UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY '2001 5otany EW SERIES, NO. 39 L Revision of the South American Ipecies of Brunfelsia (Solanaceae) imothy C. Plowmant , Knapp and J. R. Press, Editors ugust 31, 1998 ublication 1496 UBRAKY wia. Information for Contributors to Fieldiana • • i • • Referenci cs for - • uton Publ,- .. ' , and • Only -. >ciper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). FIELDIANA Botany NEW SERIES, NO. 39 A Revision of the South American Species of Brunfelsia (Solanaceae) Timothy C. Plowmant Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 U.S.A. S. Knapp and J. R. Press, Editors Department of Botany The Natural History Museum London SW7 5BD England Accepted July 2, 1996 Published August 31, 1998 Publication 1496 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1998 Field Museum of Natural History ISSN 0015-0746 PRINTED IN THE UNITED STATES OF AMERICA n,S Table of Contents FOREWORD Warfe Davis vii EDITORS' PREFACE S. Knapp and J. R. Press xi I. MATERIALS AND METHODS 1 II. TAXONOMIC HISTORY OF THE GENUS 2 III. GENERIC RELATIONSHIPS 5 IV. GENERAL MORPHOLOGY 6 V. CYTOLOGY 16 VI. HYBRIDIZATION EXPERIMENTS 19 VII. ECOLOGY AND REPRODUCTIVE BIOLOGY 20 VIII. TRENDS IN SPECIALIZATION 26 IX. GEOGRAPHICAL DISTRIBUTION AND RELATIONSHIPS 29 X. TAXONOMIC TREATMENT 34 Brunfelsia Linnaeus 34 Artificial Key to the Sections, Species, and Infraspecific Taxa of Brunfelsia 35 Brunfelsia sect. Guianenses 38 1. Brunfelsia amazonica 38 2. Brunfelsia burchellii 40 3. Brunfelsia chocoensis 40 4. Brunfelsia clandestina 43 5. Brunfelsia guianensis 46 6. Brunfelsia martiana 49 Brunfelsia sect. Franciscea 51 1. Brunfelsia australis 53 8. Brunfelsia boliviano 57 9. Brunfelsia bonodora 59 10. Brunfelsia brasiliensis 61 lOa. Subspecies brasiliensis 62 lOb. Subspecies macrocalyx 66 1 1 . Brunfelsia chiricaspi 70 12. Brunfelsia cuneifolia 72 13. Brunfelsia dwyeri 74 14. Brunfelsia grandiflora 77 14a. Subspecies grandiflora 78 14b. Subspecies schultesii 80 15. Brunfelsia hydrangeiformis 84 15a. Subspecies hydrangeiformis 84 15b. Subspecies capitata comb, et stat. nov. 86 16. Brunfelsia imatacana 89 17. Brunfelsia latifolia 91 18. Brunfelsia macrocarpa 93 19. Brunfelsia mire 95 20. Brunfelsia obovata 98 20a. Variety obovata 99 20b. Variety coriacea 101 2 1 . Brunfelsia pauciflora 103 22. Brunfelsia pilosa 108 23. Brunfelsia rupestris Ill 24. Brunfelsia uniflora 113 Dubious and Excluded Species 119 Nomina Nuda 119 XI. EDITORS' ACKNOWLEDGMENTS 119 XII. PLOWMAN'S ACKNOWLEDGMENTS 120 XIII. LITERATURE CITED 120 APPENDIX I: List of Exsiccatae 124 APPENDIX II: Brunfelsia sect. Brunfelsia 132 Description 132 Names of Brunfelsia sect. Brunfelsia 132 APPENDIX III: Additional References 133 INDEX . 134 List of Illustrations Timothy Plowman vi Brunfelsia hydrangeiformis subsp. hydran- geiformis, type xii 1. Brunfelsia americana 3 2. Root of Brunfelsia grandiflora subsp. schultesii 8 3 . Floral structure of Brunfelsia 11 4. Brunfelsia densifolia (Knapp s.n.) from Puerto Rico 12 5. Brunfelsia lactea (Knapp s.n.) from Puerto Rico 14 6. Fruit of Brunfelsia americana 17 7. Methona themisto, type specimen 24 8. Caterpillars of Methona confusa on Brunfelsia chiricaspi leaf 25 9. Distribution of section Brunfelsia 31 10. Distribution of sections Guianenses and Franciscea 32 1 1 . Brunfelsia amazonica 39 12. Brunfelsia burchellii 41 13. Brunfelsia chocoensis 42 14. Brunfelsia clandestina 44 15. Distribution of species in section Guianenses 46 16. Brunfelsia guianensis 47 17. Brunfelsia martiana 50 18. Manaca (Brunfelsia uniflora) 52 19. Brunfelsia australis 55 20. Distribution of Brunfelsia australis, B. boliviana, and B. bonodora 56 21. Brunfelsia boliviana 58 22. Brunfelsia bonodora, illustration of lectotype 60 in 23. Distribution of Brunfelsia brasiliensis and B. chiricaspi 63 24. Brunfelsia brasiliensis subsp. bras- iliensis 64 25. Brunfelsia chiricaspi 71 26. Brunfelsia cuneifolia 73 27. Brunfelsia dwyeri 75 28. Distribution of Brunfelsia cuneifolia, B. dwyeri, and B. grandiflora 77 29. Brunfelsia grandiflora subsp. grandiflo- ra and subsp. schultesii 79 30. Brunfelsia hydrangeiformis subsp. hy- drangeiformis, illustration of type 85 3 1 . Isotype of Brunfelsia hydrangeiformis subsp. capitata stat. nov 87 32. Brunfelsia imatacana 89 33. Distribution of Brunfelsia hydrangei- formis, B. imatacana, and B. mire 90 34. Brunfelsia latifolia, illustration of type 92 35. Brunfelsia macrocarpa 94 36. Brunfelsia mire 96 37. Distribution of Brunfelsia latifolia, B. macrocarpa, B. obovata, and B. pilosa 99 38. Brunfelsia obovata var. obovata 100 39. Brunfelsia obovata var. coriacea 102 40. Brunfelsia pauciflora 104 41. Distribution of Brunfelsia pauciflora and B. rupestris 105 42. Brunfelsia pauciflora cv. "Lindeniana" 109 43. Brunfelsia pilosa 110 44. Brunfelsia rupestris 112 45. Brunfelsia uniflora, illustration of type 114 46. Distribution of Brunfelsia uniflora 116 List of Tables 1 . Historical classification of Brunfelsia .... 6 2. Chromosome counts in Brunfelsia 18 3. Chromosome numbers and size in Brun- felsia 19 4. Pollinators of Brunfelsia grandiflora 23 5. Brunfelsia species recorded as larval food plants for species of Methona 26 6. Trends in specialization of Brunfelsia ... 30 7. Flowering and fruiting times of Brunfel- sia uniflora 117 IV Tim Plowman Foreword The idea for this monograph emerged from somewhat unusual circumstances. On March 29, 1942, ethnobotanist and plant explorer Richard Evans Schultes arrived at the Kofan Indian village of Conejo, on the banks of the Rio Sucumbios in the remote rain forests of southern Colombia. In the midst of his first sojourn in the Amazon, with the goal of surveying medicinal plants and arrow poisons, Schultes found himself immersed in a world of phytochemical wizardry unlike anything he had ever known. Psychoactive and toxic plants touched every aspect of the lives of the Kofan. The most important and sacred of their plants was the hallucinogen Banisteriopsis caapi, or yage, the "vine of the soul." When Schultes inquired how often the people imbibed yage, his infor- mant's response suggested that the question had no meaning. During illness, of course, and in the wake of a death. In times of need or hardship. At certain passages in life. When a young body of six has his initial haircut, or when he kills for the first time. And naturally, the shaman suggested, a youth will drink yage at puberty, when his nose and ears are pierced and he obtains the right to wear the tail feathers of the macaw. As a young man, he may drink it at his leisure to improve his hunting technique or simply to flaunt his physical prowess. The message that Schultes received was that the Kofan took yage whenever they felt like it. At least once a week, and no doubt on any occasion that warranted it, such as the eve of his own departure from Conejo a fortnight later. With the people dancing, the men facing the women in long lines moving forward and then back, turning from side to side, the dancers stamping the ground lightly in time with the drums, this solitary stu- dent of plants, a man who in time would inspire a generation of ethnobotanists, took yage with, as he would recall 50 years later, "the whole damn village." The next morning, his face still painted and his mind swirling with the sounds of chants, Schultes paddled up the Sucumbios in a dugout canoe. His head throbbed. Unbeknownst to him, the prepa- ration had contained bark of tsontinb"k'o, the cold fever tree, together with su-tim-ba-che, a root said by the Kofan to cause a "drunk worse than yage." Indeed, it had. Despite his discomfort, Schultes made a note to investigate the admixtures at length. Thirty years later he did, by dispatching his protege, Timothy Plowman, to the Putumayo. After 15 months in the field in South and Central America and the Caribbean, Tim Plowman emerged with a definitive study, a monographic revision of the genus Brunfelsia, including de- scriptions of several new species and varieties. The plant that Schultes drank at Conejo and that gave him such a headache is now known as Brun- felsia grandiflora subsp. schultesii. Tim Plowman approached botanical research with the intensity and rigor of a scientist, the cour- age of his mentor, and the flair of a poet. His first Colombian excursion set the pattern of his career. Arriving in Bogota, he traveled south to the valley of Sibundoy, where he contacted Pedro Juajibioy, a Kamsa Indian healer who as a boy guided Schultes into the upper Putumayo. Tim and Pedro retraced that journey to reach a Kofan settlement on the Rio Guamues, also visited by Schultes in 1942. The Kofan recognized B. grandiflora as tsontinba"k"d. Another of Schultes's students, Homer Pinkley, who spent a year among the tribe in 1965, had written that their shaman occasion- ally ingested the plant in order to diagnose dis- ease. This observation supported other reports dating back to the 19th century that suggested the plant might be hallucinogenic. Plowman wanted to know. At Santa Rosa on the Rio Guamues he found B. grandiflora commonly cultivated in house yards, but he also met an old shaman who brought him from the forest a rare but related plant named for the tapir because of its inordinate strength as a drug. Plowman immediately recog- nized the plant as a new species, which he later named Brunfelsia chiricaspi after the Quechua word meaning cold tree. He asked the old man to prepare it, but the shaman refused. He described the plant as a dangerous messenger of the forest and disavowed any knowledge of taking it for vi- sions. Tim persisted. Eventually the shaman agreed, although reluctantly and only on the con- dition that Pedro also drink the preparation. The drug, an extract from the bark, was murky brown and bitter to swallow. Tim felt the effects within ten minutes: a tingling sensation such as one feels when the blood rushes back to a limb that has fallen asleep. Only in this case the sen- sation grew to a maddening intensity, spreading from his lips and fingertips toward the center of his body, progressing up his spine to the base of his skull in waves of cold that flooded over his consciousness. His breathing collapsed. Dizzy vn with vertigo, he lost all muscular control and fell to the mud floor of the shaman's hut. In horror he realized that he was frothing at the mouth. An hour passed. Paralyzed and tormented by an ex- cruciating pain in his stomach, he remained only vaguely aware of where he was — on the earth, face to face with three snarling dogs fighting over the vomit that had spread in a pool around his head. The shaman, noticing his plight, did what sha- mans normally do under such circumstances: He went to bed. Desperate to escape the sensation, half-blinded by the drug, and incapable of walk- ing, Tim and Pedro stumbled and crawled through the forest for two hours until finally, toward dawn, they reached the village of San Antonio, where they were staying in an abandoned jail. As light came to the forest, they crawled into their ham- mocks, where they remained, motionless, for two days. Pedro Juajibioy, whose experience as a folk healer had taken him on a thousand flights of the spirit, summed up the experience succinctly: "The world was spinning around me like a great blue wheel. I felt that I was going to die." This story reveals something of the character of this remarkable young scholar, a man of ad- venture, grace, generosity, kindness, modesty, and honor, whose academic achievements in time might well have surpassed even those of his fa- mous mentor, Richard Evans Schultes, the "Fa- ther of Ethnobotany." Sadly, Tim's life was cut short by cruel circumstances. His untimely death at the age of 45 from AIDS on January 7, 1989, severed a career of immense promise. Already far on the way as one of the most discerning, original, and effective naturalists of the century, Tim was a gentleman, a friend of everyone, an understand- ing and devoted teacher, an intellectual of extraor- dinary depth, and a tireless and demanding re- searcher happy to share his experience and coun- sel with whomever sought his advice. Tim Plowman's interest in and love of plants developed when he was a child growing up in the temperate woodlands surrounding Harrisburg, Pennsylvania. An avid collector even as a boy, his passion for plants grew into the central metaphor of his life. After attending college at Cornell Uni- versity, he went as a graduate student to the Bo- tanical Museum of Harvard University. Such was his promise that even before enrolling in graduate school, Tim was dispatched by Schultes to the Amazon on an expedition that would define the course of his professional life. In the fall of 1966 Tim returned from Brazil flush with excitement and fully committed to spending the rest of his life in pursuit of the mysteries of the tropical rain forest. His dissertation, with its thorough bibliographic research, precise and deliberate taxonomic and ethnobotanical insights, elegant prose, and exqui- site line drawings, set the standard of excellence by which he would operate throughout his pro- fessional career. By the time his Ph.D. was offi- cially conferred, in 1974, Tim was already deeply involved in the project for which he will always be remembered, a 15 -year effort to decipher the complex taxonomy of Erythroxylum and to study the ethnobotany of coca, the sacred leaf of the Andes and the notorious source of cocaine. Of Tim's 80 published scientific papers, 46 are relat- ed to his work on Erythroxylum, and his position as the world's authority on the genus enabled him to speak eloquently and powerfully in defense of the traditional use of coca by beleaguered indig- enous peoples of the Andes and northwest Ama- zon. In 1978 Tim left Harvard for the Field Museum of Natural History, where he became tenured in 1983 and was appointed curator in 1988. If Tim grew up at the Botanical Museum at Harvard, he came into his own at the Field Museum, and his years there were both the happiest and most pro- ductive of his remarkable career. His interdisci- plinary interests in systematics, ethnobotany, and ethnopharmacology led him to interact with an in- creasingly diverse group of scholars that included not only fellow botanists but also archaeologists, phytochemists, ethnographers, and pharmacolo- gists. In addition to carrying out an active scien- tific research program as co-principal investigator of the National Science Foundations "Projeto Flo- ra Amazonica," he served on the editorial boards of numerous journals, including Flora Neotropi- cal Monographs, Advances in Economic Botany, Journal of Psychoactive Drugs, and Journal of Ethnopharmacology. Between 1984 and 1988 he was co-editor-in-chief of the Journal of Ethno- pharmacology and the scientific editor of Fieldi- ana. He was vice president of the Beneficial Plant Research Association, a fellow of the Linnean So- ciety, and a member of many professional soci- eties, including the American Society of Plant Taxonomists, the Society of Economic Botany, the Council of Biology Editors, the Society of Ethnobiology, and the New England Botanical Club. As chairman of the botany department of the Field Museum of Natural History (1986- 1988), Tim secured a substantial increase in Na- Vlll tional Science Foundation funding for the herbar- ium and developed a new facility for the curation of economic collections. His enthusiasm, spirit of cooperation, professional rigor, and passionate commitment to botany proved infectious, and un- der his leadership, morale within the botany de- partment soared. Credentials alone, however, present but a shad- ow of the man who affected so many lives in such profound ways. For Tim, life was but a vehicle for seeking understanding and for expressing free- dom. If there is a word to describe Timothy Plow- man it would be freedom, and he lived with the conviction that every person had the right to pur- sue his or her own path unshackled by the burdens of social convention. Equally at ease in the tran- quil world of plants and the society of people, Tim was charismatic, and those privileged to have spent time with him often developed a respect that bordered on reverence, for he was a true renais- sance scholar, a man out of time, whose breadth of interests and passions went far beyond the boundaries of his beloved field of botany. But it is as a botanist and intrepid plant explorer that Tim will be best remembered. He spent more than five years of his life in the most remote and inhospitable regions of the Andes and Amazon, making more than 15,000 collections of unsur- passed quality. He always considered his time in the field a privilege, and he never failed to re- member his fellow botanists toiling away in the less romantic confines of the herbaria. Tim seemed to have a Rolodex in his head that re- corded the name of every specialist in every group of plants, and he was constantly on the lookout for specimens that might prove useful to a distant colleague. He collected everything. His voucher specimens were not only complete but aesthetically beautiful, and whenever possible he augmented them with invaluable collections of live material. Living plants, many new to science and collected first by Tim, may be found in bo- tanical gardens throughout the world. In the rain forests of the Amazon Tim felt the fullness of life. He marveled at the thousand themes, the infinitude of form, shape, and texture that so clearly mocked the terminology of tem- perate botany. He always traveled in the forest as a student, and his commitment to ethnobotany grew in part from his direct experience with the indigenous peoples who understood the plants in ways that he believed he could only hope to em- ulate. To be in the forest, he said, was to be in Eden, and to say the names of the plants was to recite the names of the gods. He believed that all forms of life were manifestations of the sacred. Hence, for Tim, biological and cultural diversity represented far more than the foundation of sta- bility; they were articles of faith, fundamental truths that indicated the way things were supposed to be. Tim had a special affinity for Indians, and his uncanny ability to gain their trust and confidence was one measure of the deep respect he had for their way of life. He empathized with their world view, which defined man as but one element in- extricably linked to the whole of creation. It was this unique cosmological perspective, he believed, that enabled the Indians to comprehend implicitly the intricate ecological balance of the forest he loved so dearly. Tim viewed with pain, dismay, and increasing anger the other world view, in which man stands apart, that now threatens the forest with devastation. It was one of his fondest hopes that the lessons of ethnobotany might ulti- mately facilitate a dialogue between these two world views such that folk wisdom might temper and guide the inevitable development processes that today ride roughshod over much of Earth. The many of us who loved him as a brother and respected him as a colleague can do no better ser- vice to his memory than continue our own strug- gles to make his dream a reality. — Wade Davis IX Editors' Preface Tim Plowman not only was a "botanist's bot- anist," he also inspired a whole generation of young taxonomists with his enthusiasm for plants in both the field and herbarium. The study of the systematics of the South American species of Brunfelsia was the subject of his Ph.D. research while he was at Harvard Uni- versity. After receiving his Ph.D. in 1974, Tim worked mainly on Erythroxylum, but Brunfelsia remained an area of special interest for him. He was preparing a major revision of the group, using his dissertation as a basis and backing up with further study and extensive field work, when he died in 1989. In 1994 Tim's colleagues at the Field Museum of Natural History in Chicago sent the manu- script, together with various notes and archival material, to one of us (S.K.) to assess whether the work could be brought to completion. We are very pleased to have been able to do this, both because it will make available this extensive study of an interesting group of Solanaceae and because it will serve as a tribute to the work of a well-liked and respected colleague. In taking the working manuscript and bringing it to publication, we tried throughout to keep in mind Tim Plowman's concept of the group and changed his work as little as possible. Most of the manuscript remains essentially unaltered, but where necessary we rearranged some sections and augmented others in line with Tim's own publi- cations, notes, and records. The only major de- parture from this approach is where we felt a need to update the manuscript in the light of works published since Dr. Plowman's death. Such up- dates usually take the form of explanatory notes in the text that are clearly indicated as being ed- itors' contributions. To this end, the key was revised to incorporate additional species published (by Plowman) since the original manuscript was written. The order of specimen citations is Plowman's. In some places reasoning is not clear, but we chose not to change the order. Specimens not cited in the original work but seen by Dr. Plowman and recorded in his notes and in the herbarium at the Field Museum were added; they are marked with an asterisk and are inserted in order of collector at the end of each state or province. These later records are included in the distribution maps. Infraspecific taxa are not indicated on the maps because Plowman's later annotations were only to specific rank and could not be integrated with distributions of infraspecif- ic taxa. Additional references, some of them Plowman's own subsequent publications, were in- serted; like the additional specimens, these are marked in the bibliography with an asterisk. Several new combinations remained unpub- lished at the time of Tim's death and are validated here. The authority for these combinations should be simply Plowman, as they represent Tim's as- sessment of the status and rank of the taxa and should be attributed solely to him. The additional bibliography in Appendix II lists references to publications that postdate the origi- nal manuscript and that we believe will provide a better understanding of the current state of knowl- edge in the genus. This revision of Brunfelsia is the product of Tim Plowman's research, but we accept sole re- sponsibility for any errors of interpretation or pre- sentation. — S. Knapp J. R. Press XI Opposite: Brunfelsia hydrangeiformis subsp. hydrangeiformis (originally as Fran- ciscea hydrangeiformis), illustration of type. Illustration reproduced from J. E. Pohl, Plantarum Brasiliae Icones et Descriptiones (1826). A Revision of the South American Species of Brunfelsia (Solanaceae) Timothy C. Plowmanf S. Knapp and J. R. Press, Editors I. Materials and Methods Beginning in the fall of 1968, I began studying the genus Brunfelsia L. in the field, both in South America and in the West Indies. During trips varying in duration from a few weeks to several months, I studied many of the species of Brun- felsia in their natural habitats, giving special at- tention to population variability and ecological conditions. At this time I collected extensive ma- terial for morphological, cytological, and chemi- cal studies, including numerous herbarium speci- mens and living material in the form of seeds and cuttings for propagation in the greenhouse. I also took this opportunity to investigate the medicinal properties and uses of the plants among native people who still possess an intimate knowledge of plant medicines. I learned about these aspects of the plants not only by inquiring among native healers but also by taking the medicines myself (Plowman, 1977). Many of the species I had hoped to see in the field could not be found in localities where they were formerly collected, due to both the sporadic occurrence of the plants and the destruction of their habitats by man. At other times, the plants were found but without much sought-after flowers or fruits. To acquire a historical perspective of the genus, to properly typify specific names, and to view as much morphological variation as possible, I bor- rowed herbarium specimens from many institu- tions in Europe, South America, and the United States. These specimens, numbering about 3,000 [the number at the time of his thesis in 1974, al- though many more subsequently were examined by Plowman — Eds.}, provided the basis for the study. This material was organized according to the recommendations outlined by Leenhouts's (1968) A Guide to the Practice of Herbarium Tax- onomy, which offered many helpful suggestions. Some 55 morphological characters were ob- served and measured or scored on a character flow sheet. For many specimens it was not pos- sible to record all the characters, especially those of the fruits, which are rarely, if ever, collected, and those of the corolla, which is usually poorly pressed and often glued fast to the herbarium sheet, precluding dissection of the stamen and pis- til. The characters and their variation were ana- lyzed according to methods outlined and dis- cussed in Benson (1962), Davis and Hey wood (1963), Lawrence (1965), and Anderson (1967), among others. Descriptions of the various taxa were prepared using characters observed in her- barium specimens and in living plants. In many cases the color of the flowers and other evanes- cent characters were not known with certainty; as a result, the completeness of the description varies somewhat from species to species. Pollen was acetolyzed according to the proce- dure of Erdtman (1960). Unacetolyzed pollen grains were also examined with the aid of a scan- ning electron microscope. Cytological material was collected in the field and in the greenhouse and fixed in Carney's solution (absolute alcohol: glacial acetic acid, 3:1), then stored in 95% al- cohol. Chromosomes were stained with acetocar- mine. Bulk material for chemical analysis was col- lected in the field and dried in the open sun or in special drying ovens. This material was sent to Mr. Jan-Erik Lindgren at the Department of Tox- icology, Karolinska Institute!, Stockholm, Swe- den, and to Dr. John Leary, Massachusetts Col- lege of Pharmacy, Boston, Massachusetts. Ex- tracts of these plants were tested for alkaloids and general constituents using precipitation reactions and gas and thin-layer chromatography [see Plow- man, 1977; Lloyd et al., 1985; Schultes & Raf- fauf, 1990, 1991— Eds.]. In addition, simple tests for alkaloids were performed by Prof. Robert F. Raffauf and myself on fresh plants growing in the greenhouse. Extracts were prepared from fresh FIELDIANA: BOTANY, N.S., NO. 39, AUGUST 31, 1998, PP. 1-135 material of five species and tested for alkaloids by precipitation reactions. Sixteen species were grown in the greenhouses of the Biological Laboratories, Cambridge, Mas- sachusetts; the Arnold Arboretum, Jamaica Plain; and the Botanical Research Station of Northeast- ern University in Woburn, Massachusetts. [Other specimens were cultivated at the Field Museum of Natural History, Chicago, Illinois, in later years. — Eds.] Material was propagated from both seeds and cuttings collected in the wild or from cultivated plants. Crosses were attempted between as many species as possible. The living plants were also a valuable source of those morpholog- ical characters that are lost in dried specimens and permitted observations to be made on the growth and development of various organs, particularly the inflorescence. A considerable amount of time in this study was devoted to searching the botanical, chemical, and pharmacological literature for references to Brunfelsia. Only those works that are cited in the text are included in the bibliography. All of the specimens cited in this work were examined by me unless they are denoted by "non vidi." The use of brackets in the citation of a col- lection number indicates that some uncertainty ex- ists or, in the case of type materials, that the num- ber appearing on the sheet was not given in the original citation of the specimen. Finally, the use of taxonomic characters de- serves some explanation. Of the 24 species of Brunfelsia that are recognized in South America, four are considered to possess one variety each in addition to the typical variety. The category of subspecies is here used for geographical or eco- logical races that show morphological differenti- ation from the typical form of the species. It should be noted, however, that the degree of dif- ferentiation and isolation varies from one species to another, so that the subspecific concepts are not entirely equivalent. Furthermore, variants exhib- iting considerable differentiation exist within sev- eral subspecies (e.g., B. grandiflora subsp. schul- tesii Plowman, B. brasiliensis (Spreng.) L.B. Sm. & Downs subsp. brasiliensis). Subsequent work- ers in the genus may choose formally to recognize these as variants at infraspecific rank. II. Taxonomic History of the Genus The genus Brunfelsia was first described by Charles Plumier in 1703 in his Catalogus Plan- tarum Americanarum, commemorating the Ger- man herbalist Otto Brunfels (1464-1534). Brun- fels was a Carthusian monk and physician who, in 1530, published an herbal, Herbarum Vivae Ei- cones, that, as the name indicates, included illus- trations of living plants. This early work is out- standing for the high quality, accuracy, and nat- uralistic aspect of the woodcuts, which for the first time departed from the often copied and stylized illustrations of earlier herbals (Arber, 1912). Un- fortunately, Plumier misspelled the genus name as "Brunsfelsia" an error that has tenaciously per- sisted in the literature [see D'Arcy, 1989 — Eds.]