the floristic elements that are currently found at high elevations must have been derived during the past two to five million years, either by migration from other temperate areas or by vertical evolution from pre-existing lowland tropical elements (cf. Cleef, 1979). The first indication of open paramo-like habitats is found in the northern Andes in upper Pliocene formations and consists of a high representation of Gramineae, Compositae, and Hypericum (Hammen, 1979). All the species of Epilobium of apparently North American extraction (AA), except E. ciliatum and E. puberu- lum, occur in the paramo and puna north of central Chile and Argentina. It is unlikely that the North American progenitor of these species could have arrived and become established prior to the late Pliocene, and it may have appeared even later. Species such as E. ciliatum and E. hirtigerum probably arrived during or after the last glaciation. The ancestor of E. conjungens almost certainly arrived after the last glacial advance, since Tierra del Fuego and adjacent islands, where it is endemic, were completely ice-covered at that time (Vuilleumier, 1971). Climatic deterioration in the late Tertiary finally resulted in the drastic fluc- tuations between glacial and interglacial climates that characterized the Pleisto- cene. Tropical regions were significantly affected by alternating climates just as were more temperate areas. In the tropical Andes, there is evidence for one to four glaciations that, during the last glacial advance, resulted in the lowering of tree line from 1,200-1,500 m below its present level, and a reduction of mean temperature by 6-7°C, at least in the Andes of Colombia (Hammen, 1974). Glacial periods were colder and drier than interglacials, resulting in the downward movement of vegetation zones, gen- erally with an increase in their areal extent, and expansion of some xeric vege- tation types at the expense of moist or wet tropical ones, especially in the low- lands (Haffer, 1979). The impact of actual glacial ice during these periods was not as great in the tropics as the heavy glaciation in more temperate latitudes; although substantial glaciation took place in the Cordilleras of Peru and Bolivia and on other high mountains elsewhere (Vuilleumier, 1971). During interglacials, the climate was warmer and probably wetter, resulting in an upward movement of vegetational zones, contraction of paramo vegetation and an expansion of more mesic forest types. For paramo and dry lowland floras, the interglacials were a time of isolation and differentiation, and vice versa during glacials. In temperate South America, the impact of glacial events was in many ways much more severe than in the tropics. There is abundant evidence for four gla- ciations that covered large areas of the southern Andes with glacial ice. Between 30°S and 44°S there were extensive montane glaciers extending to progressively lower elevations. Some of these produced major ice barriers in a number of river valleys in central Chile (Simpson, 1979). From about 44°S latitude the entire Andes chain was covered with glacial ice to sea level on the western slope and 1982] SOLOMON— EP1LOBWM IN SOUTH AMERICA 261 to the base of the mountains on the east. The area covered by ice included all of Tierra del Fuego and the southern tip of Patagonia (Vuilleumier, 1971). Glacial advances effectively lowered vegetation zones in temperate South America, which allowed alpine and other high elevation vegetation to cover more extensive, con- tinuous areas and produce regions of secondary contact between formerly isolated populations. At the same time, it caused contraction and shifting of Nothofagus forest regions northward (Simpson, 1973). During interglacials these same alpine vegetation types were isolated in upper montane areas where they could undergo independent differentiation. The effects of Pleistocene climatic fluctuations on speciation patterns in Epi- lobium are not at all clear, and may have had relatively little impact, unlike patterns found in other Andean groups that have apparently diversified greatly under alternate contraction and expansion of ranges (e.g.,Perezia multiflora com- plex, Simpson, 1973; Arracacia , Lleracia, Simpson, 1975). Of the tropical species, only E. denticulatum shows extensive morphological diversity through its large range in the paramo and puna. Interestingly enough, E. denticulatum achieves its greatest variability in the altiplano and Cordilleras of Peru and Bolivia, with diminishing diversity both north and south of there. This area is the part of the range of E. denticulatum that was most heavily glaciated during the Pleistocene, so glacial advances and retreats may have influenced the variation patterns now seen. The high dispersability of Epilobium seed, however, may exclude glaciers from acting as actual barriers. Alternatively, this great diversity may be due to the large number of habitats at various elevations available for colonization by E. denticulatum in this region. Unlike Epilobium in New Zealand, which has undergone an explosive radia- tion of many closely related species adapted to a diverse array of habitats through hybridization and persistent autogamy, the largest group of closely related species in South America contains only five members (E. australe, E. barbeyanum, E. densifolium, E. glaucum, and E. nivale). Each of these is very distinctive, with mostly overlapping ranges, and no one of them can be said to have been derived by hybridization between any other two species. These species may have evolved under the isolation-habitat specialization model suggested above, but at the pres- ent time interpretation of their inter-relationships remains perplexing and awaits new insight. Pleistocene glacial advances also affected the current geographical distribution of some of the species. All of the species except three are found almost exclu- sively south of 30°S. Between the latitudes of 27°S and 3I°S there was a band of persistent aridity that crossed the Andes and was an effective barrier during both glacial and interglacial periods (Simpson, 1979). Epilobium ciliatum, E. glaucum, and E. barbeyanum all reach their northern limits at the southern edge of this zone. Only E. denticulatum has managed to cross this barrier in a narrow strip along the eastern slope of the Andes, south to about 34°S. In southern South America, the heavy glaciation south of 44°S, eliminated much of the montane vegetation. There are a large number of species that reach their southern limit at about this latitude (Simpson, 1973), including E. barbeyanum and E. densifolium, which are not known from south of about 42°S. One of the more peculiar distributions seen in Epilobium is that of E. nivale. 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69 It too reaches the southern limit of its primary range at about 42°S, but then reappears in the vicinity of Lago Argentino (5 PS), indicating a gap in its range of about 800 km. Other species, such as Isoetes savatieri Franch. (Donat, 1931), the species pair Perezia bellidifolia (Phil.) Reiche and P. megalantha Speg. (Simpson, 1973), and Gunnera tinctoria (Mol.) Mirbel (Pisano, pers. comm.), have similar distributions, which has led some authors (Auer, 1958; Skottsberg, 1916; Donat, 1931; Simpson, 1973) to suggest that there was a refugial area to the east of the Andes in the far south where these and other species managed to survive during the heaviest glaciations, while they were eliminated in between. It is possible that E. nivale does occur in the intervening area because the Pa- tagonian Andes have been little explored botanically, and E. nivale inhabits high, relatively inaccessible areas near the snow line. Reproductive Biology and Hybridization Breeding system. The majority of the species of Epilobium sect. Epilobium are autogamous, with only approximately 10% of the 170 species modally out- crossing. All are self-compatible, and, in those that are capable of self-fertiliza- tion, a full complement of seed is normally produced. Epilobium species in South America possess a diversity of breeding systems, from modally outcrossing to cleistogamous, but all of them can produce seed by self-fertilization (see review in Raven, 1979a). Nearly all the species have pale pink to rose purple flowers, including the three introduced species. Only E. hir- tigerum, E. denticulatum, E. ciliatum, and E. conjungens have petals that are white or vary from white to pink. Flower color appears to be significantly related to the degree of outcrossing, as pointed out by Raven & Raven (1976), since insects are more likely to visit the colored flowers than white ones. Epilobium densifolium has the largest number of features typical for a modally outcrossing species. The flowers are large (petals up to 1.2 cm long), with the anthers held away from the stigma at anthesis, and usually the stigma partially exserted. Each flower produces a large droplet of nectar. Epilobium australe, E. glaucum, and E. barbeyanum have smaller flowers, but in them also, the anthers are held away from the stigma at anthesis. In these four species, usually after a period of several hours, the longest stamens gradually bend inward and the de- hisced anthers make contact with the receptive stigma. Nonetheless, they are all modally outcrossing. Epilobium denticulatum has separate or even mixed populations of large- and small-flowered plants. In this species, the large flowers also have the anthers separated from the stigma, at least for a few hours. The balance of the species, including the small-flowered populations of E. denticulatum, have flowers that are more or less immediately self-pollinated at anthesis. Functional cleistogamy seems to be the general condition for E. hirtigerum, where the anthers may dehisce prior to the opening of the flower bud. Generally, the anthers dehisce at anthesis, although there may be up to a few hours delay. In addition, all species, except E. hirtigerum and some plants of E. cilia turn, produce at least a small amount of nectar that may be attractive to flower visitors. Weather conditions have a verv strone effect on the extent to which the 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 263 flowers of Epilobium open at anthesis. All species open their flowers fully on bright sunny days, including E. hirtigerum. On cloudy or rainy days, however, most plants open their flowers only partially or not at all, very strongly reducing the chances of cross-pollination or eliminating that possibility completely by being functionally cleistogamous. Despite the various features that might promote some outcrossing, insect vis- itation to any of the species appears to be quite rare. In many hours of obser- vation, at all times of the day and under various weather conditions, only two insect visitors were observed. Several unidentified medium-sized syrphid flies were seen visiting flowers of Epilobium barbeyanum in a large streamside bog near Lagunillas, Cordillera Prov. (Santiago), Chile; and a single visit by a small, unidentified bee to several plants of E. denticulatum with medium-sized flowers in a population near Rinconada, Dpto. La Paz, Bolivia was observed. Even though a number of species have floral morphologies suggestive of out- crossing, it is clear that there remains a high degree of autogamy in most species. Autogamy is probably the primary means by which populations maintain their dis- tinctness from one another, and it is certainly a strong force in limiting interspe- cific hybridization. Persistent autogamy does not prevent outcrossing, it merely reduces the frequency of an outcrossing event taking place. While cleistogamy may be the functional mode of pollination in some populations or species, oc- casional intra- and interspecific crosses between many of these species do occur, providing new genetic variability for segregation and recombination. Even crosses between groups that have developed some internal barriers to hybridization pro- duce occasional fertile seeds that may be capable of surviving. In essence, then, every population is under its own selective pressure, to which it adjusts by re- producing well adapted genotypes and occasionally, through outcrossing, obtains new genetic material. Ecological relationships. Many South American species of Epilobium grow in habitats similar to those in which species in other parts of the world are found; that is, in moist, often disturbed, open sites such as stream banks, ditches, seeps, bogs, lakeshores, etc. Other South American species exhibit a striking amount of ecological specialization, however, which further isolates a number of distinct species. The following list provides a synopsis of the ecological preferences of the various species: E. australe, almost always in or near running water, broad elevational range; E. barbeyanum, alpine bogs and very wet sites, stems often floating; E. ciliatum, any moist, disturbed site, broad elevational range; E. con- jungens, moist moss mats above timberline; E. densifolium, stabilized alpine scree, often away from water, but also along rocky stream banks; E. denticula- tum, any moist site, broad elevational range; E. fragile, moist cracks and crevices in rocks, above 4,500 m; E. glaucum, almost always in or near running water, broad elevational range; E. hirtigerum, marshes, wet places, mostly at low ele- vations; E. nivale, alpine, along rivulets and other permanently wet spots, up to permanent snow line; E. pedicel/are, steep or vertical seeps, altiplano; E. pu- berulum, moist disturbed places, mostly at low elevations. Sympatry. Despite rather strong ecological differences and geographical sep- arations, many species do occur sympatrically, at least in part of their range. 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Table 8. Sympatric occurrence of native species of Epilobium in South America. Acronyms are those given in Table 2. In addition, PED = E. pedicellare, FRAG = E. fragile, and CONJ = E. conjungens. KEY: * = species that occur together and hybridize; + = species that occur together; ? = species that probably occur together. DENT PED FRAG PUBER CIL N1V BARB DENS AUST GLAU + CONJ HIRTIG + * + * + + ? * + + * _J_ * i * * * • ■ • • « f DENT PED FRAG PUBER CIL NIV BARB DENS AUST GLAU CONJ HIRTIG This is especially true in the Andes of central Chile and Argentina at moderate elevation (1,000-1,500 m), where two or more species are often found very near each other, if not intermixed. Up to five species have been seen growing together near the Termas del Rio Blanco, Malleco Prov., Chile, at 1,300 m; namely Epi- lobium au st rale, E. ciliatum, E. densifolium, E. glaucum, and E. nivale. Table 8 summarizes the sympatric occurrence of the native species in South America. The information was obtained primarily from personal observation and mixed herbarium collections, or rarely deduced from locality information. There are a number of combinations that cannot occur because the ranges of the species involved do not overlap. Likewise, there are several sympatric combinations that should occur, but which have not been seen. These are also indicated in Table 8. One of the introduced species, Epilobium obscurum, has been found growing mixed with E. puberulum and E. ciliatum, and is known to hybridize with the latter. Natural hybridization. The occurrence of natural hybrids has been com- mented on for many years, beginning with Haussknecht (1884), although the frequency with which they are recognized depends greatly on how finely the species delimitations are drawn. Natural hybrids in Epilobium are found through- out the world, although constraints are imposed on their formation by a high degree of autogamy or internal barriers to hybridization. Very few hybrids have ever been reported from South America, primarily because of the lack of material and field examinations. Samuelsson (1923, 1930) reported hybrids between E. denticulatum and E. hirtum, now considered to be conspecific, and between E. denticulatum and E. haenkeanum (= E. pedicellare). Only one specimen of the latter combination is now considered to be of hybrid origin, although hybrids between E. denticulatum and E. pedicellare are difficult to distinguish. Eleven combinations that are known to form natural hybrids are indicated in Table 8 and documented by specimens whose pollen fertility and seed set have been estimated. These are discussed in detail under the individual species treat- 19821 SOLOMON— EP1LOB1UM IN SOUTH AMERICA 265 ments. The majority of natural hybrids have morphology intermediate between the two parents and are often characterized by reduced pollen stainability and seed set. A number of hybrid combinations would be difficult to detect, e.g., densifolium Whereve are relatively infrequent. This, in part, may be due to the ecological distinctness of the habitats in which some species grow, or to other types of barriers to the successful production of hybrid individuals. In contrast to the situation in New Zealand (Raven & Raven, 1976) where hybridization is seen as a central force in the formation of new adaptive genotypes, the South American species, because of their heterogeneous origins, and perhaps the recency of their arrival, have diversified to only a small extent, and not primarily through hybridization. Taxonomic History The first botanist to acquire specimens of Epilobium from South America was Montev guay, in 1767 when the survey ships Etiole and Boudeuse, under the command of Louis de Bougainville, stopped on their voyage to the western Pacific. Unfor- Mauritius his collections critically. These were then forwarded to France and incorporated in the collection of the Jardin du Roi. At this same time, Captain James Cook was also exploring southern South America and the Pacific. With him on his first voyage were the celebrated Joseph Banks and Daniel Solander. Among their many new collections was a second species, Epilobium australe, from Tierra del Fuego, collected in 1769. Somewhat later in the eighteenth century, two major scientific expeditions were financed by the government of Spain, the renowned travels of Hipolito Ruiz, Jose Pavon, and Joseph Dombey to Peru and Chile (1778-1788), and the Mala- spina Expedition, which was sent to survey Spanish territories in the Pacific (1789-1794). During the extensive peregrinations of the former, species of Epi- lobium were encountered in several places, such as Chancay and Tarma, Peru, and Concepcion, Chile. The specimens from Peru represented E. denticulatum, and those from Concepcion were a fourth species, E. puberulum. All of these collections, however, were published as E. denticulatum, the first name of any South American species to find its way into print (Ruiz & Pavon, 1802). The Malaspina Expedition was fortunate in having the services of two well trained botanists, Luis Nee and Thaddeus Haenke. They prepared many thou- sands of specimens from South America, Mexico, North America, and the west- ern Pacific. Among these were Epilobium pedicellare from Peru collected by Haenke and E. ciliatum, E. glaucum, and E. barbeyanum from Chile, collected by Nee. The latter species were most likely acquired when Nee left the expedition at Concepcion in late 1793 to go overland via Santiago, crossing the Andes to Mendoza and thence across the pampas to Buenos Aires. There, in May 1794, he rejoined the expedition with nearly 10,000 specimens for the return to Spain (Safford, 1905). Unfortunate political circumstances intervened, so that most of the results of the expedition were never published. A set of Haenkes material was sent to Karl Presl in Prague, who later published E. pedicellare along with 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 many other species in his Reliquiae Haenkeanae (Presl, 1831). Nee's voluminous collections went to the Royal Botanic Garden in Madrid where some plants were eventually described and published, but the majority were not. Thus, by the end of the eighteenth century, eight of the twelve native taxa recognized in this treat- ment had been collected, but the first name for any of them was not published until 1802. The time between 1820 and 1870 saw an increasing number of coastal surveys and penetrations of the interior, especially in southern South America. Most notable among the naval surveys were the United States Exploring Expedition under the command of Charles Wilkes (1838-1842), the Antarctic Expedition of James Ross (1839-1842) with J. D. Hooker as botanist, the first voyage of the Adventure and Beagle under the command of Philip King (1826-1830), and the second voyage of the Beagle commanded by Robert FitzRoy, with Charles Dar- win as naturalist (1831-1836). At the same time, a substantial number of botanists were spending extended periods of time in limited areas throughout the continent. Those who are especially important to the chronology of South American Epi- lobium were the collecting forays of E. Poeppig, T. Bridges, H. Cuming, C. Gay, and R. Spruce. During this period of active exploration, there was still a great deal of con- fusion about the limits of the species. It is not surprising, when one considers the variability of many Epilobium species, that the new