. Simultaneously with his short description and diagnosis, Plumier published a rather good illus- tration of the flower and fruit of the plant, which is now known as B. americana L. The illustration was taken from a larger, more complete drawing that, along with more detailed descriptions in manuscript form, is kept at the Museum National d'Histoire Naturelle in Paris, France. The com- plete plate was published with a more complete description in 1756 in Burman's edition of Plu- mier's Catalogus Plantarum Americanarum (Fig. 1). In the second edition of Genera Plantarum (1742), Linnaeus included Brunfelsia, based on Plumier's description. He classified Brunfelsia among the Pentandria Monogynia, indicating that the plant had five stamens (an aberrant condition that occasionally occurs in B. americana). It would thus seem that Linnaeus had access to specimens since Plumier failed to mention this character and it is not indicated in his drawing. Although there is no known specimen of B. amer- icana in extant Linnaean herbaria, he did indicate with a symbol in the fifth edition of Genera Plan- tarum (1754) that he knew the genus from her- barium material. In a letter to David Van Royen in 1767, Lin- naeus mentioned that he did, in fact, see a speci- men of the Brunfelsia of Plumier: " Brunsfelsiam Plumieri vidi exsiccatam, at ilia forte a Tua specie diversa erat, ideoque nihil certi de ea potui pron- FIG. 1. Brunfelsia americana. From Burmann's edition of Plumier's Catalogus Plantae Americanarum (1703). FIELDIANA: BOTANY TAB . LXV. PLOWMAN: REVISION OF BRUNFELSIA unciare, nee Adansonii compilationes mihi suffec- ere" (Bonnet, 1895). Van Royen had sent Linnae- us a specimen (no. 257.1, LINN) that Linnaeus recognized as being different from Plumier's spe- cies. This plant is now referable to B. undulata Swartz. The valid publication of the genus dates from 1753, when Linnaeus published B. americana in Species Plantarum, in which he followed the orig- inal misspelling of the genus. This was apparently an orthographical oversight on his part with which he was inconsistent in later works. For example, in the following year (1754), he used the correct spelling in Genera Plantarum but misspelled it in subsequent editions of Species Plantarum. In the International Code of Botanical Nomenclature (Greuter et al., 1994, p. 297), Brunfelsia is listed in Appendix IIIA as a conserved spelling [see also D'Arcy, 1989— Eds.} . The next important date in the history of the genus was 1826, when Pohl published seven new species from Brazil as the genus Franciscea, which he placed in the order Scrophularinae, close to BrowalHa. Each of Pohl's seven new species was accompanied by an excellent illustration (frontispiece, Figs. 22, 28, 31, 39, this work). Ad- ditional species were soon added to Franciscea by Chamisso and Schlechtendal (1827) and Hook- er (1828). However, as early as 1829, David Don pub- lished his observations on this group, in which he maintained that Franciscea did not differ signifi- cantly from Brunfelsia. He united the two genera under the earlier name Brunfelsia but published only one new combination, B. uniflora (Pohl) D. Don. Don also expressed the opinion, precocious for the time, that Brunfelsia belonged in the So- laneae (Solanaceae) rather than the Scrophulari- nae, an idea suggested even earlier by Antoine de Jussieu (1791). Most authors of the time ignored Don's conclu- sions and continued to recognize Brunfelsia and Franciscea as distinct genera, primarily on the ba- sis of the nature of the capsule and the length of the corolla tube (Bentham, 1835; G. Don, 1837; Endlicher, 1839; Meisner, 1840). In 1846, however, in his monograph on the Scrophulariaceae for de Candolle's Prodromus, Bentham united Franciscea with Brunfelsia, ob- serving that although the francisceas were distin- guished by some authors, especially by the dry capsule, he himself had looked in vain for defin- itive structures; B. latifolia (Pohl) Benth. and B. guianensis Benth. had small flowers like the rest of the francisceas but capsules no less fleshy than those of B. americana, and in several species the fruit was completely unknown. Bentham's treatment of Brunfelsia included 19 species, eight of which were formerly considered Franciscea. He divided the genus into three groups without rank: Multiflorae and Uniflorae (Franciscea) and Longiflorae (Brunfelsia), placed among the Salpiglossideae. This work bears the distinction of being the only revision of Brunfelsia that has attempted to encompass the entire genus. Bentham considered the Salpiglossideae to be a "suborder" (subfamily) of the Scrophulariaceae and noted their close affinity with the capsular Solanaceae, with which, he added, they might be better associated. Shortly after Bentham's revision appeared, John Miers began publishing a series of papers on the Solanaceae consisting of taxonomic treat- ments, descriptions of new genera and species, and a major reorganization of the tribes into two families, the Solanaceae and the Atropaceae. These replaced the older concept of Solanaceae and included the Salpiglossideae of Bentham. The text of Miers 's work was originally published in The Annals and Magazine of Natural History be- tween 1849 and 1857 and later appeared bound together in two volumes accompanied by 87 plates under the name Illustrations of South Amer- ican Plants (Miers, 1855, 1857). This extensive work remains a major source of information for solanaceous plants. Miers (1849) erected the new family Atropa- ceae as a distinct group with imbricate or plicate corolla estivation that was intermediate between the Solanaceae (with valvate estivation) and the Scrophulariaceae (with imbricate estivation). The group included ten tribes, the last of which he named the Brunfelsiae. Miers firmly believed that Brunfelsia and Franciscea should be maintained as distinct genera, citing some ten differences that he considered diagnostic. Among these were the habit of the plants, the relative length and shape of the corolla, the color of the corolla, the shape of the anthers, the size and structure of the fruit, and the number of seeds per fruit. Several points of difference indicated by Miers between Brunfelsia and Franciscea no longer stand when one considers the wide range of vari- ation and many additional species that are now recognized. For example, he stated (1849, 1850) that brunfelsias are trees attaining 20 ft in height whereas francisceas are small shrubs. In fact, both low shrubs and small trees occur in about equal FIELDIANA: BOTANY numbers in both groups. Miers considered the fruit of Brunfelsia a fleshy drupe containing an indehiscent putamen and that of Franciscea a de- hiscent capsule that splits along four vertical su- tures. It is now clear that within both Brunfelsia and Franciscea, dehiscent and indehiscent, thin- and thick-walled, and fleshy and dry fruits occur. On the basis of limited material, Miers failed to recognize the overall similarities between these groups. However, several differences that he pointed out remain valid and serve to distinguish Brunfelsia and Franciscea as distinct sections of the genus. Perhaps the most disturbing fact in Miers 's study of Brunfelsia is his omission of B. guianen- sis, a species that Bentham (1846) described and commented on in the Prodromus. In Miers's oth- erwise complete list of new combinations, in which he transferred most of Bentham's new brunfelsias to Franciscea, B. guianensis is con- spicuously missing. Since B. guianensis possesses several features that are intermediate between Brunfelsia and Franciscea, it seems that Miers perhaps simply ignored it because it blurred the sharp distinction he wanted to demonstrate. The view of Bentham, to unite Franciscea with Brunfelsia, eventually prevailed over Miers's ideas, perhaps because of the wide distribution and general acceptance of de Candolle's Prodro- mus. Although some of Miers's conclusions have not gained acceptance in light of more recent knowledge, his extensive observations are none- theless valuable to any taxonomist interested in the Solanaceae. For example, Bentham and Hook- er (1873) drew heavily from Miers's data in de- scribing the Solanaceae for Genera Plantarum. There has been only one other major study of Brunfelsia since the time of Miers and Bentham. This is found in J.A. Schmidt's treatment of the Scrophulariaceae in Martius's Flora Brasiliensis in 1864. Although only Brazilian species were in- cluded, then numbering about ten, Schmidt made many sound taxonomic judgments and correctly re- duced many names to synonyms. Until the present time, this work remained the standard reference for Brunfelsia in Brazil, which possesses the largest number of species of any country in South Amer- ica. However, Schmidt's treatment is now out of date, as may be expected, and not without error. In an attempt to clarify certain nomenclatural problems regarding his new species Brunfelsia mire Monach., Monachino (1953) offered a brief discussion of the names of the South American species. This was a work of limited value and provided little help in solving the taxonomic dif- ficulties in the genus, being based primarily on the erroneous conclusions of others. Besides the original descriptions of the plants, which for certain species are the only information available, only floristic works have provided ac- counts of the species in particular areas. Foremost among these, which are by no means of uniform quality, are the following works: Flora of the Brit- ish West Indian Islands (Grisebach, 1861); Botany of Porto Rico and the Virgin Islands (Britton & Wilson, 1925); Flora de Cuba (Liogier, 1957); Flora of Peru (Macbride, 1962); Flora Ilustrada Catarinense, Solandceas (Smith & Downs, 1966); Flowering Plants of Jamaica (Adams, 1972); and Flora of Panama (D'Arcy, 1974). Table 1 summarizes the history of classification of Brunfelsia, including generic, tribal, and family placement. It should be noted under the heading "Family" that the groupings of the earlier bota- nists were not equivalent to our modern families of plants. These are included for historical pur- poses only. III. Generic Relationships Brunfelsia is a distinct genus that is not readily confused with other genera. It belongs to the tribe Salpiglossideae of the Solanaceae, as circum- scribed by Bentham (1835, 1846, under Scrophu- lariaceae), Bentham and Hooker (1873), Wettstein (1895), and Baehni (1946). This group is consid- ered the most advanced tribe of the Solanaceae1 and resembles Scrophulariaceae in several charac- ters, such as the zygomorphic corolla with some- what imbricate estivation, the number of stamens reduced to four or two, and the capsular fruits. It is distinguished from the latter by intraxylary phlo- em (Metcalfe & Chalk, 1950) and from other So- lanaceae by the estivation of the corolla and the number of stamens. The closest relatives of Brunfelsia are Browal- lia L., a weedy herbaceous genus of two or pos- 1 Although traditionally placed in the tribe Salpiglos- sideae and considered closely related to Browallia and Salpiglossis, results of recent phylogenetic studies using chloroplast DNA (Olmstead & Palmer, 1992; Olmstead & Sweere, 1994; Olmstead et al., in press) indicate that Brunfelsia may be better placed in a monophyletic group with the genera Petunia, Fabiana, and Nierembergia. The morphological evidence remains to be assessed. — Eds. PLOWMAN: REVISION OF BRUNFELSIA TABLE 1 . Summary of historical classification of Brunfelsia. Author "Family" Tribe Genus Linnaeus (1753) Pentandria Brunfelsia Monogynia Swartz (1788) Didynamia Brunfelsia Angiospermae Jussieu (1791) Personatae Brunfelsia Pohl (1826) Scrophularineae Franciscea (excluding Brunfelsia) D. Don (1829) Solaneae Brunfelsia (including Franciscea) Bentham (1835) Scrophularineae Salpiglossideae Brunfelsia Franciscea G. Don (1837) Solanaceae Francisceae Brunfelsia Franciscea Endlicher (1839) Scrophularineae Salpiglossideae Brunfelsia Franciscea Bentham (1846) Scrophulariaceae Salpiglossideae Brunfelsia (including Franciscea) Miers (1849) Atropaceae Brunfelsiae Brunfelsia Franciscea Bentham and Hooker (1873) Solanaceae Salpiglossideae Brunfelsia (including Franciscea) Baillon (1888) Scrophulariaceae Salpiglossideae Brunfelsia (including Franciscea) Wettstein (1895) Solanaceae Salpiglossideae Brunfelsia (including Franciscea) Baehni (1946) Solanaceae Salpiglossideae Brunfelsia (including Franciscea) Hutchinson (1969) Salpiglossidaceae Brunfelsia (including Franciscea) Plowman (1973) Solanaceae Salpiglossideae Brunfelsia (including Franciscea) sibly three heteromorphic species, and Streptoso- len Miers, a monotypic Andean shrub genus. [Plowmania Hunz., a monotypic genus from mon- tane Guatemala, first described as a species of Brunfelsia, is also considered to be closely related to Brunfelsia (see Hunziker & Subils, 1986). — Eds.] Browallia is distinguished from Brunfelsia by its herbaceous habit; by the filaments, which are greatly dilated and ciliate at the apex; by the broad, lamellate stigma, which has recurved mar- gins, and by the dry capsule, which is dehiscent by four sutures. Streptosolen differs from Brun- felsia in having the corolla tube twisted at the base, the limb bright red-orange, and the filaments thickened at the apex. This shrub grows at high altitudes in the Andes from Colombia to Peru and is pollinated by hummingbirds. The other genera of the Salpiglossideae are pri- marily annuals found in southern South America: Salpiglossis Ruiz and Pavon, Schizanthus Ruiz and Pavon, Leptoglossis Benth., Reyesia Clos, Pe- tunia Juss., and Schwenkia L. (which also occurs in Africa). In addition, three genera of Australian shrubs are also placed in the tribe: Duboisia R. Br, with indehiscent, baccate fruits, and Antho- cercis Labill. and Anthotroche Endl., with dehis- cent, capsular fruits.2 2 Since the time of the dissertation, these genera have been segregated as the endemic tribe Anthocercideae (Haegi, 1981). and additional genera have been de- scribed.— Eds. Hutchinson (1969) elevated the Salpiglossideae to the rank of family (Salpiglossidaceae) with the explanation that the genera of this group consis- tently arrived next to Scrophulariaceae in his key to the families of plants of the world. This deci- sion does not seem justified in view of the exten- sive evidence from morphology, anatomy, and chemistry, which links the Salpiglossideae with the Solanaceae. The origin and relationships of the Salpiglos- sideae remain obscure. Baehni (1946) derived the group from two sources, the Solaneae via the sub- tribe Goetzeineae and the Atropeae via Para- bouchetia Baillon. In the past, relationships at the generic and tribal levels in the Solanaceae were studied using relatively few characters, such as the curvature of the embryo, corolla estivation, number of anthers, or type of fruit. A definitive and in-depth evaluation of these higher relation- ships is needed in which many characters are con- sidered simultaneously, including those from mor- phology, chemistry, anatomy, cytology, and other areas [see footnote 1 — Eds.} IV. General Morphology Habit All known species of Brunfelsia are woody shrubs, a few of which will become small trees. FIELDIANA: BOTANY No herbaceous members are known. In section Brunfelsia, certain species may reach 10-12 m in height (B. jamaicensis (Benth.) Griseb., B. amer- icana, B. lactea Krug & Urb.). These are some- what atypical, however, and most are only 2-3 m. Species of sections Franciscea and Guianenses are mostly medium-sized shrubs 1-3 m tall that form a component of the understory of the forest. Brunfelsia grandiflora D. Don of the western Am- azon may become a small tree 4-5 m tall or, in the Brazilian state of Acre, may become lianoid in habit, attaining up to 10 m. Brunfelsia latifolia is the most diminutive species, often flowering when only 30 cm tall and rarely reaching 1 m in height. Root The roots of Brunfelsia are diffuse in the seed- lings and young plants. However, mature plants often develop a deep, woody taproot. This is es- pecially true in species of section Brunfelsia, which often grow in rocky areas. In section Fran- ciscea, the central root may be more or less branched (Fig. 2). Branching Species of Brunfelsia may have one central trunk (B. densifolia Krug & Urb., B. mire, B. chir- icaspi Plowman) or they may be branched to a greater or lesser extent from near the base, at times producing two or more main trunks (B. aus- tralis Benth., B. grandiflora, B. brasiliensis). In some cultivated species, root suckers often appear around the main trunk and form extensive clonal plants (B. nitida Benth., B. undulata Sw., B. aus- tralis). In section Brunfelsia, the branches are often few and subvirgate, with few leaves at the end of a long, spindly branch (B. undulata, B. splendida Urb.). In sections Franciscea and Guianenses, the branches are more often diffuse or spreading and tend to arch over with age. The branching pattern in Brunfelsia is sympo- dial. When the terminal bud ceases to grow, the shoot is perpetuated from one or more subtermi- nal axillary buds or, rarely from the bracts of the inflorescence. In the new shoot, several prophylls are produced, followed by a gradual transition to mature leaves. Leaf The leaves of Brunfelsia are alternately ar- ranged with 5/13 phyllotaxy. They may be scat- tered along the branchlets or crowded toward the apex, rarely subverticillate through periodic short- ening of the internodes (B. mire, B. hydrangeifor- mis (Pohl) Benth., B. rupestris Plowman). The petiole is very short in relation to the leaf blade, varying from 1 to 10 mm in length. It is semiter- ete in section and canaliculate, and frequently be- comes transversely cracked or corrugate with age or drying. The leaves are simple with an entire margin. There is much variation in their size and shape, which provide important characters for distin- guishing the species. In form, the blade may be elliptic to lanceolate; obovate to oblanceolate; or, rarely, spathulate, ovate, linear, or linear-lanceo- late. Certain species are noted for great variability of leaf shape, sometimes in the same individual (B. uniflora (Pohl) D. Don, B. americana, B. lac- ted). In other species (B. portoricensis Krug & Urb., B. densifolia), the leaves of seedlings and young plants differ radically from the foliage of adult plants. At the apex, the leaves may be blunt, acute, or acuminate. It is of ecological interest that the leaves of species of wet, tropical forests are often acuminate in the form of a modified "drip tip" (B. grandiflora, B. macrocarpa Plowman, B. chocoensis Plowman, B. amazonica Morton). In size, leaves may be 1-30 cm long and 0.1- 15 cm wide. Leaf size may vary considerably within a species (e.g., B. grandiflora, 6-23 cm long) and is strongly influenced by local environ- mental conditions such as available light and moisture. The leaf texture is mostly firm-membranaceous to subcoriaceous, often soft and rubbery to the touch. A few species (B. splendida Urb., B. dwyeri D'Arcy, B. obovata var. coriacea J. A. Schmidt, B. rupestris) are noted for their shiny, thick-coriaceous leaves. Coriaceous leaves also occur in certain populations of B. uniflora and B. brasiliensis that grow in unusually dry or exposed situations. In color, the leaves are light to dark green and paler green beneath. Dark purple pigmentation due to anthocyanins is present in the young flush growth of several species (B. pilosa Plowman, B. plicata Urb., B. undulata). The leaves of B. splen- dida are particularly striking, being very dark green, almost black, and very shiny. The nervation in Brunfelsia consists of the mid- PLOWMAN: REVISION OF BRUNFELSIA FIELDIANA: BOTANY rib, which is completely submerged on the adaxial surface and prominent beneath, and three to 12 pairs of pinnate, lateral nerves. Each nerve spreads out from the costa and arcuately anasto- moses with the next near the margin. The veinlets are finely reticulate and may be obscure or some- what prominulous. The number of lateral nerves may be used to separate certain species (e.g., B. martiana Plowman from B. guianensis), but the angle of divergence and degree of arching are of- ten variable. Brunfelsia hydrangeiformis is unique in having the midrib, nerves, and major veinlets sunken into a shallow depression or sulcus on the adaxial surface, creating a very uneven surface texture. Indument Most Brunfelsia species are essentially glabrous throughout, although the young leaves and flower buds may be sparsely puberulent in their earlier stages, becoming glabrous with age. In the pu- bescent species, trichomes are usually encoun- tered in the twigs, leaves, pedicel, calyx, and co- rolla tube. In section Franciscea, the indument is best developed in B. boliviano Plowman, B. bras- iliensis, B. cuneifolia J.A. Schmidt, B. hydrangei- formis, and B. pilosa, and to a much lesser extent in certain populations of B. uniflora, B. pauciflora (Cham. & Schltdl.) Benth., and B. obovata Benth. In section Brunfelsia, the following species may be pubescent: B. americana, B. sinuata A. Rich., B. membranacea Urb., and B. jamaicensis. Spe- cies of section Guianenses are entirely glabrous but occasionally have sparse glandular hairs on the petioles and pedicels (B. burchellii Plowman, B. clandestina Plowman) and ciliate or villous bract margins (B. chocoensis, B. martiana). Uniseriate trichomes are found in both section Brunfelsia and section Franciscea. They may be either glandular or nonglandular, containing from two to ten cells. Glandular trichomes often have a single terminal, globose cell or may have one to several biseriate, terminal cells (B. uniflora). Several types of trichomes may occur together in the same organ, or only one type may be found. In Venezuelan populations of B. uniflora, the ca- lyx and pedicel may bear short glandular tri- chomes, while the twigs and leaves may be pilose to villous with nonglandular hairs and a sharp de- marcation at the attachment of the pedicel. In section Brunfelsia (B. americana, B. sinu- ata), simple, branched trichomes also occur on the leaves. These have not been observed in section Franciscea. The presence, location, and degree of pubes- cence appear to be highly variable in some species (B. uniflora, B. pauciflora, B. americana). At- tempts have been made in the past to distinguish varieties solely (or essentially so) on the basis of the presence or absence of trichomes (B. uniflora var. pubescens (Benth.) R. E. D. Baker & N. W. Simmonds, B. americana var. pubescens Griseb., B. pauciflora var. calycina (Benth.) J. A. Schmidt, B. brasiliensis var. acuminata (Pohl) L. B. Sm. & Downs). Numerous intergradations in degree of pubescence are found in these species, and lack- ing correlation with other, more stable characters, it would be unwise to recognize these as "varie- ties." Indeed, variation can be observed within duplicates of a single collection. [This is also commonly the case in other Solanaceae (see Knapp, 1989). — Eds.] The greater development of pubescence is sometimes associated with plants growing in drier or more exposed habitats, sug- gesting that they may be environmentally induced forms or ecotypes. In any case, the unstable nature of these variants precludes their receiving formal taxonomic status. In other species, however, the type and location of trichomes, correlated with ad- ditional characters, can be useful in delimiting subspecies (B. brasiliensis, B. hydrangeiformis). In relatively few species is the indument of di- agnostic value at the specific level. Brunfelsia pi- losa is characterized by long, pilose trichomes on the twigs, leaves, and calyces. Brunfelsia bras- iliensis has a distinctive yellow-brown pubescence that is generally distributed in the plant. Brunfel- sia cuneifolia usually has a villous calyx, a lower leaf surface, and twigs, although the type collec- tion is nearly glabrous. Brunfelsia boliviana has villous-ciliate leaf margins and villous twigs. Inflorescence The inflorescence may be strictly terminal, ax- illary, or a combination of terminal and axillary in the same plant. FIG. 2. Root of Brunfelsia grandiflora subsp. schultesii. PLOWMAN. REVISION OF BRUNFELSIA The inflorescence in Brunfelsla is basically a cincinnus or some reduction or variation thereof (Danert, 1958). In Diccionario de Botdnica, Font Quer (1965) provided a succinct definition of the cincinnus as a uniparous cymose inflorescence in which the medial plane of each branchlet is placed transversely with respect to the medial plane of the subtending bract of the main axis (pseudaxis) and with the lateral branchlets alternately dis- posed to the right and left. It is a scorpioid cyme in which the various pseudaxes (branchlets) do not all fall in the same plane. The cincinnal cyme is a primitive type of inflo- rescence in the Solanaceae in that the lateral branchlets are essentially undifferentiated from the primary inflorescence axis. In view of the other- wise advanced position of Brunfelsia in the family, this is a striking feature [but see Olmstead & Palm- er, 1992— Eds.]. In Brunfelsia species with well-developed in- florescences, the cincinnus is elaborated as fol- lows: a single flower terminates the main or lat- eral axis and is subtended by one to three (rarely more) small, caducous bracts, each of which bears a bud in its axil. From one or more of these bracts arise lateral branchlets, which are usually highly reduced, each of which bears a terminal flower and usually two tiny bracts, one on each side. Each successive branchlet is formed at right angles to its subtending bract. In secondary and higher orders of branching, only one of the two bracts forms a branchlet, alternating to the right and left. This gives the older branches of the in- florescence a zigzag pattern, produced by the re- maining pedicel scars (e.g., B. latifolia, B. gran- diflord). The most common form of inflorescence in the genus is few-flowered, compact, and sessile, with the branchlets of the cincinnus very reduced. In some species, the individual branchlets are often difficult to distinguish, so condensed are the flower clusters. Two species (B. grandiflora, B. brasiliensis) are known to develop open, lax in- florescences in which the branches may be elon- gated to greater or lesser extent. In other species (B. mire, B. hydrangeiformis), many flowers are produced but without elongation of the inflores- cence branchlets, resulting in a very dense, cap- ituliform cyme. In still others, the inflorescence consists of a single flower (B. americana, B. dwyeri, B. macrocarpa, B. uniflora) or one to three flowers (B. pilosa, B. australis, B. cunei- folia, B. rupestris). Great plasticity is found in the expression of the inflorescence in a single species and is due to both environmental and genetic factors. Spe- cies that frequently produce many flowers (B. grandiflora, B. hydrangeiformis) in a well-de- veloped inflorescence may have only a few flow- ers per cluster. Other species that usually are sin- gle-flowered (B. pilosa, B. amazonicd) will oc- casionally develop two or three flowers. Under normal forest conditions, B. chiricaspi bears up to seven flowers in the inflorescence; however, one collection (Kennedy 1386), growing in full sun in a cut-over place, had more than 20 flow- ers. Similar light-related responses have been ob- served in B. grandiflora (Plowman 2535). This variability, evident to some extent in most of the species, limits the usefulness of the number of flowers per inflorescence as a taxonomic char- acter, particularly in the construction of keys. Flower The flowers of Brunfelsia species are com- posed of the following parts: a pedicel; a five- lobed calyx; a long-tubular corolla with a five- lobed limb; two pairs of stamens inserted in the upper part of the corolla tube; and a pistil, the style of which nearly equals the tube. There are major differences among the flowers of the three sections of the genus; however, within a section, the flowers are remarkably similar with only a few minor exceptions. Differences that do occur and that are important in distinguishing the spe- cies are found in the relative sizes of the flowers and their parts. The flowers are slightly zygomorphic with bi- lateral symmetry. This is observable in the calyx, which occasionally has unequal lobes; in the cur- vature of the corolla tube; in the unequal limb; and in the didynamous stamens. The stamens and style are curved toward the "back" or posterior side of the flower, in line with the uppermost co- rolla lobe, and the anthers and stigma are brought very close to that side of the tube. This construc- tion is probably advantageous in pollination be- cause the proboscis of the pollinating insect (Lep- idoptera) is most likely to descend along this part of the tube (see Fig. 3). The flowers of section Brunfelsia are very large with a long tube and a usually showy white limb that fades to yellow (Figs. 4, 5). They are ex- tremely fragrant in the evening and apparently are pollinated by night-flying moths (Sphingidae). In section Franciscea the flowers are smaller and 10 FIELDIANA: BOTANY GUIANENSES BRUNFELSIA FRANCISCEA FIG. 3. Floral structure of the sections of Brunfelsia. have a shorter tube with a purple limb and a white eye at the center. They are usually not fragrant and are pollinated by day-flying butterflies from several different families. Species of section Gui- anenses have small whitish or greenish flowers with a short tube and a relatively narrow limb. They are said to be fragrant, but the pollinators are unknown. Calyx The calyx is actinomorphic or slightly zygomor- phic, in form tubular to campanulate and often somewhat inflated. It varies in length from 3 to 35 mm. In section Brunfelsia the calyx is very short compared to the corolla tube; in sections Francis- cea and Guianenses it is usually about half as long, PLOWMAN: REVISION OF BRUNFELSIA 11 12 FIELDIANA: BOTANY although in some species (B. pauciflora, B. hy- drangeiformis) it may equal the corolla tube. The calyx teeth or lobes are triangular, ovate or lanceolate, and valvate in bud. In section Brun- felsia they are often blunt with a few ciliate tri- chomes. In sections Franciscea and Guianenses the teeth are acute or acuminate, and the ciliate margins are much reduced or obsolete. The calyx is glabrous or may be furnished with glandular (B. boliviana, B. pauciflora, B. uniflord) or nonglandular (B. pilosa, B. brasiliensis) tri- chomes. In color it is usually some light shade of green to yellowish (B. undulata, B. grandiflord) or purplish (B. pilosa, B. jamaicensis). The size and shape of the calyx, as well as its pubescence, are taxonomically important in spe- cific delimitation. In most species the calyx is te- rete, but in a few (B. obovata, B. boliviana, B. cuneifolia, B. nitidd) it is five-angled at the sinus- es and concave at the lobes. The nature of the calyx in fruit also provides useful characters. The calyx is usually persistent in fruit but scarcely accrescent. It may become thicker and firmer in texture and in section Fran- ciscea frequently bears lenticular outgrowths near the base. It may split along one or more sides to accommodate the developing fruit, persisting only at the base, or it may increase in girth to partially or completely enclose the fruit at maturity (B. hy- drangeiformis, B. pauciflora, B. brasiliensis). Brunfelsia macrocarpa is unique in the genus in having a large, truly accrescent calyx that be- comes thick and leathery, completely enclosing the fruit. Corolla The corolla differs significantly in the three sec- tions of Brunfelsia, as mentioned above (Fig. 3). In section Brunfelsia the corolla tube is very long, from 6 to 24 times as long as the calyx. It be- comes gradually dilated near the apex but never curved or constricted at the mouth (Fig. 5). The limb usually is held at an angle to the tube. The five lobes may be narrow or broad, plane or un- dulate to crenate at the margin. In color the co- rolla tube is greenish or white; the limb is usually pure white when it first opens, turning yellow with age. Brunfelsia americana and B. lactea may be streaked with purple. Anomalous species are B. cestroides A. Rich, and B. purpurea Griseb. of Cuba, the flowers of which are said to be purplish red. Section Franciscea has a much shorter tube that is only one to five times as long as the calyx. It is briefly inflated near the apex to include the anthers and stigma, then slightly constricted to form a raised, fleshy ring. The tube may be white or violet, and the limb is deep to pale violet, fad- ing to pale lavender or pure white with age. The ring or eye at the center may be white or yellow. The limb in section Franciscea may be large and varies from 1 .5 to 8 cm in diameter. The lobes are broadly rounded, overlapping at the lateral mar- gins. The corolla tube in section Guianenses is short, only two to three times as long as the calyx, and gradually dilated or inflated toward the apex. There is no marked constriction as in section Franciscea. The flowers in section Guianenses are white or greenish white and apparently do not change color with age. The corolla limb in section Guianenses is relatively small, rarely exceeding 25 mm in diameter. The lobes are quite narrow and convex; the lateral margins appear to be re- flexed and are completely folded under in pressed specimens. This is also observed in B. densifolia of section Brunfelsia (Fig. 4). The estivation of the corolla lobes in bud is a combination of quincuncial and imbricate. In quincuncial estivation, two of the lobes lie wholly to the outside, two wholly on the inside, and one lobe has one side to the inside and one to the outside. In section Brunfelsia the uppermost lobe is wholly to the inside, one of the lower lobes is wholly to the outside, and the other three are im- bricate with one side in and one side out. Only the size of the corolla and the shape of the lobes are useful as taxonomic characters. Within each section the corolla is quite similar. Characters such as color and the shape of the cen- tral eye in section Franciscea are diagnostic in the living state, but these are unfortunately lost in preparation of specimens and rendered useless to the herbarium taxonomist. Androecium The four stamens occur in two pairs in the up- per part of the corolla tube, alternating with the FIG. 4. Brunfelsia densifolia (Knapp s.n.) from Puerto Rico. PLOWMAN: REVISION OF BRUNFELSIA 13 14 FIELDIANA: BOTANY upper three corolla lobes. A fifth stamen is oc- casionally produced in Brunfelsia americana, but this is an irregular condition. The glabrous fila- ments are fused to the corolla tube for most of their length and are free for only 2-5 mm. The anterior pair is longer, exceeding the stigma, and may be briefly exserted from the tube (B. obovata, B. chocoensis); the posterior pair, attached to the "back" of the tube, is shorter than the style. Both pairs are curved at the apex to project the anthers toward the back of the tube. In section Brunfelsia the filaments may be curved somewhat in the re- gion of the stigma (B. americana, B. lactea), The anthers are medifixed to the filaments and dehisce by a single longitudinal slit. In section Brunfelsia the anthers are oblong-reniform and may appear to be slightly bilobate, although the cells are confluent. At anthesis, each pair may tend to adhere to each other, apparently from the sticky nature of the pollen. Brunfelsia densifolia is somewhat anomalous in having both pairs of anthers reaching the mouth of the tube; the ante- rior pair is much reduced in size, although it pro- duces a small amount of pollen. They are not completely sterile, as implied in Urban's original description (Krug & Urban, 1897). Slight inequal- ity in anther size is not unknown in other species (cf., B. martiand) and hardly merits the formation of a new genus as proposed by Urban (1906). In sections Franciscea and Guianenses the anthers are unilocular and more or less semicircular, ap- pearing round or reniform in outline. Pollen The pollen grains of all three sections of Brun- felsia are spheroidal.3 Those of section Brunfelsia (B. americana, B. undulata) and section Guianen- ses (B. chocoensis, B. martiand) appear triporate or possibly tricolporate. Those of section Fran- ciscea are tricolpate or tricolporoidate (B. chiri- caspi, B. grandiflora, B. latifolia, B. pilosd). They are all medium-sized grains 30-38 to 34-42 jim in diameter. The surface of the pollen was examined by means of a scanning electron microscope using 3 Pollen grains of Brunfelsia species were acetolyzed according to the procedure of Erdtman (1960). unacetolyzed grains. These revealed low-relief scrobiculate or reticulate exine patterns. Although the sculpturing appears to differ slightly from spe- cies to species, the preparations were not suffi- ciently clear to permit a detailed analysis, nor were enough species observed to permit general- ization. Gynoecium The ovary is ovoid or conical and is inserted on a short, nectar-producing disc that may be ob- scure in dried specimens or appear merely as a slight thickening at the base of the ovary. The ovary is bilocular with axile placentation, bearing numerous anatropous ovules. The thin dissepi- ments occasionally do not extend to the apex of the ovary, breaking down early in fruit formation and leaving the placenta attached only at the base. The fruit then may have only one locule. The style is filamentous and nearly equals the corolla tube in length. It may be somewhat thick- ened and curved at the apex, especially in sections Franciscea and Guianenses, or suberect, as in several species of section Brunfelsia. The stigma is usually included between the four anthers in the uppermost part of the tube. Only rarely does it project to the mouth of the tube (B. densifolia). The stigma is short and briefly bifid in sections Franciscea and Guianenses and faces the posterior wall of the corolla tube. It has the appearance of an open forceps at anthesis with the stigmatic tissue included between the lobes. After pollination, the lobes close together. In section Brunfelsia the stigma is more shallowly lobed and may appear subcapitate and convex. Fruit There has been considerable misinterpretation of the fruit of Brunfelsia from both a morpholog- ical and a phylogenetic point of view. This has occurred, I believe, because early botanists had very little material to work with and drew their conclusions from only a few specimens. The fruit varies considerably from group to group, and dif- ferent fruit types may be found in otherwise closely related species. In some cases, the nature FIG. 5. Brunfelsia lactea (Knapp s.n.) from Puerto Rico. PLOWMAN: REVISION OF BRUNFELSIA 15 of the pericarp and surface of the fruit may be important in differentiating species; fruit charac- ters should be used whenever possible. Unfortu- nately, the fruits of many species are poorly or not at all known. The fruit of Brunfelsia is essentially a capsule, not a berry as suggested by Miers ( 1 850) and oth- ers in the past. There is, however, great variation in the texture of the pericarp, which may be thin- walled and brittle, leathery, fleshy or hard and in- durated (bony). In shape it ranges from globose to ovoid, and in size, from 1 to 5 cm long. There may be two locules or only one by degeneration of the dissepiments. The surface may be smooth or rough; reticulate or areolate; and green, brown, or bright yellow or orange. The pericarp may be differentiated into three distinct layers or may be more or less fused into one. In all cases the en- docarp is a thin, somewhat cartilaginous layer that on drying becomes brittle. The outer layers of the fruit wall often separate from the endocarp when the fruit rots. The thin endocarp may then split in half to release the seeds. The uncertainty and controversy concerning the nature of the fruit in Brunfelsia (see "Ecology and Reproductive Biology") reflect the scarcity of fruiting material for many species. In reality, there is a wide spectrum of variation in the texture of the pericarp, which has led different authors to classify the fruit variously as a berry, a capsule, or a drupe. The fruits of certain species are indeed difficult to classify, demonstrating the limitations in our terminology. When the full range of forms is seen, however, it is clear that all the fruits of Brunfelsia are modifications of the capsule. Dif- ferent types of capsule are found in each of the three sections of the genus, and no type is unique to any one section. In section Brunfelsia the fruit is usually leath- ery or fleshy and brightly colored. The calyx per- sists only at the base. In B. americana and B. densifolia, the valves are fleshy and yellow and orange, respectively. The valves split apart at ma- turity in a true dehiscence (Fig. 6). Brunfelsia ja- maicensis is green at maturity, with a thick, bony pericarp. Brunfelsia maliformis Urb. has a large, fleshy to leathery fruit with a distinct reticulate- areolate surface. It apparently does not open ex- cept by rotting. (B. lactea has orange valves that rot, allowing the seeds to be shaken out. — Eds.). Some of the Cuban species have fruits that are thin-walled, indehiscent, and usually brightly col- ored (B. nitida, B. clarensis Britton & P. Wilson). These may function as a berry in dispersal by an- imals. In section Franciscea the fruit is usually a rel- atively small capsule that is green or brown and has a coriaceous pericarp that becomes thin and brittle on drying. The persistent calyx encloses the fruit, at least in part. These capsules may split open at maturity, but only sparingly. It may be that many species are completely indehiscent and their seeds are released only upon rotting on the forest floor. Two anomalous species are B. dwyeri (from Panama) and B. macrocarpa (from South America), in which the fruits have a leathery and a fleshy pericarp, respectively. The calyx of B. macrocarpa increases greatly in size and itself be- comes fleshy, completely enclosing the bright yel- low fruits. In section Guianenses we find fleshy, yellow fruits in two of the six species, with the short ca- lyx present only at the base. The fruits of B. ama- zonica and B. burchellii are thin-walled and dry at maturity; the fruit of B. martiana is as yet un- known. Seed The seeds are oblong or ovoid and often an- gular or compressed on one or more sides. They range in size from 2 to 13 mm long and in di- ameter from 1 to 7 mm. The surface texture is reticular-pitted and reddish brown in color. The surface pitting is formed by many tightly com- pressed surface outgrowths, resembling globulose hairs, that slough off as the seeds reach maturity. [See Soueges, 1907; Edmonds, 1983; Lester & Durands, 1984. These hairs are lignified thicken- ings of the lateral testal cell walls. — Eds.] Each of these hairs leaves a tiny pit in the surface. The embryo is relatively large and straight and is em- bedded in endosperm. The cotyledons are flat and ovate to elliptic. The size of the seeds may be useful as a taxonomic character, but their shape and number per capsule appear to be quite vari- able. V. Cytology The chromosomes of very few species of Brun- felsia have been investigated. Four counts for the genus are known from the literature, but these are of limited value at the specific level because no voucher herbarium specimens were prepared. 16 FIELDIANA: BOTANY FIG. 6. Fruit of Brunfelsia americana. PLOWMAN: REVISION OF BRUNFELSIA 17 TABLE 2. Chromosome counts in Brunfelsia. Species Chromosome no. Author B. americana B. americana B. australis B. calycina (= B. pauciflora) B. dwyeri B. americana B. australis B. calycina (- B. pauciflora) B. dw\eri 2n = 22 n = 11 n = 12 In = 22 2n = 22 2n = 22 n = 11 n = 12 2n = 22 2n = 22 Bhaduri (1933) Madhavadian (1968) Ratera (1944) Darlington and Janaki Ammal (1945) Davidse (1981) Bhaduri (1933) Madhavadian (1968) Ratera (1944) Darlington and Janaki Ammal (1945) Davidse (1981) These findings are summarized in Table 2. On the basis of this evidence, Darlington and Wylie (1955) hypothesized that the base number for Brunfelsia is x = 11. The mitotic chromosomes of Brunfelsia are dif- ficult to prepare because the young root tips pro- duced in flower pots tend to be hard and brittle, making squashes very difficult. Treatment in 1% HC1 softens the material somewhat, but despite this treatment, no good mitotic figures could be found in three species examined (B. australis, B. latifolia, B. pauciflora). Sufficient seeds of Brun- felsia were not available to test the possibilities of preparing squashes from the roots of young seed- lings. A second method of obtaining chromosome preparations is to examine pollen mother cells un- dergoing meiosis. Young flower buds were col- lected both in the field and from cultivated spec- imens in the greenhouse and immediately placed in Carney's solution for at least 24 hr and refrig- erated when possible. Material was collected about midday on sunny days. It was difficult to obtain a sufficient number of buds because brun- felsias generally have few flowers per inflores- cence, and these are usually in various stages of maturation; not infrequently only one flower per inflorescence per day will be available for cytol- ogy. Compounded by the fact that most species flower only sporadically and some not at all under greenhouse conditions, securing adequate material was a task. Meiosis occurs in Brunfelsia when the buds are very small (about 1.0-2.5 mm long) and well be- fore the corolla emerges from the calyx. The young anthers were teased out in 2% acetocar- mine and gently heated, then squashed with even pressure. Relatively few countable meiotic figures were obtained out of numerous attempts. The re- sults obtained in five species and one subspecies are summarized in Table 3. All counts were obtained from meiotic figures with the exception of B. pauciflora, which appar- ently represents a somatic division in a tapetal cell. The meiotic figures obtained were fixed in late anaphase I or telophase II. The chromosomes are small, from 2.0 to 5.5 (jtm (cf., Swanson, 1957), and exhibit some size variation between the different species examined. In the case of B. nitida and B. americana, the smaller size is due to the fact that only two chromatids are present (telophase II). The somatic chromosomes of B. pauciflora appear greatly enlarged. This is prob- ably the result of endomitosis, which produces a high degree of endopolyploidy within the cell. Unfortunately, no normal mitotic cells could be obtained for this species. The chromosomes obtained from these species were very compact in the stages observed, sug- gesting a high degree of contraction due to chro- mosomal coiling. No details of chromosomal structure could be ascertained from the meiotic figures. It is possible, however, to see the centro- mere in the somatic chromosomes of B. pauciflo- ra. In an extensive investigation, Gottschalk (1954) examined the structure of the pachytene chromo- somes of many species of Solanaceae, with par- ticular reference to the presence and position of heterochromatic and euchromatic areas. Because of the high degree of contraction, he was unable to obtain pachytene chromosomes in what he re- ferred to as B. macrophylla (probably B. pauciflo- ra). He did, however, observe the presence of chromocenters in the interphase nucleus, indicat- ing that the chromosomes of this genus are dif- ferentiated into euchromatic and heterochromatic regions. Similar results were found in species in 18 FIELDIANA: BOTANY £ u Si CN 1 per inflorescence 28. Leaves congested near branch tips, with strongly revolute mar- gins, 1-4 cm long. Campo rupestre, Brazil . . 23. B. rupestris 28. Leaves scattered along branches, not strongly revolute, usually >5 cm long. Forests 29. Leaves elliptic-oblong, ovate or obovate, blunt or abruptly short acuminate. Rio de Janeiro 17. B. latifolia 29. Leaves narrow-elliptic to oblong-lanceolate, apically acu- minate 30. Pedicels 6-12 mm long; calyx in fruit nearly as long as capsule. SE Brazil 9. B. bonodora 30. Pedicels 2-6 mm long; calyx in fruit half as long as capsule or less. W South America . . 14b. B. grandi- flora subsp. schultesii PLOWMAN: REVISION OF BRUNFELSIA 37 Brunfelsia sect. Guianenses Plowman in Hawkes, J. Linn. Soc. (Bot.) 76: 294. 1978. TYPE SPECIES — Brunfelsia guianensis Benth. in DC., Prodr., 10: 200. 1846. Shrubs to 2 m tall. Inflorescence terminal and axillary. Flowers small, 1-7 per inflorescence. Corolla white or greenish white, tube 2-3 times as long as calyx, open at mouth, not constricted, the lobes narrow, usually with deflexed margins. Stigma briefly bifid, in the form of a forceps. Cap- sule dry or fleshy at maturity. South America. 1. Brunfelsia amazonica C.V. Morton, Proc. Biol. Soc. Wash. 62: 151. 1949. Type: BRA- ZIL Amazonas: Manaus, Estrada do Raiz, sil- va secundaria, non inundabilis, manacd, 24 Mar. 1937, Ducke 430 (holotype, us 1693434; isotypes, A, F-902265, K, MO- 1156896, NY, R-75434, s). Figure 11. Shrub or small tree 1.5-2.5 m tall. Branches spreading, knobby. Bark buff to reddish brown, cracked in longitudinal fissures. Brant hk-ts shiny, glabrous, light, yellowish brown. Leaves scattered on branchlets, blade 6.5-12 cm long, 2.5-5 cm wide, elliptic-oblong to lanceolate, abruptly acu- minate, acumen often oblique, broadly cuneate to narrowed at base, glabrous, dark green above, pale green beneath, firmly membranaceous, lateral nerves 5-8(10), straight; petiole 2-5(8) mm long, transversely corrugate with cracked epidermis, glabrous. Inflorescence terminal or axillary, ses- sile with 1-2(5-6) flowers. Flowers white or greenish white, fragrant. Bracts 1-2 per flower, 2-6 mm long, linear to lanceolate, glabrous, ca- ducous. Pedicel curved in bud, becoming erect, 1 3-20 mm long, 1 mm in diameter, slender, thick- ened toward apex, glabrous. Calyx 8-12 mm long, 5-8 mm in diameter, tubular-campanulate, slightly inflated, obovate in bud, glabrous, green, membranaceous, teeth 2-6 mm long, erect, some- what unequal, ovate-lanceolate, apically blunt to acuminate; calyx in fruit, enclosing capsule at base. Corolla tube 20-25 mm long, 1.5-3.0 mm in diameter, twice as long as calyx, cylindric, slightly dilated toward mouth, glabrous, minutely glandular at mouth; limb 15-25 mm in diameter, spreading, lobes 6-11 mm long, subequal or the posterior lobe somewhat larger, oblong to obo- vate, somewhat convex, blunt or rounded at apex, the sides of the lobes reflexed after anthesis, nar- rowed slightly toward base. Stamens inserted at middle of tube, included; filaments subligulate, upper pair 5 mm long, lower pair 2-3 mm long; anthers 1-1.5 mm in diameter, orbicular-reniform. Ovary 1.5 mm long, 1 mm in diameter, narrowly conical; style 16-18 mm long; stigma 1 mm long, bifid, upper lobe somewhat larger. Fruit 10 mm in diameter, globose, apiculate at apex, green, smooth, pericarp thin, about 0.2 mm thick, dry at maturity, sparingly dehiscent. Seeds 8-10, 5-6 mm long, 3 mm in diameter, oblong-reniform, flattened on 1 or 2 sides, angular, dark brown, reticulate-pitted with a medial hilum. Embryo 3 mm long, slightly curved; cotyledons about 0.5 mm long, elliptic. DISTRIBUTION — Brazil (Amazonas). See Figure 15. SPECIMENS EXAMINED — BRAZIL. Amazonas: Manaus, Estrada da Raiz, 18 Mar. 1943, Ducke 430 (paratype, MG, us); perto da Estrada da Raiz, 22 Mar. 1932, Ducke sp. nov. (GH, RB); Mata do Aleixo, 4 Mar. 1945, Froes 20535 (NY, us); Uyp- iranga, Rio Negro, perto de Manaus, 21 Feb. 1923, J.G. Kuhlmann 943 (GH, RB); road Manaus (Cacau-Pireira) to Manacapuru, km 25, 3 Jan. 1967, Prance, Pena & Ramos 3880 (F, INPA, K, MG, P, R, s, us); ad oram meridionalem Rio Negro, usque ad concursum flum. Solimoes, May 1851, Spruce 1495 (EM, BR, CGE, F, G, GH, K, LE, MG, NY, p, RB, w); *Rio Negro, Tapuruquara, Santa Izabel, Black 48-2905 (IAN). This species was first collected by Spruce in the middle of the last century and distributed under the name B. spruceana to the world's herbaria. This name was recorded but not validly published by Schmidt, who placed it in synonymy with B. maritima Benth. Brunfelsia amazonica is a very local species, known only from the environs of Manaus at the mouth of the Rio Negro in Brazil. Notwithstand- ing the intensity of collecting in this region, B. amazonica has not been collected often. It is a very distinct species of section Guianenses and is easily separated from B. guianensis and B. cho- coensis by the longer pedicels, the oblong-lanceo- late leaves, and dry, thin-walled fruit, and from B. martiana by the longer pedicels and smaller leaves. Brunfelsia amazonica grows in open secondary forests on the southern bank of the Rio Negro in white sandy soil. It is known to flower from Jan- uary to May; fruits have been collected only in 38 FIELDIANA: BOTANY FIG. 1 1 . Brunfelsia amazonica. PLOWMAN: REVISION OF BRUNFELSIA 39 May. Like many brunfelsias in Brazil, this species is locally called manacd. 2. Brunfelsia burchellii Plowman, Fieldiana Bot. n. s. 8: 9. 1981. Type: BRAZIL. Goias: Porto Real (now Porto Nacional), ford of Igarape, 1828-1830, Burchell 8527 (holotype, P; iso- types, K, L).H Figure 12. Shrub. Mature branchlets 2-3 mm in diame- ter, spreading, with shiny, yellowish to dark brownish, longitudinally cracked bark. Leaves scattered along stem, short petiolate, blade 70- 180 mm long, 30-75 mm wide, broadly to nar- rowly ovate, apically acuminate with a long, pointed, often falcate acumen, sometimes abruptly acuminate, minutely ciliolate at margin, basally rounded to obtuse, glabrous on both sides, dull or somewhat shiny, medium green above, dull, paler green beneath, firmly membranaceous to charta- ceous, the lateral nerves 5-9, strongly arcuate, anastomosing near margin, nerves prominent on lower surface; petiole 1-6 mm long, with a few scattered glandular hairs, glabrescent. Inflores- cence terminal on current year's branchlets, very briefly pedunculate, with 1-3 flowers. Peduncle 14 mm long, persistent, sparsely glandular-pubes- cent, glabrescent. Bracts 1-3 per flower, 4-20 mm long, linear to lanceolate, apically acuminate, glabrous or villous at nerves and margin, cadu- cous. Flower color unknown. Pedicel 4-8 mm long, 1 mm in diameter, sparsely glandular-pubes- cent, glabrescent. Calyx 1 1-20 mm long, 5-9 mm in diameter, tubular or tubular-campanulate, te- rete, with scattered glandular hairs, glabrescent, striately nerved, teeth 2-6 mm long, unequal, tri- angular-ovate, apically acute to acuminate, mi- nutely gland-tipped. Corolla tube 24-28 mm long, 1.5-2 mm in diameter, \Vi-2 times as long as calyx, straight, glabrous or bearing few scat- tered glandular hairs, orifice 5 mm in diameter; limb 1 8-30 mm in diameter, spreading, lobes sub- equal, rounded, overlapping at the lateral margins. Stamens inserted in upper part of corolla tube; filaments 1 mm wide, straplike, the anterior pair 3.5-6 mm long, apically slightly incurved or sub- erect, the posterior pair 3-4 mm long; anthers 1 mm in diameter, orbicular-reniform. Ovary 1.5 mm long, conical-ovoid; style 20-21 mm long, broadened and incurved at apex; stigma included between pairs of anthers, 1 mm long, briefly bifid, 11 All Burchell localities are taken from his itinerary, published by Smith and Smith (1967). upper lobe slightly larger. Fruit and seed un- known. DISTRIBUTION — Basin of the upper Rio Tocan- tins (Goias State) and adjacent Maranhao State, Brazil. See Figure 15. SPECIMENS EXAMINED — BRAZIL. Goias: Porto Real, Porto Real to Igarape, 1828-1830, Burchell 8415 (L); about village, Burchell 8494 (K); at en- trance to village, Burchell 8653 (K); "about the Manga" (cf., Rio dos Mangues, where it meets the Tocantins), Burchell 8752 (CH). Maranhao: Island of Sao Luiz, Estrada do Barreto, Feb.-Mar. 1939, Froes 11620 (A, F, NY, s, us). Brunfelsia burchellii is known from only a few collections by William J. Burchell in the vicinity of Porto Real (now called Porto Nacional) on the upper Rio Tocantins 150 years ago. Unfortunately, these specimens bear no field data, and the plant has not been recollected in this region. A modern collection by Froes from Sao Luiz Island, more than 1,000 km north of Porto Nacional, is tenta- tively assigned to B. burchellii but lacks mature flowers or fruits and differs somewhat in leaf ve- nation. Brunfelsia burchellii appears to be most closely related to B. guianensis, from which it differs by the ovate, acuminate leaves, the longer tubular ca- lyx, and the broadly rounded corolla lobes. With B. guianensis, B. burchellii is provisionally placed in section Guianenses on the basis of the gradu- ally dilated and not apically constricted corolla tube. Two additional collections with ovate acumi- nate leaves appear to be related to B. burchellii but cannot be placed with certainty. One of these, Ducke s.n. (RE 18141), was collected at Bragan?a in Para and bears small, white, terminal, solitary flowers and immature leaves. The other, Sucre & da Silva 9204 (F, RB), was collected at Buriti dos Lopes, Piaui, and has rather shiny, thick leaves and immature fruits completely enclosed in per- sistent, accrescent calyces. 3. Brunfelsia chocoensis Plowman, Bot. Mus. Leafl. 23(6): 245, t. 14. 1973. Type: CO- LOMBIA. Antioquia: Chigorodo, forest just southeast of Chigorodo, 45 km S of Turbo, alt. 50 m. Shrub 2 m high. Flowers rather conspicuous, of a peculiar flat white color, like unglazed paper, 15 Apr. 1945, Haught 4563 (holotype, uc M 048365; isotypes, NY, COL 109727). Figure 13. 