South American plants could be equated with variable European species (e.g., E. nivale with E. alpinum, £. ciliatum with E. tetragonum). J. D. Hooker was quite perplexed by the species of Epilobium from the Andes, and considered most of them to be variants of E. tetragonum, one of which he felt was sufficiently distinct to be described as a new variety, E. tetragonum /3 antarcticum (= E. australe) (Hooker, 1847, 1853). Quite frequently South American plants were rather arbitrarily given one of three names, E. denticulatum, E. tetragonum, or E. pedicellare. This state of affairs persisted even to the beginning of the twentieth century, when E. tetragonum was still being reported as the name for several different species (Macloskie, 1905; Arechavaleta, 1902). Carl Haussknecht, beginning with a series of papers in 1879 and culminating in his Monographie der Gattung Epilobium in 1884, published many new taxa. He had the advantage of studying the collections of many of the major herbaria of Europe, and thus had more South American material at his disposal than any previous author. Perhaps equally as important was his worldwide familiarity with the genus. He was able to dispel the notion that common European species occurred in southern South America, and he recognized several common taxa as separate species for the first time, e.g., Epilobium chilense {= E. ciliatum) and E. australe. Haussknecht was rather confused by the taxa allied to E. denticu- latum, but such was the authority of his work that some of his errors were perpetuated, resulting in the report of names such as E. bonplandianum and E. caesium from central Chile (Reiche, 1898). In total, Haussknecht recognized 18 taxa, representing 9 of the 12 native species presented in this treatment. Among the most important collections used by Haussknecht in the preparation of the Monographie were those of Eduard Poeppig and Rudolfo Philippi. Poeppig made extensive collections in central Chile from 1827 to 1829; R. A. Philippi 1982] SOLOMON— EPILOBI UM IN SOUTH AMERICA 267 arrived there in 1851 with his son Federico, both of whom worked intensively on the flora of Chile for more than 50 years. Subsequent to Haussknecht's work, R. A. Philippi described seven new species of Epilobium (Philippi, 1893) that are here considered conspecific with previously described taxa. Further botanical explorations in Chile and Patagonia during the early years of the twentieth century, especially by Per Dusen and Carl Skottsberg, produced much new material. Among the new species described were E. bar- beyanum by Leveille in 1907 and the highly distinctive E. conjungens by Skotts- berg in 1906. By 1923, partly under the impetus of a fine set of collections made by Erik Asplund in Bolivia, Gunnar Samuelsson had reviewed the entire genus in South America. He reworked nearly all of the material used by Haussknecht as well as more recent collections. As a result of his studies, he published many new taxa and recognized nearly all of those proposed by Haussknecht, bringing the total number of taxa to 34 species and 10 varieties. Since that time very little has been published on South American Epilobium. Samuelsson, after studying collections in North American herbaria, reduced three Munz 1934, 1974), MacBride uelsson very closely. Systematic Treatment Epilobium L., Sp. PI. 347. 1753, Gen. PL, ed. 5, 164. 1754. lectotype: Epilobium hirsutum L., Britton & Brown, 111. Fl. No. U.S. & Can., ed. 2, 2:590. 1913. (Complete synonymy not given.) Chamaenerion Seguier, PI. Veron. 3:168. 1754. lectotype: Epilobium hirsutum L., Holub, Folia Geobot. Phytotax. 7:84. 1972. Zauschneria Presl, Rel. Haenk. 2:28, pi. 52. 1831. type: Z. californica Presl. Chamerion (Raf.) Raf., Herb. Raf. 51. 1833. Based on Epilobium subgen. Chamehon Raf., Amer. Monthly Mag. & Crit. Rev. 2:266. 1818. type: E. amenum Raf. Crossostigma Spach, Ann. Sc. Nat. Bot., ser. 2, 4:174. 1835. type: C. lindleyi Spach, nom. illeg. = E. minutum Lindl. ex Lehm. Pyrogennema Lunell, Amer. Midi. Naturalist 4:482. 1916. type: P. angustifolium (L.) Lunell. Cordylophorum (Nutt. ex Torr. & Gray) Rydb., Fl. Rocky Mts. 590, 1064. 1917. Based on Epilobium sect. Cordylophorum Nutt. ex Torr. & Gray, Fl. N. Amer. 1:488. 1840. type: C. suffruticosum (Nutt.) Rydb. Erect or creeping perennial or annual (3 species) herbs, sometimes woody at the base, overwintering and reproducing vegetatively by loose, scaly rhizomes, stolons, buds in the axils of leaves, soboles, leafy rosettes, or turions (fleshy, scale-leaved buds produced at or below the ground surface). Leaves simple, petiolate or sessile, opposite, at least at the lower nodes, alternate above, or all opposite, or all alternate in sect. Chamaenerion; pubescent or glabrous, the pu- bescence often distributed in descending lines on raised decurrent petiole mar- gins. Flowers actinomorphic, or slightly zygomorphic in a few species, solitary in the axils of more or less reduced or unmodified leaves, clustered upwards, usually forming a few- to many-flowered, more or less discrete racemose, panicu- late, or corymbose inflorescence, or scattered and few. Floral tube present or absent (sect. Chamaenerion). Sepals 4. Petals 4, white, cream, pink to rose pur- 268 ANNALS OF THK MISSOURI BOTANICAL GARDEN [Vol. 69 pie, red, or yellow, emarginate or cleft. Stamens 8, the 4 antisepalous ones longer than the 4 antipetalous ones. Stigma capitate, clavate or deeply 4-lobed, protan- drous or protogynous, or the anthers dehiscing at about the same time that the stigma becomes receptive. Capsule narrow, elongate or rarely narrowly clavate, 4-locular, loculicidal. Seeds many or rarely one or two in each locule, mostly obovoid, with a coma of long silky hairs at the chalazal (terminal) end that is rarely absent ( 1 species and some populations of a second). Gametic chromosome numbers: n = 12, 13, 15, 16, and 18, with polyploidy based on 15 and 18 in some taxa. The generic description is based on the genus worldwide. As a result, some of the features given are not present in any of the South American species. The interested reader should refer to Raven ( 1976) and references therein for complete descriptions, discussion of generic and sectional delimitations, and synonymy. Since it is now fairly well understood that the Epilobium species that occur in South America have had various and diverse origins, the arrangement of the taxa in a systematic treatment cannot be purely evolutionary. Instead, the species have been placed in what I believe to be more or less phyletic groups, beginning with those having the A A chromosomal arrangement, endemic and native, then the BB, endemic and native, and then those that have been introduced recently. Epilobium paniculatum, in sect. Xerolobium, by virtue of its primitive or dis- tinctive features and lower chromosome number, might have been placed first. Because it is only sparingly introduced in South America, however, it has been placed at the end with the other introduced species. Neither descriptions nor keys have been prepared to distinguish the sections of Epilobium. There are several points that must be kept in mind in order to understand fully the nature of the taxa enumerated here. Often there are great differences in the seasonal aspect of the plants. Those collected at the end of a growing season may be more branched, with few, often withered, lower leaves; and the flowers, fruits and seeds are often smaller than those produced by the same plant earlier in the season. Overwintering structures such as turions are frequently produced only near the end of a growing season, and may not be present. Or, as is often the case, specimens have been collected without regard to underground parts, even though they are taxonomically useful. Seeds from old or well dried speci- mens may be 10% smaller in both length and width than fresh, mature seeds (Raven & Raven, 1976). The dimensions given in the descriptions attempt to cover the full range of dry and fresh seeds. Our understanding of the role of populations as evolutionary units has tended to broaden our view of what constitutes a species. In fact, considering the highly autogamous nature of most epilobiums, it is extremely questionable whether one can speak of more than a single plant as an evolutionary unit (Ehrlich & Raven, 1969; Levin & Kerster, 1974; Raven & Raven, 1976). Hence, the concept of a species or other taxon must be painted with a rather broad brush for it to be of any practical utility. This is fundamentally different from the approach taken in earlier times when many of the taxa in Epilobium were first given recognition. Plants that were at one time thought to be taxonomically distinct, I here may consider to be only a part of the "normal" variation within a polymorphic species or even a single population. 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 269 The descriptions and discussions have taken into account nearly all of the available herbarium material. With the exception of E. conjungens and E. fragile, all of the native species have been studied in the field, and one or more strains of each have been grown in greenhouses at the Missouri Botanical Garden. A note of special significance for the nomenclature concerns the formae that Haussknecht used in his Monographie (Haussknecht, 1884). According to Raven (1962), the use of the feminine ending suggests that Haussknecht thought of his formae simply as organizational devices for the presentation of variation, and not as formal taxonomic units. Following this convention, the formae have been included in the synonymy only when they were taken up by later authors. In the specimens examined, collections from Chile are placed in regions and IGM Mil in 1973, which divided the country into 13 regions and 50 provinces. Specimens from other countries are placed according to the most recent maps available. Key to the Species of Epilobjum in South America la. Stems creeping, growing and rooting beyond the region of flower production, leaves all opposite; flowers few and scattered. Tierra del Fuego and adjacent islands. 12. E. conjungens lb. Stems not creeping, erect, ascendent or decumbent, not growing and rooting beyond the flowers; leaves at least partly alternate; flowers few to many, in a more or less discrete inflorescence. 2a. Seeds broadly obovoid with a conspicuous constriction toward the micropylar end; floral bracts minute, fused to the pedicel; leaves often conduplicate, early deciduous, mostly alternate; annual. Western Neuquen and Chubut, Argentina. 15. E. paniculatum 2b. Seeds variously shaped, but without a conspicuous constriction; floral bracts leaf- like, reduced or not, but not fused to the pedicel, leaves flat, persistent, alternate or opposite; perennial, but usually flowering the first year. 3a. Plants glabrous (very rarely plants of E. ciliatum glabrous, but immediately distinguishable by their turions and longitudinally ridged seeds). Andes of Chile and Argentina. 4a. Plants low, caespitose; stems decumbent; leaves acute or obtuse, not glau- cous. 6. E. nivale 4b. Plants robust, loosely rhizomatous, more or less clumped: stems erect; leaves acuminate, glaucous. 10. E. glaucum 3b. Plants variously pubescent, the hairs sometimes very sparse or restricted to the junction of the petiole bases. 5a. Upper stems hirsute or villous, and glandular, the hairs erect or slightly spreading. 6a. Leaves mostly alternate, opposite only at the lowest few nodes, coarsely serrate; inflorescence erect; petals white. Southern Brazil, Uruguay, eastern Argentina. 11. E. hirtigerum 6b. Leaves mostly opposite, alternate or subopposite only above or in the inflorescence, denticulate; inflorescence usually nodding; petals pink. Andes, Costa Rica to northern Chile and Argentina. 1 . E. denticulatum 5b. Upper stems variously pubescent, but not hirsute or villous, the hairs not erect. 7a. Stems and ovaries densely erect glandular throughout, with some strigillose hairs; largest leaves with 3-5 teeth on each side; plants soboliferous or stoloniferous; seeds papillose. Central Andes of Chile and Argentina. 7. E. barbeyanum 7b. Stem and ovary pubescence various, if densely glandular then the leaves with more than 6 teeth on each side, or plants not soboliferous or stoloniferous; seeds variously sculptured. 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol . 69 8a. Leaves mostly alternate, opposite only at the lowest one or two nodes, or on very young shoots, coarsely and irregularly doubly serrate with uneven-sized teeth. Peru and Bolivia 2. E. pedicellare 8b. Leaves mostly opposite or subopposite, alternate only above or in the inflorescence, denticulate or once serrate. 9a. Stems nearly glabrous, the pubescence a few scattered hairs, mostly at the fused petiole bases; plants low, <15 cm, densely caespitose. Peru, Bolivia and Chile. 3. E. fragile 9b. Stems variously pubescent, the pubescence more or less dense throughout; plants various. 10a. Plants densely caespitose, usually < 15 cm tall, with a woody, contorted rootstock; flowers large, the petals 7-12 mm long; seeds 1.4-2.0 mm long. Cen- tral Andes of Chile and Argentina. ._ 8. E. densifolium 10b. Plants loosely clumped or solitary, usually >15 cm tall; flowers large or small; seeds less than 1.4 mm long. 11a. Leaves narrowly ovate to ovate, thick, coarsely serrate with 4— 8(— 1 2) teeth on each side; plants not glandular. Andes of Chile and Argentina. 9. E. australe 1 lb. Leaves narrowly lanceolate to lanceolate, ser- rulate or denticulate, or if narrowly ovate, then the inflorescence glandular. 12a. Floral tube with a few, erect, glandular hairs, otherwise the plants not glandular (rarely also with a few on the upper por- tion of the ovary); largest leaves usually with 10 or more low denticulate teeth on each side; inflorescence erect. Central Chile. 13. E. obscurum 12b. Floral tube not glandular, or if glandu- lar, then the glandular hairs also found on other plant parts, or minute and ap- pressed; leaf margins various; inflores- cence erect or nodding. 13a. Leaves serrulate, the largest usu- ally with 10 or more teeth on each side, acuminate, the blade gla- brous above and below; plants producing leafy rosettes or fleshy turions, the turions often persist- ing as a cluster of dead scales at the base of the stem. 14a. Seeds with longitudinal rows of flattened, fused papillae, at least along the sides; inflo- rescence glandular; lower stem smooth. Chile and Ar- gentina. 5. E. ciliatum 14b. Seeds papillose, the papillae distinct; inflorescence not glandular; lower stems often with exfoliating epidermis. Central Chile. 14. E. tetragonum 13a. Leaves denticulate, acute or ob- tuse, usually with fewer than 10 low teeth on each side, if more than 10, then the inflorescence nodding, the blade more or less pubescent, the 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 27 1 hairs sometimes minute and ap- pressed; plants without turions or basal rosettes, soboliferous or with leafy basal shoots. 15a. Stems with raised decurrent lines from the margins of the petioles; inflorescence and flowers nodding; younger leaves and upper stems usu- ally with appressed to erect glandular or blunt-tipped hairs under higher magnifi- cation; petals pink to rose purple. Andes, Costa Rica to northern Chile and Argen- tina. 1. E. denticulatum 15b. Stems without raised decur- rent lines from the margins of the petioles; inflorescence erect; leaves and stems not glandular; petals salmon pink. Central Chile. 4. E. puberulum 1. Epilobium denticulatum Ruiz & Pavon, Flora Peruviana et Chilensis 3:78, tab. 314 ,/. 1-la. 1802. type: Peru, Dpto. Junin, Tarma, 1779-1781, //. Ruiz & J- Pavon (MA, lectotype here designated, photograph MO; B (destroyed, pho- tographs BH, MO), BM, F, G, OXF, isolectotypes). De Candolle, Prodromus 3:42. 1828. Hausskn., Monogr. Epilobium 264. 1884. Samuelsson, Svensk Bot. Tidskr. 17:250. 1923. MacBride, Field Mus. Nat. Hist., Bot. Ser. 8(4):530. 1941. Munz, Opera Bot. Ser. B, 3:6. 1974. E. bonplandianum H.B.K., Nov. Gen. et Spec. Plant. 6:95. 1823. type: Colombia, Dpto. Cauca, Andes de Popayan, Paramo de Purace, 2,900 m, November-December 1801, A. Humboldt & A. Bonpland s.n. (P, lectotype here designated, photograph MO; P, isolectotype, photograph GH: F, HAL, P, probable isolectotypes). Humboldt & Bonpland, Synop. Plant. Aeq. 3:389. 1824. Hausskn., Monogr. Epilobium 267. 1884. E.junceum Forst. f. ex Spreng., Syst. Veg. 2:233. 1825, nom. illegit. Based on E. denticulatum Ruiz & Pavon. E. pedicellare auct. non Presl: Hook. & Arn., Bot. Misc. 3:309. 1833, pro parte. E. caesium Hausskn., Oesterr. Bot. Z. 29:91. 1879. type: Bolivia, Dpto. La Paz, Prov. Caupolican, Pelechuco, 3,900-4,200 m, March 1865, R. Pearce s.n. (K, holotype, photograph MO). Hausskn., Monogr. Epilobium 268, tab. 17, f. 75, 75a-c. 1884. E. andicolum Hausskn., Oesterr. Bot. Z. 29: 1 18. 1879. type: Bolivia, Dpto. La Paz, Prov. Larecaja, vicinity of Sorata, 1859, M. G. Mandon 626 (K, lectotype here designated, photograph MO; BM, BR, GOET, K, P, isolectotypes). Mandon 626 is apparently a distribution number. From an examination of the many sheets available, it is probable that several gatherings were included. The lectotype and isolectotypes are limited to those specimens annotated as E. andicolum by Haussknecht. Hausskn., Monogr. Epilobium 266, tab. 17, f. 76, 76a-c. 1884. Samuelsson, Svensk Bot. Tidskr. 17:255. 1923; Svensk Bot. Tidskr. 24:2. 1930. E. meridense Hausskn., Oesterr. Bot. Z. 29:148. 1879. type: Venezuela, Est. Merida, Sierra Nevada, 3,300 m, August 1842, J. Linden 418 (W, lectotype here designated, photograph MO; BM 2 sheets, BR (probable), F, G (photographs BH, MO), K, OXF, P 2 sheets, TCD, W, isolectotypes). Hausskn., Monogr. Epilobium 266. 1884. H. Lev., Iconogr. Epilobium tab. 193, 194. 1911. Sam- uelsson, Svensk Bot. Tidskr. 17:257. 1923; Svensk Bot. Tidskr. 24:9. 1930. Munz, Aliso 4:488. 1960; N. Amer. Fl. II 5:218. 1965; Opera Bot., Ser. B, 3:7. 1974. Vareschi, Flora de Los Paramos 247,/. 77. 1970. E. helodes H. Lev., Bull. Herb. Boissier, ser. 2, 7:589. 1907. type: Colombia, Dpto. Tohma, Paramo de Ruiz, high montane forest, 3,000 m, 1 1 September 1883, F. C. Lehmann 3158 (G, holotype, photograph MO; BM, US, isotypes). E. meridense var. helodes (H. Lev.) Samuelsson, Svensk 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Bot. Tidskr. 17:258. 1923. Samuelsson, Svensk Bot. Tidskr. 24:10. 1930. Munz, Opera Bot., Ser. B, 3:7. 1974. E. denticulatum var. aberans Samuelsson, Svensk Bot. Tidskr. 17:253. 1923. type: Bolivia, Dpto. La Paz, Prov. Aberoa, Challapata, 3,750 m, 31 March 1921, E. Asptund 3250 (UPS, holotype, photograph MO; S, isotype). E. denticulatum var. confertum Samuelsson, Svensk Bot. Tidskr. 17:252. 1923. type: Ecuador, Prov. Pichincha, Quito, 1847, W. Jameson 192 (W, lectotype, photograph MO; BM, G, S, TCD, US, isolectotypes; P. Munz, Opera Bot., Ser. B, 3:6. 1974). E. denticulatum var. macropetalum Samuelsson, Svensk Bot. Tidskr. 17:253. 1923. type: Peru, Dpto. Cuzco, Pacechac, near Rio Urubamba, 4,500 m, 3 March 1903, A. W. Hill 155 (K, lectotype; P. Munz, Opera Bot., Ser. B, 3:6. 1974). E. asplundii Samuelsson, Svensk Bot. Tidskr. 17:256, tab. 3J\ 2. 1923. type: Bolivia, Dpto. La Paz, Prov. Omasuyos, Challa, Isla del Sol, 3,850 m, 18 April 1921, E. Asplund 3708 (UPS, holotype, photographs MO, S). E. meridense var. condensatum Samuelsson, Svensk Bot. Tidskr. 17:258. 1923. type: Ecuador, Prov. Chimborazo, El Altar, 3,900-4,000 m, July 1903, H. Meyer 174 (JE, lectotype here designated, photograph MO; JE, isolectotype). All of the material cited by Samuelsson was located at Berlin (B) and is now destroyed. The lectotype has been selected from the duplicates of that material at other institutions. E. aequinoctiale Samuelsson, Svensk Bot. Tidskr. 17:259, tab. 2,f. /. 1923. type: Colombia, Dpto. Naririo, Tiiquerres, 1844-1847, H. Karsten (W, lectotype here designated, photographs BH, MO, S; JE, isolectotype). H. Lev., Iconogr. Epilobium tab. 198 (as E. repens). 191 1. Munz, Opera Bot., Ser. B, 3:4. 1974. E. assurgens Samuelsson, Svensk Bot. Tidskr. 17:261, tab. 5, f. 2. 1923. type: Ecuador, Prov. Tunguragua, Volcan Tunguragua, "Locis paludosis," May 1858, R. Spruce 5389 (W, lectotype here designated, photographs BH, MO, S; BM, K 2 sheets, NY, OXF, TCD, W, isolectotypes). Samuelsson, Svensk Bot. Tidskr. 24:3. 1930. E. bolivianum Samuelsson, Svensk Bot. Tidskr. 17:263, tab. 2,f. 3. 1923. type: Bolivia, Dpto. La Paz, Prov. Murillo, stream at La Cumbre, 4,600 m, 26 May 192 1 , E. Asplund 4014 (UPS, holotype, photographs MO, S; B (destroyed, photographs BH, MO, US), S, W, Z, isotypes). MacBride, Field Mus. Nat. Hist., Bot. Ser. 8(4):530. 1941. E. deminutum Samuelsson, Svensk Bot. Tidskr. 17:264, tab. 4,f. 5. 1923. type: Bolivia, Dpto. La Paz, Prov. Murillo, Nevado Huayna-Potosi, Glaciar Franz Josefs, Germann 24 (W, holotype, photographs BH, MO, US, fragment UPS). Samuelsson, Svensk Bot. Tidskr. 24:5. 1930. E. hirtum Samuelsson, Svensk Bot. Tidskr. 17:266, tab. 2 y f. 2. 1923. type: Bolivia, Dpto. La Paz, Prov. Murillo, San Jorge, now within the city of La Paz, 3,500 m, 1 November 1920, E. Asplund 666 (UPS, lectotype here designated, photographs MO, S; S, isolectotype). Samuelsson, Svensk Bot. Tidskr. 