40 FIELDIANA: BOTANY BRUNFELSIA burchellii Plowman FIG. 1 2. Brunfelsia burchellii. Reproduced courtesy of the Field Museum of Natural History. PLOWMAN: REVISION OF BRUNFELSIA 41 BRUNFELSIA chocoensis Plowman FIG. 13. Brunfelsia chocoensis. Reproduced courtesy of the Botanical Museum of Harvard University. Shrub 2 m tall. Branches sparse, terete, some- long, 9-11 cm wide, elliptic to oblong, abruptly what knobby at nodes, naked, gray or grayish to long acuminate at apex, acumen often subfal- brown. Branchlets 3 mm in diameter, glabrous, cate, obtuse at base, glabrous, dull, dark green shiny, light grayish brown. Leaves few, usually above, pale green beneath, firmly membrana- crowded at ends of branchlets, blade 18-28 cm ceous, lateral nerves 6-7, spreading; petiole 10- 42 FIELDIANA: BOTANY 15 mm long, stout, brown. Inflorescence subses- sile, terminal, 1 -flowered, or, on previous branch- es, axillary with 1-3 flowers. Flowers white, fra- grant. Bracts 2-4 mm long, ovate to ovate-lan- ceolate, concave, ciliolate at margin, caducous. Pedicel 5-6 mm long, erect, slender, glabrous. Calyx 8-12 mm long, 3-6 mm in diameter, tu- bular-campanulate, globose in bud, glabrous, firm- ly membranaceous, veinlets prominulous, reticu- late, teeth 2-4 mm long, erect, ovate, blunt to acute; calyx in fruit to 13 mm long, persistent, dotted with lenticels. Corolla tube 22-27 mm long, 2-2.5 mm in diameter, twice as long as ca- lyx, gradually dilated toward mouth, glabrous; limb 22-25 mm in diameter, spreading from rounded mouth 5 mm in diameter, the lobes 8-10 mm long, subequal, rounded at apex, rarely emar- ginate, slightly overlapping at margin above base. Stamens included in upper third of tube; filaments 5 mm long, somewhat flattened, lower pair curved at apex, upper pair suberect, glabrous; anthers 2 mm long, orbicular-reniform, light brown. Ovary 2.5 mm long, ovoid; style 10-20 mm long, slen- der; stigma 1.5 mm long, bifid, gaping, the upper lobe somewhat larger. Fruit 2-3 cm long, 2-3 cm in diameter, globose to ovoid, smooth, yellow when ripe, mesocarp soft, fleshy, up to 1.5 mm thick, endocarp thin, cartilaginous. Seeds few, 2- 5, 8-10 mm long, 6-7 mm in diameter, ovoid- ellipsoid, angular, dark brown, reticulate-pitted. Embryo 8 mm long, straight; cotyledons 3.5 mm long, widely elliptic. DISTRIBUTION — Panama, Colombia. See Figure 15. SPECIMENS EXAMINED — PANAMA. Darien: Cerro Pirre, 11 Apr. 1967, Bristan 566 (ECON, osu), Bristan 569 (ECON, osu), * Bristan 1220 (MO), *Duke 5300 (MO), ^Gentry & Clewell 7138 (MO); *along Rio Armila, WSW of Puerto Obal- dia, Mori 6828 (MO). COLOMBIA. Choco: Hy- dro Camp No. 14, Rio Salaqui, 6 days upstream from Rio Sucio, alt. ca. 200 m, 23 May 1967, Duke 11351 (ECON); *hills below first rapids on Rio Truando, Duke 13331 (MO). Brunfelsia chocoensis is restricted to the north- ernmost part of the Choco region of Colombia in the low-lying basin of the Rio Atrato, an area bound by the Serrania de Baudo in the west and the Serrania de Abibe in the east. Its range ex- tends northward to Cerro Pirre across the Pana- manian border. Although additional collecting may reveal a larger area for the species, B. cho- coensis appears to be endemic at this low eleva- tion (50-200 m) in swampy forest. In 1945 it was reported as a common species at Chigorodo, the type locality. I failed to find the plant on a col- lecting trip there, primarily because of the large- scale destruction of the forests for agriculture and pasturage. Most closely related to B. guianensis of the Guianas and northern Brazil, this species differs in having much larger leaves with longer petioles, longer pedicels, and broader corolla lobes. The fruit of B. chocoensis is very similar to the fleshy capsules of both B. guianensis and B. macrocar- pa. Although B. chocoensis was originally thought to be related to B. macrocarpa, it now seems that the resemblance is only superficial. The flowers of B. macrocarpa are very large and purple, fading to white with age in the manner of species of section Franciscea. In addition, the ca- lyx of B. macrocarpa is accrescent and encloses the mature fruit, whereas the calyx in B. chocoen- sis is small and present only at the base of the fruit. Data from herbarium labels (Bristan 566 and 569) indicate that the fleshy, yellow fruits of this species are edible and that the flowers are fra- grant. 4. Brunfelsia clandestina Plowman, Fieldiana Bot. n.s. 8: 11. 1981. Type: BRAZIL. Bahia: Municipio Itapebi, Fazenda Lombardia, BR 101 ao lado leste. Arbusto de 3 m de altura. Flor branca. Capoeira, 12 Aug. 1971, T. S. dos Santos 1777 (holotype, CEPEC 7086; iso- type, F 1849031). Figure 14. Shrub or treelet to 6 m tall. Trunk to 8 cm in diameter. Bark on trunk and branches yellowish brown, cracking longitudinally and transversely, shedding in thin, irregular flakes. It ranch lets 1 .5- 2 mm in diameter, glabrous, grayish to dark, red- dish brown, more or less shiny, cracked longitu- dinally. Leaves short petiolate, the blade 35-120 mm long, 15-50 mm wide, elliptic to oblong-lan- ceolate, rarely lanceolate or obovate, apically acute to acuminate, the apex itself obtuse, basally acute or obtuse, glabrous or rarely sparsely pu- berulent on the costa beneath, medium green above, somewhat lighter green beneath, shiny on both surfaces, chartaceous to subcoriaceous, rare- ly coriaceous, the lateral nerves 6-8, mostly straight, forming an angle of 45-70° with midrib, anastomosing with the arcuate marginal nerve 3- 6 mm from margin; petiole 26 mm long, glabrous PLOWMAN: REVISION OF BRUNFELSIA 43 BRUNFELSIA clandestiiia Plowman FIELDIANA: BOTANY or with scattered glandular hairs. Inflorescence terminal or subterminal on mature or newly formed twigs of current season, with or without a short peduncle, 1- or 2-flowered. Bracts 0-3, 2- 6 mm long, linear or cymbiform, truncate at apex, sparsely pubescent or glandular-pubescent, cadu- cous. Flowers white. Pedicel 3-9 mm long, 1 mm in diameter, glabrous, becoming thicker in fruit, to 3 mm in diameter, warty-lenticellate. Calyx 8- 16 mm long, 3-7 mm in diameter, tubular or tu- bular-campanulate, terete, glabrous, light green, membranaceous, teeth 2-5 mm long, subequal, triangular-ovate, apically acute or acuminate, the apex itself blunt and minutely glandular-papillose; calyx in fruit persistent, 11-16 mm long, shiny, coriaceous, striately nerved, tightly enclosing ba- sal half of capsule. Corolla 18-25 mm long, 1-3 mm in diameter, tube 1.5-2 times as long as ca- lyx, straight, glabrous, rarely with a few glandular hairs, orifice 3-5 mm across; limb 15-26 mm in diameter, spreading, inclined, lobes 6-10 mm long, subequal, expanding somewhat with age, broadly obovate to rounded. Stamens inserted in upper part of corolla tube; filaments 0.6-1 mm wide, strap-shaped, the anterior pair 3-5 mm long, incurved at apex, included, the posterior pair 1 .5- 3 mm long; anthers 1 mm in diameter, orbicular- reniform. Ovary 1-1.5 mm long, 0.8-1.2 mm in diameter, conical-ovoid, with about 10-12 ovules per locule; style 16-20 mm long, incurved at apex; stigma 1-1.5 mm long, briefly bifid, upper lobe slightly larger. Capsule dry at maturity, 13- 20 mm long, 13-15 mm in diameter, ovoid to subglobose, apiculate, smooth, shiny, dark green at maturity; pericarp thin, 0.5-1 mm thick, crus- taceous. Seeds ca 9-15, 5-7 mm long, 2.5-3 mm in diameter, oblong-reniform, terete, or somewhat flattened on one side, dark brown, reticulate-pit- ted. Embryo 3-6 mm long, straight; cotyledons 1-2 mm long, ovate to elliptic, radicle 2-4 mm long. DISTRIBUTION — States of Bahia and Espirito Santo, Brazil. See Figure 15. SPECIMENS EXAMINED — BRAZIL. Bahia: "Prov. Jacobina," 1841, Blanchet 3354 (G, LE, w), 1843, Blanchet s.n. (c, G, w); "Igreja Velha," 1841, Blanchet 3352 (BM, BR, G, P); Serra de Sin- cora, Brejao a Iracema, 17 Feb. 1943, Froes 20210 (IAN, NY, us); Municipio Jaguaquara, Ja- guaquara a Apuerama, 4 Oct. 1972, Pinheiro 1980 (CEPEC, F, NY); Municipio Itacare, Itacare-Ubai- taba, 14 Apr. 1980, T.S. dos Santos 689 (CEPEC, F); Rodovia BA 654, km 6 ao oeste de Itacare, approx. 14°18'S, 39°02'W, ca. 60 m, 12 Apr. 1980, Plowman, Mattos Silva & dos Santos 10066 (CEPEC, F), Plowman, Mattos Silva & dos Santos 10087 (CEPEC, F); Municipio Itambe, Itambe, 24 Nov. 1942, Froes 20067 (IAN, NY); Municipio Ita- ja do Colonia, 12 km da estrada em dire9ao a Feirinha ao lado oeste, margem esquerda do Rio Corro, 23 Oct. 1969, T.S. dos Santos 433 (CEPEC, F); Municipio Belmonte, Estaqao Experimental Gregorio Bondar, km 58 da rodovia Belmonte/Ita- pebi, 16 May 1979, Mattos Silva et al. 357 (CE- PEC); Municipio Santa Cruz de Cabralia, Reserva Biologica do Pau-Brasil, 18 Sep. 1971, T.S. dos Santos 1964 (CEPEC, F); cerca de 16 km a oeste de Porto Seguro, 21 Mar. 1978, Mori et al. 9775 (CEPEC, F); antiga rodovia que liga a Esta9&o Ecol- ogica de Pau-Brasil a Santa Cruz, 5-7 km ao NE da Esta?ao, ca. 12 km ao NW de Porto Seguro, 16°23'S, 39°8'W, ca 80-100 m, 5 Jul. 1979, Mori et al. 12082 (CEPEC, F, us); Municipio Guaratinga, rodovia Guaratinga/Sao Paulinho, km 25, 2 Apr. 1973, Pinheiro 2086 (CEPEC, F); without locality, 1857, Blanchet s.n. (G, L, LE); without locality or date, Blanchet 1455 (F), Blanchet s.n. (F, MG, NY); *Municipio de Cairu; Estrada Cairu-Itabera, 8 km S of Cairu, Carvalho & Gatti 795 (F). Espirito Santo: Linhares, Vale do Rio Doce, km 6 da ro- dovia BR 101, lado sul, 30 Sep. 1971, T.S. dos Santos 2015 (CEPEC, F, NY). Brunfelsia clandestina was first collected by the Swiss collector Blanchet in the state of Bahia more than 100 years ago. Following earlier au- thors, I originally assigned Blanchet's several col- lections of the species to B. uniflora, although I pointed out certain differences between typical B. uniflora and the Blanchet material (Plowman, 1973). These early collections lacked adequate field data and fruiting material [see Plowman, 1981— Eds.}. Brunfelsia clandestina superficially resembles several other brunfelsias and may be confused with them. Complete specimens, including field data and flower color, are essential for making positive identifications. Brunfelsia clandestina ap- pears to be most closely related to B. martiana, a FIG. 14. Brunfelsia clandestina. Reproduced courtesy of the Field Museum of Natural History. PLOWMAN: REVISION OF BRUNFELSIA 45 FIG. 15. Distribution of species in section Guianenses: Brunfelsia amazonica (open square), B. burchellii (open triangle), B. chocoensis (open circle), B. clandestina (solid triangle), B. guianensis (solid square), and B. martiana (solid circle). species that also grows in the moist forests of Ba- hia. Brunfelsia clandestina differs in having much smaller leaves with fewer lateral nerves and one- or two-flowered inflorescences that are borne ter- minally. In dried specimens, B. clandestina may also be confused with B. uniflora, a species known from Bah fa but belonging to a different section of the genus (section Francisced). Brun- felsia clandestina differs mainly in having dark brown or reddish brown branchlets, glabrous or nearly glabrous leaves and twigs, and a tubular- campanulate rather than narrowly tubular calyx. The flowers of B. clandestina are white; those of B. uniflora are violet, fading to white with age. Brunfelsia clandestina grows in the moist coastal forests of southern Bah fa and Espirito Santo. Earlier collectors in Bahia also found this species further inland in areas formerly covered with mesophytic forest. Most of these areas have now been converted to pastures (Mori & Mattos Silva, 1979). The last collection in the drier, in- terior part of Bahia was made in 1943. 5. Brunfelsia guianensis Benth. in DC., Prodr., 10:200. 1846. Lectotype (designated here): SURINAM. Without locality or date, Host- mann 1278 (lectotype, K; isolectotypes, G, s, w). Lectoparatype: FRENCH GUIANA. Cay- enne, 1820, Perrottet s.n. (G, G-DC, non vidi; photograph, NY). Figure 16. Shrub to treelet 1-3 m tall. Branches lax, spreading. Branchlets terete, becoming knobby at nodes, glabrous, rarely pubescent, epidermis split- 46 FIELDIANA: BOTANY BRUNFELSIA guianensis Benth FIG. 16. Brunfelsia guianensis. Reproduced courtesy of the Linnean Society, London. (First published in Plowman, 1978: Botanical Journal of the Linnean Society, 76: 294-295.) ting in longitudinal cracks, eventually shedding in glabrous, dull, dark green above, paler green be- flakes, light to dark brown. Leaves with blade 6- neath, firmly membranaceous, lateral nerves 4-5, 15 cm long, 2-6.5 cm wide, obovate to elliptic, arcuate; petiole 3-8 mm long, slender, glabrous, rarely lanceolate, abruptly acuminate at apex, acu- rarely pubescent. Inflorescence axillary or ter- men often subfalcate, cuneate to narrowed at base, minal, with 1-2 flowers. Flowers white, odorless. PLOWMAN: REVISION OF BRUNFELSIA 47 Bracts 1-2 per flower, 2-10 mm long, linear-lan- ceolate or squamiform, glabrous, caducous. Ped- icel 2-5 mm long, slender, short, glabrous. Calyx 7-10 mm long, 4-7 mm in diameter, ovoid-cam- panulate, inflated, rarely tubular-campanulate, ovoid in bud, glabrous, green, membranaceous, sparsely veined, teeth usually unequal, two pos- terior teeth somewhat shorter, 2-5 mm long, ovate, acute to acuminate at apex; calyx in fruit persistent, becoming somewhat coriaceous, split- ting on 1 or more sides. Corolla tube 20-26 mm long, 1-1.5 mm in diameter, 2.5-3 times as long as calyx, slender, straight or curved, gradually di- lated above middle, glabrous, greenish white to white, limb 15-22 mm in diameter, spreading, lobes 6-8 mm long, subequal, oblong-obovate to spathulate, narrowed at base, rounded or blunt at apex, lateral margins strongly reflexed, white, oc- casionally with traces of purple. Stamens includ- ed within tube; filaments slender, suberect, ligu- late, anterior pair 4-5 mm long, posterior pair 2- 4 mm long; anthers 1 mm in diameter, orbicular- reniform. Ovary 1.5-2 mm long, 1 mm in di- ameter; ovoid-conical, style slender, broadened to- ward apex; stigma included between anthers, brief- ly bifid, upper lobe slightly larger. Fruit 2.5-3.2 cm in diameter, 2.5-4 cm long, globose to ovoid, smooth, yellow when ripe, exocarp thin, rubbery, mesocarp soft, fleshy, 5-8 mm thick, endocarp thin, cartilaginous. Seeds 7-10, 10-13 mm long, 5-7 mm in diameter, oblong, flattened on 1 or 2 sides, angular, reddish brown to nearly black, re- ticulate-pitted. Embryo 8 mm long, straight; cot- yledons 2 mm long, elliptic. DISTRIBUTION — Surinam, French Guiana, Bra- zil. See Figure 15. SPECIMENS EXAMINED — SURINAM. In monti- bus Nassau, in forest near km 0.1, 17 Feb. 1949, Lanjouw & Lindeman 2137 (K, NY); near km 6, 7, 10 Mar. 1949, Lanjouw & Lindeman 2564 (K, NY); along creek with large rapids, 24 Mar. 1949, Lan- jouw & Lindeman 2888 (K, NY); Saramacca River, Jacob Kondre, 19 Jun. 1944, Maguire 23894 (NY); Jodensavanne-Mapane creek area; Dist. Broko- pondo, N of Brokopondo, 55°W, 5°N, 14 Feb. 1966, van Donselaar 3114 (NY); ad flum. Paloe- meu et Tapanahoni confl., 3°20'N, 55°27'W, 10 Apr. 1963, Wessels Boer 1195 (NY, us); without locality, Sep. 1853, Wullschlagel 1505 (w); *Brownsberg Nature Park; 90 km S of Paramar- ibo, Mori, Bolten 8394 (MO); *Lely Mountains; 175 kms SSE of Paramaribo, Mori, Bolten 8575 (MO); *near airport, road Oelemari, Boer 1047 (u); *Mapanegebied, Elburg 9819 (u); *prope Joden- savanne, Heyligers 438 (u); *Zuid River near Kayser Airstrip, Irwin et al. 55978 (u); *Forest Reserve Zanderij, Lanjouw 345 (GH); *Ribanau E of Moengo, Lindeman 5889 (u), *Lindeman 6069 (u); *S of Moengo tapoe, Lindeman 6162 (u); *W slopes of Bakhuis Mountains, Florschutz & Maas 2924 (GH, u), *Maas 3094 (GH, u); Blakawatra, Schulz 8645 (u). FRENCH GUIANA. Mathoury, Feb. 1901, Lemme s.n. (p); Karouany, 1855, Sagot 423 (s, w); without locality, 1821, Perrottet s.n. (G), 1821, Poiteau s.n. (G); *above Taruni Creek, Forestry Department of British Guiana 7495 (NY); *Tumuc-Humac, Frontiere Bresil Guyane, Gran- ville 1378 (CAY); *N de Degrad Claude, Granville 2204 (GH); *Chemin de Smerillons, a 5 km de Degrad Claude, Granville 2224 (CAY); *Basse Ouaqui, au niveau de "Degrad Roche," Granville B-4854 (CAY); *Saul, Monts La Fumee, Mori & Boom 14726 (F), *Mori & Boom 14834 (F), *Mori et al. 8767 (MO); *Haute Approuague, au conflu- ent de la crique Parepou, Oldeman B-1896 (CAY); *region of Tumuc-Humac, Massif du Mitaraka, Sastre 1649 (CAY, p); *Haut Oyapock, Peola 20 km, en aval de Trois Sauts, Sastre 4751 (P). BRA- ZIL. Amapa: Rio Oiapoque, islands of first cach- oeira on Rio laue, 2°53'N, 52°22'W, 0.5 km E of confluence with Rio Oiapoque, 28 Aug. 1960, Ir- win, Pires & Westra 47900 (MG). Para: Parque Indigena do Tumucumaque, Rio Paru de Oeste, Missao Tiriyo, Cavalcante 2518 (F); Belem, in sylvis ad Para, May 1819, Martius s.n. (M); vicin- ity of Belem, Sep.-Oct. 1961, Pires 51899 (NY, us); estrada do Cafezaldo IAN, 28 Oct. 1949, N.T. da Silva 336 (GH, NY, p); Almeirim, Rio Arrayo- los, 26 Apr. 1903, Ducke s.n. (MG, INPA); Orixi- mina, 8 Dec. 1906, Ducke s.n. (MG); Rio Cumi- namirim, 16 Dec. 1906, Ducke s.n. (MG); Obidos, 10 Jan. 1920, Ducke s.n. (RB); Rio Branco, 21 Mar. 1924, J.G. Kuhlmann 1722 (RB); Cachoeira do Rio Capim, 29 Jun. 1897, Huber 887 (MG); *Belem, IPEAN, Reserva Mocambo, Pires & Sil- va 10218 (GH, IAN), * Pires & Silva 11436 (GH); *Belem, IPEAN, Schubert 2150 (us); *Rio Trom- betas, planalto Saraca, N.T. da Silva & Santos 4631 (F). Brunfelsia guianensis was for a long time con- sidered an anomalous species among the South American brunfelsias. With the recent discovery of several related species, it is now known to be- long to a distinct section of the genus, of which it is the type species (Plowman, 1978). Brunfelsia guianensis differs from other species in this group 48 FIELDIANA: BOTANY in having obovate leaves and a somewhat inflated, ovoid-campanulate calyx. The fruit is large and fleshy and yellow at maturity. A lowland tropical forest species, B. guianen- sisis is distributed throughout the lower Amazon basin, chiefly north of the river, reaching well into the Guianas. It is known to flower from October to April, and fruits have been collected throughout the year. In the Brazilian state of Para, this species is used in local medicine under the name manaca with properties similar to those of B. uniflora (see LeCointe, 1947; Plowman, 1977). 6. Brunfelsia martiana Plowman, Bot. Mus. Leafl. 24(20): 37. 1974. Type: BRAZIL. Provinciae Rio Negro (Amazonas): habitat in sylvis ad flum. Japura prope Sao Joao do Principe, Dec. 1819, Martins 3247 (holotype, M; isotypes, M). Figure 17. Shrub to 1 m tall. Branches few, naked below, somewhat knobby at nodes, glabrous. Bark cracked longitudinally, dark reddish brown, shiny, furnished with lenticels. Leaves scattered along branchlets, subsessile, blade 10-25 cm long, 4-8 cm wide, mostly oblong, sometimes elliptic-ob- long or oblong-obovate, acuminate at apex, broad- ly cuneate to blunt at base, glabrous on both sides, upper surface dark green, nitid or dull, lower sur- face paler green, sometimes nitid, firmly membra- naceous to subcoriaceous, midrib dark reddish brown, lateral nerves 8-12, spreading, straight; petiole very short, 1-4(12) mm long, glabrous, dark brown, becoming cracked, rugose. Inflores- cence terminal or axillary in the upper leaf axils, glabrous, axis 2-10(15) mm long. Flowers 1-7 per inflorescence, sometimes with short peduncle that articulates with pedicel, greenish white to white. Bracts 1-3 per flower, 1-10 mm long, lin- ear-lanceolate, concave, ciliolate at margin, ca- ducous. Pedicel 3-6(10) mm long, short, slender, glabrous. Calyx 8-12 mm long, 3-8 mm in di- ameter, tubular to tubular-campanulate, glabrous, rarely striately veined, teeth 2-6 mm long, sub- equal, triangular-ovate to ovate-lanceolate, acute to acuminate at apex. Corolla tube 20-24 mm long, 1-3 mm in diameter, twice as long as calyx, straight, cylindric, inflated at apex, glabrous; limb 15-22 mm across, spreading, somewhat undulate, lobes 5-10 mm long, subequal, oblong-obovate, abruptly reflexed at lateral margins, rounded-sub- truncate at apex. Stamens included within upper part of corolla tube; filaments subligulate, anterior pair 4 mm long, posterior pair 3 mm long; anthers about 1 mm in diameter, orbiculate-reniform, slightly unequal, the upper pair somewhat smaller. Ovary 2 mm long, oblong-ovoid; style about 15 mm long, filamentous, a little broader and curved at apex; stigma about 1 mm long, briefly bifid, upper lobe slightly larger. Fruit and seed incom- pletely known. DISTRIBUTION — Guyana, Brazil (Amazonas, Ba- hia, Maranhao, Para). See Figure 15. SPECIMENS EXAMINED — GUYANA. Essequibo- Demerara River, Madray-Bubu Trail, Topy Trysil Forest, 8 Feb. 1944, Forest Department 4422 (K). BRAZIL. Amazonas: Manaus and vicinity, road Manaus-Caracarai, km 22, 21 Mar. 1967, Prance et al. 4704 (wis); Rio Negro, Cachoeira Baixa de Tarumo, 11-14 Apr. 1973, Schultes & Rodriguez 26132A (ECON), Schultes & Rodrigues 26133A (ECON); *Mun. de Barcelos; Rio Demeni, Froes 28393 (IAN); *vicinity of Manaus, Prance et al. 11611 (MO). Bahia: Dist. Ilheus, Ferradas, Dec. 1818, Martius s.n. (M); Feira de Santana, Apr. 1850, collector unknown (G). Maranhao: Ilha Sao Luis; estrada que vai do Rio Anil para Maio- ba, Froes 25650 (IAN); Mun. Sta. Luzia; Fazenda Cacique, W of Sta. Inez and E of Entroncameen- to, Taylor et al. 1084 (F). Para: Faro, 22 Jan. 1910, Ducke s.n. (MG); Bella Vista, Rio Tapajoz, matta das immediacoes da Campina do Perdido, 12 Feb. 1917, Ducke s.n. (MG); Regiao do Jutah y de Almerim. Palhal, 16 Apr. 1923, Ducke s.n. (RB); Gurupa, 25 Feb. 1923, Ducke s.n. (RB); Para, May 1819, Martius 3300 (M). Without state: Schott 5361? (w), Schott s.n. (w). Brunfelsia martiana belongs to section Gui- anenses. The type specimen collected by Martius (Schedulae 3247) on the Rio Japura includes a handwritten description of the plant that Martius apparently intended to publish as B. acuminata, a name that was later used by Pohl and Bentham for a form of B. brasiliensis. Brunfelsia martiana has been very poorly col- lected, as is true of many of the species of section Guianenses. The few collections available were collected over an extensive geographical area spanning the entire breadth of Brazil, and nearly all are preserved in European and Brazilian her- baria. The species has been collected only twice in the last 50 years; no collections have been made in the state of Bahia since 1850. One ap- parently quite early collection with six duplicates is preserved in Vienna. The label bears only the data "Schott 5361, Brasilia." This appears to be PLOWMAN: REVISION OF BRUNFELSIA 49 BRUNFELSIA martiana Plowman FIG. 17. Brunfelsia martiana. Reproduced courtesy of the Botanical Museum of Harvard University. 50 FIELDIANA: BOTANY the wrong label for the specimen because Schott collected only in the state of Rio de Janeiro and no other collection of the species has ever been made there. Mislabeling of the collections of Schott and Pohl has been recognized since the time of Martius, who made a special note of it in Flora Brasiliensis [1(1): 82 (1906)]: "Dolendum quod, quae specimina a cl. POHL, quae a cl. SCHOTT collecta sint, ex plagulis herbarii pala- tini Vindobonensis non semper statui potest." The origin of this collection remains question- able, yet it must be taken into consideration be- cause it represents one of the more complete gath- erings of the species and includes an immature fruit. Brunfelsia martiana is a small shrub with greenish white flowers. It is related to both B. amazonica and to B. guianensis and occurs sym- patrically with both of these species. Brunfelsia martiana differs from B. amazonica in having larger leaves (10-25 vs. 6-12 cm) with more lat- eral nerves (8-12 vs. 5-8) and much shorter ped- icels (3-6 vs. 13-20 mm). Brunfelsia martiana is closely allied with B. guianensis as well, and specimens of the former were originally considered to be exceptional forms of B. guianensis. Additional material proved the distinctiveness of B. martiana. It is now distinguished from B. guianensis by having larger (10-25 vs. 6-15 cm), oblong leaves with more lateral nerves (8-12 vs. 4-5), a calyx that is usually narrow-tubular instead of ovoid-cam- panulate, and a corolla tube that is rarely more than twice as long as the calyx. In B. guianensis it is frequently 2%-3 times as long. The flowers of all three of these related species are remarkably similar in the shape and color of the corolla, especially in the curious form of the corolla lobes, the lateral margins of which are de- flexed, giving them a rectangular appearance. The fruit of B. guianensis is a thick-walled, fleshy cap- sule; that of B. amazonica is thin-walled and dry at maturity. The collection labeled Schott 5361 mentioned above includes a small, immature fruit enclosed by a coriaceous and striately veined ca- lyx. The pericarp appears cartilaginous but not fleshy in this early stage. A second collection, which for B. martiana has exceptionally small leaves reminiscent of B. gui- anensis, was made by Martius near Belem do Para. This specimen included the remnants of a fruit that is small, definitely thin-walled, and brit- tle, perhaps the remains of the endocarp only. Martius tentatively named this plant B. flexuosa and wrote a description (Schedulae 3300) that ap- pears on the same page attached to the type spec- imen of B. martiana. The nature of the fruit in this species remains problematical. When, indeed, it is fully known, it should be very useful in de- termining the relationship of B. martiana to other species. Brunfelsia martiana is one of the most widely distributed species of section Guianenses, grow- ing in tropical lowland forests from Guyana south to Bahia in Brazil and westward to the Colombian frontier. The curious distribution between the Am- azon basin and the coastal forests of Bahia has been observed in other groups of plants (Plow- man, 1979). Brunfelsia sect. Franciscea (Pohl) Griseb., Fl. Brit. W. I.: 432. 1861. Franciscea Pohl, PI. Bras. Icon. Descr. 1: 1, t. 1-7. 1826. Lecto- type species (designated here): Franciscea uniflora Pohl. TYPE SPECIES — Brunfelsia uniflora (Pohl) D. Don, Edinburgh New Philos. J. 86. Jul. 1829. Shrubs or small trees, 0.3-10 m tall. Inflores- cence terminal, rarely axillary. Flowers large or small, 1-many per inflorescence. Corolla purple fading to pale violet or white with age, with white eye at mouth, tube 1-4 times as long as calyx, slightly inflated near apex, then slightly constrict- ed to form thickened ring, the lobes broadly rounded, overlapping at lateral margins. Stigma briefly bifid, in the form of a forceps. Anthers orbicular-reniform in outline. South and Central America. Note: Pohl, in establishing his new genus Fran- ciscea, included seven species. In recognizing this name as a section of Brunfelsia, it is necessary to designate a type species. I have chosen B. uniflora (Pohl) D. Don for the following reasons. It is the oldest known species of the genus, having ap- peared in 1 648 in the work of Piso under the name Manacd, accompanied by a good illustration showing the flowers and fruits (Fig. 18). The ho- lotype collected by Pohl and preserved in Vienna is a reasonably good specimen showing the char- acters of the species. The species is one of the most widespread of the genus and is well repre- sented in herbaria. Lastly, B. uniflora (as B. ho- peand) was the only species mentioned by Gri- sebach (1861) in setting up Franciscea as a sec- tion of Brunfelsia. PLOWMAN: REVISION OF BRUNFELSIA 51 FIG. 18. Manaca (Brunfelsia uniflora). From Piso, W., De Medicina Brasiliensis (1648). 52 FIELDIANA: BOTANY Although the date 1827 appears on the title page of Pohl's work Plantarum Brasiliae Icones et Descriptiones, it has been pointed out (Stafleu, 1967; Stafleu & Cowan, 1983) that the first part of the work, including the text and plates of Fran- ciscea, actually appeared in August 1826. 7. Brunfelsia australis Benth. in DC., Prodr. 10: 200. 1846. Lectotype (designated by Hunt, 1978): BRAZIL. Brasilia meridionalis, Rio Uruguay, Baird s.n. (K). Lectoparatypes: AR- GENTINA. Buenos Aires, in hortis Bonarien- sibus sub nomine Jasmini Paraguayensis cul- ta, Tweedie s.n. (K); Estrada de Minas, Mar- tius s.n. (?). No collection of Martius with this data has been located. Query was made by Bentham. Sellow 1573, also cited by Ben- tham, is excluded as a type; it belongs to B. pilosa Plowman. Figure 19. Franciscea australis (Benth.) Miers, Ann. Mag. Nat. Hist. ser. 2, 5: 250. 1850. Brunfelsia hopeana var. australis (Benth.) J.A. Schmidt in Mart., Fl. Bras. 8(1): 262. 1864. Brunfelsia paraguayensis Chodat, Bull. Herb. Bois- sier, ser. 2, 1: 406. 1901. Type: PARAGUAY. In silvis pr. Rio Apa, May, without year, Hassler 725a (holotype, G). Brunfelsia uniflora f. typica Hassl., Feddes Repert. Spec. Nov. Regni Veg. 15: 243. 1919. Non sensu D. Don. Type: PARAGUAY. In silvis pr. Rio Apa, May, without year, Hassler 725a (holotype, G). Brunfelsia uniflora f. obovatifolia Hassl., Feddes Re- pert. Spec. Nov. Regni Veg. 15:243. 1919. Type: PARAGUAY. Cordillera de Altos, in silvis, fru- tex vel arbor parva 3-5 m, corolla albo-coeru- lescens vel violacea in eadem planta, Sep. 1898- 1 899, Hassler 3246 (holotype, o; isotypes, A, G, NY, P, w). Brunfelsia uniflora f. intermedia Hassl., Feddes Re- pert. Spec. Nov. Regni Veg. 15: 243. 