24:4. 1930. MacBride, Field Mus. Nat. Hist. Bot. Ser. 8(4):532. 1941. Variable, clumped perennial herbs ( 1 0-)20-70(- 1 60) cm tall, reproducing veg- etatively by elongate, leafy shoots or soboles produced at or near the base. Stems erect or ascendent, several to many, mostly simple, occasionally branched above, and often from the base, strigillose to spreading hirsute, with hairs 0.1-0.4 mm long, often with an admixture of appressed to erect glandular hairs, 0.05-0.2 mm long, at least in the inflorescence. Leaves mostly opposite, alternate above and in the inflorescence, thin, green, occasionally reddish purple, especially along the veins and margins, lanceolate, occasionally ovate, rarely broadly so, 1-5.6 cm long, 0.2-1.7 cm wide, acute to acuminate, rarely obtuse at the apex, denticulate with 3— 8(— 13) low teeth on each side, acute to cuneate, occasionally obtuse or rounded at the base, strigillose to spreading hirsute on the abaxial and adaxial veins and midrib, or thinly scattered on the blade, with scattered appressed to spreading blunt-tipped or glandular hairs, 0.05-0.2 mm long, occasionally densely so, the lateral veins prominent, 2-4(-5) on each side of the midrib, on petioles 0.5-2(-4) mm long, rarely sessile. Inflorescence simple, nodding, the leaves sub- tending the flowers usually slightly reduced in size. Flowers nodding at anthesis. Ovaries often reddish or purplish, strigillose to spreading hirsute, often densely so, usually with an admixture of appressed to spreading glandular hairs, 0.9-2.2 1982] SOLOMON— EPILOBWM IN SOUTH AMERICA 273 cm long, on pedicels 2-5(-8) mm long. Floral tube often reddish purple, 0.8-2 mm deep, 1.5— 3(— 4) mm across, externally strigillose to spreading hirsute, usually with appressed to spreading glandular hairs, internally with a ring of erect villous hairs 0. 1-0.2 mm long, sometimes reduced to only a few, or occasionally gla- brous. Sepals often reddish purple, lanceolate, 2.2-5.7(-7) mm long, 1.1-2.1 mm wide, strigillose to spreading hirsute, usually with appressed to spreading glan- dular hairs. Petals pale pink to rose purple, occasionally white or nearly so, 7-4( the notch 1.1 (>- 4-0 4_)1.8-3.2(-4 (-3.2) mm long; the longer stamens, and often the shorter, shedding directly on the stigma at anthesis, or held away and bending toward the stigma after anthesis in some populations. Style white to slightly bluish, (2.2-)3.5-6.3(-9) mm long; stigma white, capitate to clavate, 1.1-2 mm long, 0.8-1.2 mm thick, occasionally exserted beyond the anthers. Capsules erect, strigillose to spreading hirsute, usu- ally with appressed to erect glandular hairs, often glabrate or glabrous, 3.5-7 (-8.5) cm long, 1 .3-1 .7 mm thick, on pedicels 0.3-2. 5(-3. 4) cm long. Seeds brown, papillose, obovoid, 0.8-1.1 mm long, 0.35-0.45 mm thick, the chalazal end with a short appendage, 0.04-0.1 mm long, 0.08-0. 14 mm wide; coma white to slightly 4-7 18. Distribution (Fig. 1): A widespread, weedy species that grows along stream banks, seeps, bogs, moist embankments, roadsides, or other open disturbed sites, mostly in the paramos and puna of the high Andes but also extends into forested zones. In South America throughout the northern and central Andes from the paramos of Venezuela and Colombia, including the Sierra de Santa Marta and the Sierra de Perija, southward through Ecuador; widely distributed in the alti- plano of Peru, Bolivia, northeastern Chile, and northwestern Argentina, continu- ing southward along the eastern side of the Andes to central Mendoza Prov., Argentina; disjunct in the Sierra de San Luis and the Sierra Grande de Cordoba. In North America restricted to the Sierra de Talamanca, Costa Rica. Found growing between 2,500 and 4,800 m throughout most of its range, descending to lower elevations along roads in forested zones, or along rivers in drier areas; at its southern limit typically between 1,800 and 3,000 m, and in the Sierra Grande de Cordoba between 1,200 and 2,500 m. Flowering throughout the year from Venezuela to Peru; however, from Bolivia southward there is a marked suppres- sion of flowering during the coldest, driest months (July and August in Bolivia) and therefore in the southernmost parts of its range, flowering occurs between December and March. Representative specimens examined: Argentina, catamarca: Dpto. Ambato, Sierra de Ambato, Casa de Cubas, 3,000 m. Hunziker & DeFulvio 19828 (CORD, MO); Dpto. Poman, Las Casitas, Sierra de Ambato, 2,500 m, Hunziker & Ariza 20424 (CORD, MO); Dpto. Capayan, Los Angeles, Russo 792 (LIL); Belen, Quebrada de Los Potrerillos, 2,600 m, Sleumer & Vervoorst 2461 (US). Cordoba: Dpto. San Javier, Quebrada del Tigre, 1 ,800 m, Bridarolli 1440 (LP); Pampa de Achala, Los Gigantes, Castellanos in 1924 (BA); Sierra Achala, Cuesta de La Sala Grande, Hieronymus 856 (CORD, F, US); Sierra Grande, Cuesta de Copina, Hunziker 8738 (CORD, MO, RSA); Sierra Grande, E slope Cerro Champaqui, Hunziker 9618 (CORD, MO, RSA); La Cumbrecita, 1,450 m, 31°55'S, 64°15'W, Solomon 4192 (MO), jujuy: Dpto. Yavi, Quebrada de Toquero, 3,600 m, 22°18'S, 65°36'W, Cabrera et al. 15380 (BAB, LP, M); Dpto. Tumbaya, Volcan Chilcayo, 23°43'S, 65°40'W, Fabris et al. 6290 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN I Vol. 69 57°W 40°S 800 Km Figure 1. Distribution of Epilobium denticulatum (LP). Dpto. Tilcara, Quebrada del Abra de la Cruz, Fabris et al. 6353 (BAA, LP); Dpto. Valle Grande, Caspala, 3,000 m, 23°I9'S, 65°08'W, Fabris & Crisci 6925 (LP); Dpto. Tumbaya, Angosto del Chani, 3,900 m, Kiesling et al. 298 (LP); Dpto. Humahuaca, Rio Despensas, 4,000 m, Ruthsatz 1V60 (BAA, MO), la rioja: Dpto. Famatina, Los Corrales, Cabrera et al. 27254 (BAB, SI); Dpto. Famatina, Sierra Famatina above La EnCrucijada, 3,200 m, 28°58'S, 67°42'W, Hawkes et al. 3404 (C); Sierra de 1982) SOLOMON— EPILOBWM IN SOUTH AMERICA 275 Famatina, Cueva de Medina, 3,100 m, Krapovickas 6163 (BAB, CORD), mendoza: Quebrada de Alvarado, 3 km S of Chilicito, 1,550 m, Hjerting 6322 (C); Dpto. Tunuyan, 10 km W of Campo de Los Andes, 1,500 m, Metis & Barkley in 1950 (LIL); Dpto. Tupungato, Rio La Carrera, 2,350 m, Sleumer427 (B). salta: Dpto. Calayate, Sierra de Los Quilmes, 2,400 m, Castellanos in 1943 (BA, RSA); Nevado Castillo, 25°1 1'S, 65°21'W, Lorentz & Hieronymus 88 (CORD, F, GOET, JE, S, US); Dpto. Guachipas, Pampa Grande, 1,600 m, 25°40'S, 65°30'W, Hunziker 1820 (POM); Dpto. Santa Victoria, Santa Victoria, 2,385 m, Meyer 4962 (F, UC). san juan: Dpto. Iglesia, Cordillera Agua Negra, Piedras Negras, 30°16'S, 69°47'W, Castellanos in 1950 (US); Dpto. Iglesia, Vega de Santa Rosa, 3,300 m, 29°00'S, 69°15'W, Hunziker & Caso 4791 (BAB, CORD); Cienaga de Las Cabeceras, 2,700 m, 31°47'S, 69°07'W, Kurtz 9808 (BAF, CORD), san luis: Pancanta, 32°59'S, 66°11'W, Cas- tellanos in 1925 (BA); Sierra de San Luis, Arroyo de Las Aguilas, Vignati 101 (LP), tucuman: Dpto. Tan, Lara, 3,200 m, Rodriguez 310 (BA, BAF, GH, MO, RSA, S, SI); Dpto. Chicligasta, Estancia Las Pavas, Rio Cascada, 2,700 m, Venturi 3033 (BA, BAB, DS, GH, MO, SI); Dpto. Tafi, Cerro San Jose, 2,700 m, Venturi 3573 (BA, BAB, DS, GH, RSA, SI, US); Dpto. Chicligasta, Estancia Santa Rosa, 3,300 m, Venturi 4787 (BA, BAB, LP, MO, SI). Bolivia, cochabamba: Cona-Cona, 4,000 m, 18°00'S, 66°45'W, Brooke 5181 (BM); Vila-vila, 18°00'S, 65°30'W, Brooke 5859 (BM, F, NY); Between Chimore and Cochabamba, Km 80, 3,000 m, 16°42'S, 64°49'W, Cardenas 760 (LIL, US). Prov. Ayopaya, Sailapata, 2,700 m, I6°30'S, 66°35'W, Cardenas 3057 (BR, F, P, S, US). Prov. Cercado, Sierra Tunari near Lago Huara-Huara, I7°I2'S, 66°I0'W, Hawkes & Hjerting 6606 (C, MO). Prov. Quillacollo, 28 km from Quillacollo on road to Morochata, 3,850 m, 17°19'S, 66°22'W, Hawkes & Hjerting 6692 (C). 35 km W of Comarapa on road to Cochabamba, 2,700 m, King & Bishop 7654 (MO. US). Prov. Carrasco, 2 km above Siberia, 3,200 m, Steinbach 210 (F, GH, MO, NY, US, WIS). Prov. Ayopaya, Puente San Miguel, 25 km NNW of Cochabamba, 3,800 m, 17°10'S, 66°25'W, Ugent478l (DS, GH, MSC, UC, US, WIS), la paz: Prov. Murillo, entre Pongo and La Rinconada, 4,000 m, Asplund 1818 (S, UPS). Prov. Ingavi, Guaqui, Asplund2305 (S, UPS, Z). Prov. Pacajes, General Campero, 3900 m, 17°27'S, 68°58'W, Asplund 2791 (UPS). Prov. Omasuyos, Isla del Sol, Challa, 3,850 m, Asplund 3709 (S, UPS). Vicinity of La Paz, 3,000 m, Bang 75 (E, G, GH, MO, NY, PH, US). Prov. Murillo, Palca, 5 km hacia la mina San Francisco, 4,800 m, Beck 2223 (MO). Prov. Ingavi, Huacullani, 3,840 m, Beck 2470 (MO). Mina La Fabulosa, 4,600 m, 16°00'S, 68°00'W, Brooke 6329 (BM, NY). La Paz, Buchtien in 1919 (ARIZ, BM, BP, C, LD, E, F, G, GH, M, MO, NY, S, US, Z); in 1930 (BM, C, E, G, K, M, MO, S, Z). 30 km 1 • ♦ SW of Yoloso on road to Unduavi, 2,900 m, Davidson 4968 (LAM). Prov. Larecaja, < Mandon 626 (F, G, GH, GOET, JE, MICH, MO, MPU, NY, PR, S, US, W); Williams 1549 (BM, NY). Yungas, 1,800 m, Rusby in 1885 (F, GH, MICH, NY, PH, US, WIS). Valle de Hichucota, Laguna Kkota, 4,500 m, 16°07'S, 68°20'W, Solomon 4938 (MO). Valle de Hichucota, 4,150 m, Sol- omon 5007 (MO); 11.3 km NE of La Paz on road to Unduavi, 4,400 m, Solomon 5013 (MO). La Cumbre on road to Unduavi, 4,600 m, 16°20'S, 68°03'W, Solomon 5028 (MO). 6 km W of Palca on Placa-La Paz road, 3,800 m, 16°32'S, 67°58'W, Solomon 5139 (MO). Upper Valle de Zongo, 3,000 m, 16°10'S, 68°09'W, Solomon 5243 (MO). Prov. Murillo, Valle de Achocalla, 3,800 m, 16°35'S. 68°1 1'W, Solomon 5298 (MO), potosi: Between Quechisla and Chorolque, 3,600 m, 20°56'S, 66°01'W, Car- denas 304 (US). Lagunillas, 3,800 m, Cardenas 410 (US), santa cruz: Near Valle Grande, 2,600 m, Cardenas 5138 (US), tarija: Iscayache, 3,600 m, Fiebrig 3015 (BM, E, G, CH, GOET, L, M, P, PR, PRC, S, US, W). Rincon de la Victoria, 17 km W of Tarija, Krapovickas et al. 18880 (CTES). Chile, i region (tarapaca): Arica, Camino de Arica al Portezuelo de Chapiquiha, Km 99, 3,500 m, I8°I9'S, 69°31'W, Marticorena et al. 59 (CONC, MO), 98 (CONC); Al interior de Chitita, 2,500 m, I8°49'S, 69°41'W, Schlegel5100 (MO). Iquique, Al interior deCoscaya, 3,360 m, 19°51'S, 69°09'W, Ricardi et al. 418 (CONC, MO); Quebrada de Quipisca, Noasa, 3,500 m, 19°59'S, 67°08'W, Werder- mann 1060 (BM, CAS, E, F, G, GH, LIL, MO, NY, S, SI, U, UC, US, Z); Chusmiza, 19°41'S, 69°1 l'W, Zollner 2947 (MO). Parinacota, Socoroma, 3,300 m, 18°I5'S, 69°37'W, Richardi et al. 190 (CONC, MO). II region (antofagasta): Antofagasta, Sierra Almeida, Aguada Chocas, 3,700 m, 24°15'S, 68°35'W, Biese 2337 (LIL), 2388 (LIL). El Loa, Caspana, 22°20'S, 68°14'W, Mahu in 1969 (LD); Toconoa, 23° 1 l'S, 68°01'W, Pfister in 1950 (CONC). Colombia, antioquia: Paramo Frontino, Cerro de Campanas, 3,650 m, 6°28'N, 76°04'W, Boeke & McElroy 302 (MO, NY), boyaca: Sierra Nevada del Cocuy, Laguna de San Pablo, 3,850 m, Grubb et al. 107 (COL, K, MSC, US); Quebrada Surbar, between Duitama and Charala, 3,273 m, Langen- heim 3537 (COL, UC, US), caldas: Nevado del Ruiz, entre Termales y Libano, Barclay & Juajibioy 6380 (COL, MO, RSA); Pinares, above Salento, 2,800 m, Pennell 9347 (GH, NY, PH, US); Paramo del Quindo, 3,700 m, Pennell & Hazen 9989 (GH, NY, PH, S, US, WU). cauca: Between Guachicono and Rio Putis, 2,500 m, Core 933 (RSA, US); Paramo de Las Papas, El Boqueron, Idrobo et al. 3014 (COL); San Antonio, 2,300 m, Pennell 7653 (GH, NY, PH, US); Entre Popayan y Purace, Rio Anambio, 2,500 m, Perez & Cuatrecasas 5869 (COL, US), cundinamarca: 15 km N of Usaquen, 2,600 m, Haught 6205 (COL, US); 20 km NE of Bogota, 2,470 m, Luteyn et al. 4809 (COL, F, MO, NY, US); Zipaquira, 2,650 m, Pennell 2542 (F, GH, MO, NY, US), huila: Moscopan, Santa Leticia, 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 2,230 m, Garcia-Barriga & Hawkes 12893 (COL, LIL, RSA, US); Balsillas, on Rio Balsillas, 2,(KH) m. Rushy & Pennell 728 (F, GH, MO, NY, US), magdalena: Sierra Nevada de Santa Marta, Hoya del Rio Donachui, 3,600 m, Cuatrecasas & Castaheda 24538 (COL, US); Sierra de Santa Marta, Paso de Mamancanaca-Cambiremena, 3,800 m, Weston 10391 (MO), meta: Paramo de Sumapaz, 4,100 m, Cleef7987 (COL, MO); Rio Arroz above Quebrada Pedregal, 3,445 m, Fosberg 20911 (RSA, US). narino: 31 km E of Pasto on road to Sibundoy, Berry 3253 (MO); Volcan Galera, Hacienda Obonuco, Espinosa 2777 (CAS, DS, RSA); Volcan de Cumbal, near Laguna Bolsa, 2,930 m, Ewan 16138 (BM, POM, RSA, US); Paramo del Tabano-Putum, Garcia-Barriga 4552 (COL); Camino de Tiiquerres a La Chorrera, 3,000 m, Mora 755 (AAU, COL), norte de santander: Valley of Rio Chitaga, SW of Pamplona, 2,600 m, Fassett 25958 (RSA, US); Laso, N of Toledo, 2,400 m, Killip & Smith 20391 (F, GH, NY, S, US), putumayo: Valley of Sibundoy, Portachuelo, 2,400 m, Schultes & Villarreai 7736 (COL, ECON, F, GH, US); Between La Maria and Paramo de San Antonio, 2,900 m, Schultes & Villarreai 7832 (COL, GH, LIL, US); Paramo de Quilisayaco, 3.3(H) m, Urihe 5380 (COL, NY). santander: Berlin, 3,100 m, Enrique et al. 577 (COL, MO); Paramo de Las Vegas, 3,300 m, Killip & Smith 15715 (BM, F, GH, NY, P, S, US); Paramo de Mogotocoro, 3,700 m, Killip & Smith 17647 (GH, NY, US); Alto del Almorzadero, 3,840 m, 6°59'N, 72°39'W, St. John 20802 (GH, MICH, NY, RSA, S, UC, US), tolima: 41 km E de Manizales, 3,250 m, Forero et al. 3662 (COL, MO); Quindio Highway, W of Cajamarca, 3,200 m, Killip & Varela 34554 (BM, COL, GH, S, US); Paramo de Ruiz, 3,400 m, Pennell 3070 (GH, NY, US), valle: Hoya del Rio Tulua, entre Las Vegas y La Ribera, 3,300 m, Cuatrecasas 20430 (F, RSA); Rio Cali, Pichinde, 1,700 m, Duque-J aramillo 4183 (COL). Costa Rica, cartago-san jose: Cerro de la Muerte, ca. 5 km above Villa Mills, 3,400 m, Allen & Dodge 5680 (F, G, US), Holm & litis 469 (A, F, G, UPS), Whitmore 39 (F, MO, NY); La Asuncion, 32 km N of San Isidro del General, 3,335 m, 9°34'N, 83°45'W, Burger & Stolze 5989 (F), Wilbur 21237 (DUKE); 8 km S of Taja, Tessene 1474 (WIS); Cerro Buena Vista, Weaver 1413 (DUKE). Ecuador, azuay: Nudo de Portete, 2,700 m. Camp E-2169 (NY); Lago Zurru Cuchu, 3.700 m. Prescott 829 (NY); Nabon, 3°20'S, 79°04'W, Rose 23016 (US), bolivar: Western slopes of Volcan Chimborazo, 3,600 m, Harling et al. 9632 (GB, MO, RSA); Simiatug. Hacienda Talahua, 3,200 m, Penland & Summers 628 (F, GH). canar: El Tambo, 69 km S of Sibambe, 3,000 m. Camp E-3982 (G, GH, MO, NY, P, RSA, UC, US); Cerro Bueran, 5 km S of Canar, 3,600 m, Fosberg & Giler 22639 (NY. US), carchi: Valle de Maldonado, 53 km W of Tulcan, 3,100 m, 0°5()'N, 78°03'W, Holm- Nielsen et al. 5570 (AAU, F, MO, NY, S, U); Paramo El Angel, 3,500 m, 0°4I'N, 77°54'W. Holm- Nielsen et al. 5289 (AAU, COL, F, MO, NY, S, U). chimborazo: Guamote, 3,000 m, Asplund 6878 (CAS. G, S, UPS, US); Between Chunchi and Sibambe, 2,200 m, Fagerlind & Wihom 783 (S); Paramo de Tililac, 3,600 m, Harling et al. 6573 (GB, RSA); 14 km S of Mocha, 3,800 m, 1°29'S, 78°43'W, Otis E-481 (MO, WIS), cotopaxi: 29 km E of Salcedo on Salcedo-Napo road, 3,860 m, Boeke 778 (MO, NY); Pilalo-Latacunga road, 3,400 m, Holm-Nielsen & Jeppesen 1477 (AAU, C, DS, NY, S); Volcan Cotopaxi, Hacienda Yacu-Tambo, 3,659 m, Ugent & Alhornoz 5672 (DS, MSC, UC, US, WIS), imbabura: Lago Cuicocha, Islote Chica, 3,080 m, Asplund 7161 (S); Rio San Marcos, E of Volcan de Cayambe, 3,350 m. Drew E-183 (C, COL, RSA, US); Mojanda, 10 km SSW of Otavalo, 2.900 m, 0°I0'N, 78°I8'W. Sparre 13463 (S). loja: Loja, Angelica, 2,300 m, Espinosa 363 (RSA); Lado sur de Saraguro, 2,300 m, Espinosa 1386 (POM); Near Yangana, 2,000 m, Hart 1032 (MO). morona-santiago: Entre Gualaceo y General Plaza, Km 29, 3,000 m, Sparre 18777 (S). napo: Km 216 on Quito-Papallacta road, 3,700 m. Berry 2522 (MO); Cerro Antisana, 4,000 m, 0°30'S, 78°00'W, Gruhh et al. 575 (DS, MSC, NY); 10 km E of Papallacta, 2,800 m, 0°2I'S, 78°0I'W, Holm-Nielsen et al. 6839 (AAU, MO. NY, S). pichincha: Tumbaco, 2,450 m, Asplund 6554 (S); Pichincha, 3,650 m, Asplund 6581 (S); Valle de La Magdalena, Chillogallo, 2.800 m, Firmin 99 (F, S, US); Quito, Humboldt & Bonpland in 1801 (F, P, RJ); Volcan Pichincha, Sodiro in 1871 (BP, JE); Lagunita Linda, 3 km W of Paso de Guamani, 3,900 m, Sparre 15063 (S); W of Mt. Iliniza, 3,320 m, Weydahl 153 (S). tungurahua: Patate, Hacienda Leito, 2,900 m, Asplund 7961 (S); 10 km S of Mocha. 3.500 m, Harling 6938 (GB); Vicinity of Ambato, Ficoa, Pachano 145 (GH, NY, US); Rio Colorado, Volcan Chimborazo, 4,200 m, Sparre 18364 (S). zamora chinchipe: 20 km E of Loja on road to Zamora, 2,350 m, Mathias & Taylor 5195 (RSA). Peru, ama/onas: Chachapoyas, 6°13'S, 77°5I'W, Matthews in 1834 (BM. BR, E, K, OXF); Prov. Bongara, hills WNW of Pomacocha, 2,700 m, 5°50'S, 77°55'W, Wurdack 924 (F, NY, RSA, UC, US), ancash: Cordillera Negra, Callan, 4,200 m, Bernardi 16676 (G); Prov. Bolognesi, Parien- tana, E of Chiquian, 3,420 m, 10°09'S, 77°1 1 'W, Ferreyra 5659 (MO. US, USM); Prov. Carazi, Laguna Paron, 4,100 m, 9°I0'S, 77°35'W, Mostacero et al. 547 (MO, NY), apurimac: Prov. Abancay, 16 km E of Abancay on Cuzco road, 3,000 m, litis & Ugent 64 1 (DS, WIS); Prov. Andahuaylas, Mollebamba, 3,650 m, Vargas 8721 (LIL). arequipa: 20 km Wof Arequipa, Averett 1004 (MO); S slopes of Nevado Chachani, Hinklev 65 (CAS, GH, NY, US); Prov. Cailloma, 15 km NE of Chivay, Mueller et al. 2221 (LZ); Arequipa, Rio Chili, 2,500 m, Munz 15511 (F, GH, NY, POM); Pichu-Pichu, 3,000 m, I6°29'S, 7I°15'W, Sandeman 3758 (K, OXF); Prov. Condesuyos, Chuquibamba. 3,350 m, Stafford 1165 (BM, F, K). ayacucho: Prov. Huanta, mountains NE of Huanta, 3,200 m, Weberbauer 7505 1982] SOLOMON— EP1LOBWM IN SOUTH AMERICA 277 (F, GH, NY, US, WIS), cajamarca: Prov. Contumaza, Lleden, 2,600 m, Sagdstegui et al. 9408 (MO): Prov. Cajamarca, Granja Porcon (SIPA), 28 km NW of Cajamarca, 3,260 m, Ugent 5454 (DS, UC, WIS), cuzco: Prov. Quispicanchis, Marcapata, 3,000 m, I3°30'S, 70°55'W, Berry & Aronson 3020 (MO); Sicuani, 3,551 m,Hicken in 1903 (BAF, SI); Pampade Anta, 7 km W of Anta, Ancahuasi, 3,150 m, litis & Ugent 783 (DS, MSC, UC, WIS); Cordillera Vilcabamba, Rio Raccachaca, 4,500 m, McSweeney in 1976 (MO); Ollantaitambo, Munz 15544 (NY, POM, US, WTU), Pennell 13670 (F, GH, NY, PH, S, US); La Raya, 4,400 m, Pennell 13514 (F, PH); Entre Paucartambo y Tres Cruces, Cerro de Cusilluyoc, 3,400 m, Pennell 14157 (F, GH, PH); Ausangate, 4,700 m, Rauh-Hirsch PI 151 (RSA); 4 km NE of Cuzco on road to Pisac, 3,500 m, Ugent 4098 (DS, GH, MICH, MSC, UC, US, WIS); Prov. Paucartambo, 5 km NE of Paucartambo, 2,900 m, Ugent & Vargas 4407 (DS, US, WIS); Prov. Espinar, Huaillapacheta, 14°45'S, 71°25'W, Vargas 6507 (CAS, LIL); Prov. Canas San Andres de Checca, 3,860 m, 14°25'S, 71°20'W, Vargas 1 1014 (F, NA, UC); Prov. Urubamba, Canon Chicon, 2,900 m, Vargas 11054 (F, GH, NA, UC). huancavelica: San Jose de Acobambilla, 3,700 m, 12°40'S, 75°22'W, Lloyd et al. 536 (K); Prov. Tayacaja, Pampas, 3,250 m, Stork & Morton 10236 (F, G, GH, NA, UC); Prov. Huancavelica, Machajhuay between Conaica and Tinyajlla, 3,800 m, Tovar 801 (MO), huanuco: 9 km from Chirlin, 3,660 m. Gentry et al. 19214 (MO); Tambo de Vaca, 3,900 m, 9°42'S, 75°47'W, MacBride 4373 (F, GH, S, US); Pillao, 2,700 m, 9°40'S, 75°58'W, Woytkowski 34119 (F, GH, MO, UC). juni'n: Oroya, 3,700 m, Kalenborn 19 (DS, GH, MO, US). Prov. Yauli, Lago Huacracocha, 3,750 m, 1 1°40'S, 76°10'W, Lopez & Riccio 10820 (RSA); Tarma, Hacienda Maco, 3,600 m, Ochoa 238 (CORD, NY), la libertad: 70 km E of Trujillo, between Pampa and Yamo- bamba, 3,050 m, Conrad 2715 (MO); Entre Motil y Quiruvilca, Km 104-105, 3,300 m, Ferreyra 3026 (MO, USM); Prov. Patazi, entre Huancaspata y Tayabamba, 3,800 m, Lopez & Sagdstegui 8262 (MO), lima: Prov. Yauyos, Huancracha, 3,300 m, Cerrate 1180 (MO, USM); Rio Blanco, 3,000 m, Killip & Smith 21579 (F, NY, S, US); MacBride & Featherstone 677 (F, GH, K, S, US): Prov. Huarochiri, Casapalca, 4,700 m, MacBride & Featherstone 857 (F, GH, S, US), moquegua: Saylapa, near Carumas, 3,300 m, 16°49'S, 70°43'W, Weherhauer 7337 (F, S). pasco: La Quinua, 3,600 m, 10°33'S, 76°10'W, MacBride & Featherstone 2023 (F, GH, S, US); Paucartambo, 2,800 m, Woyt- kowski67ll (GH); Cerro de Pasco, 4,130 m. Ellenherg 411 (MO), puno: Amantani, 3,900 m, I5°32'S, 70°01'W, Aguilar234 (MO, USM); Prov. Azangaro, Cala-Cala, 4,100 m, 15°14'S, 70°33'W, Bemardi 16745 (G): between Ollachea and Macusani. Dillon et al. 1266 (MO); Prov. Sandia, 2-6 km S of Limbani, 3,550 m, Metcalf 30452 (UC, US); Santa Rosa, 80 km SSW of Have, 4,300 m, Pearson 33 (PH); Araranca, 4,100 m, Pennell 13432 (F, GH, NY, PH, S, US); Prov. Huancane, Occa Pampa, 3,965 m, 15°10'S, 69°35'W, Shepard 78 (GH, NY, US); 16 km NE of Macusani, 4,600 m, Webster 2 (K). san martin: Prov. Huallaga, valley of Rio Apisoncho, 30 km above Jucusbamba, 2,800 m, 7°55'S, 77°10'W, Hamilton & Holligan 1411 (S, UC). Venezuela, lara: Las Sabanetas, above Los Aposentos, 2,530 m, Steyermark 55311 (F, MO, NY, VEN). merida: La Carbonera, 2,200 m, Aristeguieta 2478 (F, MER, NY, US, VEN); Pico El Aguila, Carretera Chachopo, 3,600 m, Badillo 5176 (MY); Paramo de Los Conejos, 3,500 m, Bemardi 1257 (MER, NY); Paramo de Mucuchies, 3,750 m, Humbert 26304 (P, RSA, US); Laguna Mucubaji, 3,650 m, Steyermark 57508 (F). tachira: Cabeceras del Rio Quinimari, 15 km S de San Vicente de La Revancha, 2,400 m, Steyermark et al. 100637 (VEN); Quebrada Agua Azul, 14 km SE de Delicias, 2,300 m, 7°3I'N, 72°24'W, Steyermark & Liesner 118373 (MO, VEN). trujillo: Near Guirigay, 3,300 m, Aristeguieta & Medina 3647 (NY, VEN); Dist. Bocono, Paramo de Tufiame, 3,120 m. Berry 3122 (MO); Entre La Pena y Agua del Obispo, 28-34 km de Carache, 2,400 m, Steyermark 104954 (M, S, VEN). zulia: Dist. Perija, Serrania de Los Motilones, 10°00'N, 72°58'W, Tillett & Honig 747-834 (MO); Dist. Maracaibo, Serrania de Valledupar, 3,100 m, 10°25'N, 72°52'W, Tillett 747-1132 (MO). Epilobium denticulatum is a common, widespread, highly variable species that has its primary distribution throughout the northern and central Andes. It may be distinguished from other closely related species by its nodding inflorescence, bluish or lavender staminal filaments, leaves with relatively few denticulate teeth, and strongly appressed, or more rarely, slightly spreading, minute blunt-tipped hairs on the surfaces of the leaf blades. This latter feature and colored staminal filaments are shared with E. pedicel hire, but they are unknown in any other species. Epilobium pedicellare is immediately separable, however, by its alternate serrate leaves and erect inflorescences. fragile fragil its densely caespitose habit, thin wiry stems mostly less than 10 cm tall, sparse 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 69 pubescence, and few flowers. Some plants of E. denticulatum approach E. fragile in stature, but they generally have larger leaves, denser pubescence, more flowers and the appressed hairs mentioned above. Epilobium ciliatum is sympatric in a very limited area at the southern end of the range of E. denticulatum, and can be separated immediately by its white flowers, fleshy turions, and longitudinally ridged seeds. Epilobium denticulatum was the first species described from South America. Its original circumscription and distribution, however, were based on two species, as recognized here, E. denticulatum and E. puberulum. From the beginning, the name E. denticulatum was used for the widespread northern Andean species. Specimens of E. puberulum have often been misidentified as E. denticulatum. These two species are allopatric, and, while they may be closely related, E. puberulum differs significantly in its erect inflorescences, complete lack of glan- dular hairs, distinct floral morphology, and petal color. The long persistent confusion concerning the assignment of plants to the var- ious taxa surrounding Epilobium denticulatum is well illustrated by the problems encountered by Haussknecht (1884). He recognized seven species in his group Denticulatae, one of which is a synonym of E. pedicellare . A careful reexami- nation of the specimens he used showed that five of the remaining six species are heterogeneous, containing components from two or more species. Thus, his treat- ments of E. denticulatum, E. meridense, and E. caesium include collections of E. puberulum (e.g., Coronel, Arauco Prov., Ochsenius, GOET; Hacienda San BREM;andQuebrad Ma Mexico Prov., Chile, Gay in 1837, P). Part of the confusion with this last name is probably due to the fact that the five specimens of E. denticulatum cited, including type material of E. bonplandianum, are all late season, flowered-out plants that are superficially very similar to some specimens of E. ciliatum. Epilobium repens Me to E. ciliatum (Hoch, 1978). Haussknecht, however, included four collections of E. denticulatum in the specimens cited under that species. Superficially, the Mexican denticulatum from South America in stature, loose innovations, and the nodding inflorescences, but are distinguish- able by other features. Although Haussknecht was a very astute observer of epilobiums and had a broad understanding of them on a world wide basis, the species that he included in his Denticulatae were apparently very troublesome to him. His Monographic (Haussknecht, 1884) formed the source for the report of E. denticulatum, E. bonplandianum, E. caesium, and E. meridense in the Flora de Chile (Reiche, 1898). The most recent treatment, and the only subsequent work of any consequence, for Epilobium denticulatum is that of Samuelsson (1923). He recognized ten species and five varieties in this complex, of which six species and four varieties were proposed as new. Many of his new taxa were based on unique morphological types, which because of the relatively few collections available to him seemed amply distinct, e.g., E. hirtum, E. bolivianum. It is interesting to note that even though Samuelsson described a large number of new taxa, he discussed in several 1982) SOLOMON— EPILOB1UM IN SOUTH AMERICA 279 places the difficulty of separating the species and their polymorphic nature (Sam- uelsson, 1923; p. 251, 255, 257). In a later publication (Samuelsson, 1930), he rejected two of these new species, E. andicolum and E. asplundii, considering them to be robust variants of E. denticulatum, and synonymized E. denticulatum var. aberans with E. meridense var. helodes. In delimiting most of his taxa, Samuelsson placed great emphasis on the density and distribution of pubescence, especially on the ovaries, pedicel length, and flower size, all of which are rela- tively variable characters. Patterns of morphological variation within E. denticulatum, as circumscribed here, are extremely reticulate and diverse, and reflect the variety of selection pressures encountered throughout its enormous range. The same types of mor- phological features appear numerous times throughout its range, with some geo- graphic differentiation. Superimposed on these patterns is variation because of elevational and microclimatic conditions, as well as interpopulational differences related to the high degree of autogamy and consequent fixation of certain char- acters. The size, habit and branching patterns of E. denticulatum are extremely vari- able and are strongly influenced by exposure or elevation. The smallest plants are often found at higher elevations or in open or drier sorts of places. At these same elevations, however, much more robust plants can usually be found in more sheltered areas. There is also a pattern of increasing size at lower elevations. Most plants that exceed 80 cm in height come from areas below 3,000 m. Branch- ing patterns are dependent on elevation and microclimate, but also on the age of the plants. Often stems are simple or with a few branches. As the plant ages, they may branch profusely or not at all. This feature is highly variable within populations and shows no geographic correlation. Many times, older stems that have nearly completed flowering may lose the nodding aspect of the inflorescence and appear erect. Leaf size and, to a lesser extent, shape vary considerably and usually in parallel with the size of the plants, the smaller leaves on small plants, and larger ones on large plants. There are sporadic collections of extremely small-leaved plants from a number of different localities. Similarly, large-leaved, usually more robust plants are also found throughout the range of the species with increasing frequency in southern Peru, Bolivia, and Argentina (e.g., Between Trujillo and Bocono, Edo. Trujillo, Venezuela, Alston BM Miranda MO Prov., Argentina, Hjerting et al. 9457, C). Plants of this type formed the basis for the names E. andicolum and E. asplundii. Plants with very large leaves (greater than 5 cm) are found only in the altiplano and again in the Sierra Grande de Cordoba. Throughout the altiplano and south- ward in San Juan and Mendoza provinces, Argentina, there is a mixture of small and large-leaved plants, often within the same population. The populations from the Sierra Grande de Cordoba consist of predominantly large-leaved and usually robust plants. This pattern is modally different from that found in the rest of the range of E. denticulatum but is most likely the result of the derivation of the 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Voi . 69 Sierra Grande populations from more or less large-leaved plants from the southern altiplano to the west and northwest by long-distance dispersal. The primary leaf shape in E. denticulatum is lanceolate, with acute apices. Rarely individuals may have nearly ovate leaves with subcordate bases (e.g., La Cumbre, Dpto. La Paz, Bolivia, Asplund4014, the type of E. bolivianum; Nevado del Ruiz, Dpto. Caldas-Tolima, Colombia, Barclay & Juajibioy 6380, RSA; Calca, Dpto. Cuzco, Peru, Hammerlund 620, S). More frequently the larger-leaved plants mentioned above have somewhat elongate apices, so that the leaves are markedly acuminate. Pubescence density and distribution is also extremely variable. In general, the upper stems and ovaries are strigillose with an admixture of appressed or slightly spreading glandular or blunt-tipped hairs. These hairs also occur on the surfaces of the leaves, a feature shared only with E. pedicellare. Often the leaves become glabrate as they mature, so that only a few or none of the appressed hairs are found on the surfaces of older leaves. Density of both hair types is highly variable throughout the range of the species, although there is a trend toward denser, almost canescent, strigillose pubescence in the southern paramos and altiplano. The increased pubescence may in part be due to the drier atmospheric conditions found throughout much of the altiplano, in contrast to the moist paramos found in Costa Rica, Venezuela, and Colombia, where this pubescence type is not common. The most striking qualitative difference in pubescence type seen in E. den- ticulatum is the erect or spreading hirsute hairs that characterize plants that have gone under the name E. hirtum (e.g., Pichupichu, Dpto. Arequipa, Peru, Sande- man 3758; Chuquibamba, Dpto. Arequipa, Peru, Stafford 1165; La Paz, Bolivia, Buchtien 531; Socoroma, Tarapaca Prov., Chile, Ricardi et al. 190). The distri- bution of this character is limited to the vicinity of La Paz, Bolivia, Arequipa, Peru, and a single collection from Tarapaca Prov., Chile. In addition to the trans- formation of strigillose hairs to longer erect ones, the usually appressed glandular hairs of the leaves, stems, and ovaries are also erect and longer than normal. Epilobium hirtum was originally described from robust, fairly large-leaved plants from La Paz, Bolivia. Except for pubescence, most of these plants are indistinguishable from those called E. andicolum (e.g., Ollantaitambo, Dpto. Cuz- co, Peru, Pennell 13670). Mixed populations of plants with appressed and erect pubescence are indistinguishable except on the basis of the hair type (e.g., Rio Chili, Arequipa, Dpto. Arequipa, Peru, Munz 15511). The hirsute pubescence gives the plants such a distinctive appearance that those with this feature are immediately identified as E. hirtum. This is an excellent example of the artificial separation of plants into different taxa based on a single character in a highly polymorphic group. Flowers in E. denticulatum vary considerably in size, but exhibit no geograph- ic patterns in their size distribution. Plants with very large flowers or very small ones are found throughout the range of the species, although the majority have petals of intermediate size. Even within populations flower size can be variable, with mixtures of large and small flowered plants (e.g., La Oroya, Dpto. Junin, Peru, Kalenhorn 19; and Araranca, Dpto. Puno, Peru, Pennell 13432). At anthesis most flowers, as well as the inflorescence, as a whole nod. In smaller flowers the anthers of the longer stamens, and often the shorter, usually 1982| SOLOMON— EPILOBIUM IN SOUTH AMERICA 281 dehisce directly on the stigma, sometimes even before the flower fully opens. In larger flowers, the stamens are normally held away from the stigma at anthesis. After several hours, the longest bend inward, shedding their pollen on the stigma. Only rarely is the stigma completely exserted (Valle Paucartambo, Dpto. Cuzco, Peru, Herrera 3358, F); more frequently in the larger flowers, the stigma is par- tially exserted, but the longest stamens are able to reach at least the lower portion of the stigma, so some measure of self-pollination can take place (e.g., Ollantai- tambo, Dpto. Cuzco, Peru, Pennell 13670, PH, GH, F, NY; Huanta, Dpto. Ay- acucho, Peru, Weberbauer 7505, F, GH, NY, US). On cloudy or rainy days even the largest flowers may open only partially, or not at all, making them functionally cleistogamous. Epilobium denticulatum is sympatric with at least E. pedicellare, E. hirti- gerum. fr fi within that of E. denticulatum. Epilobium denticulatum is sympatric with E. hirtigerum only in the Sierra Grande de Cordoba and with E. ciliatum in this same mountain range, and also in the Sierra de San Luis and in a narrow band in southern San Juan and eastern Mendoza Province, Argentina. Epilobium glaucum overlaps with E. denticulatum only in this last named area. Natural hybridization involving E. denticulatum and any of these other species is apparently relatively rare. Several hybrids have been reported involving E. pedicellare (Samuelsson, 1923, 1930), but it is believed that all except one were based on non-hybrid specimens of E. pedicellare (q.v. for discussion), although hybrids of this combination would be very difficult to distinguish. Two specimens of what are possibly hybrids between E. denticulatum and E. ciliatum have been seen (Copina, Sierra Grande, Cordoba Prov., Argentina, Grassi 2280, LIL; and in the vicinity of Mendoza, Mendoza Prov., Argentina, Carette in 1906, BA). Both have intermediate leaf shape, appressed hairs on the leaf surfaces, strongly reduced seed set, and pollen stainability of 27% and 23% respectively. Although E. denticulatum and E. ciliatum both have the same chromosome arrangement, AA, the depressed pollen stainability is not unusual. Artificial hybrids between these species had pollen stainability in the range of 37-65%. fragil fi MO Rio Despensas, Dpto. Cochimoca, Jujuy Prov., Argentina, Boelcke et al. 770, Mar MO paca Prov., Chile, Richardi et al. 418, MO, CONC) that approach E. fragile. They are tentatively retained in E. denticulatum because of their larger size, dense strigillose pubescence, and appressed hairs on the leaf surfaces. These plants will deserve careful reconsideration when more material becomes available from these poorly explored regions. 2. Epilobium pedicellare Presl, Reliquiae Haenkeanae 2:30. 1831. type: "Chile," without specific locality, 1790, T. Haenke (PR-495768, holotype, photograph MO). Probably collected in Peru, and the label information is in error. E. haenkeanum Hausskn., Oesterr. Bot. Z. 29:118. 1879. type: Peru, without specific locality, 1790, 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 T. Haenke (PR, holotype, photograph MO, fragment JE). Hausskn., Monogr. Epilobium 268, tab. 16, f. 72, 72a. 1884. H. Lev., Iconogr. Epilobium, tab. 208. 1911. Samuelsson, Svensk Bot. Tidskr. 17:262. 1923. MacBride, Field Mus. Nat. Hist., Bot. Ser. 8(4):532. 1941. E. peruvianum Hausskn., Monogr. Epilobium 263. 884. type: Peru, Dpto. Lima, "ditches and Rio Rimac, near Lima," W. Nation (K, holotype, photograph MO). H. Lev., Iconogr. Epilobium, tab. 210. 1911. Robust perennial herb, the stems (30-)4Q-120 cm long, reproducing vegeta- tively by elongate leafy shoots from near the base. Stems decumbent to pendent, or ascendent, sparsely branched above, usually with an abrupt bend leading to the erect inflorescence, thick, hollow, lustrous reddish brown, strigillose all around the stem in the inflorescence, below pubescence restricted to raised descending lines from the decurrent petiole bases. Leaves mostly alternate, opposite only on young shoots or near the base, thin, dark green, narrowly ovate to ovate, 1.2- 4.3(-5) cm long, 0.5—1 .4(— 1 .8) cm wide, acute to acuminate, rarely obtuse at the apex, finely serrate or doubly serrate with (7-)16-30(-43) irregularly sized teeth on each side, acute to cuneate at the base, sparsely strigillose on the margins and the abaxial midrib, both surfaces with appressed blunt-tipped or glandular hairs, 0.05-0.1 mm long, lateral veins prominent beneath, 3-4(-5) on each side of the midrib, on petioles 1— 3(— 4) mm long, rarely sessile. Inflorescence erect, simple, the flowers congested at anthesis, the internodes elongating somewhat in fruit. Flowers erect. Ovaries often reddish, strigillose, sometimes densely so, usually with scattered appressed glandular hairs, 0.7-1.5 cm long, on pedicels 0-1 mm long. Floral tube often reddish purple, 1.1-1.5 mm deep, 1.7-2.4 mm across, externally strigillose with scattered appressed glandular hairs, internally glabrous or with a few erect villous hairs, 0. 1-0. 15 mm long, in a ring near the base. Sepals often reddish purple, lanceolate, 2.5-3.4 mm long, 1.0-1.3 mm wide, strigillose with scattered appressed glandular hairs. Petals pale pink to rose purple, obovate, 3.5-5.5 mm long, 1.7-2.5 mm wide, the notch 0.9-1.3 mm deep. Anthers cream, 0.6-0.9 mm long, 0.4-0.6 mm wide; filaments bluish purple to pale lavender, those of the longer stamens 1.7-2.6 mm long, those of the shorter 0.7-1.8 mm long, the longer, and often the shorter, stamens shedding directly on the stigma at anthesis. Style white, 2.4-3.1 mm long; stigma white, clavate, 1.0-1.4 mm long, 0.6-0.8 mm thick. Capsules erect, with scattered strigillose hairs, 3.1-4.8 cm long, 1.1- 1 .5 mm thick, on pedicels 0.3- 1 .5(-2.5) cm long. Seeds brown, papillose, obovoid, 0.8-1.0 mm long, 0.3-0.4 mm thick, the chalazal end with a short appendage 0.04-0.08 mm long, 0.08-0.12 mm wide; coma white, 4.5-7 mm long, readily detaching. Gametic chromosome number, n = 18. Distribution (Fig. 2): On moist stream embankments or seeps, widely scattered in the altiplano of central and southern Peru and western Bolivia at elevations of 2,400-3,600 m. Flowering throughout the year, but perhaps with reduced flow- ering from June through August, the coolest, driest months. Specimens examined: Bolivia, la paz: Prov. Murillo, La Paz, Asplund 58 (UPS); Bang 75a (NY, PH, US); Buchtien in 1907 (C, F, US), in 1910 (BAF, E, F, GB, GH, JE, L, NY, UPS, Z), in 1910 (BM), in 1912 (K), in 1913 (C, JE, LD, MO, S, UPS), in 1921 (GH), 8431 (US), 8433 (B, BM, BR, C, E, G, K, M, MO, S); Pennell 14217 (F, GH, NY, PH); Rusby 1806 (April 1885) (NY, WIS), 1806 (Oct, 1885) (F, GH, MICH, NY, PH, US); Shepard 157 (CAS, GH, NY, P); Solomon 5147 (MO); Thuegsegges & Feurer 4098 (MO); Weddell in 1851 (JE, P). Peru, Without locality, MacLean (K); Nee in 1790 (F, MA), ancash: Prov. Cajatambo, Ocros, 3,400m, I0°24'S, 77°24'W, Weberbauer 2706 (G, UPS), arequipa: Arequipa, Rio Chili, Averett 1007B 1982] SOLOMON— EPILOB1UM IN SOUTH AMERICA 283 800hm 800 km Figures 2-3. — 2. Distribution of Epilobium pedicellare. — 3. Distribution of Epilobium fragile. (MO); Harvard Observatory, Castle in 191 1 (GH); without locality, Harrison in 1896 (GH); Prov. Cailloma, Chivay, 3,500 m, 15°38'S, 71°36'W, Mueller et al. 2134 (LZ). huanuco: Tambo de Vaca, 9°42'S, 75°47'W, Martinet 168 (P). junin: Abocongo, Infantes 160 (LIL). lima: Prov. Huarochirf, Viso, 2,700 m, Asplund 11137 (S); Prov. Yauyos, Tupe, 2,950 m, 12°40'S, 75°46'W, Cerrate 1116 (MO); Prov. Huarochirf, Picoy arriba de Surco, 3,200 m, 11°52'S, 76°28'W, Ferreyra 6086 (MO, US, USM); Prov. Canta, entre Huascoy y Cormo, 3,000 m, 11°25'S, 76°45'W, Ferreyra 18405 (MO); Matucana, MacBride & Featherstone 90 (F, G, GH, S, US); Martinet (P); Rio Chillon, Km 123 on road Canta-Culluhuay, Mueller et al. (LZ); Rio Chillon, Obrajillo, 11°54'S, 77°09'W, Pennell 14384 (PH); Rio Huaura, 11°06'S, 77°39'W, Ruiz & Pax on in 1778 (P). Epilobium pedicellare is easily distinguished from the other South American species by its mostly alternate, finely serrate leaves, the teeth of uneven size, thick hollow stems, and an erect, congested inflorescence at least in early flow- ering. Only E. hirtigerum from eastern South America and the introduced E. paniculatum have mostly alternate leaves, but their ranges are widely separated from that of E. pedicellare, which is confined to the altiplano of Peru and Bolivia. Presl originally described this species from a specimen collected by T. Haenke during the Malaspina Expedition (1789-1793). The herbarium sheet at Prague (PR) is labeled as coming from Chile, although this species is not known to grow within Chile as it was constituted in 1790, nor did Haenke collect in northern Chile, the only areas where E. pedicellare might be found (Kiihnel, 1960). Undoubtedly the plant is mislabeled and probably came from Peru, since the many collections from the Malaspina Expedition are notorious for their incorrect labels (cf. Hitchcock, 1 909) . Shortly after the appearance of Reliquiae Haenkeanae, Hooker & Arnott (1833) used the name Epilobium pedicellare to which they referred several Chil- ean and Argentinean collections of John Gillies. These proved to be a mixture of E. denticulatum, E. ciliatum, and E. glaucum, thus obscuring the identity of E. pedicellare early on. The next botanist who dealt with any material referable to E. pedicellare was Haussknecht, who described E. haenkeanum, also from a Haenke specimen at 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Prague (PR), but not the same one used by Presl (Haussknecht, 1879). Apparently Haussknecht never saw the type of E. pedicellare , although his annotations