1919. Lec- totype: PARAGUAY. In regione lacus Ypacaray, Cordillera do Altos, arbusto 2-2.5 m, fl. lila- morado, monies y orillas, Cordilleras y bajos, azucena, Sep. 1913, Hassler 12257 (lectotype, G; isolectotypes, c, GH, K, L, MO, NY). Lectopar- atypes: PARAGUAY. In regione lacus Ypacaray. Picada a Bernalcue, Mar. 1913, Hassler 12147 (A, G); San Bernardino, orillas monies, fl. morado despues blanca, arbusto 3-4 m, Aug. 1915, Hassler (cited as Rojas) 1333 (G). Shrub to small tree 2-4 m tall. Trunk often branched from near base, to 8 cm in diameter. Bark rugose, yellowish brown. Branches ascend- ing and spreading, leafy, glabrous, green with yel- lowish brown cracks, furnished with lenticels. Branchlets glabrous, green. Leaves scattered along branchlets, blade 4.0-12.5 cm long, 2.5-6.0 cm wide, broadly elliptic to obovate, rarely orbic- ular or rhombic, blunt to rounded at apex, occa- sionally acute, cuneate at base, often briefly pu- bescent-ciliate when young, becoming glabrous, or sparsely glandular beneath at costa, dull dark green above, pale green beneath, membranaceous to subcoriaceous, lateral nerves 5-7, prominulous beneath, straight or somewhat arching, veinlets often obscure; petiole 3-8 mm long, glabrous or sparsely pubescent. Inflorescence terminal at the tips of the branchlets, sessile, with 1-4 flowers. Bracts present or absent, to 2 mm long, lanceo- late, concave, ciliolate, early caducous. Flowers showy, violet, fading to pure white with age, fra- grant. Pedicel stout, 4-7 mm long, glabrous, be- coming thicker and corky-verrucose in fruit. Ca- lyx 7-12 mm long, 4-10 mm diameter, campan- ulate, more or less inflated, glabrous, rarely pa- pillose-punctate, pale green, firmly membrana- ceous, reticulately veined, teeth 3-6 mm long, ovate-lanceolate, acute; calyx in fruit coriaceous, partially enclosing fruit, smooth. Corolla tube 25-32 mm long, 1.5-2.0 mm in diameter, 2-3 times as long as calyx, glabrous, often sparsely glandular in bud, glabrescent; limb 30-40 mm in diameter, plane, spreading, thickening at mouth prominent, round, white or yellowish, lobes equal, broadly rounded, overlapping laterally, abruptly narrowed at base. Stamens included; filaments slender, upper pair 2-3 mm long, lower pair 3-5 mm long; anthers about 1 mm in diameter, reni- form-orbicular. Ovary 2.5 mm long, 1 mm in di- ameter, ovoid-conical; style 20-28 mm long; stig- ma about 1 mm long. Capsule 10-22 mm long, 10-20 mm in diameter, subglobose to ovoid, apic- ulate at apex, smooth, glabrous, dark green, pericarp to 0.5 mm thick, dry at maturity, inde- hiscent or tardily dehiscent. Seeds 20-40, 4-6 mm long, 3-4 mm in diameter, oblong-ellipsoid, angular, dark brown. Embryo 3-4 mm long, slightly curved; cotyledons ovate. DISTRIBUTION — Brazil (Parana, Rio Grande do Sul, Sao Paulo), Paraguay, Argentina (Chaco, Corrientes, Formosa, Misiones, Santa Fe), and Uruguay; cultivated in some areas, especially in Central America. See Figure 20. SPECIMENS EXAMINED — BRAZIL. Parana: *Municipio Foz do Igua9u; Vila Postes, Buttura 157 (F); *Ceu Azul, Hatschbach 43192 (F). Rio Grande do Sul: Neu Wurttemberg, 450 m, 15 Oct. 1904, Bornmuller 219 (A, G, GH, M, w); Sao Leopoldo, 16 Oct. 1934, Rambo 321 (C); Porto Alegre, 12 Sep. 1945, Rambo 37819 (A, c, w); PLOWMAN: REVISION OF BRUNFELSIA 53 Morra da Gloria, Rambo 29147 (BR, MO). *Mun- icipio Itaqui, Dobereiner & Tokarnia 841 (RB). Sao Paulo: *Parque Estadual do Turvo, Tenente Portela, Paiva, Stehmann s.n. (F). PARAGUAY. Cordillera de Altos: Cerros de Tobati, 18 Aug. 1902, Fiebrig 49 (A, F, G, L, M, PR); Hassler 725 (G); Sep. 1913, Hassler 12559 (A); Villarrica, Sep. 1931, Jorgensen 3661 (A, c, F, MA, MO, NY, PH, s, us); Dep. Cantera, Zona Grande Bosques, Herrera Vegas, 220 m, 11 Aug. 1939, Monies 4703 (BAB); Dep. San Pedro, Colonia Primavera, 11 Sep. 1955, Woolston 571 (c, GH, NY, s, uc); without locality, 26 Aug. 1874, Balansa 2240 (G, GOET, K, LD, LE, PE, s); 4 Sep. 1854, Palmer 155 (us); *Par- aguari, Cerro San Jose, a civitate Ybicui, Bernardi 18138 (F); *Neembucu, Curupaity ad Laureles, Bernardi 20495 (F). ARGENTINA. Chaco: Dep. Resistencia, Margarita Belen, 8 Sep. 1946, Agui- lar 841 (s), 18 Sep. 1948, Aguilar 1302 (w); Dep. Primer de Mayo, Colonia Benitez, Oct.-Nov. 1928, Schulz 85 (BAB), 26 Nov. 1906, Stuckert 16307 (G); Las Palmas, Jorgensen 2098 (GH, MO, uc); Rio de Oro, km 140 de Resistencia, 29 Aug. 1944, Rojas 12133 (uc). Corrientes: Capital, Feb. 1821, Bonpland 528 (P); 1908, Llamas s.n. (BAB); Quinta La Eloisa, 77 m, 6 May 1969, Plowman 2713 (ECON, GH). Dep. Conception, Tabay, 1 Nov. 1965, Krapovickas & Cristobal 11706 (uc); 7 June 1969, Krapovickas & Cristobal 15385 (c, MO); 7 May 1969, Plowman 2723 (ECON, GH); Dep. Mburucuya: Estancia "Santa Teresa," 6 Sep. 1949, Pedersen 418 (BR, c, GH, p, s, us); Estancia "Santa Maria," Campo "Dios Gracia," 18 Dec. 1949, Pedersen 418 (BR, c, NY, p, s); Estancia Santa Ana, 21 Oct. 1944, Schwarz 93 (A); Canada Paso Clarios, 11 Nov. 1949, Schwarz 8661 (BH, G, w). Dep. Bella Vista, Bella Vista, Oct. 1904, N. Rojas Acosta 13004 (BAB); Dep. Saladas: La- guna Salada, 28 Sep. 1944, Schwarz 36 (A); La- guna Soto, 28 Sep. 1944, Schwarz 48 (A); Paso del Deseo, 1 Nov. 1944, Schwarz 139 (A, uc); 2 Dec. 1949, Schwarz 8972 (BH, G, w). Dep. Itu- zaingo, Salto Apipe, 6 Oct. 1949, Schwarz 8114 (BR); Dep. General Paz, Laguna Rincon, 2 Nov. 1949, Schwarz 8547 (LD); Dep. San Roque, San Roque cercanias, 20 Apr. 1945, Ybarrola 2963 (uc, w); Dep. San Luiz del Palmar, Colonia J. M. Llano, 19 Sep. 1945, Ybarrola 3270 (B, w); Cer- rudo Cue, 5 Sep. 1946, Ybarrola 3430 (G, p); vi- cinity of Goya, 15-30 m, 2-10 Nov. 1913, Curran s.n. (us); *Dep. Santo Tome, Garruchos, Krapov- ickas et al. 26092 (MO); *Dep. San Martin, Carlos Pellegrini, Krapovickas 29261 (MO); *Dep. San Martin, Sussini, Schinini et al. 18580 (F). For- mosa: Dep. Pilcomayo, Estancia Riacho Negro, 3 Oct. 1947, Morel 3783 (BR, MO); Salvation, 17 Nov. 1947, Morel 4144 (c); S.O. a 12 km de Fi- lipina, Est. Salaberry, 21 Nov. 1949, Morel 8921 (BH, c, G, w); Formosa, 29 Oct. 1900, Kermes 1363 (BAB); Fontana, Aug. 1933, Meyer 166 (GH). Misiones: Dep. Iguazii, Salto Iguazu, 2 Oct. 1947, Pierroti 6563 (G); 6 Oct. 1910, Rodriguez 484 (A, us); Dep. Candelaria, Martires, 24 Oct. 1917, Ber- toni 3434 (w); Loreto, 26 Sep. 1932, Muniez 99 (BAB); Dep. San Javier, Acaragua, 16 Sep. 1946, Bertoni 2939 (B); Dep. G. M. Belgraw, "El Ca- burei," 450 m, 22 Sep. 1957, Monies 27471 (M, NY); without locality, Fox s.n. (CGE). Santa Fe: Dep. General Obligado, Florencia, 30 Oct. 1923, Chiar s.n. (BAB); Villa Guillermina, 14 Feb. 1939, Ragonese 3646, (F); Indio Muerto, 6 Mar. 1939, Meyer 3076 (A). URUGUAY. Banda Oriental, 1816-1821, St. Hilaire C-2197 (p). CULTIVATED SPECIMENS OF INTEREST — BRA- ZIL. Rio Grande do Sul: Porto Alegre, 20 Mar. 1893, C. Lindman A597a (s); Santa Maria, Sil- vacultura, 14 Oct. 1955, Michel 8 (B). Rio de Ja- neiro: Rio de Janeiro, Quinta de Boa Vista, 13 Oct. 1879, Glaziou 12110 (BR, c, G, LE, p, R); Rio de Janeiro, Oct. 1915, Vincent s.n. (L). Santa Ca- tarina: Brusque, 4 Oct. 1961, Reitz & Klein 11250 (us). ARGENTINA. Tucuman: Capital, Institute Lillo, 9 Sep. 1947, Meyer 12621 (B); Dep. Trancaas, San Pedro de Colalao, 1,200 m, Apr. 1926, Venturi 4408 (A). Bentham described B. australis from three specimens, two of which are included in the pres- ent concept. Baird s.n. was chosen (by Hunt, 1978) as the lectotype. One of the other speci- mens, collected by Tweedie, was made from hor- ticultural material in Buenos Aires. The third col- lection cited by Bentham, Sellow 1573, is here referred to B. pilosa, a closely allied species. A fourth collection by Martius (with a query by Bentham) cannot be located. The locality "Estra- da de Minas" indicates another species was in- volved because B. australis does not occur north of Rio Grande do Sul in Brazil. FIG. 19. Brunfelsia australis. Reproduced courtesy of Curtis 's Botanical Magazine and Christabel King. 54 FIELDIANA: BOTANY Brunfelsia australis CHRISTABEL KING del. PLOWMAN: REVISION OF BRUNFELSIA 55 FIG. 20. circle). Distribution of Brunfelsia australis (solid triangle), B. boliviano (solid square), and B. bonodora (solid Schmidt in Martius (1864) considered B. aus- tralis to be a variety of B. uniflora (B. hopeana}. He cited the Sellow collection mentioned above and two additional specimens. One of these, Rie- del s.n., collected near Macae, Rio de Janeiro, agrees with B. bonodora. The other collection, Martius s.n., from Para, is referred to B. guianen- sis, although most of this material is sterile. Brunfelsia australis is closely related to B. un- iflora and B. pilosa and by some authors (Schmidt 1864; Hassler, 1919) has been considered synon- ymous with B. uniflora. However, these workers suffered from the disadvantage of working exclu- sively with dried herbarium specimens (Schmidt) or with local populations without understanding the entire morphological range of the species in- volved (Hassler). Studies of both B. australis and B. pilosa in the field and of all three entities in cultivation and from herbarium specimens reveal that these plants are morphologically, ecological- ly, and geographically distinct and merit the rank of species. Experimental crosses suggest that re- productive isolation exists between B. pilosa and B. australis. Brunfelsia australis is distinguished from B. uniflora and B. pilosa by the broadly elliptic to obovate leaves, the variable number (one to four) of flowers per inflorescence and the uniformly glabrous, campanulate calyx. Even though their ranges overlap, almost no in- termediates have been found between B. australis and B. pilosa. One collection from Rio Grande do Sul, Bornmuller 219, has oblong leaves and a more deeply incised calyx and may represent 56 FIELDIANA: BOTANY some introgression from B. pilosa, which also oc- curs in this region. Commonly growing as a tall shrub or treelet, B. australis is found primarily in the relatively recent drainage systems of the Rio Parana, Rio Uruguay, and Rio Jacui (Rio Grande do Sul), growing in the understory of low-elevation forests near the rivers and in adjacent uplands. It occurs from sea level to 450 m altitude. In the low-lying, often swampy river basin of the Rio Parana, B. australis is found in the scattered woodlands that appear on higher and drier ground. Two reports of widely disjunct populations are now known to represent cultivated plants. Venturi 4408 from Tucuman, Argentina, was distributed to some herbaria without the notation "culta." Specimens of Glaziou 12210 were distributed with the fictitious provenance "Rio de Janeiro, Serra da Palmital." However, a duplicate speci- men at the Museu Nacional de Rio de Janeiro bears the correct locality, "Quinta da Boa Vista (Rio), Arbusto cult." This erroneous data may be the result of direct falsification by Glaziou, which was pointed out in the Melastomataceae by Wur- dack (1970). Brunfelsia australis flowers from August to November; mature fruits appear from April to June. Frequently the entire plant is covered with fragrant, showy purple and white flowers at the same time, making this species a valuable orna- mental. It is widely cultivated in gardens between latitudes 35°N and 35°S (San Francisco, Califor- nia, to Buenos Aires, Argentina). 8. Brunfelsia boliviano Plowman, Fieldiana, Bot. n.s. 8:1. 1981. Type: BOLIVIA. Santa Cruz: Prov. de la Cordillera, region of Lagunillas, Cordillera of Incahuasi, 900 m, "at dry and sandy slopes; shrub 1-2 m; flowers white or light violet; when cattle eat the leaves of this plant [they] die." N. v. bella union. Aug. 1934, Cardenas 2813 (holotype, F 756420). Figure 21. Shrub 1-2 m tall (fide Cardenas). Bark on branches thin, yellowish to grayish brown, lon- gitudinally rugose, not exfoliating. Branchlets 2- 3 mm in diameter, more or less villous, becoming glabrous. Leaves scattered along branchlets or crowded at tips of lateral short shoots, 40-135 mm long, 23-58 mm wide, blade obovate, rarely elliptic-obovate, apically rounded with a short acumen 5-10 mm long, the acumen itself blunt to acute, somewhat revolute at margin, basally atten- uate, glabrescent on both surfaces except at the midrib, which bears villous and glandular hairs, ciliate-villous on the adaxial surface of the revo- lute margin, dull green above, paler, yellowish green beneath, firmly membranaceous or some- what coriaceous, the lateral nerves 4-9, straight, arcuately anastomosing toward the margin, prom- inulous beneath; petiole short, 2-6 mm long, more or less villous, more densely so on upper side. Inflorescence corymbiform, terminal on last sea- son's branchlets, short pedunculate, branched, with 6-15 flowers. Peduncle 5-12 mm long, more or less villous. Bracts small, 14 mm long, lan- ceolate or cymbiform, sparsely to densely villous especially at margin, caducous. Flowers light vi- olet fading to white (fide Cardenas). Pedicel short, 2-6 mm long, 1 mm in diameter, scarcely thick- ened at apex, glabrous or with sparse glandular hairs. Calyx 9-15 mm long, 3-4 mm in diameter, tubular, truncate at base, 5-angled in cross section, appearing plicate in pressed specimens, glabrous or with scattered glandular hairs, teeth 2-4 mm long, subequal, ovate, apically short acuminate, the acumen itself blunt and glandular-papillose; fruiting calyx persistent, withering. Corolla tube 21-25 mm long, 2-3 mm in diameter, about twice as long as calyx, straight; limb 18-30 mm in di- ameter, spreading, lobes 8-14 mm long, subequal, rounded. Stamens inserted in upper part of co- rolla tube; filaments strap-shaped, the anterior pair 3 mm long, suberect, briefly exserted from mouth of corolla tube, the posterior pair 2 mm long, in- cluded; anthers 1-1.2 mm in diameter, orbicular- reniform, subequal or the upper pair slightly larg- er. Ovary 2-3 mm long, 1.8-2.2 mm in diameter, narrowly ovoid, with about 30 ovules; style 19 mm long, incurved at apex; stigma 2 mm long, briefly bifid. Capsules 3-6 per infructescence, 20-25 mm long, 20-25 mm in diameter, dry at maturity, subglobose, smooth, green; pericarp thin, 1-1.5 mm thick, endocarp thin, crustaceous, with 22-28 seeds per capsule. Seeds 5-8 mm long, 2-3 mm in diameter, oblong to subreniform, subterete, dark brown, reticulate-pitted. Embryo 4 mm long, straight; cotyledons 1.5 mm long, ovate; radicle 2.5 mm long. DISTRIBUTION — The eastern Andes of southern Bolivia. See Figure 20. SPECIMENS EXAMINED — BOLIVIA. Chuquisa- PLOWMAN: REVISION OF BRUNFELSIA 57 BRUNFELSIA boliviana Plowman FIG. 21. Brunfelsia boliviana. Reproduced courtesy of the Field Museum of Natural History. 58 FIELDIANA: BOTANY ca: mountain above Bartolo, on road from Mon- teagudo to Sucre, 20°00'S, 64°45'W, 5000 ft, 20 Sep. 1949, Brooke 5653 (BM). Santa Cruz: Prov. Cordillera, Nov.-Dec. 1845, Weddell 3621 (p, 2 sheets); 5 km N of Yatarenda, 63°32'W, 19°12'S, 17 Apr. 1977, Krapovickas & Schinini 31476 (CTES, F); *Prov. Luis Calvo: Monteagudo 26 km hacia Camiri, Beck & Liberman 9379 (F). Brunfelsia boliviano is known from only five col- lection^,', all from a relatively small area in the foothills/ of the Andes in southern Bolivia. The early collection of Weddell in 1845 is labeled merely "Provincia de la Cordillera," referring to the large province in the department of Santa Cruz. However, Urban (1906) mentions that in the months given, Weddell traveled and collected from Santa Cruz da la Sierra in Santa Cruz south to Sauces (now Monteagudo) in the department of Chuquisaca. His collection was probably made near the border between the two departments. Brunfelsia boliviana is most closely related to B. cuneifolia, which occurs in southern Brazil from the state of Parana south to Rio Grande do Sul. Like the present species, it is known from only a few specimens. Brunfelsia boliviana differs in both leaf and inflorescence characters as sum- marized in Plowman (1981). The geographical distribution of these two closely related species deserves some comment because it exemplifies further a pattern observed in other brunfelsias in which vicarious species pairs are found in southeastern Brazil and the eastern Bolivian Andes. These include B. bono- dora-B. grandiflora subsp. schultesii and B. hy- drangeiformis-B. mire. In addition, disjunct and somewhat distinct populations of B. uniflora, a species primarily of eastern Brazil, have also been found in the Bolivian Andes. In each of these cases, vicarious species or populations are now separated by more than 2,000 km in which no intervening populations are known to occur. This phytogeographical pattern has been discussed by Smith (1962) and by Plowman (1979). Interestingly, the leaves of B. boliviana are re- puted to be lethal to cattle, a feature that has been reported for other Brunfelsia species (Plowman, 1977). 9. Brunfelsia bonodora (Veil.) J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13, 5-B(l): 152. 1962. Figure 22. Besleria bonodora Veil., Fl. Flumin. 261. 1829 ("1825"). Ic. 6, t. 80. 1831 ("1827"). Lectotype (designated here): Vellozo, J.M. da C., Fl. Flu- min. Ic. 6, t. 80. 1831 ("1827"). Representative collection: BRAZIL. Rio de Janeiro, Serra de Es- trella, Mar. 1833, Riedel 84 (LE). Shrub 1-2 m tall. Branches ascending, spread- ing, naked, glabrous. Bark thin, light brown. Branchlets glabrous or sparsely puberulent, gray- ish brown. Leaves scattered along branchlets, blade 5-14.5 cm long; 2-5 cm wide, narrowly el- liptic to lanceolate, acuminate at apex, blunt to cuneate at base, glabrous, at times puberulent at midrib on lower surface, shiny, dark green on up- per surface, pale green beneath, firmly membra- naceous, lateral nerves 5-8, more or less arcuate; petiole 2-6 mm long, slender, glabrous. Inflores- cence terminal, compact, unbranched, axis 3-8 mm long. Flowers 3-15, rarely reduced to 1 or 2; blue fading to white with age, fragrant. Bracts 2- 12 mm long, lanceolate, glabrous or puberulent, caducous. Pedicel 6-12 mm long, slender, thick- ened slightly at apex, glabrous. Calyx 11-14 mm long, about 5 mm in diameter, tubular-campanu- late to campanulate, ellipsoid to obovoid in bud, rarely somewhat inflated, glabrous, firmly mem- branaceous, striately veined, teeth 3-4 mm long, triangular to ovate, acute to blunt at apex, rarely apiculate, calyx to 15 mm long in fruit, subcor- iaceous, conspicuously veined, enclosing fruit. Corolla tube 17-25 mm long, 2 mm in diameter, twice as long as calyx, glabrous or sparsely glan- dular; limb 20-35 mm in diameter, plane spread- ing, lobes rounded to oblong, subtruncate or rounded at apex, rarely emarginate, overlapping at sides, narrowed abruptly at base. Stamens in- cluded in upper part of corolla tube; filaments about 4 mm long, subligulate; anthers 1.5 mm long, reniform. Ovary 1.5 mm long, ovoid-coni- cal; style slender, equaling the filaments; stigma 1 mm long, briefly bifid, upper lobe slightly larger, obtuse. Capsule 8 mm long, 6-8 mm in diameter, partially enclosed by calyx, ovoid-turbinate, apic- ulate at apex, smooth, pericarp thin-walled, drying crustaceous, sparingly dehiscent. Seeds about 5, incompletely known. DISTRIBUTION — Brazil (Rio de Janeiro, Sao Pau- lo). See Figure 20. SPECIMENS EXAMINED — BRAZIL. Rio de Ja- neiro: Serra de Estrella, 700 m, 29 Oct. 1946, Brade 18643 (RB), 12 Nov. 1823, Riedel 18 (A, LE, mixed collection; NY, mixed collection; us), 14 Dec. 1823, Riedel 59 (A, LE, mixed collection); PLOWMAN: REVISION OF BRUNFELSIA 59 ( Tal) 60 FIELDIANA: BOTANY Macahe, May 1832, Riedel sp. nov. (LE); Barra Mansa, Fazenda Paraiso, 3 Dec. 1960, Duarte 5477 (RB, us). Sao Paulo: Taubate, Dec. 1817, Martius 541 (M). Without state: Riedel 112 (us). Brunfelsia bonodora was first described in the ge- nus Besleria by Vellozo in Flora Fluminensis, a work that appeared in 1829 although 1825 ap- pears on the title page (Carauta, 1973). Vellozo's scanty description was accompanied by a some- what stylized illustration. This drawing must now be considered the lectotype of the species since no specimens of Vellozo exist (Fig. 22). In his monograph of Brunfelsia in 1846, Ben- tham erroneously placed Vellozo's species in syn- onymy with his B. latifolia (based on Franciscea latifolia Pohl). This concept also included mate- rial now recognized as B. grandiflora subsp. schultesii of the western Amazon and Andes. This problem was discussed by Monachino (1953) and Macbride (1962), but neither of these workers were able to resolve the issue, primarily because they saw neither the type materials nor represen- tative collections from southeastern Brazil. It is now clear that the name B. latifolia and its synonym, B. maritima Benth., apply to a distinct species known only from the sand restingas around the city of Rio de Janeiro. Brunfelsia bon- odora is represented by several collections from the mountains of the state of Rio de Janeiro that agree with Vellozo's description and plate. This concept is, however, not identical with Macbride's interpretation of the species since he included ma- terial here referred to as B. grandiflora subsp. schultesii. Nevertheless, B. bonodora is related to both B. latifolia and B. grandiflora subsp. schultesii, which is in part responsible for the confusion of these concepts. Brunfelsia bonodora is distin- guished from B. latifolia in having a more robust, upright habit; longer, lanceolate, and apically acu- minate leaves; and longer pedicels. In addition, B. bonodora differs ecologically, growing in the montane rain forests of the Serra do Mar and not in open, coastal formations (restingas). There seems to be some overlap in these two species, but the status of possible hybrids cannot be de- termined with so few specimens at hand, most of which lack adequate collection data. In fact, only two modern collections of B. bonodora are known. Macbride (1962) considered B. bondora to be conspecific with the plant here treated as B. gran- diflora subsp. schultesii. On close examination, however, there are several points of difference be- tween them. Brunfelsia bonodora has longer ped- icels (6-12 vs. 2-5 mm), a more compact inflo- rescence, larger bracts, and the capsule almost completely enclosed by the calyx in fruit. Ac- cording to the specific epithet, B. bonodora is fra- grant; B. grandiflora almost never is. Geographically, B. bonodora and B. grandiflo- ra are separated by several thousand miles. They exhibit, however, a distribution similar to those of B. hydrangeiformis and B. mire and may be con- sidered to be vicarious species that arose from a common ancestor. Brunfelsia bonodora is known only from the coastal montane rain forests in the state of Rio de Janeiro, at elevations up to 700 m. It flowers from October to December and fruits during May and June. Additional collections of this plant with complete field data will greatly aid in understanding its specific characters and rela- tionships. 10. Brunfelsia brasiliensis (Spreng.) L. B. Sm. & Downs in Reitz, Fl. Ilustr. Catar., pt. 1, Sola: 303. 1966. Gerardia brasiliensis Sprang., Syst. Veg. 2: 806. 1825. Type: BRAZIL. Brasilia meridionalis, without locality, Sellow s.n. (type destroyed at Berlin; representative collection and possible isotype, M).12 Shrub 0.3-2 m tall. Bark roughish, reddish to light brown. Branches numerous, arising from near base, leafy, yellowish brown, pilose, es- pecially so when younger, or glabrous. Branch- lets of current year leafy, slender, erect, densely pubescent with yellowish brown, nonglandular hairs. Leaves scattered along branchlets, some- times more crowded toward apex, blade 3-13 cm long, 1-4.5 cm wide, lanceolate to oblong-lanceo- 12 In the manuscript Plowman gave no obvious indi- cation that he wished to lectotypify this species as cited. We have thus left the lectotypification of this species to later Brunfelsia specialists. — Eds. FIG. 22. Lectotype of Brunfelsia bonodora (originally as Besleria bonodora). From Vellozo, J. M. C, Flora Fluminensis (1881). PLOWMAN: REVISION OF BRUNFELSIA 61 late, more rarely obovate-oblong, acute to acu- minate at apex, occasionally blunt, cuneate to nar- rowed at base, the upper surface sparingly pubes- cent, dark to light green, rarely nitid, subscabrous or glabrate, the lower surface variously pubescent, often villous at midrib, rarely glabrous, paler green, firmly membranaceous to coriaceous, lat- eral nerves 6-10, mostly straight; petiole 1-5 mm long, puberulent to villous. Inflorescence terminal and subterminal, corymbiform, compact and near- ly sessile or lax-elongate, with 5-many flowers, rarely reduced to 1-2, the axis 1-2 cm long, pu- bescent-villous. Bracts small, 2-15 mm long, lan- ceolate, ciliolate at margin, pubescent, caducous. Flowers often showy, violet fading to pale lav- ender or white with age, odorless, short pedun- culate. Peduncles 2-8 mm long, thickened at apex and articulating with pedicel, persistent, villous or glandular-pubescent, bearing 1-3 bracts. Bracts minute, 1-2 mm long, linear-lanceolate. Pedicel 2-15 mm long, erect, pubescence yellowish to yellowish brown, sparsely pilose to villose, rarely glandular, thickening in fruit. Calyx 8-22 mm long, 3-8 mm in diameter, tubular to tubular-cam- panulate, with variable yellow to yellowish brown pubescence, sparsely to densely villous, rarely pi- lose or glandular, light green to purple, firmly membranaceous, teeth 1-5 mm long, erect, some- times recurved in fruit, triangular to ovate, acute at apex; calyx in fruit campanulate to urceolate, becoming coriaceous, striate-nerved. Corolla tube 17-25 mm long, 1-3 mm in diameter, 1-2 times as long as calyx, sparsely pilose or glabrous, pale violet below, white toward apex, limb 18-32 mm in diameter, plane spreading, lobes subequal, broadly rounded to truncate at apex, lateral mar- gins. Stamens included in upper part of tube; fil- aments slender, slightly broader at base, glabrous, white, lower pair 2-4 mm long, upper pair 4-5 mm long; anthers 1 mm long, oblong-reniform. Ovary 1-5 mm long, ovoid-conical, green; style 17-18 mm long, filiform, bright green; stigma 1- 2 mm long, briefly bifid, the upper lobe somewhat larger, bright green. Capsule 8-13 mm long, 8- 10 mm in diameter, enclosed by calyx, ovoid to subglobose, apically apiculate, smooth, shiny, light to dark green, pericarp thin, 0.5 mm thick, cartilaginous to crustaceous, dry at maturity, spar- ingly dehiscent. Seeds (2)7-10, 4-6 mm long, 3 mm in diameter, oblong-ovoid, more or less an- gular, dark reddish brown, reticulate-pitted. Em- bryo 4 mm long, very slightly curved; cotyledons 1.5 mm long, ovate-elliptic. DISTRIBUTION — Brazil. See Figure 23. Key to the Subspecies of Brunfelsia brasiliensis (Spreng.) L. B. Sm. & Downs 1 . Inflorescence terminal, nearly sessile, compact, with 1-8 flowers; peduncles shorter than ped- icels; pedicels 5-15 mm long; calyx 15-22 mm long; leaves often glabrescent lOa. subsp. macrocalyx 1. Inflorescence terminal and subterminal, lax or dense, often branched, with 8-many flowers; peduncles equaling or longer than pedicels; pedicels 2-5 mm long; calyx 7-17 mm long; leaves usually pubescent-villous beneath lOb. subsp. brasiliensis lOa. Brunfelsia brasiliensis subsp. brasiliensis. Figure 24. Franciscea acuminata Pohl, PI. Bras. Icon. Descr. 1: 4, t. 3. 1826. Type: BRAZIL. Rio de Janeiro: in umbrosis inter frutices ad Mandioca, Schott 6163 (holotype, BR). Brunfelsia acuminata (Pohl) Benth. in DC., Prodr. 10: 199. 1846. Brunfelsia brasiliensis var. acuminata (Pohl) L. B. Sm. & Downs in Reitz, Fl. Ilustr. Catar, pt. 1, Sola: 305. 1966. Franciscea ramosissima Pohl, PI. Bras. Icon. Descr. 1: 5, t. 4. 1826. Type: BRAZIL. Rio de Janeiro: ad Serra Tingua, Schott 3720 (holotype, w; iso- types, GH, w). Brunfelsia ramosissima (Pohl) Benth. in DC., Prodr., 10: 199. 1846. Franciscea confertiflora Pohl, PI. Bras. Icon. Descr. 1: 6, t. 5. 1826. Type: BRAZIL. Minas Gerais: inter frutices locis umbrosis in via de Villa Fan- ado vel Bom Successo dicta ad Villa do Principe, Oct.-Nov. 1820, Pohl 3478 (holotype, w; iso- type, w). Brunfelsia confertiflora (Pohl) Benth. in DC., Prodr., 10: 199. 1846. Brunfelsia ramosissima var. confertiflora (Pohl) J. A. Schmidt in Mart., Fl. Bras. 8(1): 260. 1864. Franciscea divaricata Pohl, PI. Bras. Icon. Descr. 1: 6, t. 6. 1826. Type: BRAZIL. Minas Gerais: inter frutices, locis umbrosis in via de Sao Joao d'El Rey, ad Villa Paracatudo Principe, floret ab Octubri usque ad Martium, 1818, Pohl 593, 6165 (holotype, w; isotypes, F, GH LE, w). Brunfelsia ramosissima var. laxiflora J. A. Schmidt in Mart., Fl. Bras. 8(1):260. 1864. Type: BRA- ZIL. Brasilia meridionalis, without locality, Sel- low s.n. (holotype, M). Brunfelsia ramosissima var. parcifolia J. A. Schmidt in Mart., Fl. Bras. 8(1):260. 1864. Type: BRAZIL. Minas Gerais: without locality, Claus- sen 171 (holotype, BR). 