are found on a number of other specimens from Presl' s herbarium. As a result, in the preparation of his Monographic Haussknecht was uncertain as to the exact identity of E. pedicellare and tentatively assigned it as a synonym of E. dentic- ulatum (Haussknecht, 1884). To confuse the issue further, the type of Epilohium haenkeanum, at the time Haussknecht described it, was annotated as E. denticulatum with reference to Presl' s brief description of that species in Reliquiae Haenkeanae (Presl, 1831). This description, however, differs substantially from the characters exhibited by the specimen. Haussknecht (1884) pointed out this discrepancy and provisionally placed Presl' s description of E. denticulatum with his E. caesium without having seen the specimen from which Presl drew the description. Among the extant specimens at Prague (PR), there is a single sheet annotated as E. denticulatum in the same hand as the label for E. pedicellare, presumably that of Presl. This specimen is a small piece of a probable hybrid between E. ciliatum and E. glau- cum, which matches the description of E. denticulatum as given by Presl and is probably the one on which it was based. Unfortunately, this specimen is labeled as coming from Chile, not Peru, as given in Reliquiae Haenkeanae, so some doubt still remains. In his Monographic, Haussknecht (1884) also described a third taxon referable to Epilohium pedicellare as E. peruvianum, which he allied with E.franciscanum Barbey (E. ciliatum subsp. watsonii (Barbey) Hoch & Raven) to which it bears a vague resemblance but is not directly related. As a result of Haussknechf s interpretation of E. pedicellare and the difficult access to PresFs material, subsequent authors have used the name E. haenkea- num, with E. pedicellare considered as a synonym of E. denticulatum. Epilohium pedicellare has been relatively rarely collected and is apparently much more scattered in its distribution than the widespread E. denticulatum with which it often grows. This may be due to some ecological specialization because the only habitats where this species has been seen are very steep, almost vertical seeps that are continuously wet. In contrast, E. denticulatum is much more weedy, growing in ditches, seeps, stream banks, or almost any habitat that has a per- manent supply of water. In places where E. pedicellare is found, it often grows in a pendent or reclining fashion with an abrupt U-shaped bend before the begin- ning of the erect inflorescence. The stems are often more than I m long and form dense masses. This unusual habit can often be seen on herbarium material and is very different from the ascending or erect stems and nodding inflorescences of E. denticulatum. The nearest relative of E. pedicellare is probably E. denticulatum, with which it shares two unique characters, unknown in any other South American species: the presence of minute appressed or slightly spreading blunt-tipped hairs, usually on both surfaces of the leaves, and pale lavender to bluish staminal filaments. Because the range of Epilohium pedicellare is completely contained in that of E. denticulatum and they often grow together, it would seem likely that hybrids would occasionally be produced. Samuelsson (1923, 1930) reported a number of plants as hybrids, but only one specimen of these seems likely to be of truly 1982) SOLOMON— EPILOBIUM IN SOUTH AMERICA 285 hybrid origin (La Paz, Dpto. La Paz, Bolivia, Buchtien 37, US, mixed with E. pedicellare). The others fall within the variation seen in E. pedicellare, although hybrids of this combination are very difficult to distinguish. Based on the mor- phology of the two putative parents, the specimen indicated above is inter- mediate with leaves having fewer, less sharp teeth, thinner stems, and larger flowers than is typical for E. pedicellare. This specimen had a pollen stain- ability of 81%. 3. Epilobium fragile Samuelsson, Svensk Bot. Tidskr. 17:291, tab. 4, f. 3a-c. 1923. type: Bolivia, Dpto. La Paz, Prov. Murillo, "in rupibus irrigatis," La Cumbre, 4,500 m, 26 May 1921, E. Asplund40l5 (UPS, lectotype, photograph MO; B (destroyed, photographs BH, MO, US), LD, S, W, Z, isolectotypes). Samuelsson, Svensk Bot. Tidskr. 24:7. 1930. MacBride, Field Mus. Nat. Hist., Bot. Ser. 8(4):531. 1941. E. nivale auct. non Meyen: Samuelsson, Svensk Bot. Tidskr. 17:290. 1923, pro parte; MacBride. Field Mus. Nat. Hist., Bot. Ser. 8(4):532. 1941. Low, densely caespitose perennial herb, 3— 10(— 1 5) cm tall. Stems many, thin, usually reddish brown, decumbent or erect, mostly simple or few-branched from a fibrous, or somewhat woody rootstock, subglabrous, with a few strigillose hairs, 0.1-0.2 mm long, at the petiole bases, or occasionally scattered in descending lines from the petiole bases, rarely with a few erect glandular hairs, 0.1-0.2 mm long. Leaves all opposite or alternate in the inflorescence, thick, occasionally reddish purple, narrowly ovate to elliptic, rarely lanceolate, 0.4-1.2 cm long, 1.5- 4 mm wide, obtuse to rounded, rarely acute at the apex, subentire with 0-3 obscure teeth on each side, acute to cuneate at the base, glabrous, or with a few scattered strigillose hairs on the margins, lateral veins obscure, none or 1-2 on each side of the midrib, on poorly defined petioles, 0-2.5 mm long. Inflorescence erect, simple, few-flowered. Flowers erect at anthesis. Ovaries reddish purple, 0.4-1 cm long, glabrous, or with a few strigillose or erect glandular hairs, on pedicels 1-9 mm long. Floral tube often reddish purple, 0.8-1.2 mm deep, 1.1- 2.4 mm across, externally glabrous or with a few strigillose or erect glandular hairs, internally with a sparse ring of a few erect villous hairs, 0.08-0.13 mm long, near the base. Sepals often reddish purple, lanceolate, 2.5-3.3 mm long, 1.1-1.4 mm wide, glabrous. Petals pink to white, obovate, 3.0-5.5 mm long, 1.9- 3 mm wide, the notch 0.4-1 mm deep. Anthers 0.6-0.8 mm long, 0.5-0.7 mm wide; filaments of the longer stamens 1.2-3 mm long, those of the shorter 0.7-2 mm long, at least the longer, and usually the shorter, shedding directly on the stigma. Style 1.7-2.6 mm long; stigma subcapitate, 0.8-1.2 mm long, 0.5-1.1 mm thick. Capsules erect, glabrous, 1.8-2.3 cm long, 1.1-1.5 mm thick, on pedicels 2-5 mm long. Seeds pale brown, minutely papillose, obovoid, 1.2-1.3 mm long, 0.4-0.5 mm thick, the chalazal appendage 0.04-0.08 mm long, 0.08-0.12 mm wide; coma pale yellow brown, 3.5-5 mm long. Distribution (Fig. 3): Typically inhabiting moist crevices in rock outcrops or cliff faces; scattered throughout the Altiplano of Peru, from Dpto. Ancash south to western Bolivia and extreme northeastern Chile at elevations of 4,400-5,000 m. Flowering specimens have been collected from May, June, July and Novem- ber. 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 Specimens examined: Bolivia, la paz: Ravin de Chuquiaguillo, Weddell in 1851 (P). Chile, i region (tarapaca): Parinacota, Lago Cotacotani, 4,500 m, I8°I2'S, 69°I5'W, Sudzuki 480 (K). Peru, ancash: Huaraz, 4,600 m, Weberbauer 2969 (G). cuzco: Ausangate, 4,800 m, Rauh- Hirsch PI258 (RSA). huancavelica: San Jose de Acobambilla, Nahuincocha, 12°40'S, 75°22'W, Lloyd & Marshall 100 (K). junin: La Oroya, 35 km W of Hacienda Cochas, 5,000 m, Gutte 3256 (LZ); Casa Cancha, Cerro La Viuda, 4,400 m, 1 1°05'S, 76°35'W, mikes (U.S. Exploring Expedition) in 1839 (GH, K, US), lima: Prov. Huarochiri, Casapalca, 4,700 m, MacBride & Feather stone 867 (F, GH, S, US), puno: Caccachara, 80 km SW of Have, 4,900 m, Pearson 107 (PH); San Antonio de Esquilache, 4,500 m, 16°06'S, 70°18'W, Stafford 729 (BM, F), Tutin 1181 (BM). Epilobium fragile is a diminutive, high Andean species that has been collected rarely, so little is known of its biology or relationships. It forms low, densely caespitose clumps, with the thin, wiry stems seldom more than 10 cm long. Plants are characteristically glabrous, except for a few strigillose hairs on the leaf mar- gins, at the junction of the fused petiole bases, and rarely in scattered, descending lines, with or without a few scattered erect glandular hairs. Epilobium fragile has usually been confused with E. nivale, even by Samu- elsson, who described the former (Samuelsson, 1923). As a result, E. nivale has been erroneously reported from Peru (MacBride, 1941, and Samuelsson, 1923). Samuelsson considered E. fragile to be most closely related to E. nivale, both of which he segregated in his group Nivalia, based on their compact growth form, small leaves, and the more or less glabrous nature of the plants (Samuelsson, 1923). Superficially both species are quite similar in habit and overall morphology. However, E. fragile differs from E. nivale by the presence of strigillose and glandular hairs, thin, wiry stems, ovate to elliptic, blunt-tipped leaves, and slightly larger, often white flowers, although there is some overlap in all the characters except pubescence. Ecologically they are also separated, with E. fragile growing in crevices in rocks, and E. nivale inhabiting moist places near bogs or along streams. The present ranges of E. fragile and E. nivale are separated by about 1 ,500 km. Additional exploration of the Andes in northern Chile and northwestern Argentina, however, may expand the range of both species. The chromosomal arrangement of E. fragile is unknown, so the possibility that it may be more closely related to E. nivale, which has the BB arrangement cannot be ruled out. A second, and what is considered more likely, hypothesis is that E. fragile is most closely allied to E. denticulatum, a species with the AA chromosomal arrangement, which is widespread in the northern and central An- des. In addition to having strigillose and glandular hairs in common, some of the smaller specimens of E. dentieulatum approach E. fragile in habit, although they are larger, more densely pubescent, and usually have more and larger flowers. The geographic range of E. fragile is completely contained in that of E. den- ticulatum, and, considering the very recent nature of the high montane habitats where E. fragile is found, it may have been derived directly from E. dentieulatum. 4. Epilobium puberulum Hook. & Arn., Hook. Bot. Misc. 3:309. 1833. type: Chile, X Region (Los Lagos), Prov. Chiloe, Isla Chiloe, 1831, H. Cuming 36 (K, lectotype here designated, photograph MO; BM, BP, BR (probable), E 2 sheets (photographs A, K), GH, OXF (probable), PRC, TCD, W (photographs BH, GH, MO), isolectotypes). Hausskn., Monogr. Epilobium 257. 1884. Reiche, Fl. Chile 2:245. 1898. H. Lev., Iconogr. Epilobium tab. 192, 197 (as E. bar- 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 287 beyanum), 211 (as E. denticulatum). 1911. Samuelsson, Svensk Bot. Tidskr. 17:248. 1923. E. denticulatum Ruiz & Pavon, Fl. Peruv. et Chil. 3:78. 1802, pro parte, as to plants from Conception, Chile. E. denticulatum auct. non Ruiz & Pavon: Gay, Hist. Fis. Chile 2:247. 1846. E. pedunculatum R. Phil., Anal. Univ. Chile 41:713. 1872. type: Chile, VI Region (OHiggins), Cordillera de Colchagua, November 1860, C. Landheck (SGO-531 14, lectotype here designated, photograph MO; SGO-41459, isolectotype, photographs GH, MO). E. gracile R. Phil., Anal. Univ. Chile 84:748. 1893, non Bruegg., Jahres-B. Naturf. Ges. Graub. Ser. 2, 25:70. 1882. type: Chile, X Region (Los Lagos), Prov. Valdivia, Cordillera de Valdivia, Pi- rihuaico, January 1887, O. Philippi (SGO-53083, lectotype here designated, photograph MO: SGO-53046, isolectotype, photographs GH, MO). Perennial herb, 20-60 cm tall, overwintering and reproducing vegetatively by elongate leafy shoots produced at or near the base. Stems erect, usually branched above, and often below, or simple, terete, strigillose to spreading hirsute through- out, the hairs 0.15-0.4 mm long, with obscure raised descending lines from the decurrent petiole bases, these often poorly developed or absent. Leaves mostly opposite, alternate above and in the inflorescence, often with fascicles of small leaves in the axils, thin, bright green, lanceolate to rarely narrowly ovate, 0.8- 2.8(-3.3) cm long, 0.2-K-1.5) cm wide, acuminate to acute at the apex, regularly and remotely denticulate with 3-6 teeth on each side, cuneate to acute, occa- sionally obtuse or rounded at the base, strigillose to spreading hirsute on both surfaces, usually glabrate with age and then the pubescence restricted to the margin and the abaxial and adaxial midribs and lateral veins, the lateral veins obscure, 2-4 on each side of the midrib, on more or less well defined petioles, 0.5-2 mm long. Inflorescence erect, simple or branched, the leaves subtending the flowers usually reduced in size. Flowers erect. Ovaries often reddish purple, densely strigillose to spreading hirsute, (0.8-)1.2-2.5(-3) cm long, on pedicels 0.4- 1. 3(- 1.7) cm long. Floral tube 1-2 mm deep, 1.4-2.1 mm across, externally strigillose to spreading hirsute, internally with a ring of erect villous hairs, 0.2- 0.3 mm long, near the base. Sepals lanceolate, 2-3.1 mm long, 0.9-1.3 mm wide, strigillose to spreading hirsute. Petals clear salmon pink, broadly obovate, 3.5-5 mm long, 2.5-3.2 mm wide, the notch 0.8-1.5 mm deep. Anthers cream to white, 0.6-0.8 mm long, 0.4-0.6 mm wide; filaments cream to white, those of the longer stamens 0.9-1.3 mm long, those of the shorter 0.3-0.6 mm long; only the longer shedding directly on the stigma at anthesis. Style cream to white, 2. 1-2.8 mm long; stigma cream to white, clavate, 0.8-1.6 mm long, 0.5-0.8 mm thick. Cap- sules erect, strigillose or spreading hirsute, (2-)3.2-5(-6.6) cm long, 1-1.4 mm thick, on pedicels 0.5-2 cm long. Seeds brown, papillose, obovoid, 0.7-0.9 mm long, 0.3-0.4 mm thick, the chalazal end with a short appendage 0.04-0.08 mm long, 0.06-0.1 mm wide; coma white, 5-7 mm long. Gametic chromosome num- ber, n = 18. Distribution (Fig. 4): Frequent and somewhat weedy, along roadsides, gravelly or sandy stream beds, embankments, and other open, moist, usually disturbed sites. Widely distributed in central Chile from Valparaiso to Isla Chiloe, and Aisen (probably introduced at the latter). In Argentina, found only in a narrow band along the eastern slopes of the Andes in southern Neuquen, Rio Negro, and northeastern Chubut provinces. Typically encountered from near sea level to 800 m rar<*1v as hiph as 1.000 m. Flowerine October to March. 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol.. 69 <=D '<: 100 200 100 400 S00 hm • « -»(M i " ii ■ ► ■ Figures 4-5. Distribution of Epilobium species. — 4. E. puberulum.S. E. nivale. 1982] SOLOMON— EPILOBWM IN SOUTH AMERICA 289 Representative specimens examined: Argentina, chubut: Cholila, Crovetto 3084 (SI), neuquen: San Martin de Los Andes, Bridarolli 2087 (LP); Hua-Hum, Dawson & Schwabe 2352 (BAB mixed with E. ciliatum); Lago Lacar, Peninsula Pucara, Dawson & Schwabe 2724 (BAA, BAB, MO). Rio negro: Bariloche, Burkart 6026 (RSA), Maldonado in 1941 (F), Parodi in 1934 (BAA, MO, S); Cerro Otto, 1,000 m, 4I°09'S, 71°23'W, Hosseus 38 (BAF, CORD); Lago Nahuel Huapi, 770 m. Ljungner 158 (GB). Chile, v region (aconcagua): Valparaiso, Vina del Mar, Hacienda Las Siete Hermanas, Bultmann 23523 (CONC); Valparaiso, Claude-Joseph 3565 (US); Quebrada Verde near Valparaiso, Maximow in 1854 (JE). vi region (o'higgins): Cachapoat, Termas de Cauquenes, Gay 187 (F, P, RJ: mixed with E. ciliatum); Reid (K mixed with E. glaucum). Colchagua, without locality, Philippi (JE, SGO). vii region (maule): Curico, Vichuquen, 34°53'S, 72°00'W, Philippi 627a (SGO): 5 km above Los Queries, 760 m, 35°00'S, 70°45'W, Solomon 4335 (MO). Linares, Quinamavida, Philippi in 1893 (SGO); Linares, Quezada in 1975 (MO). Talca, Cordillera de Talca, El Picazo, Barros82 (GH): Constitucion, Reiche in 1890 (SGO). vm region (bio-bio): Arauco, Contulmo, Behn in 1919 (M); Lanalhue, 37°56'S, 73°I7'W, Claude-Joseph 5980 (US); Arauco, 50 m, Penned 12966 (PH); Llico, Philippi in 1861 (SGO); Boca del Rio Tubul y Raqui, Ricardi in 1949 (CONC); Isla Mocha, Laguna Hermosa, Weldt-Rodriquez 1166/461 (CONC). Bio-Bio, Camino de Bio-Bio a Santa Barbara, Estero Pirquinco, 400 m, 37°49'S, 7I°40'W, Marticorena et al. 1015 (CONC, MO); Cordillera de Antuco, La Cueva, Philippi in 1882 (G, SGO). Concepcion. Tome, Collen, Junge in 1934 (CONC); Gualqui, Kuntze in 1892 (NY); Florida, Merxmidler 24865 (M); Nonguen, near Concepcion, Moore 296 (LA); Coronel, Ochsenius in 1866 (BR, GOET); Concepcion, Dombey in 1782 (P), Elliot 148 (BM, E, NY), Ruiz & Pavdn in 1782 (MA). Nuhle, Entre Cabrero y Salto del Laja, Fundo Trilahue, 130 m, Mar- ticorena et al. 855 (CONC); Cuchacucha, Nee in 1793 (MA); Montahas de Chilian, Philippi (SGO); Pilmaiquen, 36°40'S, 71°51'W, Philippi (G); Bulnes, Philippi in 1878 (JE, SGO); 8 km E of San Rafael on road to San Ignacio, 36°35'S, 72°45'W, Solomon 4394 (MO), ix region (araucani'a): Cautin. Laguna Lorilon, Fundo Flor, Behn in 1946 (CONC); Villarrica, Burger in 1965 (GOET); Freire, Claude-Joseph 5882 (US); Pemehue, 39°29'S, 72°30'W, Germain in 1894 (SGO); Truf-Truf, 150 m. 38°44'S, 72°34'W, Gunckel 16908 (US); Peliin, 38°30'S, 72°20'W, Lanfurgo in 1885 (SGO); 2 km S of Metrenco, Moore 310 (LA, MO); Termas de Palguin, 700 m, 39°22'S, 7I°45'W, Solomon 4525 (MO). Malleco, Cordillera de Nahuelbuta, Los Alpes, Fundo Solano, Eyerdam 10296 (F, NY, SGO, UC, US); Curacautin, Maldonado 39 (L); Curacautin, Rio Blanco, Penned 12726 (F, GH, NY, PH, SGO); Pidima, Fundo Chequenco, 38°01'S, 72°26'W, Pfister in 1946 (CONC); 16 km N of Curacautin, 800 m, 38°20'S, 70°50'W, Solomon 4489 (MO), x region (los lagos): Chiloe, Puerto Carmen, Quellon. 43°08'S, 73°46'W, Marticorena 1766 (CONC); Camino de Caicumeo, Philippi in 1886 (SGO): 14 km S of Castro, Weldt-Rodriguez 734-29 (CONC). Llanquihue, Colegual, 41°22'S, 73°1 1'W, Klenner in 1952 (CONC); Hautrunes-Maullin, Pfister 289 (CONC); Lago Todos Los Santos, Petrohue, Stubbe in 1961 (VALD); 5 km N of Puerto Montt, Wall & Sparre in 1947 (S). Osorno, Cuinco, 40°32'S, 73°I0'W, Rudolph in 1933 (VALD); Chuyaca, Rudolph in 1944 (VALD); 3 km E of Puyehue, 40°40'S, 72°I8'W, Solomon 4596 (MO); Centinela, Sparre 4399 (S); Osorno, Wall & Sparre in 1947 (S). Palena, Rolecha, 4I°55'S, 72°50'W, Pfister in 1951 (CONC). Valdivia, Valdivia, Buchtien in 1901 (BAF, BM. BREM. E, M,P, S, US); Lechler44l (G,GH,GOET, K), Philippi 514 (BM. BREM, G, GOET, JE, P, UPS); Panguipulli. Claude-Joseph 2556 (US); Corral, Gunckel 15424 (GH); 5 km NE of La Union, Moore 297 (LA, MO); Hacienda San Juan, 40°I5'S, 73°05'W, Philippi in 1886 (JE, SGO); Cordillera de Ranco. Huahum, Philippi in 1887 (SGO); Cordillera Pelada, Philippi in 1889 (JE, SGO). xi region (aisen): Aisen, Aisen, Andreas 509 (U). Epilobium puberulum is a distinctive species of south central Chile. Its densely strigillose stems and ovaries with generally small lanceolate, few-toothed leaves, erect habit, and salmon-pink flowers make it easily recognized from all its congeners. In fact, the flower color, unknown in any other species of Epilobium, is sufficient to identify living material immediately. Superficially some plants of E. puberulum have a strong resemblance to some individuals of E. denticulatum. There are, however, a number of striking morphological differences; in addition, these two species are allopatric, with a gap of several hundred kilometers between them at their closest point. Epilobium puberulum is much more strict in habit and lacks the glandular hairs and nodding inflorescence of E. denticulatum, and the flower color is distinctive. The nearest relative of E. puberulum is probably E. denticulatum. Epilobium 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 puberulum may have the same chromosomal arrangement as E. denticulatum (A A), although this has not yet been confirmed. The first collections of E. puberulum were made by Ruiz, Pavon, and Dombey, from the vicinity of Conception, Chile, during their stay there in 1782 and 1783. Because of the similarity of these specimens to collections of E. denticulatum from Peru, they were all described under the name E. denticulatum. Not until some thirty years later were the Chilean plants recognized as a separate species. In subsequent years, R. A. Philippi published two additional taxa based on trivial variants. Variation within this species is primarily in the size and density of the leaves and the branching pattern. These have no geographic basis, but are mostly related to the age of the plant and the season in which it was collected. Early season plants tend to be simple with fairly large leaves (mostly up to 2.5 cm), with fascicles of small leaves in their axils. As the growing season progresses, these axillary shoots may elongate, producing densely branched plants. The leaves of the secondary shoots are often smaller than those of the primary branches pro- duced earlier. Pubescence also varies, not only in density but in type. For ex- ample, a collection from La Union, Valdivia Prov., Chile (Moore 297, MO) is finely strigillose throughout with mostly glabrate leaves, while one from Vegas Blancas, Malleco Prov., Chile (Solomon 4459, MO) is densely spreading hirsute with both surfaces of the leaves pubescent. Ecologically, E. puberulum is distinct from most other species. Although it does grow sympatrically with three species, its distribution is the inverse of the normal pattern. As was indicated earlier, except for E. hirtigerum and E. pu- berulum, all the South American species are montane, with a few descending to much lower altitudes at higher latitudes. In contrast, E. puberulum grows at lower elevations in the coast range, central valley, and the lower slopes of the Andes in Chile, seldom extending above 1,000 m. Its limited and scattered distribution in Argentina suggests that it is a relatively recent introduction to the eastern slopes of the Andes, perhaps appearing there only with the clearing of the forest land and increased communication across the Andes during the last century. The single collection from Aisen, Chile, is undoubtedly a recent introduction. The high degree of enforced autogamy mentioned earlier makes hybridization with other species an extremely rare event. With one exception no evidence was seen in any of the populations examined in the field or from herbarium material that would indicate that hybridization takes place, although it is not an absolute impos- sibility. There is one set of very unusual collections made by Bridges at Valparaiso, Chile during the 1830s (Bridges 179, BM, E, K, W), apparently all from the same plant or population, which can only be explained as being of hybrid origin, with Epilobium ciliatum as the other likely parent. While the upper inflorescence is very E. puberulum-Wk^, the lower leaves are several times longer and broader (4.5 x 2 cm), with many more teeth, than is known in any other collection of E. puberulum. The leaves approach a size and shape common for E. ciliatum, which also occurs in the same area. The flowers are borne singly in the axils of leaves throughout the length of the simple stems, with the antipetalous staminal filaments ca. 0.5 mm long, as is normal in E. puberulum. The average pollen stainability of these plants was 74% (range 67-88%). This unusual set of specimens was 1982] SOLOMON— EPILOBWM IN SOUTH AMERICA 291 identified by Haussknecht as belonging to E. caesium, the type of which came from northern Bolivia. 5. Med. Repos. II 5:361. 1808. subsp. ciliatum. type: 7. S. Rafinesque. No specimens of this species col- Rafinesque South American material is given; for a complete discussion of North Amer- ican names see Hoch, 1978.) E. pedicellare auct. non Presl: Hook. & Arn., Bot. Misc. 3:309. 1833, pro parte. E. pedicellare fi latifolium Walp., Nov. Act. Acad. Caes. Leopold. 19, Suppl. 1:328. 1843. type: Chile, VI Region (OHiggins), Cordillera de San Fernando, January-March 1831, F. Meyen (B, holotype, destroyed). No other authentic material has been seen. Haussknecht places E. pedi- cellare j8 latifolium as a synonym of E. chilense and cites a collection of Meyen from the Cor- dillera de San Fernando at Berlin (B) (Haussknecht, 1884, p. 272). Based on this placement, the brief description and locality, this variety probably belongs here. E. tetragonum auct. non L.: Hook, f., Fl. Antarct. 2:270. 1847, pro parte; Fl. N. Z. 1:60. 1853, pro parte, as to the plants from the Falkland Islands. E. chilense Hausskn., Oesterr. Bot. Z. 29:118. 1879. type: Chile, without specific locality or date (1826-31), H. Cuming (W, lectotype here designated, photograph MO; BM, BR, M, OXF, PR 2 sheets, isolectotypes). Hausskn., Monogr. Epilobium 272. 1884. Reiche, Fl. Chile 2:249. 1898. H. Lev., Iconogr. Epilobium, tab. 168. 1911. Samuelsson, Svensk Bot. Tidskr. 17:269. 1923. E. cunninghamii Hausskn., Oesterr. Bot. Z. 29:118. 1879. type: Chile, XII Region (Magallanes), Prov. Magallanes, Punta Arenas (Sandy Point), 28 February 1863, R. Cunningham (LE, holotype, not seen; K, JE (probable), isotypes). Samuelsson, Svensk Bot. Tidskr. 17:276. 1923. E. valdiviense Hausskn., Oesterr. Bot. Z. 29: 1 18. 1879. E. chilense var. valdiviense (Hausskn.) Hoss., Trab. Inst. Bot. Farm., Buenos Aires 33:57. 1915. type: Chile, X Region (Los Lagos), Valdivia, R. A. Philippi (W, holotype, photograph MO). Hausskn., Monogr. Epilobium 271. 1884. Reiche, Fl. Chile 2:249. 1893. H. Lev., Iconogr. Epilobium, tab. 166. 1911. Auct. non Hausskn.: Skotts- berg, Kungl. Sv. Vet. Akademiens Handlinger 50(3):42. 1913. Samuelsson, Svensk Bot. Tidskr. 17:272. 1923. E. bonplandianum auct. non H.B.K.: Hausskn., Monogr. Epilobium 267. 1889, pro parte as to the plants from Mexico and Chile. Reiche, Fl. Chile 2:246. 1898. E. magellanicum R. Phil. & Hausskn. ex Hausskn., Monogr. Epilobium 271. 1884. type: Chile, XII Region (Magallanes), Prov. Magallanes, without precise locality, summer 1864-65(7), R. A. Phi- lippi (W, lectotype here designated, photograph MO; W, JE, SGO-41443, SGO-53016, W (last 4 probable), isolectotypes). The date is taken from SGO-53016. Reiche, Fl. Chile 2:248. 1898. H. Lev., Iconogr. Epilobium, tab. 171. 1911. Samuelsson, Svensk Bot. Tidskr. 17:278. 1923; Svensk Bot. Tidskr. 24:9. 1930. E. aconcaguinum R. Phil., Anal. Univ. Chile 84:745. 1893. type: Chile, V Region (Aconcagua), Prov. Los Andes, Los Andes, on the banks of the Rio Aconcagua, December 1885, R. A. Philippi (SGO-53036, lectotype here designated, photographs GH, MO; SGO-53010, isolectotype). E. albiflorum R. Phil., Anal. Univ. Chile 84:745. 1893. type: Chile, X Region (Los Lagos), Prov. Valdivia, Hacienda San Juan, 40°15'S, 73°05'W, January 1886, R. A. Philippi (SGO-53044, lec- totype here designated, photograph MO). E. glandulosum auct. non Lehm. Reiche, Fl. Chile 2:250. 1898. E. valdiviense var. alboffii Macloskie, Rep. Princeton Univ. Exped. Patagonia 8:611. 1905. type: Argentina, Terr. Tierra del Fuego, Ushuaia, N. Alboff. No authentic material has been seen. This name was originally published without an indication of rank. The varietal level has been chosen because this was the generally used concept at the time of publication. E. santa-cruzense Dusen, Kungl. Sv. Vet-Akadis. Arkiv. f. Botanik 7(2):208, tab. 3,f. 2-4, tab. 7, f t 29-34. 1907. type: Argentina, Prov. Santa Cruz, upper valley of the Rio Santa Cruz, 11 February 1905, P. Dusen 5842 (S, lectotype here designated, photograph MO; K, SI, UPS, isolectotypes). Samuelsson, Svensk Bot. Tidskr. 17:278. 1923. E. hookerianum Hausskn. ex Skottsberg, Kungl. Sv. Vet. Akademiens Handlinger 56(5):271. 1916. type: Falkland Islands, East Falkland, between Port Darwin and North Arm, 27 January 1908, C. Skottsberg 122 (UPS, lectotype here designated, photograph MO; BA, GB, LD, S, SGO, isolectotypes). E. chilense var. latifolium Samuelsson, Svensk Bot. Tidskr. 17:270. 1923. type: Chile, V Region 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol . 69 (Aconcagua), Valparaiso, T. Bridges (K, lectotype here designated, photograph MO; G, isolec- totype). Based on E. chilense f. "latifolia" Hausskn., Monogr. Epilobium 272. 1884. E. chilense var. maerum Samuelsson, Svensk Bot. Tidskr. 17:271. 1923. type: Chile, V Region (Aconcagua), Prov. Quillota, Campana de Quillota, 1856-57, Ph. Germain (W, lectotype here designated, photograph MO; K, SGO-53057, SGO-53098, isolectotypes). Based on E. chilense f. "macro" Hausskn., Monogr. Epilobium 272. 1884. E. longipes Samuelsson, Svensk Bot. Tidskr. 17:271, tab. 4, f. 2. 1923. type: Argentina, Prov. Chubut, Lago Buenos Aires, mouth of Rio Fenix, 11 December 1890. C. Skottsberg 946 (UPS, lectotype here designated, photographs MO, S). E. argentinum Samuelsson, Svensk Bot. Tidskr. 17:273, tab. J,/. 5. 1923. type: Argentina, Prov. Mendoza, Rio Salado Superior, Cajon de Las Aguas Amarillas, 22 January 1893, F. Kurtz 7621 (JE, lectotype here designated, photographs MO, S). E. constrictum Samuelsson, Svensk Bot. Tidskr. 17:275, tab. 4, f. 4. 1923. type: Argentina, Prov. Santa Cruz, Lago Argentino, Valle del Rio Santa Cruz, 19 January 1905, P. Dusen 5614 (S, lectotype here designated, photographs MO, S; BAF, SI, UPS, isolectotypes). E. caesiovirens Samuelsson, Svensk Bot. Tidskr. 17:279, tab. 4, f. I. 1923. type: Argentina, Prov. Mendoza, Las Cuevas, Puente del Inca, 2,700-2,800 m, 13 February 1903, G. O. Malme 2889 (S, lectotype here designated, photograph MO; B (destroyed, photographs BH, MO, US), S, UPS, W, isolectotypes). E. leiophyton Samuelsson, Svensk Bot. Tidskr. 17:280, tab. 3,f. 1. 1923. type: Chile, VII Region (Maule), Cordillera de Curico, 2,500 m, January 1897, C. Reiche (UPS, lectotype here designated, photograph MO; B (destroyed, photographs BH, GH, MO, NY, US), isolectotype). Variable perennial herbs (5-)20-60(-90) cm tall, overwintering and reproduc- ing vegetatively by fleshy-leaved turions or compact leafy rosettes produced at or below ground level, the leaves of the perennating structures subentire, obovate to depressed obovate, or ovate to depressed ovate, 0.5-1.5 cm long, 0.5-2 cm wide. Stems erect, occasionally ascendent, often well branched from the base or above, or sometimes simple, terete or quadrangular, reddish brown or purplish above, strigillose, the hairs 0.1-0.2 mm long, throughout in the inflorescence, glabrate below, then strigillose in raised descending lines from the decurrent petiole bases, with an admixture of erect to slightly appressed glandular hairs, 0. 1-0.2 mm long, mostly in the inflorescence, occasionally throughout, rarely glabrous. Leaves opposite, alternate above, thin, green, lanceolate to narrowly ovate (0.6-)l.3-6.5(-7.5) cm long, (0.3-)0.6-2.6(-3. 1) cm wide, acuminate to acute at the apex, remotely, irregularly serrulate with (8-) 10-22 teeth on each side, obtuse to rounded, subcordate or occasionally cordate at the base, glabrous or with a few scattered strigillose hairs on the margins, the lateral veins 4-5(-7) on each side of the midrib, on petioles 0-3 mm long. Inflorescence erect, often branched, rarely somewhat nodding, the leaves subtending the flowers much reduced, acuminate, and usually narrower. Flowers erect. Ovaries often reddish purple, strigillose, usually with an admixture of erect to appressed glandular hairs, (0. 8-) 1-2.2 cm long, on pedicels 2-5 mm long. Floral tube often reddish, 0.9-1.5 mm deep, (0. 8-) 1. 2-2. 2 mm across, externally strigillose, usually with an admix- ture of erect to appressed glandular hairs, internally usually with a ring of erect villous hairs, 0. 1-0.2 mm long near the base, often with only a few hairs present and occasionally glabrous. Sepals often reddish, lanceolate, 1.8-2.8 mm long, 0.8-1.2 mm wide, strigillose, usually with an admixture of erect to appressed glandular hairs. Petals white, often flushed with pink after anthesis, or pink, obovate, 3.0-4.5 mm long, 1.5-2.5 mm wide, the notch 0.7-1.5 mm deep. Anthers cream to white, 0.5-0.7 mm long, 0.4-0.5 mm wide; filaments cream to white, those of the longer stamens 1.5-2.2 mm long, those of the shorter 0.8-1.5 mm 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 293 long; the longer, and usually the shorter, stamens shedding pollen directly on the stigma before or at anthesis. Style cream to white, 1—1 -8(— 2.3) mm long; stigma clavate to subcapitate (0. 8-) 1.5-2. 5 mm long, 0.4-0.8 mm thick. Capsules erect, with scattered strigillose and glandular hairs, 3-6.2 cm long, 1-1.6 mm thick, on pedicels 0.4-1. 5(-2. 2) cm long. Seeds gray brown, narrowly obovoid, 0.9-1.4 mm long, 0.3-0.5 mm thick, with conspicuous longitudinal rows of flattened, fused papillae, at least along the sides, the micropylar end acuminate, the chalazal end with a short appendage, 0.04-0.08 mm long, 0.08-0.2 mm wide; coma white to slightly yellowish, 5-8 mm long, readily detaching. Gametic chromosome num- ber, n = 18. Distribution (Fig. 6): Common and weedy in open, often disturbed areas, usually permanently moist, or at least moist for part of the growing season, such as roadsides, sandy and gravelly stream beds, embankments, boggy areas, and banks of lakes and ponds. In South America, throughout the Andes of Chile and Argentina from central IV Region (Coquimbo), Chile, and San Juan Prov., Ar- gentina, southward to Tierra del Fuego, in the coast range between Santiago and Valparaiso, generally throughout the central valley and coastal hills south of Concepcion, scattered eastward on the Patagonian plain in southern Chubut and Santa Cruz Prov., Argentina, and in the Sierra de San Luis and Sierra Grande de Cordoba; also in the Falkland Islands. In North America from Alaska to British Columbia, eastward throughout much of Canada to Labrador, southward through New England to Pennsylvania and parts of the Appalachians, south westward from M Mexico Mexico Mexico eastern China, and eastern Siberia. Introduced in Hawaii, Europe and European U.S.S.R., New Zealand, and parts of Australia and Tasmania (Raven & Raven, 1976). In the northern part of its South American range found from 1,200-2,900 m, gradually descending to sea level on the western side of the Andes near Concepcion. In the southern part of the range mostly between 200 and 1,900 m, and in Tierra del Fuego from sea level to 350 m. Flowering primarily from De- cember to March throughout, but as early as September in the northernmost parts of the coastal mountains of Chile. Representative specimens examined: Argentina, chubut: Colonia Sarmiento. Biraben 555 (LP); Dpto. Cushamen, 30-40 km NE of Esquel, 42°49'S, 71°05'W, Boelcke et al. 16036 (BAA, MO); Dpto. Futuleufii, Esquel, La Hoya, Cabrera et al. 25987 (LP); Lago Fontana, Castellanos in 1932 (BA, MO, RSA); Lago Futalaufquen, Castellanos in 1945 (F, LIL); Cholila, Crovetto 3009 (SI); Golon- drinas,42°01'S, 7I°31'W, Crovetto 3 197 ( BAB, LIL); Vallede Las Plumas, Lago General Paz, Gerling 43 (BAF); 72 km S of Tecka, crossing Rio Cherque, 800 m, Hjerting 629 (C); Valle del Lago Blanco, 45°54'S, 71°15'W, Koslowsky 172 (BA); Trevelin, Estancia Rio Frio, Krapovickas 4000 (BAB); Lago Puelo, desembocadura del Epuyen, Perez-Moreau in 1940 (BA, LIL, RSA); Parque Nacional Los Alerces, Lago Cisnes, Perez-Moreau in 1944 (BA); Lago Epuyen, Soriano 1374 (BAA, SI); Rio Senguer. Estancia Laurita, 44°43'S, 70°04-24'W, Soriano 1480 (BAA, SI); Dpto. Tehuelches, Arroyo Temenhuao, 44°13'S, 71°16'W, Soriano 2651 (BAB), cordoba: Dpto. San Alberto, Pampa de Achala, 2,000 m, Cabrera et al. 16620 (LP); Sierra Grande, cerca del Rio Yuspe detras del Cerro Blanco, L650 m, Hunziker 10253 (CORD, MO, RSA mixed with E. hirtigerum); Sierra Achala, Cerro de Los Gigantes, Kurtz 3879 (CORD, RSA); Sierra Grande, Cuesta de Copina, 1,610 m, 31°50'S, 64°10'W, Solomon 4149 (MO); Sierra Grande, La Cumbrecita, 1,450 m, 31°55'S, 64°I5'W, Solomon 4199 (MO). mendoza: Dpto. Las Heras, puente del Inca, Laguna de Los Horcones, Boelcke et al. 9823 (BAA, 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 57°W 40"S 50 100 200 300 400 500 km Figure 6. Distribution of Epilobium ciliatum ssp. ciliatum in South America 1982] SOLOMON— EPILOB1UM IN SOUTH AMERICA 295 BAB, MO). Dpto. Malargiie, Arroyo Las Mangas, 2,000 m, Curette 284 (LP, SI); Dpto. Malargiie, Arroyo del Cordon del Cura, Castellanos in 1941 (BA, RSA); Dpto. San Carlos, Refugio Gral. Al- varado, 2,000 m, Cuezzo & Barkley 20MZ45I (NY); Arroyo Chacayco, above Laguna Carilauquen, 36°32'S, 70°H'W, Kurtz 6086 (JE); Rio Salado Superior, Los Molles, Kurtz 7550 (JE); Dpto. San Rafael, Rio Atuel, entre Puesto de Ubilla y Paso del Pico Plateado, Kurtz 7592 (CORD, JE, LP); Dpto. San Carlos, Arroyo de Yaucha, 33°46'S, 69°02'W, Kurtz 111 53 (CORD); Dpto. Tupungato, Finca Los Helechos, 33°20'S, 69°H'W, Leal 1259 (LIL); Dpto. Tunuyan, Rincon Colorado, 2,800 m, Leal 1289 (RSA); Dpto. Tupungato, Estancia La Carrera, 2,350 m, Metis & Paci in 1949 (BR, G, W); Dpto. Las Heras, Uspallata, 2,000 m, Semper 540 (LIL, NY); Valle del Atuel, Cajon del Burro, 2,400 m, Wilczek 412 (G). neuquen: Lago Nahuel Huapi, Isla Victoria, 770 m, Boelcke 1795 (BAA, SI); Dpto. Minas Las Ovejas, 37°01'S, 70°45'W, Boelcke et al. 10752 (BAA, BAB, MO); Dpto. Minas, Laguna Epulauquen, 1,300 m, 36°50'S, 71°04'W, Boelcke 10856a (BAA, BAB, MO); Dpto. Chos Malal, Vegas del Pelan, 1,700 m, 36°54'S, 70°20'W, Boelcke et al. 11140 (BAA, BAB, MO); Dpto. Minas, Varvarco Campos, Arroyo Benitez, 36°17'S, 70°39'W, Boelcke et al. I4224V2 (BAA, MO); confluencia de los Rios Neuquen y Varvarco, Invernada Vieja, Boelcke et al. 14439 (BAA, BAB); Arroyo Chacayco, 37°05'S, 69°45' W, Chicchi 91 (LP); Rio Limay, Chicchi 193 (LP); Dpto. Huiliches, Volcan Lanin, Arroyo Rucu-leufu, Correa et al. 5596 (BAB mixed with E. australe); Parque Nacional Lanin, Bahos de Epulafquen, Correa et al. 5843 (BAB); Pampa del Malleo, 39°48'S, 70°58'W, Crovetto AN-128 (CTES); Rahue, Estancia Ochoa, 39°23'S, 70°49'W, Dawson & Schwabe 2129 (BAA, BAB); Hua-Hum, Dawson & Schwabe 2352 (BAA, BAB; mixed with E. puberulum); 10 km from Junin de Los Andes towards Zapala, 900 m, Hjerting 6306 (C); Neuquen, Jergensen 636 (BAB); Lago Lolog, Kalela 1756 (H); Dpto. Norqufn, Cajon de Trolope, 37°46'S, 71°04'W, Kurtz 6184 (CORD, JE); Parque Nacional Lanin, San Martin de Los Andes, Cerro Chapelco, 1,700 m, Leon & Calderon 978 (BAA); Cerro de Copahue, Fuente Zulena, O'Donell 2072 (LIL); Dpto. Los Lagos, Villa La Angostura, Pedersen 1547 (BR, C, MO, P, US); Lago Nahuel Huapi, Peninsula Quetrihue, Perez-Moreau in 1940 (BA, RSA); Dpto. Catan-Lil, Estancia Bernal, Arroyo del Canadon, 39°34'S, 70°36'W, Perez-Moreau 3077 (BAB); Dpto. Chos-Malal, Canchahuinganco, 37°01'S, 70°23'W, Ragonese 166 (BA, RSA); Dpto. Loncopue, Cajon Chenque Pehuen, 38°06'S, 70°55'W, Rugolo & Agrasar 130 (BAA); Dpto. Catan- Lil, Cerro Chachil, 39°05'S, 70°38'W, Rugolo & Agrasar 372 (BAA, MO); Lago Nahuel Huapi, Brazo Huemul, Arroyo Huelta, 40°58'S, 71°22'W, Solomon 4623 (MO); Camino El Huecu a Loncopue, 1,000 m, Spegazzini 175 (BAB); Dpto. Picunches, Pino Hachado, Arroyo Haichol, Valla et al. 3009 (BAA, CTES, MO); Lago Quillen, Valla et al. 3204 (BAA, CTES, MO). Rio negro: Parque Nacional Nahuel Huapi, Laguna Frias, 41°05'S, 71°50'W, Boelcke & Correa 5379 (BAA, BAB, SI); Bariloche, 770 m, Buchtien in 1905 (BAF, BP, BREM, E, F, GB, GH, JE, L, M, PR, S, SI, US, W, Z), Cabrera 5942 (F, GH, LP, NY, POM, US); Parque Nacional Nahuel Huapi, Lago Fonck, Perez-Moreau in 1942 (BA, RSA); Cascada del Mallin Ahogado, El Bolson, Scolnik 318 (LIL); Cerro V. Lopez, W of Bariloche, 1,000 m, 41°03'S, 71°35'W, Solomon 4611 (MO); Villa Cerro Catedral, 1,030 m, 41°10'S, 7l°28'W, Solomon 4613 (MO); Rio Llogdconto, 41°22'S, 71°30'W, Solomon 4630 (MO); Cerro Tron- ador, 1,200 m, Solomon 4637 (MO); 3.9 km S of Campamento La Veranada, 1,000 m, 41°28'S, 7P28'W, Solomon 4642 (MO); Dpto. Pilcaniyeu, Pilcaniyeu, Estancia Raylluso, Vallerini 424 (BAA). san juan: Dpto. Iglesia, El Rodeo, 30°12'S, 69°06'W, Cuezzo 1848 (LIL); Dpto. Calingasta, Cordillera del Espinazito, Los Manantiales, 2,770 m, 32°14'S, 69°58'W, Kurtz 9610 (BAF, CORD); Dpto. Cal- ingasta, Las Lumbreras, 2,000 m, Spegazzini 655 (BAB), san luis: Sierra de San Luis, Cerro Canutal. 