62 FIELDIANA: BOTANY FIG. 23. Distribution of Brunfelsia brasiliensis (solid square) and B. chiricaspi (solid circle). Leaves 3-13 cm long, 1-4.5 cm wide, puber- ulent or scabrous above, often glabrescent, vari- ably villous beneath, often densely so at midrib, firmly membranaceous to subcoriaceous. Inflo- rescence variable, dense to lax, occasionally elon- gate and well-developed, with 10-many flowers, rarely reduced to about 5. Peduncles equaling or longer than pedicels, 2-10 mm long, rufo- villous, not glandular. Pedicels 2-5 mm long, villous. Ca- lyx 7-15 mm long, variably villous with dense, yellowish brown hairs, rarely pilose or glabres- cent, eglandular. Corolla tube 17-22 mm long, 1.5-2 times as long as calyx; limb 16-30 mm in diameter. DISTRIBUTION — Brazil (Distrito Federal, Goias, Minas Gerais, Parana, Rio de Janeiro, Sao Paulo). SPECIMENS EXAMINED — BRAZIL. Distrito Federal: gallery forest ca. 10 km NW of Planal- tina, 950 m, 27 Feb. 1966, Irwin et al. 13194 (us); gallery forest ca. 10 km E of Lagoa Paranoa, near DF-6, 1000 m, 25 Feb. 1970, Irwin et al. 26556 (wis); *Brasilia; Bacia do Rio Sao Bartolomeu, Corrego Forquilha, Heringer et al. 6148 (F). Goias: *25 km NW of Brasilia, Irwin et al. 15799 (F). Minas Gerais: Mun. Santo Luzia, Fazenda da Chicaca, 1 100 m, 20 Nov. 1945, Assis 102 (GH, R, us), Assis 137 (GH), Assis 142 (GH), 13 Dec. 1945, Assis 181 (GH, uc, us); Lagoa Santa, Oct.- Nov. 1863, Engler 1006 (c), 14 Jan. 1864, Engler 1009 (c), 30 Oct. 1863, Engler sp. nov. (c), 25 Mar. 1933, Mello Barreto 7786 (F), Warming 103 (NY, s); Serra do Cipo, km 120, 2 Sep. 1933, Mel- lo Barreto 7788; Mun. Belo Horizonte, Morro do Candido, Jan. 1934, Atamp. 6560 (R); Matta da Caixa d'Areia, 15 Dec. 1918, Gehrt 3276 (SP); PLOWMAN: REVISION OF BRUNFELSIA 63 FIG. 24. Brunfehia brasiliensis subsp. brasiliensis (originally as Franciscea confertiflora). From Pohl, J. E. Plantarum Brasiliae Icones et Descriptiones (1826). 64 FIELDIANA: BOTANY estrada para Lagoa Santa, 15 Jan. 1951, Joly 1133 (SP); Villa Paraiso, 20 Nov. 1922, Mello Barreto 7789 (F); Barreiro, 21 Jan. 1933, Mello Barreto 7791 (F); Serra do Taquaril, 18 Jan. 1933, Mello Barreto 7791 (F), 29 Mar. 1933, Mello Barreto 7793 (F); Jardim Botanico, 17 Nov. 1933, Mello Barreto 7794 (R); Serra da Mutuca, 27 Jan. 1945, Moreira 5775 (GH); Fazenda da Baleia, 4 Mar. 1940, Oliveira 21 (us); Morro das Pedras, 3 km S of Belo Horizonte, 1000 m, 12 Feb. 1945, Wil- liams & Assis (GH); near Lagoa Pampulha, 1000 m, 8 Mar. 1945, Williams & Assis 6088 (F, GH, MO, NY); Mun. Carandai, Carandai, km 416, 25 Nov. 1946, Duarte 631 (GH, RB); Hermilo Alves, 10 Jan. 1965, Duarte 8741 (GH, RB); Mun. Ouro Preto, Itacolomi, 1400 m, 31 Dec. 1950, Macedo 2829 (MO, NY, s); Mun. Araxa, Barreiro, 10 Feb. 1951, Macedo 3131 (s, SP, us); Mun. Santa Bar- bara, Serra do Cara9a, 14 Apr. 1933, Mello Bar- reto 7784 (F); Mun. Tombos, Fazenda das Antil- has, 21 Jan. 1936, Mello Barreto 7767 (F, R); Mun. Diamantina, Serra dos Crystais, 6 Nov. 1937, Mello Barreto 9603 (F, R); Po de Araca, 17 Nov. 1937, Mello Barreto 9833 (F, R); Mun. Ja- boticatubas, Lagoa de Dona Ignacia, 6 Jan. 1940, Mello Barreto 10530 (R); Mun. Vi£osa, state ag- ricultural school, 17 Dec. 1958, Irwin 2269 (NY, R, us); Fazenda de Dr. Christioma, 700 m, 30 Dec. 1930, Mexia 5467 (BM, BH, F, G, GH, MICH, MO, NY, PH, s, uc); Mun. Serro, 3 km from Serro, 12 May 1945, Williams & Assis 6859 (GH); Aguas Virtu- osas, 15 Jan. 1919, A. Amaral s.n. (SP); So^ego, Jan. 1921, A. Amaral s.n. (SP); Carmo do Rio Cairo, Fazenda Corrego Bonito, 5 Sep. 1961, Andrade 1024, Emmerich 985 (us); Campanha, Nov. 1896, Brandao 2066 (R); Serra do Cipo, km 131-132, Palacio, 4 Dec. 1949, Duarte 2029 (GH, RB); Ar- raial do Pinheiro, Oct. 1840, G. Gardner 5063 (BM, CGE, F, G, GH, K, NY, p, us, w); Serra do Pal- mital, 24 Jun. 1884, Glaziou 15311 (c, K, R); Santa Barbara do Mato Dentro, 12 Jan. 1921, Hoehne 4920 (SP, us); Caeti, Nov. 1915, Hoehne 6129 (R); Sabara, Jan. 1916, Hoehne 6846 (us); Serra de Grao Mogol, 1,100 m, 12 Nov. 1938, Markgraf 3495 (RB); prope Pitangui, Cueillie a Sabana, 11 Mar. 1862, Netto s.n. (R); Barbacena, 21-24 Jun. 1879, Netto et al. s.n. (R); BR-4, km 938, 40 km depois de Medina, 16 Jan. 1965, Pabst 8340, Pereira 9451 (HB, RB); Serra do Cipo (Concei9ao), 1,200 m, 22 Dec. 1948, Palacios, Balegno & Cuezzo 3467 (B, w); Caldas, 3 Feb. 1857, Regnell 1-375 (BR, c, M, p, R, s, us); Itabura, 1816-1821, St. Hilaire D-80 (MO, NY); Penha, St. Hilaire s.n. (us); Sitio, 20 Nov. 1905, Sampaio 68, 74 (R, us); Serra de Itabira do Campo, 1 1 Sep. 1887, Schwacke s.n. (R, us), 20 Dec. 1888, Schwacke s.n. (R); Congonhas do Campos, 1843, Stephan s.n. (BR); Serra do Frio, 1833, Vauthier 541 (GH, K, L, w); Serra da Mantiqueira, Passa Quatra, Esta9ao Florestal da Mantiqueira, 31 Jun. 1949, Vidal s.n. (R); without locality, Capanema s.n. (GH, RB), 1842-1843, Claussen 146 (BM, CGE), Claussen 147 (BM, CGE), Claussen 157 (w), 1840, Claussen 334 (BR), Claussen s.n. (BM, BR, GH, K, w), Weddell 1648 (p); *Serra do Espinha9o, SW of Rio Jequiti and Mendanha on road to Diaman- tina, Anderson 8949 (F, MO); *Municipio Tombas; Fazenda das Antilhas, Mello Barreto 7787 (F, R); *Belo Horizonte; Serra do Taquaril, Hab. Canga, Barreto 7790 (F); *ca. 17 km SW of Minas Ger- ais-Bahia state border, Davidse et al. 11611 (MO); *13 km W of Lavras along Highway 265 to Bar- bacena, Davidse & Ramamoorthy 10709A (F); *Municipio Pato de Minas; Corrego Barreiras, Hatschbach 42939 (F); *Municipio Itamonte; Ser- ra da Mantiqueira, Hatschbach & Kummrow 45569 (F); *Municipio Diamantina; Biri-Biri, Hatschbach & Pelanda 27993 (GH); *Serra do Es- pinha9o, Serra do Itabirito ca. 45 km S of Belo Horizonte, Invin et al. 19549 (MO); *Serra do Espinha9o, Serra do Cipo, Irwin et al. 20150 (MO); *Serra do Espinha9o, Irwin et al. 29189 (MO); *Serra de Espinha9o, S of Ouro Preto, Irwin et al. 29407 (MO); *Serra do Espinha9o, ca 34 km E of Belo Horizonte, Irwin et al. 30628 (MO); *Municipio Lima Duarte; Conceicao do Ibitipoca, Parque Florestal Estadual de Ibitipoca, Souza et al. 521 (F). Parana: *Municipio Senges, Rio Ca- juru, Hatschbach 43398 (F); *Ponte Nova-Minas Gerais, Hatschbach & Ahumada 31333 (MBM). Rio de Janeiro: Monte Corcovado, 300-700 m, 6 Mar. 1924, L.H. & E.Z. Bailey 758 (BH); by way of Laranjeiras, 31 Dec. 1825, Burchell 1372 (p), 1841, Claussen s.n. (w), 26 Dec. 1920, Ducke & J.G. Kuhlmann s.n. (RB); summit, 1836, Gardner 248 (BM, CGE, G, K, w), 10 Jan. 1865, Glaziou 812 (BR, c, R), May 1838, Guillemin 825 (P), Langs- dorff s.n. (us), Nov.-Dec. 1832, Luschnath s.n. (BR), 1834, Luschnath s.n. (BR), 700 m, 22 May 1969, Plowman 2786, Sucre 5086, (GH, ECON, PH, RB), Feb. 1832, Riedel 467 (L, GOET, LE), Nov.- Dec. 1832, Riedel 1190 (L, LE), 12 Jul. 1915, Rose & Russel 20221 (GH, us); Pico da Tijuca, 18 Jan. 1931, Brade 10569 (R), Brade 10583 (GH, R), 1838, Guillemin 41 (P); Excelsior, 3 Feb. 1930, J.G. Kuhlmann s.n. (GH, RB), Dec. 1824, Riedel 1249 (LE), 23 Aug. 1896, Ule s.n. (R); mata da Tijuca, 10 Nov. 1896, Ule s.n. (R), 1836, Vauthier PLOWMAN: REVISION OF BRUNFELSIA 65 42 (F, c, P). Novo Friburgo, 9 Dec. 1918, Curran 619 (GH, s, SP), Curran 623 (F, GH); Morro Pos- sole, Sitio Dr. Dungs, 1300 m, 28 Jan. 1968, Pabst 9115 (HB), 23 Dec. 1887, Glaziou 17170 (BR, c, LE, P, us); Petropolis, 1883, Glaziou 14171 (BR, c, c); pres de la roche de Juraracacu, 25 Mar. 1879, Glaziou 11394 (c, P); Carangola, Sep. 1943, Goes & Constantino 543 (RB); Retire, Nov. 1943, Goes & Constantino 710 (RB); Caetetu, Dec. 1943, Goes & Constantino 883 (RB); Estrada de Con- torno, 650 m, 2 Feb. 1968, Sucre 2360, Braga 203 (GH, RB); Mata do Judeo, 700 m, 7 Dec. 1968, Sucre 4213, Braga 1169 (RB), 1858-1860, Wawra & Maly 426 (LE, w); Terexopolis, Parque Nacional da Serra dos Orgaos, 16 Dec. 1942, Duarte de Barros 1195 (RB); Posso, Morro das Antenas de Televisao, 12 Feb. 1968, Sucre 2339, Braga 182 (GH, RB); Serra dos Orgaos, 3000 ft, 7 Jan. 1837, Gardner 564 (BM, CGE, G, GH, K, NY, us, w), 1885, Glaziou 15132 (c, G, K, LE), May 1839, Guillemin 954 (P); Imbuhy Lane, Jan. 1838, Miers 4495 (us), 900 m, Dec. 1952, Vidal 11-5784 (R); Santa Ana, 1822-1823, Beyrich s.n. (P, s); Montagnes pres Sumidoro, Laland s.n. (P); Cantagallo, Peckolt 181 (w); ad confines Rio de Janeiro-Minas Ger- ais, in graminosis partis superioris montis Itatiaia, 2000-2500 m, Sep. 1901, Wettstein & Schiffner s.n. (w); without locality, 1834, Gaudichaud 442 (G, P), Raben 703 (BR), 1840, Regnell 173 (s), 1832, Riedel s.n. (GH, GOET, L, LE, NY, P, s, us, w), St. Hilaire A-746 (P), 1838-1842, U.S. South Pa- cific Exploring Expedition s.n. (NY), Oct. 1916, Vincent s.n. (L), Widgren 486 (s); *Morro Quei- mado, Almeida 1286 (F); *Floresta da Tijuca; Ped- ra do Archer, Angeli 250 (F); *Morro Queimado, Brade 18785 (F); *Rio de Janeiro, Pedra da Gav- ea, base da Chamine, Carauta 2420 (F); *Gua- nabara; Pedra da Gavea, Mesa, Carauta and de Oliveira 1523 (F); *Guanabara; Tijuca, Castellan- os 23973 (F); *Municipio de Macae, Pico de Fra- de, Farney et al. 584 (F); *Municipio de Macae, Pico de Frade, Farney et al. 597 (F, RB); *Rio de Janeiro; Alto de Boa Vista, Estrada da Vista Chi- nesa, mata atras do Decam, Landrum et al. 4150 (F); *Tijuca, Bom Retire, Pabst 7411 (F); *Estrada da Vista Chinesa, Strang 172 (F); *Pedra da Gav- ea, Sucre 7446 (F); *Morro Queimado, Sucre 8002 (F); *Serra da Carioca, no topo do Morro Queimado, Vianna et al. 1574 (F). Sao Paulo: Jundiahy, region of Morro da Mursa, 1000 m, 4 Apr. 1915, Brade 7020 (SP); Sao Paulo, Vila Ema, Dec. 1932, Brade 12119 (R), Jan. 1952, Brade 21218 (RB); Campos do Jordao, 10 Sep. 1937, Campos Porto 3381 (R), Campos Porto s.n. (GH, RB), 22 Oct. 1938, Hashimoto 1 (RB), 27 Jul. 1898, Lofgren 3986 (SP); Atibaia, Nov. 1931, Constan- tino s.n. (RB); Guaratingueta, 27 Jan. 1920, Gehrt 3667 (SP); Mogi das Cruzes, 18 Apr. 1921, Hoeh- ne 5481 (SP, us), Oct. 1833, Riedel 1466 (G, LE, NY, us); Sao Jose dos Campos, 17 Dec. 1908, Lof- gren 86 (RB, s); ca. 7.7 km SW em linha reta da pra9a principal de Sao Jose dos Campos, 300 m, NW da Via Dutra, 600 m, 25 Oct. 1961, Mimura 53 (us); Ipiranga, Dec. 1912, Luderwaldt 1086 (RB, SP), Jun. 1907, Usteri 23/306 (NY); Taubate, Lund 754, Lund 756 (c); inter Lorena et Taubate, Apr. 1833, Riedel 1465 (K, LE, NY); Serra da Bo- caina, Jan. 1925, Lutz s.n. (R); in nemorosis editis retro Bananal in via publica versus Sao Paulo, Dec. 1817, Martius s.n. (M); Morro Formozo, Dec. 1817, Martius s.n. (M); Serra da Mantiqueira, Fazenda de Cruzeiro do Dr. Major Novaes, 3-12 Jan. 1884, Saldanha 8587 (R); *Municipio Sao Jose de Barreiro; Parque Nacional da Serra da Bo- caine, Fazenda das Garrafas, Lima, H.C. de, & Ramamoorthy 1224 (F); *Municipio Registro; Rod. Registro-Sete B arras, Hatschbach & Gui- mardes 45004 (F); *Serra da Bocaina, W. Hoehne 6154 (F), *W. Hoehne 6155 (F); *Salesopolis; Es- ta£ao Biologica de Boracea, perto do Rio Coruja, Mattos, J., & Mattos, N., 14284 (F); * Serra da Bocaina; Parque Nacional, Occhioni 9174 (F). Without state: Bowie & Cunningham 228 (BM, s), 1842, Dupre s.n. (BR, NY, P), Germain 171 (F), Guillemin 171 (G), Lobb s.n. (K), Martius 1292 (G, L, M, NY), Pohl 263 (PR), 265 (PR), 266 (PR), Schiich s.n. (BR, w), Sellow s.n. (BM, BR, CGE, F, K, L, M, s), Weir 23 (CGE, K). lOb. Brunfelsia brasiliensis subsp. macrocalyx (Dusen) Plowman comb, et stat. nov. Brunfelsia hopeana van macrocalyx Dusen, Archiv. Mus. Nac. Rio de Janeiro 13: 94. 1905. Type: BRAZIL, Rio de Janeiro, na mata da encosta da Serra do Itatiaia, in silva, 1600 m, 5 Jan. 1896, Ule 636 (holotype, R 66551). Leaves 3-8 cm long, 1.2-2.2 cm wide, gla- brous and often nitid above, glabrescent beneath or with pubescent midrib and nerves, subcoria- ceous to coriaceous. Inflorescence short, subses- sile, terminal, with 2-8 flowers, rarely reduced to 1 . Peduncles 2-5 mm long, usually much shorter than pedicel, sparsely long-pilose or densely glan- dular-pubescent. Pedicel 5-15 mm long, villous or glandular-pubescent. Calyx 15-22 mm long, pilose or glandular-pubescent or glabrescent. Co- rolla tube 1 8-25 mm long, scarcely exserted from 66 FIELDIANA: BOTANY calyx or up to H/2 times as long; limb 25-32 mm in diameter. DISTRIBUTION — Brazil (Minas Gerais, Parana, Rio de Janeiro, Santa Catarina, Sao Paulo). SPECIMENS EXAMINED — BRAZIL. Minas Ger- ais: Mun. Diamantina, Guinda, 5 Nov. 1937, Mel- lo Barreto 9479 (F, R). Parana: Ipiranga, Monte Alegre, 1100 m, Dusen 3388 (R); Bauhado, 13 Dec. 1911, Dusen 13562 (GH, NY, s); Guaratuba, 21 Feb. 1952, Reitz 4375 (us); *Palmeira, Hatsch- bach 11878 (MBM); *Barra Rio Papagaios (Balsa Nova), Hatschbach 18747 (MBM); *Municipio Guaratuba; Rio Sai, Hatschbach 19447 (MBM); *Municipio Jaquariaiva; Fazenda Chapada S. An- tonio, Hatschbach 20997 (GH); *Municipio Pir- quara; Serra do Emboque, Hatschbach 26532 (MBM); *Municipio Pirai do Sul; Joaquim Murtin- ho, Hatschbach 39209 (GH, MBM); *Municipio Palmeira, Rio Tibagi, Kummrow 970 (GH); *Bran- dalize, Lindeman and Haas 1174 (u). Rio de Ja- neiro: Alto Macahe de Nova Friburgo, 22 Jan. 1874, Glaziou 6908 (BR, c, s), Glaziou 13478 (c, F, LE); 1882, Glaziou 14172 (c, G, LE); Petropolis a Itamaraty, 25 Mar. 1879, Glaziou 11395 (BR, c, G, LE, R); Serra dos Orgaos, 18 Nov. 1887, Gla- ziou 17169 (BR, c, p, R), 2,200 m, Jun. 1915, Liitz- elburg 6187 (M); Morro Assu, 2400 m, Jan. 1916, Lutzelburg 6559 (M, NY), Feb. 1838, Miers s.n. (BM); Serra do Itatiaia: 1900 m, Sep. 1913, Brade & Tamandare 6576 (SP), 1915, Campos Porto 173 (R, RB), 2100 m, Jun. 1902, Dusen 569 (R), 18 Oct. 1903, Dusen 2023 (K, R, s, us), 1750 m, 28 Oct. 1927, Ginzberger 93 (F), Moreira s.n. (R); Parque Nacional de Itatiaia, km 15, 17 Aug. 1948, Occhioni 1131 (GH, RB); prope Registro, 1800 m, 14 Nov. 1954, Pabst & Brade 10283 (us); Ma- cieras, 1900 m, Sep. 1934, Brade 14074 (B, R, RB), 1800-2000 m, 23 Oct. 1927, Zerny s.n. (w); entre Maromba e Macieiras, 1820 m, 29 May 1969, Plowman 2891, Sucre 5191 (ECON, GH, MICH, PH, RB, uc, YEN), 1,700 m, Plowman 2906, Sucre 5206 (GH, MICH, PH, RB), 1200-1900 m, 22°27'S, 44°39'W, 10 Jan. 1929, L.B. Smith 1788 (GH, s, us), 26 Feb. 1965, Vogel 677 (us); *Parque Na- cional de Itatiaia, Lindeman & Haas 5157 (u); *Parque Na9ional de Itatiaia; entre Maromba e Macieras, Sucre 5806 (F). Santa Catarina: Mun. Salete, Taio, Ribeiro Grande, 16 Dec. 1950, Reitz 3985 (G, NY, uc, us); Mun. Rio do Sul, Alto Mat- ador, 800 m, Reitz & Klein 7601 (us). Sao Paulo: Alto da Serra, 2 Feb. 1913, Brade 6037 (s, SP), Brade 7625 (us), 2 Dec. 1920, Hoehne 4595 (SP, us), 22 Dec. 1919, Hoehne 8250 (SP, us), 27 Nov. 1902, Puttemans 5890 (SP); Serra da Bocaina, 1650 m, 12 May 1951, Brade 20953 (RB); Fazen- da Campo Grande, Linha Ferrea Ingleza, 17 Nov. 1892, Edwall 1981 p.p. (B, us); Esta?ao Biologica, 800-900 m, lat. 23°46'S, long. 46°20'W, 23 Feb. 1929, Smith 1989 (GH, us); Campos do Jordao, Umuarama, 1,750 m, 22 Nov. 1949, M. Kuhlmann 2338 (SP), 22 Nov. 1957, M. Kuhlmann 4309 (SP); proximo do Rio Guaratuba, 18 Mar. 1958, M. Kuhlmann 4359 (SP). A name that has been in the synonymy of this species, Gerardia brasiliensis Spreng., was first published in 1825. Although it was not mentioned by Pohl, both Bentham and Schmidt placed it in synonymy with B. ramosissima. This specific ep- ithet was only recently recognized by Smith and Downs (1966) as the first legitimate name and basionym for the species, which now must be called B. brasiliensis (Spreng.) L. B. Sm. & Downs. Sprengel described G. brasiliensis from a spec- imen collected by Sellow in Brazil, and his de- scription, albeit brief, more or less agrees with the present concept of the species. There is, however, one point of difference in Sprengel's description, where he writes "pedunculis axillaribus solitar- iis." Usually this species has terminal, branched inflorescences, although there may be subterminal axes produced in the uppermost leaf axils. Unfortunately, Sprengel's herbarium was dis- mantled and sold after his death by his son. Some of the specimens survived and were eventually deposited at the Berlin herbarium, where they were destroyed in the Second World War (Stafleu, 1967; Stafleu & Cowan, 1985). Sellow apparently made several collections of this species with nu- merous duplicates, none of which bears collection numbers or localities. It is thus impossible to de- termine which specimens might be isotypes of the original specimen that Sprengel described. As a representative collection, I selected another spec- imen of Sellow that was sent to Munich from Ber- lin in exchange [this is not to be taken as neotyp- ification of the name (see footnote 11) — Eds.}. In 1826 Pohl published seven new species of his new genus Franciscea, including F. ramosis- sima, F. acuminata, F. confertiflora, and F. di- varicata. He stated at that time that these "spe- cies" were all closely related but believed that their differences were great enough for them to be considered separate entities. Pohl's specific concepts were generally followed by Bentham, who in 1846 transferred all of Pohl's species of PLOWMAN: REVISION OF BRUNFELSIA 67 Franciscea to the genus Brunfelsia, omitting only F. divaricata, which he placed in synonymy with B. confertiflora. According to Bentham, B. con- fertiflora differed from B. ramosissima in having a many-flowered, crowded inflorescence, an in- flated calyx, and the corolla tube only W2 times as long as the calyx. Brunfelsia acuminata as maintained by Ben- tham was merely a glabrous form of B. ramosis- sima. The type specimen from which Pohl's orig- inal drawing was made is preserved in Brussels (BR). A specimen at Vienna (w), collected and ap- parently annotated by Schott as F. acuminata, has a well-developed indument and does not conform to the description of this species. As mentioned above in the discussion of B. martiana, there has been considerable confusion of collection data and numbers in the Brazilian collections of Pohl and Schott. For example, the same collection in different herbaria may bear the same number or numbers with either or both names. For this rea- son the numbers of these two collectors should be used with some caution. Schmidt (1864), in his account of Brunfelsia for Flora Brasiliensis, treated this group of "species" as a single variable species, B. ramosissima, with three varieties. Variety laxiflora included B. ra- mosissima and B. acuminata of Bentham, and va- riety confertiflora conformed more or less with his B. confertiflora. Schmidt described a third va- riety, parcifolia, for a form with small, coriaceous leaves. Annotations in Schmidt's handwriting on the type and other specimens read "parvifolia," indicating either a typographical error in publi- cation or that he changed his mind. The former is more likely since these plants have quite small, but not few, leaves. This form is now considered only a minor variant. Brunfelsia brasiliensis is here treated as a sin- gle polymorphic species, widely distributed in southern Brazil, with one subspecies, macrocalyx, occurring at higher elevations from the Serra do Mar and Serra da Mantiqueira southward in the mountains to Santa Catarina. Brunfelsia bras- iliensis exhibits a wide range of morphological variation throughout its area of distribution, which consists mostly of mountainous, ecologically di- versified terrain. In this region, primarily in the states of Minas Gerais, Rio de Janeiro, and Sao Paulo, there exist ample opportunities for geo- graphical and ecological isolation of populations with subsequent morphological differentiation. Environmental factors also play an important part in the phenotypic variability seen in this spe- cies, and different forms may be interpreted as ecotypes. For example, in exposed, well-drained places such as the crest of a peak or rocky out- crop, the plants are frequently low in stature with horizontal branches, small, often coriaceous leaves, and fewer and smaller flowers. Lower down the same slope, in shaded, more protected, and probably more fertile places, the plants may attain a height of up to 3 m with well-developed inflorescences and larger flowers and leaves. The great morphological variation in B. bras- iliensis occurs chiefly in the presence, type, and location of indumentum, in the size and form of the inflorescence, and to a lesser extent in the size and form of the flowers and leaves. An important diagnostic feature of this species is the presence of a persistent peduncle, a short stalk that articulates with the pedicel of each flow- er. This structure is scarcely, if ever, developed in any other species of Brunfelsia. Some populations of B. uniflora, however, may have a similar stalk in its solitary flowers. This "peduncle" is perhaps more accurately described as an ultimate branch of the inflorescence that immediately precedes a flower. It commonly bears 1-3 small bracts near the base, the axillary buds of which may produce additional flowers, thereby proliferating the inflo- rescence. The peduncle may be shorter or longer than the pedicel or, more rarely, obsolete. The dis- tal end is dilated into a short, flat torus that be- comes conspicuous when the flower falls. The inflorescence arises at the tips of the bran- chlets and is usually much-branched and many- flowered. It may be strictly terminal and compact to lax and open with several subterminal branches also developing into flowering axes. Alternatively, the flowers may be reduced to a few, often sub- sessile ones and rarely to a single flower in sub- species macrocalyx. The calyx in B. brasiliensis may be tubular to nearly campanulate in form and varies greatly in size. The corolla tube may be scarcely exserted from, or up to twice as long as, the calyx. The presence of a well-developed indument, particularly in the inflorescence, is also diagnostic in B. brasiliensis. The trichomes are nonglandular, uniseriate, long or short, and straight or curved. In dried material, they are frequently yellowish or yellowish brown. Often the indument may be densely villous to tomentose. More rarely it ap- pears quite sparse and pilose, or the plants may be glabrescent. The upper surface of the leaves is frequently glabrescent, with short, subscabrous hairs that break off near the base. The lower sur- 68 FIELDIANA: BOTANY face of the leaves is usually pubescent, especially at the nerves and midrib. Brunfelsia brasiliensis is easily distinguished from most other species of the genus by the reg- ularly oblong-lanceolate leaves, the distinctive yellowish brown pubescence, and the presence of the short peduncle articulating with the pedicel. Rare glabrous forms of B. brasiliensis (= B. ac- uminata (Pohl) Benth.) resemble B. bonodora but are distinguished by their stalked pedicels. Brun- felsia brasiliensis appears to be most closely re- lated to the B. bonodora-B. grandiflora complex by the nature of the more or less branched inflo- rescence and floral and fruit morphology. Brunfelsia brasiliensis subsp. brasiliensis is distributed in southeastern Brazil from Minas Gerais south to Sao Paulo, between 700 and 1400 m elevation. The species shows a fairly wide eco- logical tolerance but prefers well-drained sites. It is frequently encountered in the mares de morros, "the seas of hills," which occur in southeastern Brazil, particularly in Minas Gerais. Here the plant grows on granitic peaks and rocky outcrops, often in dry, exposed habitats. It seems to occur less commonly in the humid forests on the slopes of the Serra do Mar. It also grows in association with the campos, or open grassland communities, and in capaes, small patches of woodland found scattered in the campos. Brunfelsia brasiliensis is also encountered in capoeiras, a general term for cut-over, secondary woods, and is one of the few brunfelsias that survive cutting of the primary for- ests. Around Brasilia, it has been reported from gallery forests. Brunfelsia brasiliensis is a freely flowering plant with the flowers appearing pri- marily from September to March. The fruits have been collected from October to March. This spe- cies has been cultivated in European conservato- ries since its discovery, although it is much rarer in cultivation than other species. To my knowl- edge, it is not planted in the American tropics outside of Brazil. Peckolt (1909) says that it is a common garden plant in that country. Brunfelsia brasiliensis is known as manacd, manacd da ser- ra, erva de macaco, and manacd assu (Peckolt, 1909). From herbarium labels (Occhioni s.n., V. Assis 102, 142) we know that it is considered poi- sonous. At higher elevations in the mountains of Rio de Janeiro, Sao Paulo, and south to Santa Catarina, there occurs a distinct subspecies of B. bras- iliensis. This was first described by Dusen in 1905 as B. hopeana var. macrocalyx from a specimen collected on Mount Itatiaia. This type superficial- ly resembles B. uniflora (=B. hopeana) in having the inflorescence reduced to a single flower, but this is somewhat exceptional for the subspecies as now circumscribed. Brunfelsia brasiliensis subsp. macrocalyx dif- fers from B. brasiliensis subsp. brasiliensis in having fewer flowers per inflorescence (one to eight), a very short peduncle in which the flowers may appear sessile, a longer pedicel (5-15 mm) that always exceeds the peduncle, and a usually longer calyx (15-22 mm). The corolla tube of subspecies macrocalyx is short relative to the ca- lyx and is just barely exserted. The indumentum is frequently less well-developed, especially in the leaves, which may be glabrescent and bright green. In addition, the trichomes may be glandu- lar, especially in the calyx and pedicels, a char- acter not normally encountered in subspecies brasiliensis. Morphologically intermediate populations are known to exist between subspecies brasiliensis and subspecies macrocalyx. In collections from the Serra dos Orgaos and Alto Macahe in the state of Rio de Janeiro, subspecies brasiliensis shows a tendency to develop a large calyx (to 17 mm) similar to that of subspecies macrocalyx. How- ever, because these plants are, in general, larger in all parts, they may be considered exceptionally vigorous individuals perhaps growing in a partic- ularly favorable place (cf. Glaziou 6908, 13478, 17169). Other collections (cf. Occhioni 1131) from Mount Itatiaia are referable to subspecies macrocalyx, although they exhibit a multiflorous inflorescence. These plants may indeed suggest putative hybridization between the two subspe- cies; however, this cannot be proved at this stage of knowledge of the group. Several collections from the states of Parana and Santa Catarina have been referred to B. brasiliensis subsp. macrocalyx (e.g., Dusen 13562, Reitz 3985). These popula- tions are somewhat heterogeneous and in some characters resemble B. cuneifolia (see discussion under that species), such as the indument and the reduction in number of flowers, often to one. Fur- thermore, the calyx tends to be smaller than in more northern populations of this subspecies, somewhat resembling that of subspecies bras- iliensis. Because of the complex pattern of variation that is encountered in this region, I believe this problem cannot be resolved completely without additional materials and a knowledge of the plants in the field. Brunfelsia brasiliensis subsp. macrocalyx in the PLOWMAN: REVISION OF BRUNFELSIA 69 northern part of its range occurs at altitudes be- tween 1650 and 2400 m on relatively few moun- tain peaks and high campos: Mount Itatiaia, Alto Macahe, and Serra dos Orgaos in Rio de Janeiro; and Alto da Serra, Serra de Bocaina, and Campos do Jordao in Saao Paulo. In Sao Paulo (Campo Grande) and southward, subspecies macrocalyx occurs at somewhat lower elevations (800-900 m). Ecologically, it grows in wet, montane cloud forests in association with Podocarpus and also occurs in moist, high-elevation campos. In the Parque Nacional de Itatiaia, plants have been fre- quently collected between Maromba and Maci- eiras at 1600-1900 m elevation and are repre- sented by two distinct forms. One of these (cf. Plowman 2891, Sucre 5191) is sparsely branched with dark leaves and pubescent calyx with erect lobes. The other (cf. Plowman 2906, Sucre 5206), which grows sympatrically a few hundred yards away, is profusely branched with bright green, glabrous leaves and a glabrous calyx with strong- ly recurved lobes. Though strikingly different to the casual observer, these plants must be regarded as minor phenotypic variants of a single subspe- cies. Analysis in the field of situations such as this gives a measure of the kind of variability that is present in a population and a better basis for un- derstanding comparable problems where field studies are not possible. Efforts were made to rear these plants in the greenhouse from seed to study the nature of the characters and their inheritance. However, the seedlings proved to be exceedingly sensitive to fluctuations in temperature and humidity and soon died. 11. Brunfelsia chiricaspi Plowman, Bot. Mus. Leafl. 23(6): 255, t. 17. 