32°46'S, 66°05'W, Castellanos in 1929 (BA); Cerro Sololosta, 32°51'S, 66°00'W, Pastore 84 (SI). santa cruz: Chonque-Aike, 49°16'S, 69°45'W, Ameghino in 1898 (LP); Dpto. Lago Argentine Es- tancia Maria Elisa, 50°20'S, 71°60'W, Ancibor & Vizinis 4626a (BAA, MO); Dpto. Rio Chico, Lago Pueyrredon, Rio Oro, 500 m, Boelcke et al. 12897 (BAA, BAB); Dpto. Lago Buenos Aires, 29 km NW of Perito Moreno, 46°25'S, 71°09'W, Boelcke et al. 16097 (BAB, MO); Camino a Paso Roballos, 47°1 1 'S, 71°36'W, Boelcke et al. 16176 (BAA, MO); Dpto. Lago Argentino, Lago San Martin, Estancia Cancha Rayada, 48°54'S, 72°26'W, Boelcke et al. 16293 (BAB, MO); Tehuelches, 250 m, 46°50'S, 67°27'W, Donat 283 (BM, CAS, F, G, GB, GH, K, MO, NY, S, SI, U, UC, Z); Cerro Buitreras, 51°43'S, 70°09'W, Hauthal in 1899 (CORD); Between Cerro del Fraile and Cordon de Los Cristal 50°32'S, 72°39'W, James 471 (BM, DS, SI); Cerro Buenos Aires, 350 m, 50°24'S, 72°59'W, James 3004 (BM); Rio Coyle, Hotel Las Horquetas, 51°24'S, 70°17'W, Rahn 4396 (C); Rio Los Antiguos, 300 m, 46°32'S, 71°38'W, Roivainen 2526 (H). El Calafate, 190 m, 50°20'S, 72°15'W, Solomon 4653 (MO); Glaciar Perito Moreno, 190 m, 50°28'S, 73°02'W, Solomon 4658 (MO); Lago Volcan, Estancia El Rincon, 47°46'S, 72°15'W, Spegazzini 324 (BAB); Dpto. Guer Aike, Estancia Stag River, 51°34'S, 7 1°57' W, T.B.P.A . (Boelcke et al.) 3280 (BAB, MO, RNG); Pto. Tres Marias, 700 m, 51°28'S, 72°05' W, T.B.P.A. (Ambrosetti <£ Mendez) 3680 (MO); Lago Argentino, Brazo Onelli, 300 m, Vervoorst 4560 (LIL, S, W). tierra del fuego: Rio Grande, Expedition Fac. de Ciencias E.F.y N. 140 (BA); Estancia Harberton, 30 m, Goodall 71 (LP, RNG); Rio Lashifashaj, Brown's sawmill on Ruta O, 40 m, Goodall 585 (E, LP, MICH, NA, RNG, RSA, UC, US); Estancia Moat, Goodall 641 (MICH, NA, RNG, SI, UC, US, WIS; mixed with E. australe); Estancia Viamonte, Estero Manantiales, 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. W Goodall 906 (NA, RNG); Estancia Cullen, Rio Cullen, Goodall 3180 (LTR. NA); Estancia Cullen. Arroyo Beta, Goodall 3224 (MICH, NA, RNG, UC); Lago Fagnano, Hunziker 6711 (BAB, UC). Bahia Lapataia, Perez-Moreau in 1948 (BA, MO, RSA); 1 km NW of Ushuaia, 50 m, Solomon 4712 (MO); Ushuaia, Arroyo Buena Esperanza, 150 m, Solomon 4741 (MO). Chile, iv region (coquimbo): Choapa, Cuncumen, 1,050 m, 31°55'S, 70°35'W, Solomon 4273 (MO). Limari, Cordillera de Hurtado, 30°30'S, 70°30'W, Gov 523 (P); Rio Gordito, 3,000 m, 31°02'S. 70°20'W, Jiles 2548 (CONC). v region (Aconcagua): Los Andes, 10 km W of Rio Blanco, 32°55'S, 70°I9'W, Hutchison 131 (UC, US); Los Andes, Philippi in 1882 (SGO). Petorca, Cerro Chache. IX km E of La Ligua, 1,900 m, Morrison 17050 (G, GH, K, MICH, MO, NA, UC, WTU). Quillota, Limache, Cajon Grande, Boelcke 432 (BAA, SI). San Felipe, San Felipe, Claude-Joseph 2488 (US). Valparaiso, Camino a Laguna Verde, 33°06'S, 71°41'W, Behn in 1930 (CONC); Concon, Garaventa 500 (BH, CONC); Cerro del Colliguay, 33°12'S, 7I°07'W, Jaffuel 656 (GH); Valle de Marga-Marga, Jaffuel & Pirian 3094 (GH). region metropolitana (Santiago): Cordillera, Rio Yeso, Laguna Los Piuquenes, 2,500 m, Biese 808 (LIL, NY); El Volcan, 1,300 m, 33°50'S, 70°12'W, Burkart 9346 (SI); San Jose de Maipo, 1,050 m, 33°40'S, 70°20'W, Solomon 4313 (MO). Melipilla, Las Vizcachas, 10 km from La Dormida, 1,860 m, Morrison 16779 (G, GH, MO, NA, S, SI, UC). Santiago, 5 km above Farellones, La Parva, 2,700 m, Moore 397 (LA, MO); Maitenes, Perez Caldera, 1,800 m, 33°M'S, 70°29'W, Skottsberg & Sparre 11082 (CONC, S); Junction of road to La Disputada with road to Farellones, 1,400 m, 33°20'S, 70°20'W, Solomon 4291 (MO), vi region (o'higgins): Cachapoal, Rio Coya above El Teniente, 2,900 m, Pennell 12307 (GH, PH, SGO); Banos de Cauquenes, Perez- Moreau in 1948 (BA, MO, RSA). Colchagua, San Fernando, Bertero 237 (P); Termas Vegas Del Flaco, 1,750 m, Mahu 9747 (H, MO, UC); Rincon de Tinguiririca, Ricardi in 1950 (CONC, LIL); 19.3 km above La Rufina, 1,000 m, 34°55'S, 70°30'W, Solomon 4330 (MO), vn region (maule): Curico, La Montana, 34°54'S, 70°54'W, Mahu 10756 (MO); Cordillera del Planchon, Nee in 1793 (MA, mixed with E. glaucum); 6.2 km above Los Ouefies, 780 m, 35°00'S, 70°45'W, Solomon 4332 (MO). Linares, Panimavida, 35°45'S, 7I°25'W, Philippi in 1883 (BM). Quinamavida, prope Parral, Philippi in 1893 (JE, SGO); Cajon Troncoso, 36°20'S, 70°45'W, Schlegel 3695 (CONC). Talca, Cor- dillera de Talca, El Picazo, Barms 84 (GH); 52.4 km above El Colorado, 900 m, 35°50'S, 70°55'W, Solomon 4338 (MO), vm region (bio-bio): Arauco, Laraquete, 37°I0'S. 73°l l'W, Hosseus 85 (CORD); Contulmo, Lago Lanalhue, Ricardi in 1949 (CONC, LP; mixed with E. puherulum). Bio-Bio, Entre Cabrero y Salto del Laja, 37°07'S, 72°23'W, Marticorena et al. 852 (CONC, MO); Dpto. La Laja, Cunibal Oriente, 150 m, 37°3I'S, 72°15'W, Marticorena et al. 874 (CONC, MO); Camino de Santa Barbara a Rio Huequecura, Puente Piulo, 290 m, 37°42'S, 72°00'W, Marticorena et al. 891 (CONC); Estero Epiin, 1,260 m, 37°52'S, 71°28'W, Marticorena et al. 970 (CONC, MO); On road to Laguna La Laja, 1,400 m, 37°25'S, 7I°20'W, Solomon 4434 (MO); Antuco, 530 m, 37°20'S. 7I°40'W, Solomon 4445 (MO). Concepcion, Tome, Junge in 1935 (CONC); Concepcion, Junge 3016 (US); 5 km SW of Concepcion, 200 m, Skog 1030 (CONN); 9.7 km W of San Rafael, 450 m, 36°35'S, 72°45'W, Solomon 4403 (MO). Nuhle, Termas de Chilian, Garganta del Diablo, Gleisner 162 (CONC, MO); Termas de Chilian, 1,720 m, Moore 412 (LA, MO), 2,200 m, Werdermann 1314 (E, K, NY, S, UC, US, Z; mixed with E. australe); 8.2 km E of San Rafael, 300 m, 36°35'S, 72°45'W, Solomon 4395 (MO), ix region (araucania): Cautin, Desembocadero del Rio Tolten, 39°15'S, 73°I4'W, Friedrich in 1934 (CONC); 4 km SW of Termas de Palgufn, 780 m, Moore 302 (LA, MO); Temuco, banks of Rio Cautin, Roivainen 2994 (H); Puente Correntoso, 15 km E of Villarrica, 210 m, 39°16'S, 72°00'W, Solomon 4536 (MO); Pitrufquen, 100 m, 38°59'S, 72°39'W, Sparre 3454 (S). Malleco, Cordillera de Nahuelbuta, Los Alpes, Fundo Solano, 590 m, Eyerdam 10296 (NY); Parque Nacional de Nahuelbuta, Pino de Canales, Mahu 5860 (MO); Arroyo Lumaco, Capitan Pastene, 100 m, 38°U'S, 73°00'W, Marticorena et al. 1224 (CONC); Entre Pino Hachado y Liucura, 1,080 m, Marticorena et al. 1434 (CONC); Dpto. Victoria, Termas de Tolguaca, 1,180 m, Morrison & Wagenknecht 17500 (G, GH, MO, S, SI, UC); Curacautin, 700 m, Pennell 12715 (F, GH, NY, PH, SGO); Buenavista below Volcan Tolguaca, 1,100 m, Pennell 12815 (F, GH, NY, PH, SGO); Pidima, Fundo Chequenco, Pfister in 1946 (CONC); Laguna San Pedro, Lonquimay, Pinto in 1953 (CONC); 2.8 km below Vegas Blancas, 37°45'S, 72°58'W, Solomon 4472 (MO); 30 km W of Curacautin, 500 m, 38°20'S, 72°05'W, Solomon 4472 (MO); 30 km W of Curacautin, 500 m, 38°20'S, 72°05'W, Solomon 4473 (MO); Termas del Rio Blanco, 1,300 m, 38°35'S, 7I°35'W, Solomon 4496 (MO); Lago Icalma, 1,000 m, 38°48'S, 7I°16'W, Zollner 7868 (MO), x region (los lagos): Chiloe, Castro, Gay 63 (P); Piruquina, Junge 291 (CONC). Llanquihue, Paso Perez Rosales, 980 m, Moore 325 (LA); Camino Ensenada a Raliin, Pfister in 1946 (CONC); Calbuco, Philippi in 1893 (JE, SGO); Rio Maullin, Wall in 1947 (GB). Osorno, Salto Pilmaiquen, Rudolph in 1931 (VALD); Chuyaca, 40°34'S, 73°07'W, Rudolph in 1933 (VALD); Nadi Caipulli, Rudolph in 1951 (VALD); 7.7 km E of Puyehue, 200 m. 40°40'S, 72°18'W, Solomon 4592 (MO); Lago Constancia. 1.000 m, Sparre & Smith 353 (CONC. mixed with E. australe). Palena, Rio Palena, Delfin in 1887 (SGO). Valdivia, Cordillera de Ranco, without collector (844) in 1852 (GOET, JE); Corral, 30 m, Gunckel 3195 (BH); Lago Puyehue, Isla Fresia. Mahu in 1956 (MO); Camino de Conaripe a Puerto Fui, Km 6, Marticorena et al. 473 (CONC); San Juan. 40°I5'S, 73°05'W, Philippi in 1887 (SGO); 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 297 Valdivia, Philippi in 1888 (CORD, K, US), xi region (aisen): General Carrera, Valle Leon, Lago Buenos Aires, 550 m, von Rentzell 6242 (GH, SGO, SI); Ventisquero Soler, 500 m, 46°52'S, 73°08'W, Seki 547 (CONC). Coihaique, Lago Seco, 45°35'S, 72°02'W, Schlegel 2345 (CONC). xn region (magallanes): Antdrtiea Chilena, Isla Navarino, 3 km W of Puerto Williams, 100 m, Moore 353 (LA, MO); Isla Navarino, Puerto Douglas, 55°10'S, 68°08'W, Vervoorst 385 (LIL). Magallanes, Morro Chico, 52°02'S, 7I°26'W, without collector or date (WU); Seno Otway, Rio El Canelo, Biese 1249 (LIL); Rio Tres Brazos, 10 m, 53°I6'S, 71°02'W, Cekalovic 90 (CONC); Bahia Santiago, Cun- ningham in 1867 (S); Neighborhood of Monte Dinero, 52°I9'S, 68°36'W, Cunningham in 1868 (NY): 80 km NE of Punta Arenas, 25 m, Eyerdam et al. 23935 (G, GH, NA, UC); Mina Loreto, near Punta Arenas, 150 m, Mexia 7979 (BM, GB, GH, K, MO, NY, S, UC); Isla Riesco, Estancia Rio de Los Palos, 53°26'S, 73°30'W, Pfister & Ricardi in 1952 (CONC); Fiordo Silva Palma, Angostura Titus, Rio Raul, Pisano 3752 (HIP, MO, RNG); Seno Skyring, Estancia Skyring, Pisano 4383 (HIP, MO); 8 km W of Aguas Frescas, 100 m, 53°20'S, 71°05'W, Solomon 4685 (MO); Laguna Parrillar, 350 m, 53°20'S, 7I°08'W, Solomon 4686 (MO); 2 km W of Puerto Hambre, 20 m, 53°40'S, 70°55'W, Solomon 4689 (MO). Tierra del Fuego, Bahia San Felipe, Cunningham in 1867 (K); Estancia Vicuna, Cerro Bahamondes, 300 m, Goodall 1756 (MICH, NA, RNG, UC, US); South of Cerro Sombrero, 52°46'S, 69°18'W, Goodall 2060 (LTR, NA); Bahia Inutil, Estancia Cameron, 53°38'S, 69°39'W, Moore 1076 (CAS, GH, RNG); Lago Blanco at Rio Blanco, Moore 1098 (CAS, RNG); Aserradero Rio Bueno, 53°40'S, 69°55'W, Pisano 2486 (CONC, HIP, RNG); Rio Fontaine, 54°29'S, 68°59'W, Skottsherg 122 (UPS). Ultima Esperanza, Estancia Cerro Castillo, Lago Porteno, Moore 1023 (DS, RNG, S, SGO); Estancia Bories, Lago Escondido, 51°55'S, 72°09'W, Moore 1028 (DS, RNG, US); Rio Rubens, Kalela 2088 (H); 2 km E of Puerto Condor, 51°38'S, 72°40'W, Solomon 4703 (MO); Estancia Cerro Castillo, Lago Sofia, 5I°32'S, 72°37'W, T.B.P.A. (Latour et al.) 1508 (BAB, MO); Seno Ultima Esperanza, Puerto Toro, 51°25'S, 73°04'W, T.B.P.A. (Moore & Pisano) 1896 (BAB, MO, RNG). Falkland Islands, Without locality, Hooker in 1842 (BM, E, G, K, P). east Falkland: Goose Green, ca. 3 mi SE of Bodie Creek House, Moore 612 (RNG); Darwin, Moore 640 (DS, GH, LP, S, UC). Port Harriet, Wright (K). west Falkland: Fox Bay, Vallentin in 1910 (K). In southern South America, Epilobium cilia turn subsp. ciliatum is an ex- tremely widespread, weedy, highly polymorphic taxon. Despite its variable na- ture, this species possesses many features that clearly distinguish it from other taxa. Most plants produce turions or leafy rosettes below or at the ground surface as overwintering structures, a feature that is not present in any other native South American species. This, combined with the presence of seeds with longitudinal lines of fused, flattened papillae, erect glandular-pubescent inflorescences, and small white to pink flowers, serves to separate E. ciliatum from other species with which it might be confused. Two species in particular, E. ohscurum and E. australe, are often mistaken for E. ciliatum, especially when the specimens lack underground parts. Epilobium ohscurum produces leafy basal shoots and has a densely strigillose inflorescence with glandular hairs restricted to the floral tube, rose purple petals, and narrowly lanceolate, denticulate leaves. Some plants of E. ciliatum from higher elevations or exposed sites are especially easy to confuse with E. australe, because their leaves are relatively small with few teeth, more congested, thicker, and the plants often are low in stature (less than 20 cm). Epilobium australe is shortly rhizomatous with leafy basal shoots; it completely lacks glandular pubescence and has thick, few-toothed, ovate leaves. Epilobium ohscurum and E. australe have seeds without lines of fused papillae. Many names have been given to the forms of E. ciliatum in South America. Most of these were proposed at a time when the number of collections was small, and each of the taxa appeared more or less homogeneous. For example, E. chi- lense was based on large-leaved, robust, densely glandular plants from central Chile, while E. valdiviense was drawn from specimens from the same area that were more graceful, less pubescent, with the leaves shorter than the internodes. In his monograph, Haussknecht (1884) recognized only three taxa in this complex 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 in South America. Subsequently, other names were proposed piecemeal for more or less distinctive populations or because of a lack of literature or understanding of previously published names. Samuelsson (1923) recognized all of Haussknechfs taxa plus two others and, in addition, proposed five new species. These new taxa were either from remote areas or had distinctive morphological features that separated them from the other taxa with which Samuelsson was familiar. He placed a great deal of emphasis on the density of pubescence, leaf size and disposition, and branching patterns in separating his taxa; these features are extremely plastic within this subspecies. The relationship of South American collections to North American taxa has been uncertain historically. Haussknecht, in his original description of E. ade- nocaulon, listed a collection from Aconcagua Prov., Chile (R. Philippi, W) among the four collections cited, with the other three from North America (Haussknecht, 1879). Later, in his monograph (Haussknecht, 1884), he re-identified this speci- men as E. chilense but then cited a collection from Missoula, Montana (Krause, JE) under the latter name. Samuelsson (1923) later reassigned the Montana plants to E. adenocaulon. Apparently after publication of his monograph, Hauss- knecht annotated a few additional collections from Chile as E. glandulosum (Reiche, 1898) and E. adenocaulon (e.g., Castro, Chiloe Prov., Chile, Gay 63, P). The relationship of the South American populations to those in North America was brought up again when it was learned that the seeds of Epilobium chilense shared a peculiar feature, the presence of longitudinal ridges of flattened, fused papillae, with the E. ciliatum complex (£. ciliatum, including E. glandulosum and E. watsonii; and E. oreganum; Seavey et al., 1977). This, combined with fleshy-leaved turions in both South and North American plants, strongly sug- gested that E. chilense was very closely related to E. ciliatum, if not conspecific (Seavey & Raven, 1977c). Hoch (1978) examined the Epilobium ciliatum complex in North America, especially E. ciliatum itself. Plants of this species had long gone under a plethora of names, in particular E. adenocaulon and E. glandulosum. Hoch's study made it clear that the many taxa recognized from North America could not be recog- nized and still maintain a functional, information-rich taxonomy. As a result, E. ciliatum was reduced to a series of three modally and geographically distinct but intergrading subspecies. In light of the careful work of Hoch and an examination and comparison of the bulk of the available South American material with a broad spectrum of North American populations, it is clear that the many taxa proposed from South Amer- ica can be reduced to the single variable taxon E. ciliatum subsp. ciliatum as circumscribed by Hoch (1978). Indeed, some plants are almost indistinguishable (e.g., Yellowknife, Mackenzie Territory, Canada, Cody & McCanse 3327, MO, and near Paso Puyehue, Osorno Prov., Chile, Solomon 4593, MO; Kyle Canyon, Charleston Mtns., Clark Co., Nevada, Goodman & Hitchcock 1707, MO, and Vegas Blancas, west of Angol, Malleco Prov., Chile, Solomon 4450, MO; Rat- tlesnake valley, Missoula Co., Montana, Barkley 1887, MO, and Cerro Tronador, Rio Negro Prov., Argentina, Solomon 4640, MO; Drummond Is., Chippewa Co., Michigan, McVaugh 9143, MO, and Rio Blanco, north end of Lago Blanco, Tierra 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 299 del Fuego Prov., Chile, Moore 1098, RNG; Hawks Ranch, 25 mi S of Laramie, Albany Co., Wyoming, Churchill in 1918, MO, and between Cerro Blanco and La Hollada, Cordoba Prov., Argentina, Solomon 4096, MO; Ruby River, 30 mi S of Varney, Madison Co., Montana, Parker 7033, MO, and 8 km W of Aguas Frescas, Magallanes Prov., Chile, Solomon 4685, MO). These examples cover the full range of overlapping morphological characters found in North and South American plants, from simple to branched, densely glandular-pubescent to sparsely strigillose, and in size from 4 cm to over 40 cm. Although there is considerable overlap in morphological features, two com- ponents, leaves and turions, are modally slightly different from populations in North America. The majority of South American plants produce fleshy-leaved turions just below the ground surface. This is especially prevalent at higher ele- vations and latitudes. Plants with leafy rosettes at the ground surface are found throughout the range. However, they form only a small portion of the plants from the southern part of the distribution of Epilohium ciliatum and are only frequent in the northernmost populations and in the coastal mountains and at low eleva- tions (i.e., below about 1,000 m). Similarly, the leaves of plants from higher elevations and latitudes in South America show a tendency to be slightly thicker, generally shorter and proportionately broader, with fewer marginal teeth than many North American populations. Again, this is only a trend, with plants of very similar morphology appearing in North America, but not as often. The same trend from leafy rosettes toward condensed turions is also seen in North Amer- ican populations, especially in cold or extreme habitats (Hoch, 1978). In general, although E. ciliatum subsp. ciliatum in South America is quite variable, the overall diversity of morphological types represented is not as great as that found in North America and many of the conspicuously distinctive mor- phological types now included in subsp. ciliatum of North America are not pres- ent (e.g., E. holosericeum, E. ecomosum). It seems most likely that E. ciliatum is a fairly recent, possibly late Pleistocene or even Holocene, arrival in South America by means of long-distance dispersal. While it has diversified to some extent, the reduced variability of morphological types encountered probably re- flects the relatively limited genotypic diversity of its founding population. The most extreme range of variability in E. ciliatum is found in stature and branching patterns. Not only do these differ from one population to the next, but they are usually conspicuous within populations, depending on the location of an individual: in shade or full sun, at the edge of a stream, in a bog, or on a drying gravel bar. Plants are often simple, especially when small, but generally branch above or from below later in the growing season. Secondary branching can be extremely dense and prolific when the terminal shoot is damaged, giving the plants a bushy appearance. Low plants, ca. 15 cm tall, with this type of branching formed the basis of the name E. magellanicum. Since E. ciliatum is perennial, turions or leafy rosettes as overwintering struc- tures are produced late in the season. The dead scales of the previous year's turions usually persist around the base of the stem, often several centimeters under the ground surface, but these are frequently lost when specimens are gath- ered. Because E. ciliatum, like most other Epilohium species, will germinate, grow, and reproduce in a single season, many plants collected early in the growing 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 6^ season or on newly formed, disturbed sites may appear to be annual, without any indication of perennating structures. It is certainly possible that individuals in particularly marginal habitats may be facultatively annual and do not produce overwintering structures at all. Within a given population there is frequently an enormous disparity in size of the individuals. For example, an extensive population at the Termas de Chilian, Nuble Prov., Chile {Solomon 4373, 4376) produced plants from less than 6 cm to more than 60 cm tall, all actively flowering and producing fully mature capsules. There were many more small and medium-sized individuals than large ones. This pattern is frequent throughout the range of E. ciliatum although not necessarily in every population. Variation in the size of herbaceous plants has always posed a problem for taxonomists, especially when dealing with limited material that does not cover the full range of morphological diversity. Per Dusen described Epilobium santa- cruzense from several dozen plants between 5 and 10 cm tall, collected fcW in uli- ginosis^ from the upper valley of the Rio Santa Cruz. He compared them with E. anagallidifolium, but suggested that their seeds were very similar to E. sa.xi- montanum, a member of the E. ciliatum complex. Although these plants are quite distinctive when taken by themselves, they have been retained in E. ciliatum because populations from slightly further west around Lago Argentino (Solomon 4653, 4658, MO) contain plants of very similar morphology as well as a contin- uously intergrading series of larger plants. The seeds of E. santa-cruzense have the conspicuous ridges typical of E. ciliatum. The extent to which size differences, as seen in Epilobium ciliatum, are ge- netically controlled is not known. In a recent study of an annual, out-crossing species of Stephanomeria, Gottlieb (1977) showed that there was no apparent genetic difference between small and large members of a single population, and that size and seed production were regulated by environmental factors. It is certainly possible that a similar situation occurs in Epilobium ciliatum. In a series of collections from Villa Cerro Catedral, Rio Negro Prov., Argentina {Solomon 4613, 4614, 4615, MO), plants from a level sandy area, which had once been wet, were mostly less than 7 cm tall, with small flowers and densely leafy stems. Only a few meters away, at the edge of a small stream, plants 30 to 40 cm tall, with long internodes and larger flowers were found. In between were plants of intermediate stature, their size apparently depending on the available soil moisture. The high rate of autogamy in E. ciliatum undoubtedly must have some impact on the genotypes available for each succeeding generation, but its significance is not known. A study similar to that conducted on Stephanomeria exigua Nutt. would be well worth the investigation. Stem vestiture also shows significant variation. In general, Epilobium ciliatum has both glandular and strigillose pubescence, although both types may be sparse, or the glandular hairs may be restricted to just the ovaries. Only rarely are com- pletely glabrous plants encountered, such as those on which Samuelsson based his E. leiophyton. The density of the glandular pubescence seems to have some geographical basis. Plants from lower elevations in the central valley and coastal range in the northern portion of the Chilean distribution (mostly north of Valdivia) are often densely glandular-pubescent (e.g., Concon, Valparaiso Prov., Chile, 1982] SOLOMON— EPILOBIUM IN SOUTH AMERICA 301 GaraventaSOO; Limache, Quillota Prov., Chile, Looser 2007, 2011 ; Corral, Valdiv- ia Prov., Chile, Gunckel 15169; 44 km E of El Colorado, Talca Prov., Chile, Solo- man 4339; 30 km W of Curacautin, Malleco Prov., Chile, Solomon 4473). Within this distribution, densely glandular plants are frequent and often characterize entire populations. At higher elevations and further to the south, glandular hairs are less dense, and the dense type does not appear in Argentina at the same latitude. Although pubescence is not an insignificant feature and does apparently have a geographical basis, it is not useful to recognize these plants in a formal way, because the variation in other characters is indistinguishable from that found in other populations, and pubescence density is often a quantitative and plastic character. The only evident correlation for the presence of densely glandular plants may be due to their occurrence in the warmest and driest portions of the range of E. ciliatum in South America, although the functional significance of most hair types in plants is very poorly known (Ehleringer et al., 1976; Levin, 1973). Epilobium ciliatum typically has small, highly autogamous flowers that show little variation in size. Like the turions and leaves discussed earlier, the color of the petals also varies geographically. Low-elevation plants from the central valley and coast range of central Chile have white flowers that flush only a faint pink, or not at all, after anthesis. Those from higher places in the Andes at this same latitude are also usually white at anthesis, but these may fade to a darker pink afterward. As one moves southward along the Andes, beginning in the vicinity of San Carlos de Bariloche, plants with pink flowers at anthesis become increas- ingly common, so that populations from Santa Cruz Prov., Argentina, adjacent Chile, and Tierra del Fuego are all pink flowered. Pink flowered populations may also be found further north, but only at higher elevations. The pink petal color is produced by malvidin-3-5-diglucoside (Harborne, 1967), the presence and abun- dance of which, in the case of E. ciliatum, is possibly environmentally controlled. The genes that control pigment production are perhaps induced under the influ- ence of decreasing temperatures, or increased ultraviolet radiation at high ele- vations or latitudes. Transplants of turions from pink flowered plants (Ushuaia, Tierra del Fuego, Argentina, Solomon 4712; grown as Ml 856 in 1979), under greenhouse conditions, produced white flowers. Seed from two other pink flow- ered populations (near Ventisquero Perito Moreno, Prov. Santa Cruz, Argentina, Solomon 4656, 4658; grown as M1850, M1851, in 1979, respectively) also pro- duced only white flowers under greenhouse cultivation. It is not surprising that a weedy, autogamous species such as Epilobium cil- iatum should exhibit a substantial amount of variation, both between populations and within a single population. The often disturbed habitats where E. ciliatum occurs, such as stream beds, seep, or embankments, over a broad elevational range, are ephemeral, being newly created or altered from one year to the next, and often in a drastic manner, by such agents as flooding. The ability to suc- cessfully colonize these localities requires not only high dispersability, but also great genotypic and phenotypic plasticity to cope with a broad spectrum of en- vironmental parameters. It may be safely said that E. ciliatum is a superlative example of such an opportunistic species. Disjunct populations of Epilobium ciliatum are found in the Sierra de San Luis 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 and the Sierra Grande de Cordoba. These undoubtedly originated by long-dis- tance dispersal of seed from the main range of the Andes via the westerly winds. Some of the most robust and largest leaved individuals seen in any population came from the Sierra Grande, where they were found growing in moist ravines (Solomon 4074, 4096). All of the populations examined in the Sierra Grande had white flowers. Disjunct populations also occur in the Falkland Islands and must have been derived in a similar fashion from populations to the west. Apparently, E. ciliatum is rare and scattered along streams on these islands (Moore, 1968). Epilobium ciliatum is by far the most common species encountered in the central and southern Andes of Chile and Argentina, where it grows sympatrically with at least nine other species. It overlaps with E. hirtigerum and E. denticu- latum, in the Sierra Grande, and with the latter also in a narrow band along the east side of the Andes in Mendoza and San Juan provinces, Argentina, and in the Sierra de San Luis. Despite the highly autogamous or even functionally cleistogamous flowers, occasional hybrids with Epilobium australe, E. barbeyanum, E. denticulatum, E. glaucum, E. hirtigerum, E. obscurum, and possibly E. puberulum do occur. Only hybrids with E. australe and E. glaucum are discussed here, because they are the most frequently encountered, in part due to their broadly sympatric range with E. ciliatum throughout the southern Andes. Details on the other hybrids are found under each of the other species listed above. All of the species that hybridize with Epilobium ciliatum, except E. dentic- ulatum and E. puberulum, have the BB chromosome arrangement. Hybrids be- tween the two chromosomal groups are usually easily distinguished because of the intermediate morphology and reduced seed set and pollen stainability. Hybrids between E. ciliatum and E. glaucum (e.g., Arroyo Huelta, Lago Nahuel Huapi, Solomon 4626, 4627, MO; and Cerro Lopez, Rio Negro Prov., Argentina, without collector in 1953, BAB) generally have narrow, few-toothed leaves, strigillose pubescence, sometimes with glandular hairs, and loose, elon- gate turions. The average pollen stainability for Solomon 4626, 4627, and the Cerro Lopez collection was 21%, 19%, and 22%, respectively. Hybrids between Epilobium ciliatum and E. australe are more difficult to distinguish, but the reduced seed set or poor capsule development is an indicator of hybrid origin. In hybrids of this combination (e.g., Ea. Achalay, Santa Cruz Prov., Argentina, T.B.P.A. 2349, MO, RNG; Rio Coyle, Santa Cruz Prov., Ar- gentina, Dauber 4, BAA; and Arroyo Beta, Tierra del Fuego, Argentina, Cas- tellanos in 1942) have intermediate leaf morphology, usually a few glandular hairs, larger pink flowers, good seeds with few, imperfectly formed ridges, and loose, somewhat elongate leafy basal shoots. The average pollen stainability for Dauber (MO, RNG), and Castellanos in 1942 was 19% 30% 6. Epilobium nivale Meyen, Reise um die Erde 1:315. 1834. Walp., Reliq. Meyen, 327. 1843. type: Chile, VII Region (Maule), Cerro Imposible, 3,800 m, Jan- uary-March 183 1 , F. Meyen (JE, lectotype here designated, photograph MO; B (destroyed, photographs BH, GH, MO, NY), BR, K, P, W, isolectotypes). Gay, Hist. Fis. Chile 2:349. 1846. Walp., Repert. Bot. Syst. 5:666. 1846. Hausskn., 1982| SOLOMON— EPILOB1UM IN SOUTH AMERICA 303 Monogr. Epilobium 251. 1884. Reiche, Fl. Chile 2:244. 1898. H. Lev., Iconogr. Epilobium, tab. 205. 1911. Samuelsson, Svensk Bot. Tidskr. 17:290. 1923. E. alpinum auct. non L.: Hook. & Arn., Hook. Bot. Misc. 3:390. 1833. E. andinum R. Phil., Anal. Univ. Chile 84:747. 1893. type: Chile, VII Region (Maule), Cordillera de Talca, Cuesta de Las Animas, February 1879, F. Philippi (SGO-53014, lectotype here designated, photographs GH, MO; SGO-41463, SGO-53021 (specimen in upper right portion of sheet), prob- able isolectotypes). Philippics handwritten annotation of Epilobium andinum has been seen on six herbarium sheets at SGO, representing three different species, E. nivale, E. australe, and E. ciliatum subsp. ciliatum, all potential types for the name E. andinum. That all three species were included in the protologue is evidenced by: 1. fck El ejemplar mas grande del herbario mide 19 centimetros, . . . M (The largest specimen in the herbarium measures 19 cm, . . .). There is one specimen, SGO-53019, which is about 19 cm tall, a collection off. ciliatum subsp. ciliatum. 2. "Los ejemplares de la Cordillera de Talca y de La Cueva (Sierra Velluda) son menos altos y echan muchas ramas en su base" (The specimens from the Cordillera de Talca and La Cueva (Sierra Velluda) are smaller and produce many branches from the base). The specimen from La Cueva, SGO-53020, is an example of E. australe. 3. In the Latin diagnosis, R. Philippi begins with, "Epilobium glaberrimum; caule tereti; foliis omnibus oppositis, . . . ." There are three specimens of £*. nivale, SGO-53014, SGO-41463, and SGO-53021 in upper right, all of which are glabrous and have consistently opposite leaves. Among native South American Epilobium, only E. nivale and the distinctive E. glaucum are glabrous. Based on the Latin diagnosis, it appears that Philippi had the specimens of E. nivale specifically in mind, even though he included specimens of two other species in his discussion. Low, caespitose perennial herb, (5-)10-20(-30) cm tall, from a fibrous root system. Stems numerous, decumbent, terete, simple, occasionally branched above, usually reddish purple throughout or bicolored reddish purple on the upper sur- face and green on the lower, glabrous. Leaves mostly opposite, alternate only in the inflorescence, thick, dark green, often reddish purple along the margins, nar- rowly ovate to lanceolate, 0.5-1.6(-2.5) cm long, 1.5— 5(— 8) mm wide, acute or occasionally obtuse at the apex, remotely denticulate with 1-4 low teeth on each side, acute to cuneate at the base, glabrous, the lateral veins very inconspicuous, none or 1, rarely 2, on each side of the midrib, on petioles 0.5-1 mm long. purpl rpl 1.2 cm long, on pedicels 1-6 mm long. Floral tube often reddish-purple, 0.5-0.8 mm deep, 1.2-2 mm wide, glabrous. Sepals often reddish purple, lanceolate, 1.9-2.8 mm long, 0.8-1.3 mm wide, glabrous. Petals pale pink, 2.8-4.5 mm long, (1-)1.4- 2.2 mm wide, the notch 0.5-0. 9(-l. 2) mm deep. Anthers cream, 4-0 4-0 1.6 mm long, 4-0 the stigma at anthesis. Style cream to white, 1.2-1.6 mm long; stigma cream to (>- purpl 1-4 1.2(-1.8) cm long. Seeds pale brown, papillose, obovoid, 1.1-1.4 mm long, 0.3-0.5 mm thick, the chalazal end with an appendage, 0.04-0. 1 mm long, 0.08-0. 15 mm wide; coma white, 2.5-7 mm long. Gametic chromosome number, n = 18. Distribution (Fig. 5): Along rivulets, boggy places, stream banks, and other permanently wet sites in alpine areas, often extending upward to the snow line, occasionally on stable scree, and sometimes extending to slightly lower eleva- tions, especially in the forest regions, along rocky stream beds. In the central 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 and southern Andes of Chile and Argentina from Paso Portillo (33°S), southward along the mountains to the vicinity of Epuyen, Chubut Prov., Argentina, disjunct to the Torres del Paine region of Chile and Lago Argentino, at elevations of 2,000- 3,500 m in the northern portion of its range, gradually descending to 1,200-2,000 m in Rio Negro and Chubut, and in the southernmost populations from 350 to 1,200 m. Flowering December to March. Representative specimens examined: Argentina, chubut: Epuyen, Cuartel Epuyen, Lourteig & Buchinger 22 (P). mendoza: Valle Atuel, Lago Atuel, 3,100 m, Backer et al. 1972 (C, DS); Dpto. Las Heras, Las Cuevas, 3,220 m, Boelcke 9766 (BAA, BAB, MO); Dpto. Malargue, Paso Pehuenche, Boelcke et al. 10358 (BAA, BAB, MO); Arroyo en el alto valle, Calmuco, 36°24'S, 69°50'W, Burkart et al. 14178 (LIL); Tres Esquinas, 33°50'S, 69°00'W, Curette 285 (LP, SI); Laguna Diamante, 3,200 m, Curette 286 (LP, SI), Serru 39 (LP); Valle del Atuel, Cajon del Burro, Gerth 108 (L, SI); Valle Hermoso, Kurtz 5860 (CORD, G, JE, NY), Sosa 13 (SI); Dpto. Malargue, entre Arroyo Alverjalito y Arroyo Lerias Amarillas, Kurtz 7160 (CORD, JE, SI); Dpto. San Rafael, Arroyo El Indigeno, 10 km N of Volcan Overo, Lagiglia 2218 (LP); Dpto. Tunuyan, Valle del Alto Tunuyan, Paso Hondo, Leal 2059 (LIL); Dpto. San Rafael, Cerro de Los Molles, Kurtz 7514 (CORD, JE, LP), Rossi 297 (LIL); Sleumer 637 (B, LIL). neuquen: Parque Nacional Nahuel Huapi, Cerro Colorado, entre Pto. Manzano y Lago Traful, Boelcke & Correct 6909 (BAA, BAB); Norquinco, 2,200 m, 39°08'S, 7 1°18' W, Comber 539 (K); Dpto. Lacar, Cerro Chapelco, 1,800 m, Cabrera 23009 (LP), Correa et al. 5886 (BAB, MO); Copahue, 1,000 m, Kraftsik in 1968 (BAB). Paso Pino Hachado, Parodi 2215 (BAA); Lago Villarino, 40°26'S, 7I°36'W. Roth in 1896 (LP). Rio nfx;ro: Cerro Ironador, Rio Manso Su- perior, Gentili in 1975 (MO); Cerro Utne, 41°19'S, 71°21'W, Hosseus 456 (CORD), 463 (CORD); Parque Nacional Nahuel Huapi, Cerro Catedral, 2,000 m, Pedersen 1488 (C); Cerro V. Lopez, W of Bariloche, 1,500 m, 41°05'S, 71°35'W, Solomon 4620 (MO), santa cruz: Estancia Lago Roca, Cerro Fraile, 50°32'S, 72°39'W, James 427 (BM, DS); Lago Argentino, Cerro Buenos Aires, 50°24'S, 72°59' W, James 711 (BM), Skottsberg 945 (UPS); Dpto. Guer Aike, Estancia Las Viscachas, 1,250 m, 50°46'S, 72°08'W, T.B.P.A. (Arroyo et al.) 2671 (RNG). Chile, v region (aconcagua): Los Andes, Juncal, 32°52'S, 70°10'W, Buchtien in 1903 (US); Laguna del Inca, Kurtz in 1886 (JE, LP, NY); Paso Portillo, 2,800 m, Milner in 1935 (CONC), Sparre 1687 (S), Wall in 1946 (A, S). region metropolitana (Santiago): Cordillera, Vegas del Tupungato, Behn in 1930 (CONC), Melin in 1930 (UC); Cerro de San Pedro Nolasco, 33°48'S, 70°16'W, Gillies (BM, E, OXF); Laguna Negra, Vidal in 1873 (SGO); Ski village La Parva, 2,850 m, 33°20'S, 70°I8'W, Moore 395 (LA, MO), Solomon 4361 (MO), vi region (ohiggins): Cachapoal, Sewell, 2,400 m, Bastin 16 (US); Cauquenes, Gay 188 (P); Rio Coya above El Teniente, 2,700 m, Pennell 12292 (F, GH, NY, PH, SGO); Ventisquero Los Cipreses, 2,000 m. Reed in 1872 (POM). Colcha^ua, Cordillera de Colchagua, 2,000 m, Pirion 107 (GH). vn region (maule): Curico f Cajon del Azufre, Albert in 1891 (SGO); 20 km oeste del Paso Vergara, Calderdn in 1967 (CONC); Laguna de Teno, 2,500 m, Marticorena & Matthei 841 (CONC). Linares, Cordillera de Linares, Philippi in 1862 (G, HAL, S); Termas de Longavi, Schumann in 1888 (SGO). Talca, Laguna del Maule, Aravena 1021 (UC), Ricardi et al. 969 (CONC, MO), vm region (bio-bio): Bio-Bio, Cordillera de Antuco, La Cueva, Rahmer in 1887 (SGO). Nuble, Termas de Chilian, Jaffuel 3728 (GH), Pennell 12413 (F, GH, NY, PH), Solomon 4372 (MO), ix region (araucania): Malleco, Termas del Rio Blanco, 1,300 m, 38°35'S, 71°35'W, Solomon 4504 (MO), xn region (magai lanes): Ultima Esperanza, Estancia Cerro Paine, 50°59'S, 72°58'W, Pisano 4344 (HIP, MO). Epilobium nivale is a distinctive, high montane species, readily recognized by its glabrous, delicate, decumbent, clumped or somewhat caespitose stems, with few, small pink flowers. Only E. glaucum and some individuals of the introduced E. paniculatum are also glabrous, but they are much more robust and erect. The small pink flowers are often functionally cleistogamous, because they open par- tially or not at all on cloudy days, and only for a few morning hours on sunny days. Epilobium nivale grows sympatrically with four other species, but only rarely is there evidence to suggest interspecific hybridization. Hybrids with £. austral e and E. densifolium would be difficult to distinguish from small-leaved forms of those species, except perhaps in flower size. Recognizable hybrids involving E. 1982] SOLOMON— EP1LOBIUM IN SOUTH AMERICA 305 ciliatum have not been found, and only a single collection from Termas de Chilian, Prov. Nuble, Chile {Solomon 4372, MO) exhibits a tendency toward larger, acu- minate leaves and the more erect stems characteristic of E. glaucum. This spec- imen, however, had a pollen stainability of 96%, which is substantially different than the stainability of artificial hybrids between these species. Ecologically, E. nivale occurs in the same habitats and community associa- tions in the southernmost disjunct populations as it does in, for example, the vicin- ity of Lago Nahuel Huapi; that is, in wet places at the upper limits of Nothofagus pumilio (Poepp. & Endl.) Krasser scrub and at higher elevations. In the Torres del Paine region, at elevations approaching 1,000 m, E. nivale is apparently the only species found in permanently wet places (Pisano, 1974, as E. conjungens). Farther north, beyond the limits of the Nothofagus forest, E. nivale continues to appear in permanently wet sites, but at increasing elevations. 7. Epilobium Bull. Herb. Boissier, ser. 2, 7:589. 1907 type: Chile, IV Region (Coquimbo), Vegas del Toro, Cordillera de Coquimbo, 2,500 m, 1 February 1883, F. Philippi (?) (G, holotype, photograph MO; JE, SGO-53122, isotypes). The specimen at G is accompanied by a letter from F. Philippi dated 29 May 1883, and the specimens were probably collected by him. H. Lev., Repert. Spec. Nov. 9:19. 1910-11. Samuelsson, Svensk Bot. Tidskr. 17:265. 1923; Svensk Bot. Tidskr. 24:4. 1930. Perennial herbs (8-) 15-40 cm tall, overwintering and reproducing vegetatively by elongate leafy shoots or soboles produced from the base. Stems erect ascen- dent or decumbent, terete, usually simple or sparingly branched throughout, abundantly branched from the base, densely pubescent throughout with erect glandular hairs, 0.1-0.2 mm long, with an admixture of strigillose hairs, 0.1-0.2 mm long, in descending lines from the petiole bases or scattered in the inflores- cence. Leaves mostly opposite, usually alternate in the inflorescence, the inter- nodes often elongate and the leaves remote in fruit, thin, bright green, lanceolate to narrowly ovate, (0.6— )1— 2 cm long, 0.2-1 cm wide, acute to acuminate, rarely obtuse at the apex, remotely denticulate with 3-5 teeth on each side, the margin frequently undulate, acute to obtuse or occasionally rounded at the base, with erect glandular hairs on both surfaces, usually with an admixture of strigillose hairs on the adaxial midrib and margins or thinly scattered on both surfaces, glabrate with age, the lateral veins obscure, 2-4 on each side of the midrib, on poorly defined petioles 0-1 mm long. Inflorescence erect, simple, the leaves sub- tending the flowers undifferentiated or slightly reduced in size. Flowers erect. Ovaries densely covered with erect glandular hairs, with an admixture of strigil- lose hairs, these also often dense, 0.9-1.5 cm long on pedicels 1-5 mm long. Floral tube 0.8-1.5 mm deep, 1.3— 2(— 2.7) mm across, externally with erect glan- dular and scattered strigillose hairs, internally with a conspicuous ring of erect villous hairs, 0. 15-0.2 mm long near the base. Sepals lanceolate, 2.5-3.2(-4) mm long, 0.9-1.5 mm wide, with erect glandular and scattered strigillose hairs. Petals pale pink, obovate, 4-6 mm long, 2.5-3.4 mm wide, the notch 1-1.7 mm deep. Anthers cream to white, 0.6-1 mm long, 0.5-0.7 mm wide; filaments cream to white, those of the longer stamens 1.5-2(-3) mm long, those of the shorter 0.6- 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 69 1(— 1.9) mm long; the longer stamens shedding directly on the stigma at or shortly after anthesis, occasionally the shorter ones also. Style cream to white, capitate, 0.6-1 mm long, 0.6-1 mm thick. Capsules strictly erect, sparsely covered with erect glandular and scattered strigillose hairs, 3.4-5.2 cm long, 1.3-1.6 mm thick, on pedicels (0. 5-) 1.2-2. 9 cm long. Seeds brown, papillose, obovoid, 0.85-1.3 mm long, 0.4-0.5 mm thick; coma white to slightly yellowish brown, 5-8 mm long. Gametic chromosome number, n = 18. Distribution (Fig. 7): In very wet places in bogs, and along streams, sometimes partially submerged or floating; throughout the central Andes of Chile and Ar- gentina, from southern III Region (Atacama), Chile, and central San Juan Prov., Argentina, southward to the vicinity of Lago Nahuel Huapi; also in the coast range (Cuesta La Dormida) between Santiago and Valparaiso, at elevations of 2,000-3,500 m in the north, gradually descending to 900-1,700 m in the southern part of its range. Flowering November to March. Specimens examined: Argentina, mendoza: Dpto. Malargiie, camino a Paso Pehuenches, Km 50, 1,800 m, Cabrera et al. 22831 (LP), neuquen: Dpto. Los Lagos, Estancia Fort in Chacabuco, Boelcke 4547 (BAA); Dpto. Minas, Arroyo Las Bandurria, entre Las Ovejas y Laguna Epulauquen, 1,250 m, 36°55'S, 70°56'W, Boelcke et al. 10788 (BAA, BAB, MO), Ragonese 225 (BA, RSA); Dpto. Minas, Laguna Epulauquen, 1,300 m, 36°50'S, 71°05'W, Boelcke 10906 (BAA, BAB, MO); Dpto. Chos Malal, Vegas del Pelan, 1,700 m, 36°54'S, 70°20'W, Boelcke et al. II 134 (BAA, BAB, MO); Dpto. Minas, Laguna Varvarco Campos, Arroyo Benitez, 36°17'S, 70°39'W, Boelcke 14224 (BAA, BAB); Traful, Estancia La Primavera, Castellanos in 1938 (BA, RSA), Gerrovia in 1938 (LIL); Rahue, camino a Alumine, Dawson & Schwabe 2180 (BAA, BAB, CTES); Paso Pino Hachado, Hauman in 1920 (BA); Dpto. Loncopue, Cajon Chenque-Pehuen, 38°06'S, 70°55'W, Rugolo & Agrasar 172 (BAA, MO), san juan: Dpto. Calingasta, Manantiales, 3,500 m, Fabris & Zuloaga 8449 (LP), Zardini 180 (LP); Dpto. Iglesia, Rio Blanco, Sierra San Guillermo, 3,200 m, 29°08'S, 69°30'W, Hosseus 1443 (CORD); Dpto. Calingasta, Cienaga de Las Cabeceras, 2,700 m, 3I°47'S, 69°07'W, Kurtz 9806 (BAF, CORD), 9815a (CORD), Spegazzini in 1937 (BAB); Dpto. Iglesia, Tudcum, Perrone in 1950 (BA, MO). Chile, hi region (atacama): Huasco, Quebrada Alfalfa, 2,800 m, 28°52'S, 69°49'W, Johnston 5996 (GH, S); Rio de la Laguna Grande, 2,750 m, 28°44'S, 69°57'W, Johnston 6009 (GH, S). iv region (coquimbo): Choapa, Rio Illapel, without collector or date (K), Volkmann in 1860-61 (SGO); Quebrada La Vega Escondida, E of Cuncumen, 2,630 m, Morrison & Wagenknecht 17433 (GH, NA, UC), Worth & Morrison 16556 (NA, UC). Elqui, Laguna de Elqui, Barros 6616 (US); 6 km al oeste del Embalse La Laguna, 3,100 m, 30°13'S, 70°04'W, /?/car