1973. Type: CO- LOMBIA. Putumayo: Umbria, 0°54'N, 76°10'W, 325 m, forest, shrub 1.5 m, flowers sky blue, zanango, medicinal, Oct. -Nov. 1930, Klug 1810 (holotype, A; isotypes, F, s, us 1456539). Figure 25. Shrub or treelet 1-3 m tall. Trunk to 5 cm in diameter near base. Bark thin, roughish, grayish brown. Branches few, lax, spreading, naked. Branchlets glabrous, light brown to ochraceous, shiny, outer bark thin. Leaves scattered along branchlets or somewhat crowded, blade 20-30 cm long, 7-12 cm wide, elliptic to lanceolate, some- times obovate, apically obtuse with short, subfal- cate acumen or acuminate, cuneate to blunt at base, glabrous, dull, dark green above, pale green beneath, smooth, subcoriaceous, lateral nerves 8- 10, straight; petiole 5-10 mm long, short, stout, glabrous, dark brown, roughish. Inflorescence corymbiform, terminal or axillary, usually few- flowered with 4-7(20) flowers, puberulent or gla- brous. Bracts 1-2 mm long, lanceolate, concave, glabrous. Flowers sky blue to violet, fading white, with 5-angled white spot at mouth. Pedi- cels 6-13 mm long, slender, glabrous. Calyx 10- 13 mm long, 4-6 mm in diameter, tubular-cam- panulate, slightly inflated, glabrous, purplish to pale green, teeth short, broadly triangular, acute to blunt with short glandular acumen; calyx in fruit coriaceous, striately nerved, dotted with len- ticels. Corolla tube 22-25 mm long, 3 mm in di- ameter, twice as long as calyx, straight, glabrous; limb 25-30 mm across, thickening at mouth prominent, fleshy, 5-angled, lobes subequal, strongly deflexed at anthesis, the uppermost slightly larger, rounded. Stamens included in up- per part of corolla tube; filaments subligulate, lower inner pair 2.5 mm long, upper outer pair 3.5 mm long, reaching the mouth; anthers to 1.5 mm long, orbicular-reniform, light brown. Ovary 2 mm long, ovoid-conical; style filamentous, equaling filaments; stigma briefly bifid, obtuse, the upper lobe somewhat larger. Capsule about 10 mm long, 8 mm in diameter, dry at maturity, subglobose. Seeds few, 6 mm long, 2.5 mm in diameter, ellipsoid-reniform, reticulate-pitted. DISTRIBUTION — Colombia, Ecuador. See Figure 23. SPECIMENS EXAMINED — COLOMBIA. Putu- mayo: Orito, near Texaco drilling site, 11 Feb. 1972, Kennedy 1386 (GH); alrededores de Puerto Limon, 300-400 m, 15 Feb. 1953, Mora 1040 (COL); El Whiskey at Finca "Santa Marta," 13 km S. of Umbria near road, 400-500 m, 27 Nov. 1968, Plowman 2080 (ECON, GH); forest about 2 km SW of San Pedro, N of Puerto Asis, 400-500 m, 28 Nov. 1968, Plowman 2081 (ECON, GH); Rio Guamues, San Antonio, 5 Sep. 1966, Pinkley 420 (ECON); Santa Rosa, 28 Nov. 1966, Pinkley 563 (ECON); trail between Santa Rosa and road to Hor- miga, 2 Dec. 1968, Plowman 2092 (ECON, GH); *Rio Putumayo, Puerto Ospina, Schultes 3422 (GH). ECUADOR. Napo: * Along Rio Aguarico upstream from Santa Cecilia, Duke 16047 (MO); *Coca (Puerto Francisco de Orellana), S side of Rio Napo, Marling & Andersson 11771 (GB); *Coca (Puerto Francisco de Orellana), Marling & Andersson 11804 (GB, GH); *Hacienda Cotapino 70 FIELDIANA: BOTANY BRUNFELSIA chiricaspi Plowman FIG. 25. Brunfelsia chiricaspi. Reproduced courtesy of the Botanical Museum of Harvard University. (Conception), Marling et al. 7021 (GB, GH); *Path Cotapino (Conception), Rio Bueno, Marling et al. 7151 (GB, GH); *Coca de Orellana, lower Rio Pay- amino, Marling et al. 7772 (GB, GH); *Hacienda San Carlos at Rio Napo, ca. 15 km E of Coca (Puerto Francisco de Orellana), Lugo 2724 (GB, GH); *Tierra Colorada, ca. 3 km N of Coca (Puerto Francisco de Orellana), Lugo 2942 (GB, GH); *Ca- non de los Monos, road Coca (Puerto Francisco de Orellana), Lago Agrio, ca. 12 km N of Coca, PLOWMAN: REVISION OF BRUNFELSIA 71 Lugo 2981 (GB, GH); *road from Lago Agrio-E 1 Chaco, Lugo 3476 (GB, GH); *7 km N of Coca; Estacion Experimental de INIAP-Payamino, Zar- uma et al. 56 (F). Brunfelsia chiricaspi was originally known only from a few collections from a small area of southwestern Colombia on the eastern flank of the Cordillera Occidental. [Plowman predicted that the distribution would prove to be much greater, and this species is now also known from many recent collections in adjacent Ecuador (see above). — Eds.] It grows as an understory shrub in humid, primary forests at elevations of 325-500 m. The fruit is as yet imperfectly known because of a lack of material. Brunfelsia chiricaspi is related to B. grandiflora, which is widespread in the western Amazon, and to B. mire of Bolivia and adjacent Peru and Brazil. Brunfelsia chiricaspi differs from B. grandiflora in having larger leaves, longer pedicels (6-13 vs. 2- 6 mm) and usually fewer flowers. It differs from B. mire in the basally obtuse, elliptic to lanceolate leaves; the few-flowered cyme; and smaller flowers (corolla tube 22-25 vs. 25-38 mm long). Known generally as chiricaspi in the Colom- bian Putumayo, this species is used as a strong medicine and narcotic by the Kofan Indians (see Plowman, 1977). 12. Brunfelsia cuneifolia J. A. Schmidt in Mart., Fl. Bras. 8(1): 259. 1864. Type: BRAZIL. In Brasilia australiore, Sellow 4016 (lectotype, designated here, destroyed at Berlin; photo- graph of lectotype, F 621824, NY, us; isolec- totypes, F, M).13 Riedel 1467, also cited by Schmidt, is excluded as a type. It belongs to B. obovata Benth. var. obovata. Figure 26. Shrub to about 1 m tall, much branched. Branches terete, naked. Bark grayish to yellow- ish brown. Branchlets short, leafy, sparsely to densely villous, rarely glabrescent, hairs often yel- lowish to reddish brown, sometimes glandular. 13 It is clear that Plowman was intending to lectotypify this species using the destroyed B sheet as good photo- graphs of it exist. He appears to specifically not consider the unnumbered Sellow sheets at F and M as good can- didates for lectotypes (in the strict sense). — Eds. Leaves scattered or crowded near the tips of the branchlets, blade 3.5-10 cm long, 1.8-3 cm wide, oblong-obovate to elliptic-lanceolate, cuneate to blunt at apex, rarely acuminate, narrowed to broadly cuneate at base, the upper surface gla- brescent or with long, scattered hairs, the lower surface velutinous, often densely so at midrib and nerves, rarely glandular or glabrescent, somewhat shiny above, dark green, pale green beneath, firm- ly membranaceous to subcoriaceous, lateral nerves 4-7; petiole 2-4 mm long, velutinous or glandular. Inflorescence terminal on tip of branchlets, sessile, with 1-3 flowers. Bracts 2-8 mm long, small, lanceolate, glandular, ciliolate, caducous. Flowers violet fading to white with age. Pedicel 3-5 mm long, slender, thickened at apex, velutinous or glandular pubescent. Calyx 12-18 mm long, 4-6 mm in diameter, tubular, in- flated, 5-angled, drying plicate, pubescent with short glandular hairs, especially near base, or vil- lous, pale green, membranaceous, teeth 3-7 mm long, ovate-lanceolate, acute to acuminate at apex. Corolla tube 18-28 mm long, 1.5-2 mm in di- ameter, \Vi times as long as calyx, pentagonal at base, pilose or minutely glandular in upper part; limb 25-37 mm in diameter, spreading, lobes 10- 15 mm long, undulate, broadly rounded. Stamens inserted in upper part of tube; upper pair of fila- ments 3-4 mm long, suberect, exserted slightly from mouth; lower pair 2-3 mm long, included, curved at apex; anthers 1.5 mm long, orbicular- reniform. Ovary 2 mm long, conical-ovoid, style about 23 mm long, filamentous; stigma 1 mm long, briefly bifid. Fruit and seed unknown. DISTRIBUTION — Brazil (Parana, Rio Grande do Sul, Santa Catarina, Sao Paulo). See Figure 28. SPECIMENS EXAMINED — BRAZIL. Parana: Mun. Mangeirinha, Lageado San Antonio, 20 Oct. 1966, Hatschbach 15159 (US); *Municipio de Guarapuara; Guarapuara, Frenzel 666 (MBM); *In- acio Martins, Hatschbach & Guimardes 30302 (MBM); *Municipio Campo Largo; Serra do Pu- runa, Kummrow 2432 (F). Rio Grande do Sul: Farroupilha, 15 Oct. 1957, Camargo 2123 (B), 3 Oct. 1957, Camargo s.n. (s); *Municipio Cacequi, Dobereiner & Tokarnia 732 (RB); *Esmeralda; Estacion Ecologica de Aracuri, Jarenkow 132 (F). Santa Catarina: Mun. Lajes, Passo do Socorro, FlG. 26. Brunfelsia cuneifolia. 72 FIELDIANA: BOTANY I \ N? 1995963 FIELD MUSEUM OF NATURAL HISTORY ~t» PREFEITURA MUNICIPAL DE CURITIBA MUSEU BOTANICO MUNICIPAL i io li -a ZT- A- icuaa Sta. Clara d 50573 Arbuato 2 B| flor lllas. Interior aata te gal«ria. PMC • Off 004 PLOWMAN: REVISION OF BRUNFELSIA 73 900 m, 31 Oct. 1963, Klein 4343 (us); Mun. Novo Horizonte, Lauro Miiller, 400 m, 24 Oct. 1958, Reitz & Klein 4076 (NY, us); Campos Novos, 12 Sep. 1963, Reitz & Klein 16158 (GH, us); Mun. Campos Alegre, lower fazenda of Ernesto Schei- de, Campo Alegre, 900 m, 11 Dec. 1956, Smith & Klein 8567 (us); Mun. Chapeco, Guatambu, ca. 27°06'S, 52°45'W, 350-400 m, Smith & Reitz 12541 (GH, R, NY, us); Mun. Irani, Campo de Irani, ca. 26°57'S, 51°50'W, 700-900 m, 8 Nov. 1964, Smith & Klein 13044 (GH, MO, p, R, uc, us). Sao Paulo: Ilha do Cardoso, perto de Cananeia, Prance et al 6933 (F). In his original description of B. cuneifolia in Martius's Flora Brasiliensis (1864), Schmidt cited two syntypes. One of these, Riedel 1467, is clear- ly B. obovata of Bentham. The other, which now serves as the type, was collected by Sellow in southern Brazil, but no specific locality was giv- en. The specimen cited by Schmidt (Sellow 4016) was destroyed at the Berlin herbarium; two un- numbered Sellow duplicates, at Chicago (F) and Munich (M), which conform in every way with the photograph of the type, may be considered isolectotypes. Known from only a few collections, the avail- able material of B. cuneifolia shows a consider- able amount of variation in the form of the leaves and type and amount of indument. Brunfelsia cu- neifolia is distinguished from other species of southern Brazil by the apically cuneate leaves and a five-angled tubular calyx that appears plicate when pressed and dried. The leaves and calyces are always pubescent with either of two types of trichomes present. The type collection and mate- rial from Rio Grande do Sul have short glandular hairs. Most specimens from Santa Catarina have a dense villous to velutinous indument consisting of longer, nonglandular hairs. The form of the leaves and calyx associated with both types is fairly uniform. Brunfelsia cuneifolia is not easily distinguished from other species of southernmost Brazil. It su- perficially resembles B. pilosa, which is sympatric with B. cuneifolia. Brunfelsia pilosa, however, usually has a more pronounced acumen in the leaf and a terete, tubular calyx that is purple and bears only a few, long pilose hairs. In addition, the flowers of B. pilosa are somewhat larger than those of B. cuneifolia. The villous pubescence of B. cuneifolia is very reminiscent of the yellowish indument character- istic of B. brasiliensis. Populations interpreted here as B. brasiliensis subsp. macrocalyx extend as far south as Santa Catarina and overlap with the range of B. cuneifolia. Collections of these two species in this region are morphologically very similar and may be easily confused. Brun- felsia brasiliensis subsp. macrocalyx may be sep- arated by the presence of the persistent peduncle, which articulates with the pedicel of the flowers, and by the terete, campanulate or tubular-campan- ulate calyces. This species usually has longer ped- icels and shorter calyx teeth than B. cuneifolia. Nevertheless, the intermediate nature of these populations indicates that some hybridization may have occurred between these two species and pos- sibly involved B. pilosa as well. The true affinities of B. cuneifolia appear to lie with B. obovata var. obovata, which occurs far- ther north in Sao Paulo and Minas Gerais [but see also B. boliviana. — Eds.]. This plant resembles B. cuneifolia in the tubular pentagonal calyx, which also nearly equals the corolla tube in length, and in the obovate leaves. In both species the upper- most pair of anthers may be briefly exserted from the corolla tube, a character that is unusual in sec- tion Franciscea. Brunfelsia cuneifolia differs in having the leaves cuneate at the apex, fewer flow- ers per inflorescence, and a pubescent calyx that is much more slender than in B. obovata (4-6 vs. 6-10 mm). These two species also differ striking- ly in their ecological preference. Brunfelsia obov- ata is a plant of marshes and flooded areas, whereas B. cuneifolia occurs in uplands. Brunfelsia cuneifolia is a relatively low shrub that is found in habitats apparently similar to those of B. pilosa. It is specifically reported from Araucaria forests (arau-carieto) and also from gallery forest and capoeiras, or secondary forest, and between 400 and 900 m altitude. It flowers in October and November and is known by the com- mon name flor de Trovoada, or "thunderstorm flower." 13. Brunfelsia dwyeri D'Arcy, Ann. Missouri Bot. Gard. 57:259. 1971 ("1970"). Type: PANAMA. Panama: Cerro Jefe in "Cleusia" FIG. 27. Brunfelsia dwyeri. 74 FIELDIANA: BOTANY 1709790 FIELD MUSEUM OF NATURAL HISTORY PLANTS OF PANAMA Brunfelsia dwyeri D'Arcy det. by R.L. Wilbur, 1971. Provincia de Panama': slopes of Cerro Jefe beyond Cerro Azul between 4-8 miles in mostly heavily wooded slopes. Common shrub 2-5 m. tall. Corolla deep violet blue becoming paler with age. R.L. Wilbur R.E. Weaver #11358 25 January 1970 PLOWMAN: REVISION OF BRUNFELSIA 15 (sic) forest, 2700-3000 ft, shrub, flowers pur- ple to whitish purple, 27 Jan. 1966, Tyson, Dwyer & Blum 3312 (holotype, MO 1820936; isotype, MO 1917560). Duke & Dwyer 13933 (MO) is excluded from the cited paratypes. Figure 27. Shrub or small tree 2-10 m tall. Trunk to 6- 8 cm in diameter, branched from near the base. Bark gray to grayish brown. Branches terete or somewhat angular, naked. Branchlets leafy. Leaves with blade 5-11 cm long, 2.5-5 cm wide, obovate, elliptic or oblong-lanceolate, acuminate at apex, slightly revolute at margin, cuneate or narrowed at base, glabrous, shiny and light to very dark green above, pale green beneath, cori- aceous, lateral nerves 5-7, veinlets reticulate, prominulous, especially on lower surface; petiole 3-10 mm long, becoming transversely corrugate- rugose. Flowers solitary, terminal or axillary in the upper leaf axils, purple, fading to white with age, with irregular white spot at mouth. Peduncle 1-5 mm long, sometimes obsolete. Bracts 1-3 mm long, linear to ovate, puberulent, ciliate-mar- gined, caducous. Pedicel 5-15 mm long, about 2 mm in diameter, glabrous. Calyx 12-22 mm long, 5-10 mm in diameter, tubular to tubular-campan- ulate, inflated or not so, glabrous, subcoriaceous, reticulately veined, yellowish green or purplish, teeth incumbent or erect, triangular to ovate-lan- ceolate, at apex acute or blunt; calyx in fruit co- riaceous, often becoming dotted with lenticels, verrucose near base, splitting completely along one or two sinuses. Corolla tube 25-40 mm long, 2-4 mm in diameter, 2-3 times as long as calyx, glabrous; limb 30-80 mm in diameter, spreading, lobes 10-20 mm long, broadly rounded, the upper lobe slightly larger. Stamens inserted in upper part of corolla tube; filaments slender, terete, up- per pair about 3 mm long, lower pair about 5 mm long; anthers included within corolla tube, 2 mm in diameter, orbicular-reniform. Ovary about 2 mm long, conical, ovoid, weakly 4-angled; style filiform; stigma briefly bifid, the lobes subequal, blunt. Capsule 1.5-2.5 cm long, 1-1.5 cm in di- ameter, globose to ovoid, usually 2-celled, at ma- turity often appearing 4-lobed with 4 prominent cruciform grooves, pericarp to 4 mm thick, tough and leathery, drying hard, surface smooth and green at first, becoming rugulose-verrucose, light brown. Seeds 3-15, 8-10 mm long, 3-4 mm in diameter, oblong-ovoid, prismatic, dark reddish brown, reticulate-pitted. Embryo straight, coty- ledons ovate. DISTRIBUTION — Panama, known only from Cer- ro Jefe, Provincia de Panama. See Figure 28. SPECIMENS EXAMINED — PANAMA. Panama: Cerro Jefe, beyond Goofy Lake, 16 Nov. 1967, Correa & Dressier 471 (MO); along main road before turnoff to summit, 6 Jan. 1971, Croat 13136 (GH, MO); Cerro Jefe, 8 Apr. 1970, D'Arcy, Dressier & Correa 3948 (MO), D'Arcy 3959 (MO), 21 Jan. 1967, Duke 9429 (MO), 2 Apr. 1969, Dwyer, Durkee & Castillon 5051 (MO), 12 Mar. 1968, Dwyer 7280 (GH, MO), ca. 2,900 ft, 29 Jul. 1967, Dwyer & Ganger 7349 (GH, MO); along road from Cerro Jefe to La Eneida (Cerro Azul), Croat 13072 (GH, MO), 12 Feb. 1971, Escobar 359 (COL, ECON, GH), 26 Jul. 1969, Kennedy 316 (ECON), 12 Apr. 1972, Plowman 3160 (COL, ECON, GH, MICH, uc), 9 Jul. 1966, Tyson, Dwyer & Blum 4335 (MO), 2300-2700 ft, 19 Jan. 1969, Tyson 5315 (MO), 25 Jan. 1970, Wilbur & Weaver 11358 (GH). Brunfelsia dwyeri was discovered and de- scribed during studies on the flora of Panama. So far as is known, it occurs only on Cerro Jefe in central Panama between 700 and 1,000 m eleva- tion, an area now famous for its high floristic en- demism (Lewis, 1971). The collection Duke & Dwyer 13933 (det. W. G. D'Arcy, 1973) from Cerro Pilon in Provincia Cocle, which was cited in the original description as B. dwyeri, is actually Schultesianthus megalandrus (Dunal) Hunz. The thick-walled fruit and shiny, coriaceous leaves are distinctive features of B. dwyeri. It ap- pears to be most closely related to B. macrocarpa of coastal Colombia and Ecuador, the flowers of which are similarly large and solitary and which also possesses a thick-walled fruit. The two spe- cies are unique in section Franciscea in possess- ing a fleshy pericarp. The resemblance of B. dwyeri to B. lactea of Puerto Rico, pointed out by D'Arcy (1971), is indeed "superficial" because these species are placed in separate sections of the genus and have no close affinities with one an- other. Brunfelsia dwyeri is a beautiful shrub with at- tractive, shiny foliage and large, showy flowers. It is locally common in the cloud forests on the upper slopes of Cerro Jefe, occurring in secondary woods with seasonal rainfall. This locality and species represent the northernmost extension of Brunfelsia sect. Franciscea. As pointed out by Lewis (1971), the entire forest of Cerro Jefe and surrounding areas is rapidly being destroyed for raising poultry and cattle. Brunfelsia dwyeri and 76 FIELDIANA: BOTANY FIG. 28. square). Distribution of Brunfelsia cuneifolia (solid triangle), B. dwyeri (solid circle), and B. grandiflora (solid many other endemic species of plants are in grave danger of extinction. This species flowers and fruits intermittently throughout the year, although I failed to find flow- ers in mid-April. Local people reported that the bark and roots of this plant are used for a "rem- edy." 14. Brunfelsia grandiflora D. Don, Edinburgh New Philos. J.: 86. Jul. 1829. Type: PERU. San Martin: Uchiza, 1798, Tafalla s.n. (ho- lotype, BM; isotype, MA). Figure 29. Franciscea grandiflora (D. Don) Miers, Ann. Mag. Nat. Hist., ser. 2, 5:250. 1850. Shrub or small tree 1-6(10) m tall. Trunk to 7 cm in diameter near base, much branched. Bark thin, roughish, light to dark brown. Branches slender, ascending or spreading, often subvirgate and arching, leafy, glabrous. Branchlets glabrous, rarely pubescent, green. Leaves more or less two- ranked, scattered along branchlets, blade 6-23 cm long, 2-8 cm wide, lanceolate to oblong, often arched with ascending laminas, apically long to short acuminate, the acumen subfalcate, cuneate to narrowed at base, glabrous or sparingly pubes- cent beneath at midrib, dull, dark green above, paler green beneath, rarely glaucescent or nitid, firmly membranaceous to subcoriaceous; petiole 2-6 mm long, glabrous or minutely pubescent, be- coming corky, transversely corrugate with age. Inflorescence terminal and subterminal, simple or branched, dense or lax, the axis 5-45 mm long. Flowers 5-many, showy, scentless, violet fading to white with age, with rounded, white ring at PLOWMAN: REVISION OF BRUNFELSIA 77 mouth. Bracts 1-3 per flower, 1-5(10) mm long, lanceolate to ovate, ciliolate, pubescent or glabra- te, caducous. Pedicel 2-5(10) mm long, glabrous or with few sparse glandular hairs, becoming thicker and corky- verrucose in fruit. Calyx 5-13 mm long, 3-7 mm in diameter, tubular-campanu- late, somewhat narrowed toward base, globose to obovate in bud, somewhat inflated or not so, gla- brous, rarely punctate or sparsely glandular with- in, smooth or striate-nerved, light yellow-green to gray-green, firmly membranaceous to subcoria- ceous, teeth 1-5 mm long, triangular to ovate, blunt to short acuminate, erect or incumbent, re- curved slightly with age; calyx in fruit persistent, coriaceous, becoming corky-verrucose especially near base, often splitting on one or more sides. Corolla tube 15-40 mm long, 1.5-3 mm in di- ameter, 2-5 times as long as calyx, glabrous, pale violet to white; limb 20-50 mm in diameter, spreading, violet fading to white with age, lobes 7-15 mm long, subequal or uppermost somewhat larger, rounded or emarginate at apex, overlapping at sides, abruptly narrowed at base. Stamens completely included in upper part of corolla tube; filaments thin, upper pair 4 mm long, lower pair 3 mm long, white; anthers 1-1.5 mm long, orbicular-reniform, light brown. Ovary 1.5-2 mm long, sessile, conical to ovoid, pale yellow; style slender, slightly dilated at apex; stigma about 1 mm long, briefly bifid, unequal, the up- per lobe somewhat larger, obtuse, green. Fruit 8-20 mm long, 8-20 mm in diameter, ovoid to subglobose, obtuse or apiculate at apex, smooth, nitid, dark green turning brownish, with corky- punctate or -verrucose outgrowths, pericarp thin, 0.3 mm thick, crustaceous, drying brittle, tardily dehiscent. Seeds 10-20, 5-7 mm long, 2-3 mm in diameter, variable in shape, ellipsoid to ob- long, angular, dark reddish brown, reticulate-pit- ted. Embryo about 4 mm long; slightly curved, cotyledons 1.5 mm long, ovate-elliptic. DISTRIBUTION — Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia. See Figure 28. Key to the Subspecies of Brunfelsia grandiflora 1. Corolla tube 30-45 mm long, 2-3.5 mm in diameter, opening at mouth linear-oblong; limb 35-42 mm in diameter, capsule 17-22 mm long 14a. subsp. grandiflora 1. Corolla tube 15-30 mm long, 1-2 mm in di- ameter, opening at mouth elliptic-obovate; limb 20-40 mm in diameter; capsule 11-16 mm long 14b. subsp. schultesii 14a. Brunfelsia grandiflora subsp. grandiflora Brunfelsia calycina f. grandiflora (D. Don) Voss in Vilm., Blumengartner, ed. 3, 1: 743. 1896. Brunfelsia tastevinii Benoist, Bull. Soc. Bot. France 75: 295. 1928. Type: BRAZIL. Acre: region de Rio Jordao, Tastevin s.n. (holotype, p; isotypes, F, P). Leaves 10-23 cm long, 3-8 cm wide, glabrous or sparingly pubescent at midrib; petiole 3-12 mm long. Pedicel 2-10 mm long. Calyx 9-13 mm long, teeth 2-5 mm long, ovate-lanceolate. Corolla tube 30-45 mm long, 2.5-3.5 mm in di- ameter, the mouth 6-9 mm long; limb 35-52 mm in diameter, spreading. Capsule 17-22 mm long. DISTRIBUTION — Colombia, Ecuador, Peru, Bra- zil, Bolivia. SPECIMENS EXAMINED — COLOMBIA. Meta: *25 km S from Granada toward San Juan, Luteyn el al. 4769 (F, MO). ECUADOR. Loja: *City of Loja, Zarucchi & Andrade 2316 (F). Napo: Mera, 4 Mar. 1940, Lugo 23 (s); Canton Napo, near Achidona, 650 m, 19 Apr. 1935, Mexia 7275 (F, s, uc, us); *Costado S del Rio Napo, E de Puerto Napo, por carretera nueva hacia Rio Arajuno, Neill 6546 (F); *Rio Napo 8 km abajo de Puerto Misahualli; Reserva Biologica Jatun Sacha, Neill & Palacios 7101 (F). Zamora-Chinchipe: Reg. Oriental, Huamboya, 1500-1600 m, 15 Feb. 1944, Acosta-Solis 7576 (F); Camino Gualaquiza al Rio Zamora (Las Jibarias), 850 m, 25 Dec. 1948, Scolnik 1495 (us); *Zamora-Zumba (along Rio Jamboe), Marling & Andersson 13906 (GB, GH); *Guadalupe-San Jose de Yacuambi (28 de Mayo), along Rio Yacuambi, Harling & Anders- son 13934 (GB, GH). PERU. Cusco. *Prov. Pau- cartambo; Callanga, Woytkowksi 514 (F). Huan- uco: woods above Pozuzo, 6000 ft, Aug. 1863, Pearce 290 (K); Monzon, 950 m, 17 Jun. 1958, Woytkowski 5026 (F, LE, P, s); Divisoria, 1700 m, 16 Sep. 1946, Woytkowski 34536 (F, MO, uc); *Prov. Tingo Maria; Rio Huallaga, Boeke 1224 (F). Loreto: Pongo de Manseriche, Colon, 1924, Tessmann 5541 (o, s); Rio Ucayali, Canchahuayo, 20 Nov. 1898, Huber 1490 (MG); Santa Catalina, 11 Dec. 1898, Huber 1562 (MG); Prov. Coronel Portillo, Plantacion Margarita, cerca a la Diviso- ria, 1500-1600 m, 14 Aug. 1946, Ferreyra 985 (B, M, us). San Martin: Rio Huallaga, Rio Chi- purana, Tarapoto, 6 Dec. 1898, Huber 1556 (MG); 78 FIELDIANA: BOTANY BRUNFELSIA grandiflora D.Don subsp. grandiflora ; subsp. sell ul te si i FIG. 29. Brunfelsia grandiflora subsp. grandiflora and subsp. schultesii. Reproduced courtesy of the Botanical Museum of Harvard University. PLOWMAN: REVISION OF BRUNFELSIA 79 *Prov. Mariscal Caceres; Dist. Campanilla, Rio Sion, Caserio de Sion, Schunke, V. 3480 (NY); *Prov. Mariscal Caceres; Dist. Tocache Nuevo, Quebrada de Huaquisha, Schunke, V. 4051 (NY); *Prov. Mariscal Caceres; Dist. Tocache Nuevo, Quebrada de Huaquisha, Schunke, V. 4034 (NY). Without department: E of the Andes, 4000- 6000 ft, Jul. 1884, Pearce 295 (K); woods near Rio Bambusi, one of the tributaries of Maranon, Mar. 1868, Pearce s.n. (K); in flum. Maranon ripis inundatis, Apr. 1855, Spruce 3988 (K); 1909- 1914, Weberbauer 6151 (F). BRAZIL. Acre: ba- sin of Rio Purus near mouth of Rio Macauhan (tributary of Rio Yacu), lat. 9°20' S, long. 69°W, 3 Aug. 1933, Krukoff 5284 (F, G, GH, K, LE, M, MICH, MO, NY, RB, s, sp, uc, us); Rio Moa, 10 km above and below Maita, 16 Apr. 1971, Prance et al. 11998 (wis); *Municipio Rio Branco; Rio Branco-Porto Velho Highway, Albuquerque et al. 1269 (F); *Municipio Rio Branco; Seringal Cor- redeira, Colocac. ao Terra Alta, H. C. de Lima et al. 2122 (F); *Cruzeiro do Sul, Rio Jurua and Rio Moa, Maas et al. 12963 (MO); *Cruzeiro do Sul, Rosa 622 (MO); *Rio Branco, Rod. Rio Branco- Porto Velho, J.U. Santos et al. 48 (F). Amazonas: Rio Purus, Monte Verde, 18 Apr. 1904, Huber 4575 (MG), 29 Apr. 1904, Huber 4589 (MG); Rio Purus-Rio Ituxi, opposite Labrea, 23 Jun. 1971, Prance et al. 13471 (wis); Serra da Moa near school, 1 May 1971, Maas et al. P 12709 (wis); *region of Rio Jordao, Tastevin s.n. (F, P). With- out locality: 1840-1848, Lobb 65 (K). BOLIVIA. Beni: Tumi Chucua 30 km S of Riberalta along Rio Beni, Solomon 7609 (F); *5 km SW of Ri- beralta, Solomon 7905 (F). La Paz: *Prov. Nor Yungas: 12 km hacia el Literal, W del Rio Beni, Beck 13335 (F). CULTIVATED SPECIMENS OF INTEREST — ECUA- DOR. Napo-Pastaza: Rio Aguarico, Dureno, 27 Dec. 1965, Pinkley 43 (ECON), 24 May 1966, Pinkley 202 (ECON); Tena, 28 Sep. 1966, Pinkley 460 (ECON). PERU. Loreto: Maynas, Iquitos, 1924, Tessmann 3541 (G, s). San Martin: Mon- son River, 6 km from Tingo Maria, 625-1100 m, 30 Oct. 1949-19 Feb. 1950, Allard 21176 (F, us). BRAZIL. Para: Belem, Jardim Botanico do Mu- seu Goeldi, Apr. 1908, BAKER 61 (B, c, G, GH, K, LE, M, NY, us). Rio de Janeiro: *Municipio Rio de Janeiro; Jardim Botanico do Rio de Janeiro, Ferreira & Costa 16 (F). 14b. Brunfelsia grandiflora subsp. schultesii Plowman, Bot. Mus. Lean*. 23(6):259, t. 18. 1973. Type: COLOMBIA. Amazonas: Rio Amazonas, near mouth of Rio Loretoyacu and Puerto Narino, bush 12 ft, flowers white or purple, poisonous to cattle, sanango, very positive for alkaloid, 13-15 Sep. 1966, Schul- tes, Raffauf & Soejarto 24108 (holotype, GH; isotype, COL). Leaves 6-20 cm long, 2-8 cm wide, lanceolate, glabrous; petiole 2-6 mm long. Inflorescence variable, compact or lax. Pedicel 2-6 mm long. Calyx 5-10 mm long, teeth 1-3 mm long, trian- gular to triangular-ovate. Corolla tube 15-30 mm long, 1-2 mm in diameter, curved toward apex; limb 20-40 mm in diameter, spreading, mouth 3- 5 mm long. Capsule 11-16 mm long, 10-16 mm in diameter. DISTRIBUTION — Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil. SPECIMENS EXAMINED — COLOMBIA. Amazo- nas: Leticia, about 1 km NE of town on road to brick factory, 30 Jan. 1969, Plowman & Kennedy 2310 (ECON); Rio Amazonas, forest along river about 2 km downriver from Puerto Narino near mouth of Rio Loretoyacu, 5 Feb. 1969, Plowman et al. 2404 (GH, INPA, uc); *Rio Amazonas, down- river from Puerto Narino near mouth of Rio Lor- etoyacu, Plowman et al. 2407 (F). Caqueta: Rio Orteguaza, La Maria (Hacienda de Ramon Achiardy), 23 km SE of Venecia, 400 m, 7 Mar. 1944, Hermann 11237 (NY); La Rastra, 22 Apr. 1956, Vogel 23 (us); Caqueta, Dec. 1930, Uribe Uribe P. s.n. (us); Florencia, 420 m, Feb. 1930, Perez-Arbelaez 688 (COL, us). Meta: llanos of San Martin, 30 Sep. 1916, Da\ve 241 (K, us); San Martin, 400 m, 2 Aug. 1946, Uribe Uribe 1353 (COL). Putumayo: selva higrofila del Rio Putu- mayo, Puerto Porvenir, arriba de Puerto Ospina, hacia la Loma, 230-250 m, 17 Nov. 1940, Cua- trecasas 10611 (COL); Puerto Ospina, 230 m, 25 Nov. 1940, Cuatrecasas 10795 (COL, us); Mocoa, bosque higrofilo en la Quebrada del Rio Mulato, 570-600 m, 26 Dec. 1940, Cuatrecasas 11275 (COL, F); camino entre Mocoa y Puerto Asis, 500 m, 21 Jun. 1935, Garcia Barriga 4586 (COL, us), 500-700 m, 28 Aug. 1963, Juajibioy Chindoy 277 (ECON); Pepino, 20 Nov. 1968, Plowman 2039 (ECON), 1800-2400 ft, 16 Mar. 1953, Schultes & Cabrera 19115 (GH); Rio Caqueta, Floresta, about 15 km downriver from Puerto Limon, 20 Dec. 1968, Plowman 2183 (ECON); Uchupayaco, en la planada entre Urcusique en las orillas del Rio Uchupayaco, 300 m, 22 Feb. 1942, Schultes 3340 (GH); San Antonio del Rio Guamues, 7 Sep. 1966, 80 FIELDIANA: BOTANY Pinkley 444 (ECON); along trail between San An- tonio and Santa Rosa, 1 Dec. 1968, Plowman 2090 (ECON). VENEZUELA. Barinas: Barinitas, Mar. 1953, Aristeguieta 1669 (NY, YEN); Parque Barinitas, 600 m, 22 Jun. 1956, Bernardi 3310 (G). ECUADOR. Napo: Rocafuerte, high forest of Hacienda Arcadia, Rio Curaray, 200 m, 7 July 1933, Heinrichs 496 (F, G), Rio Aguarico, Dureno, 8 Dec. 1966, Pinkley 575 (ECON); Confluence of Quiwado and Tiwaneo Rivers, Davis & Yost 958 (F); *Limoncocha on Rio Napo, Drummond 7330 (MO); *Nuevo Rocafuerte, Rio Yasuni, Marling et al. 7474 (GB, GH); *Rfo Aguarico, Dureno (30 km E of Santa Cecilia), Marling et al. 7700 (GB, GH); *Canon de Los Monos, ca. 12 km N of Coca (Puerto Francisco de Orellana), Marling & An- dersson 11724 (GB); *environs of Limoncocha, Madison et al 5390 (F); *Archidona, Pinkley 457 (F). Pasta/a: *basin of Rio Pastaza, Pacayacu- Sarayacu region, Gill 45 (NY). PERU. Cuzco: Prov. Paucartambo, Villa Carmen, 720 m, 1 May 1965, Vargas 16301 (us). Huanuco: *Prov. Leon- cio Prado; Tingo Maria, Castillo, Plowman 5825 (F); *Prov. Pachitea; Dist. Honoria, Bosque Na- cional de Iparia, Schunke V. 1480 (NY). Loreto: Prov. Maynas, Iquitos region, ca. al Cementerio de Iquitos, 120 m, 18 Sep. 1964, Dodson 2822 (MO); Mishuyacu near Iquitos, 100 m, Jan. 1930, Klug 741 (us); Rancho Indiana, 110 m, 24 Jan. 1932, Mexia 6444 (BM, F, G, GH, K, MO, NY, PH, s, uc, us); Rio Nanay, upland forest near Samito, 19 Feb. 1969, Plowman 2494 (ECON, GH); Rio Mazan, Gamitanacocha, 100-125 m, 12 Jan. 1935, Schun- ke 8 (A, F, NY, uc, us); Masisea, 275 m, Killip & Smith 26844 (us); Yurimaguas, lower Rio Hual- laga, 135 m, Aug.-Sep. 1929, Killip & Smith 27667 (us), 25056 (NY), 281 78 (us), Jan. 1831, Poeppig 2206 (w); Balsapuerto (lower Huallaga Basin), 150-350 m, 28-30 Aug. 1929, Killip & Smith 28644 (us), 220 m, Feb. 1933, Klug 2880 (A, F, G, GH, MO, NY, s); Pucallpa, 200 m, 4 Aug. 1946, Soukup 3041 (F); Pongo de Manseriche, 1924, Tessmann 4921, Rio Maranon, Tessman 5005 (G); Prov. Ucayali, lower Pisque, Nueva Be- liza, 1923, Tessmann 3243 (G, NY, s); *Prov. May- nas; Moena Cano between Iquitos and Rio Itaya, Gentry et al. 15707 (MO); *IVITA, km 59 Pucall- pa-Tingo Maria road, Gentry et al. 18573 (MO); *Bosque Nacional von Humboldt, km 86 Pucall- pa-Tingo Maria road, Gentry et al. 18664 (MO); *Prov. Maynas; Negro Urcu, Rio Napo, McDaniel 15328 (MO); *Prov. Maynas; Pebas, mouth of Rio Ampiyacu, Plowman et al. 6455 (F); *Prov. May- nas; Pebas, Plowman et al. 6737 (F); *Prov. May- nas; Pebas, Plowman et al. 6939 (F); *Prov. May- nas; Dist. Iquitos, Caserio Momoncillo, Revilla 304 (MO); *Prov. Maynas; Dist. Iquitos, cerca a la boca del Rio Nanay, Revilla 1724 (MO); *lower Rio Nanay, Williams, LI., 584 (F). Madre de Dios: *ca. 5 km from Puerto Maldonado near Rio Tambopata, Gentry 16249 (MO). San Martin: Tar- apoto and vicinity, W side of Rio Huallaga, S of Shapaja 1-4 km, 900 ft, 28-30 Jul. 1937, Belshaw 3130 (NY, uc, us); Pucaloma, 330-400 m, 12 Jul. 1950, Ferreyra 7778 (us), 1855-1856, Spruce 3973 (BM, BR, CGE, G, MG, NY, w), 350 m, Oct. 1902, Ule 6481 (G, MG, L), 750 m, Dec. 1929, LI. Williams 5482 (F), 830 m, 6 Feb. 1947, Woyt- kowski 35008 (F, G, MO, uc); entre Bellavista y Banos 200-300 m, 5 Sep. 1948, Ferreyra 4760 (us); Zepelacio near Moyobamba, 1100-1200 m, Oct.-Nov. 1933, Klug 3326 (A, F, G, GH, MO, NY, s); Roque, 19 Jul. 1925, Melin 186 (s); Saposoa, 400 m, 14 Oct. 1959, Woytkowski 5525 (F, LE, p, us); Rioja, 800 m, 9 Jan. 1961, Woytkowski 6170 (GH); Boqueron Pass, 92 km from Tingo Maria on highway to Pucallpa, ca. 400 m, 16 Dec. 1949-5 Jan. 1950, Allard 22096 (us); *Prov. San Martin; Tarapoto, above Hotel Turista, Plowman 5984 (F). Without state: Peru, 1835, Matthews 1320 (CGE, K). BRAZIL. Acre: Rio Jurua-Mirim, vicinity of Porangaba, 21 May 1971, Maas et al. PI 3228 (wis); Rio Acre, Xapury, Jan. 1911, Ule 9746 (K, MG); Paraguassu, Seringal, Nov. 1911, Ule 9749 (K, MG); Rio Branco, 26 Feb. 1962, Vasconcelos & Coelho s.n. (INPA). Amapa: Rio Oiapoque in shade of virgin forest, 5 km SE of Clevelandia, 3°48'N, 51°53'W, 5 Aug. 1960, Irwin et al. 47359 (MG, NY); lower slopes and base of Mount Tipac, 0-100 m, 3°36'N, 51°19'W, 16 Oct. 1968, Irwin 48753 (NY). Amazonas: Esperan9a at mouth of Rio Javary, 26 Jan. 1952, Ducke 865 (F, GH, MG, MO, R, us); Camatian, 30 Jan. 1949, Froes 24035 (RB); Rio Purus, Cachoeira, 21 Mar. 1904, Huber 4204 (MG); 28 Apr. 1904, Huber 4377 (MG); Rio Purus, Rio Ituxi, Rio Curuquete, Cachoeira Re- publica, 24 Jul. 1971, Prance et al. 14551 (wis); upper Amazon, in forest near Tabatinga, 30 Nov. 1874, Traill 599 (K, p); Tocantins, 24 Nov. 1874, Traill 600 (K, p); Rio Jurua, Bocca do Paranan, Apr. 1901, Ule 5453 (G, K, MG, L); *Camatian, Froes 23974 (GH). Rondonia: basin of Rio Ma- deira, 1 km S of Ribeirao, road Abuna to Guaja- ra-Mirim, 27 Jul. 1968, Prance et al. 6570 (wis); *30 km NE de Guajara-Mirim, Krapovickas & Schinini 35155 (F). BOLIVIA. Beni: Rio Iboy, Beni, 800 ft, 14 Feb. 1922, White 2386 (NY, us); Beni River, Jul. 1886, Rusby 2122 (NY). La Paz: PLOWMAN: REVISION OF BRUNFELSIA 81 Guanai-Tipuani, Bolivian Plateau, Apr.-Jun. 1892, Bang 1398 (A, BM, F, c, GH, K, M, MICH, MO, NY, PH, R, us, w); Mapiri, 550 m, 9 Apr. 1927, Buchtien 1299 (us); Mapiri region, San Carlos at Sarampiuni, 600 m, 28 Feb. 1927, Buchtien 1300 (us); Charopampa, 570 m, Nov. 1907, Buchtien 2029 (us), 5,000 ft, Apr. 1886, Rusby 621 (BM, G, GH, K, LE, MO, NY, PH, us, w); Tumu-pasa, 1800 ft, 7 Jan. 1902, White 1840 (BM, NY, us). Pando: bank of Rio Madeira, 6 km above Abuna, 1 1 July 1968, Prance et al. 5854 (wis); Rio Acre, Cobija, Dec. 1911, Ule 9747 (K, MG), Jan. 1912, We 9748 (G, K, MG). Santa Cruz: Prov. Gutierrez, Canton Buenavista, Dolores, 450 m, Steinbach 1805 (A, uc); Buenavista, 500 m, 14 Dec. 1920, Steinbach 5487 (GH), 450 m, 2 Jan. 1926, Steinbach 7399 (A, K); *Buena Vista, Bella Union, Steinbach 7936 (A, BM, F, K). Brunfelsia grandiflora was first collected dur- ing the Ruiz and Pavon expedition in 1798. Ben- tham (1846) considered it to be very similar to his B. latifolia and perhaps also to its variety. Most material that Bentham referred to B. latifolia is now considered B. grandiflora subsp. schultesii, indicating that he recognized the affinities of the plants concerned, although he misinterpreted B. latifolia, which is endemic in Rio de Janeiro (Bra- zil). Brunfelsia grandiflora consists of two subspe- cies: subspecies grandiflora and subspecies schul- tesii. Subspecies grandiflora is much more limited in its distribution than is the second subspecies. It occurs in the drainage systems of the Ribs Mar- anon, Huallaga, and Ucayali in Peru; at higher elevations in eastern Ecuador; and in the upper- most tributaries of the Rios Purus and Jurua in Brazil. It grows primarily at elevations of 650- 2000 m in the region known as the montana, hu- mid, montane rain forest that stretches from Co- lombia to northeasternmost Argentina on the east- ern slopes of the Andes. Brunfelsia grandiflora subsp. grandiflora is characterized by its large flowers (to 50 mm across) and fruits (to 22 mm in diameter), as well as by its higher altitudinal preference. Subspecies schultesii looks very similar but is generally smaller in all its parts (see key to subspecies). From the relatively few collections available, sub- species grandiflora appears to be variable and ex- hibits some exceptional forms. For example, col- lections from the northern part of the range (Ec- uador and northern Peru) often have divaricating, not spreading, lateral nerves in the leaves and longer pedicels (Pearce 290, Lugo 23). The latter feature also appears in high-elevation populations of B. uniflora in Bolivia. In the 1920s, a Catholic missionary named Tas- tevin collected this subspecies in the Rio Tarauaca in the state of Acre, Brazil. He stated that it was a high-climbing liana (Tastevin, 1922) and con- fused the plant with the psychotropic Banisteriop- sis. Nevertheless, other collectors have indicated that the plants may reach 10 m in height, an ex- ceptionally large size. Whether B. grandiflora subsp. grandiflora may actually become lianoid in habit has yet to be confirmed by field observa- tions. Brunfelsia grandiflora subsp. grandiflora is well-known in cultivation, especially in the New World tropics, as an ornamental. In the Amazon region, it is also cultivated for its narcotic and medicinal properties [see Plowman, 1977; Schul- tes & Raffauf, 1990, 1991— Eds.] and is known by several common names: sanango, chici-qui- sanango, chiricsanango, and sananguillo (Peru); and manacd grande (Brazil). Brunfelsia grandiflora subsp. schultesii has long been recognized in the literature under sev- eral misapplied names, including B. bonodora, B. latifolia, and B. maritima. This was primarily the result of "follow-the-leader" taxonomy in which no botanist consulted the type material on which the original descriptions were based. It is now clear that these other names all apply to species that are found only in southeastern Brazil. Brunfelsia grandiflora subsp. schultesii is wide- ranging and polymorphic, occurring in western South America from Venezuela southward to Bo- livia. In addition, disjunct populations are known from the Territory of Amapa in northern Brazil. It grows commonly as an understory shrub in pri- mary and secondary forests between 100 and 900 m elevation. Because this subspecies is widely cultivated in South America for medicinal and or- namental purposes, some activity of man has un- doubtedly influenced its present distribution. The two subspecies of B. grandiflora differ pri- marily in the size and form of the flowers and in the size of the fruits. Subspecies schultesii also tends to be smaller in habit and leaf size, but there is considerable overlap between the subspecies in these characters. The two subspecies are also ecologically and, to some extent, geographically isolated. The typ- ical subspecies is usually found between 900 and 2000 m elevation in the inter-Andean valleys of Peru. Brunfelsia grandiflora subsp. schultesii nor- 82 FIELDIANA: BOTANY mally occurs at much lower elevations, between 100 and 900 m, throughout the upper Amazon and its tributaries, although isolated populations in Bolivia may reach 1800 m. From morphological and geographical evidence, it appears that subspe- cies grandiflora has arisen from its much more widespread and variable counterpart through iso- lation in the intermontane valleys. Intermediates between B. grandiflora subsp. grandiflora and B. grandiflora subsp. schultesii are known from areas where the two coexist. This occurs in the Brazilian state of Acre and in the lower Maranon valley of Peru near Pongo de Manseriche (cf. Tessmann 4921, Krukoff 5284, Vie 5453). These specimens are intermediate in the size of their flowers and fruits, which suggests hybridization between the two subspecies in these areas. No intermediate forms have been found in the higher montane valleys or in the tropical low- lands. As would be expected from such a wide-rang- ing plant, B. grandiflora subsp. schultesii shows considerable morphological variation. Some of this variation may be due to environmental fac- tors such as those that exist between the llanos of Venezuela and the cloud forests of Bolivia, but it seems that some populations have become genetically differentiated as well since morpho- logically distinct groups can be readily and con- sistently separated. These variants, however, are relatively slight and in my judgment should not receive formal names. I will instead discuss some of the more prevalent variations that occur. The most common form of subspecies schultesii, represented by the type collection (Schultes et al. 24108), is found from the Colombian Putumayo south to Tarapoto in Peru and eastward to Leticia in the Colombian Amazon. These plants have large, more or less oblong-lanceolate leaves and many flowers in a compact inflorescence. It is this form that occurs disjunct in the north Bra- zilian state of Amapa (Invin 47359, 48753). This is the only known collection east of Leticia, a disjunction of some 2300 km. The species is ap- parently absent from the lower Amazon, al- though additional collecting in this region may prove otherwise. In the northern part of its range, in the llanos of Venezuela and Colombia and south to the Co- lombian Putumayo, another form of B. grandiflo- ra subsp. schultesii occurs. These populations have large, oblong, subcoriaceous leaves and more lax, branched inflorescences (cf. Ariste- guieta 1669, Perez-Arbelaez 688). The leaves in some specimens approach those of B. chiricaspi in their large size and ochraceous coloring on drying. These plants apparently survive in scat- tered patches of woodland in the llanos and in gallery forests along the rivers. A third form of B. grandiflora subsp. schul- tesii may be distinguished among populations occurring on the eastern slopes of the Bolivian plateau (cf. R. S. Williams 425) in wet, montane forests up to 1600 m altitude. These plants have rather small, lanceolate leaves and few flowers per inflorescence, rarely reduced to three. These populations show strong similarities to B. bono- dora of the coastal forests of Rio de Janeiro, a species still very poorly known. Brunfelsia gran- diflora subsp. schultesii differs primarily in hav- ing shorter pedicels and a smaller calyx, al- though additional characters may be found when more material becomes available. The floristic affinities between the Yungas regions of Bolivia and Rio de Janeiro have been pointed out else- where (Plowman, 1979). These differing forms of B. grandiflora subsp. schultesii intergrade completely with adjacent populations and should not be considered taxo- nomically significant. They are mentioned here to point out the degree of variation that exists in this subspecies, which may, in some cases, lead to confusion with other species. Brunfelsia grandiflora subsp. schultesii grows usually as a shrub 1-2 m tall, at times becoming a treelet to 5-6 m. It lives in the understory of shady, moist forests; at the margins of forest clearings; and along rivers. It commonly persists after the forest has been cut for lumber or shift- ing agriculture, sprouting easily from the roots. Considered a valuable ornamental and medicinal plant, subspecies schultesii is much planted in Indian houseyards and fields and often persists after the inhabitants move elsewhere. In this way, the plant has probably increased its distri- bution under the influence of man. There is some indication that its occurrence as far east as Le- ticia in the Colombian Amazon may be due to man's activities. Populations in this area have so far been located only near the major rivers in places most likely inhabited at some time in the past. Brunfelsia grandiflora subsp. schultesii flow- ers and fruits sporadically throughout the year over most of its range, with two main periods of flowering: December to February and July to August. The flowers are usually unscented, al- though they may occasionally contain the odor PLOWMAN: REVISION OF BRUNFELSIA 83 of jasmine (Plowman 2019, Ducke 865). This plant is known by a number of vernacular names that are indicative of its usefulness in native medicine [these are given, along with an account of the medicinal and narcotic uses, in Plowman, 1977— Eds.]. 15. Brunfelsia hydrangeiformis (Pohl) Benth. in DC., Prodr., 10: 198. 1846. Franciscea hydrangeiformis Pohl, PI. Bras. Ic. 1: 7, t. 7. 1826. Type: BRAZIL. Rio de Janeiro: ad Olaria et Sumido rio in via ad Rio Parahybuna, habitat inter frutices locis umbrosis, floret Sep- tembri et Octubri, 1818, Pohl (115) (holotype, w; isotypes, F-875036, M, w). Shrub 1-2 m tall. Stem erect, sparsely branched at intervals. Bark grayish brown. Branches few, appearing at intervals, 4-6 mm in diameter, stout, naked, glabrous, the epidermis splitting longitudinally. Leaves crowded at ends of branches, alternate, subverticellate with 5-10 together, blade 13-32 cm long, 3-15 cm wide, oblanceolate to subspathulate, rarely lanceolate to oblong, abruptly to long acuminate with straight or subfalcate acumen, rarely acute, narrowed-cu- neate at base, the upper surface glabrous, often nitid, dark green, the lower surface sparsely pu- bescent mostly at nerves, or glabrous, pale green, membranaceous to subcoriaceous, reticulately ru- gose, with sulcate nerves, lateral nerves 7-11, spreading or diverging, depressed in sulci; petiole 2-13 mm long, glabrous. Inflorescence terminal, forming in uppermost whorl of leaves, densely capituliform to corymbiform, rarely lax, hemi- spheric, sessile, the axis 1-3 cm long, persistent. Flowers 5-50, purple with white spot at mouth, fading to nearly white with age. Bracts paired, 2- 10 mm long, lanceolate to linear-lanceolate, pu- bescent or glabrous, ciliolate at margin. Pedicel 2-8 mm long, glabrous or rarely pubescent. Calyx 15-35 mm long, 3-10 mm in diameter, tubular to tubular-campanulate, oblong in bud, slightly in- flated, variably pubescent, densely pilose, glan- dular, or glabrous, light green to ferrugineous, thin to firmly membranaceous, teeth 2-10 mm long, erect, ovate to linear-lanceolate, acute to acumi- nate at apex; calyx in fruit, thin, completely en- closing fruit. Corolla tube 20-35(45) mm long, 1-2 mm in diameter, 1-1 Vi times as long as calyx, rarely longer, slender, glabrous or glandular pu- bescent; limb 18-32 mm across, spreading, lobes 7-12 mm long, rounded, overlapping at lateral margins. Stamens included in upper part of co- rolla tube; filaments slender, upper pair 5 mm long, lower pair 3 mm long; anthers 1.5-2 mm long, reniform. Ovary 2-2.5 mm long, ovoid; style slender, equaling the filaments; stigma brief- ly bifid, lobes unequal, about 1 mm long. Fruit completely enclosed by calyx, 12-20 mm long, 9-15 mm in diameter, ovoid-subglobose, apicu- late at apex, smooth, greenish brown, pericarp thin-walled, dry at maturity, crustaceous, sparing- ly dehiscent. Seeds ca. 5-10, 5 mm long, 2 mm in diameter, oblong, irregularly angular, dark brown, deeply reticulate-pitted. Embryo un- known. DISTRIBUTION — Brazil. See Figure 33. Note: According to the International Code of Botanical Nomenclature (Article 60.8, Recom- mendation 60G) (Greuter et al., 1994), it is rec- ommended that in compound names of Latin or- igin, the final vowel of the first part be reduced to i before a consonant. Thus, correctly, the orig- inal orthography of hydrangeaeformis is correct- able to hydrangeiformis. Key to the Subspecies of Brunfelsia hydrangei- formis (Pohl) Benth. 1. Leaves subspathulate to broadly oblanceolate, reticulately rugose with sulcate nerves; pedicel 2-4 mm long; calyx 3-6 mm in diameter with yellowish brown pubescence; calyx teeth 4-10 mm long, linear to linear-lanceolate 15a. subsp. hydrangeiformis 1 . Leaves oblong to oblong-lanceolate, smooth or sulcately nerved; pedicel 4-8 mm long; calyx 5-10 mm in diameter, glabrous or with sparse glandular hairs; calyx teeth 2-4 mm long, ovate to ovate-lanceolate 15b. subsp. capitata 15a. Brunfelsia hydrangeiformis subsp. hydran- geiformis. Figure 30 and frontispiece. FIG. 30. Brunfelsia hydrangeiformis subsp. hydrangeiformis (originally as Franciscea hydrangeiformis). From Pohl, J. E., Plantarum Brasiliae Icones et Descriptiones (1826). 84 FIELDIANA: BOTANY PLOWMAN: REVISION OF BRUNFELSIA 85 Franciscea macrophylla Cham. & Schltdl., Linnaea 2: 601. 1827. Type: BRAZIL. Sellow sp. nov. (holotype destroyed at Berlin; fragment of ho- lotype, F-643247; photograph of holotype, F-621825). Brunfelsia macrophylla (Cham. & Schltdl.) Benth. in DC., Prodr. 10: 198. 1846. Leaves with blade 18-32 cm long, 6-15 cm wide, subspathulate to broadly oblanceolate, abruptly acuminate, glabrous on both sides, retic- ulately rugose with sulcate nerves depressed from above. Pedicel 2-4 mm long. Calyx 18-35 mm long, 3-6 mm in diameter, tubular, scarcely in- flated, sparsely pubescent to velutinous with yel- lowish hairs, otherwise purple to brownish, dry- ing ferrugineous, teeth 4-10 mm long, linear to linear-lanceolate. Corolla tube briefly exserted from calyx. DISTRIBUTION — Brazil (Bahia, Minas Gerais, Rio de Janeiro, Sao Paulo). SPECIMENS EXAMINED — BRAZIL. Bahia: 1857, Blanchet s.n. (G). Minas Gerais: Pirange, Oct. 1840, Gardner 5065 (BM, CGE, F, G, GH, K, NY, p, us, w); Sao Francisco de Paula, Rio do Peixe, Aug. 1896, Magalhaes 1643 (R); Mun. Serro, Bo- cada Mata, 1400 m, 6 Oct. 1945, Williams & As- sis 7928 (GH, R, uc, us); Sabara, Nov. 1839, Claussen s.n. (BM, BR, G, p); without locality, Claussen s.n. (CGE), Claussen 201 (BR), Langs- dorffs.n. (us), Sep. 1836, Raben 14 (c), Schiich 591 (CGE, w), 1858, Weddell 878 (G); *km 30-32 on the road from Batinga to Jequitinhonha, Bun- ting et al. 10351 (F). Rio de Janeiro: Floresta da Tijuca, 7 Aug. 1879, Glaziou 11393 (K, p), Nov. 1894, Ule s.n. (R, us); Serra de Estrella, 1841, Claussen s.n. (w); estrada entre a Estrella e a Es- trada Rio-Petropolis, 24 Oct. 1932, J.G. Kuhl- mann s.n. (RB); Nova Friburgo, Oct. 1947, Leite 4188 (NY); Serra de Friburgo, Sitio do Gaucho, Oct. 1964, Duarte 8445 (HB); Cachoeira de Ma- cacu, 22 Oct. 1964, Duarte & Pereira s.n. (GH, RB); Serra Tingua, Rio San Antonio, 24 Aug. 1879, Glaziou 11392 (c, NY, p); Rio Parahyba, 1887, Moura s.n. (R); Ipiahos, Aug. 1934, O. Peckolt s.n. (R); Magdalena, Marreiras, Oct. 1933, Santos Lima 200 (RB); without locality, Langs- dorff43 (LE), 1816-1821, St. Hilaire A-555 (p); *Nova Friburgo, Parada do Gaucho, Pereira 9241 (F); *Municipio Santa Maria Madalena; Parque do Desengano, Santa Clara, Araujo 954 (F); *near Mury, Maas et al. 3326 (u). Sao Paulo: *Muni- cipio de Banana]; Parque Nacional da Serra Bo- caina, Martinelli 1115 (F). Without state: 1814- 1817, Bowie & Cunningham 100 (BM), Sellow s.n. (F, M, MO, s). 15b. Brunfelsia hydrangeiformis subsp. capitata (Benth.) Plowman comb, et slat. nov. Figure 31. Brunfelsia capitata Benth. in DC., Prodr., 10: 198. 1846. Type: BRAZIL. Rio de Janeiro: Serra dos Orgaos, shrub about 4 ft high growing in rather moist places in virgin forests reaching an ele- vation of about 4500 ft, manaca dobrado, Feb. 1837, Gardner 563 (holotype, K; isotypes, BM, F 678955, G, NY, P, w). Franciscea capitata (Benth.) Miers, Ann. Mag. Nat. Hist. Sen 2, 5:250. 1850. Brunfelsia capitata var. angustifolia Benth. in DC., Prodr., 10:198. 1846. Type: BRAZIL. Rio de Ja- neiro: Organ Mountains, 1841, Gardner 5826 (holotype, K). Brunfelsia hydrangeiformis var. glabriscula J. A. Schmidt in Mart., Fl. Bras. 8(1):256. 1864. Lec- totype (designated here): BRAZIL, prope Rio, Gardner 563 (isotypes, BM, F, G, K, NY, p, w). Lectoparatype: BRAZIL. Rio de Janeiro: in montibus prope Nova Friburgo, Jul. 1832, Riedel 85 (LE). Leaves with blade 13-26 cm long, 3-11 cm wide, oblong to oblanceolate, rarely lanceolate, acute to acuminate at apex, glabrous on upper sur- face, puberulent beneath at nerves or glabrous, smooth to sulcately nerved, rarely reticulate ru- gose. Pedicel 4-8 mm long. Calyx 14-35 mm long, 5-10 mm in diameter, tubular to tubular- campanulate, often somewhat inflated, glabrous or sparsely glandular pubescent, light green, teeth 2- 4(7) mm long, ovate to ovate-lanceolate. Corolla tube rarely exserted from calyx. DISTRIBUTION — Brazil (Espirito Santo, Minas Gerais, Rio de Janeiro, Sao Paulo). SPECIMENS EXAMINED — BRAZIL. Espirito Santo: Santa Barbara de Caparao, 4 Dec. 1929. Mexia 4089-a (uc, us). Minas Gerais: Estasoao Experimental de Cafe, Coronel Pacheco, 29 Nov. 1940, Heringer 460 (RB); without locality, Ker- FIG. 31. Brunfelsia hydrangeiformis subsp. capitata comb, et stat. nov. Isotype specimen in the herbarium of The Natural History Museum, London (BM). 86 FIELDIANA: BOTANY Del. bl T. PlowiMB Wl - ;^HtLsS-n?:°d- rJ^llia^o^roaiTto'Bclo Horizonte (Rio ). Elevation 1175 ">• i. KelS OOS OUHl^Ja / 1 ^1H" ..••rujrr^ J- . • — i -,d ™k ~.-~^ «* °- t'srsSrti'ri.'s A«";"«"'^' A4sft s«riKTRsr* sssMts^esai 102 FIELDIANA: BOTANY well to B. silvicola published in 1896 by Taubert, which is here placed in synonymy. The type of B. silvicola was destroyed in Berlin in 1945, but an- other collection (Ule 77) from the same locality is extant and may serve as a representative col- lection and may possibly be a topotype. Brunfelsia obovata var. coriacea flowers in the month of September, somewhat earlier than var. obovata; fruits have been collected in March. Va- riety coriacea appears to grow under the same ecological conditions as var. obovata (swampy ar- eas) but is found in a generally much drier region of cerrado vegetation, probably being confined to localized wet areas. 21. Brunfelsia pauciflora (Cham. & Schltdl.) Benth. in DC., Prodr. 10: 199. 1846. Figure 40. Franciscea pauciflora Cham. & Schltdl., Linnaea 2: 600. 1827. Type: BRAZIL. E Brasilia meridion- ali, without locality or date, Sellow s.n. (holotype destroyed at Berlin; fragment, F 621699; photo- graphs, F, NY; isotypes, M). Besleria inodora Veil., Fl. Flum.: 261. 1829 ("1825"), Ic. 6, t. 81. 1831 ("1827"). Lectotype (designated here): plate 81 of Vellozo. Brunfelsia calycina Benth. in DC., Prodr. 10: 199. 1846. Lectotype (designated here): BRAZIL. Santa Catarina: Insula Santa Catarina, 1832, Baron de Colchester s.n. (K). Lectoparatype: BRAZIL. Santa Catarina: Insula Santa Catarina, Bade s.n. (specimen not located). Lund 755, also cited by Bentham, is excluded as a type. It is B. obovata Benth. var. obovata. Franciscea calycina (Benth.) Miers, Ann. Mag. Nat. Hist. ser. 2, 5: 250. 1850. Brunfelsia pauciflora var. calycina (Benth.) J. A. Schmidt in Mart., Fl. Bras. 8(1): 257. 1864. Franciscea eximia Scheidw. ex Moore & Ayres, Card. Mag. Bot. 1: 16. 1850. Lectotype (desig- nated here): Moore & Ayres, Gard. Mag. Bot. 2: 177 + t. 1850. Representative collection: culti- vated at Royal Botanic Gardens, Kew, Herbari- um Hookerianum, 1 867, without collector (K). Brunfelsia eximia (Scheidw.) Bosse, Handb. Blu- meng. 524. 1859. Brunfelsia calycina var. eximia (Scheidw.) L. H. Bai- ley & Raffill, Stand. Cyclop. Hort. 1: 581. 1914. Franciscea macrantha Lem., Jard. Fleur. 3: 349, t. 249. 1853. Lectotype (designated here): Lemai- re, Jard. Fleur. 3: t. 249. 1853. Representative collection: Royal Botanic Gardens, Kew, culti- vated in temperate house, 1 1 Nov. 1952, Souster s.n. (K). Brunfelsia calycina var. macrantha (Lem.) L. H. Bai- ley & Raffill, Standard Cyclop. Hort. 1: 581. 1914. Franciscea lindeniana Planch., Belgique. Horticole. 15: 100, t. 16. 1865. Type: Planchon, Belgique. Horticole. 15: 100, t. 16. 1865. Representative collection: Royal Botanic Gardens, Kew, culti- vated in temperate house, 16 Jun. 1953, Souster s.n. (K) (see Fig. 42). Brunfelsia lindeniana (Planch.) G. Nicholson, 111. Diet. Gard. 1: 216. 1884. Brunfelsia calycina var. lindeniana (Planch.) Raffill, Gard. Chron. ser. 3, 83: 370. 26 May 1928. Brunfelsia calycina var. floribunda L. H. Bailey & Raffill, Standard Cyclop. Hort. 1: 581. 1914. Representative collection: New Jersey, Ruther- ford, cultivated at Julius Roehrs Co., 29 Mar. 1918, Bailey s.n. (BH). Shrub 1-3(5) m tall. Bark somewhat smooth or longitudinally cracked, grayish brown. Branches sparse, erect or spreading. Bra nth lets stout, gla- brous, rarely pubescent or glandular, smooth, dark green. Leaves scattered or crowded near branch tips, blade 6.5-16 cm long, 2-6.5 cm wide, ob- long to oblong-lanceolate, more rarely elliptic-ob- long or obovate-oblong, apically acute to short acuminate, sometimes blunt or emarginate, cune- ate to attentuate at base, glabrous, punctate or rarely glandular pubescent beneath, chiefly at midrib, upper surface dark green, dull to nitid, lower surface pale green, firmly membranaceous to subcoriaceous, lateral nerves 5-11, spreading, straight or broadly arching; petiole 5-12 mm long, glabrous or pubescent. Inflorescence terminal, sessile or subsessile, with 1-11 flowers. Flowers showy, deep, reddish purple fading to pale lav- ender or white with age, with white spot at mouth sometimes edged with violet. Bracts small, 1-3 per flower, 1-8 mm long, linear-lanceolate, con- cave, narrowly acute apically, glabrous or glan- dular ciliolate, caducous. Pedicel 1 1-25(35) mm long, stout, broader toward apex, glabrous or sparsely glandular, in fruit becoming thicker and corky-verrucose. Calyx 18-32 mm long, 6-10 mm in diameter, tubular, rarely tubular-campanu- late or inflated, punctate to glandular pubescent or completely glabrous, firmly membranaceous to subcoriaceous, pale to dark green, teeth 3-8 mm long, ovate to ovate-lanceolate, incumbent, acute to acuminate at apex; calyx in fruit campanulate- urceolate, completely enclosing fruit, becoming slightly larger and thick-coriaceous, conspicuous- FlG. 39. Brunfelsia obovata var. coriacea. PLOWMAN: REVISION OF BRUNFELSIA 103 3 cm BRUNFELSIA pauci flora (Cham.K Schlecht] Benth. FIG. 40. Brunfelsia pauciflora. Reproduced courtesy of the Linnean Society, London. 104 FIELDIANA: BOTANY FIG. 41. Distribution of Brunfelsia pauciflora (solid circle) and B. rupestris (solid square). ly striate-nerved with light brown, lenticular-ver- rucose outgrowths. Corolla tube 28-36 mm long, 1.5-3 mm in diameter, 1-2 times as long as calyx, reddish purple above, paler beneath, ring at mouth 4-5 mm long, elliptic, white, sparsely glandular; lobes 15-30 mm long, broadly rounded to subel- liptic, rounded-truncate or blunt at apex, some- what accrescent with age, more or less overlap- ping laterally. Stamens included in upper part of tube; filaments subterete, anteriorly canaliculate, white, upper pair 4-6 mm long, lower pair 2-4 mm long; anthers 1.5-2 mm long, orbicular-reni- form, greenish brown. Ovary 2-3 mm long, 1.5 mm in diameter, conical-ovoid, pale green; style 25-30 mm long, filamentous, lavender; stigma 1 mm long, briefly bifid, lobes subequal, white. Capsule included in calyx, 20-22 mm long, 15- 18 mm in diameter, subglobose to ovoid, smooth, light green, pericarp thin, dry at maturity, spar- ingly dehiscent. Seeds 12-30, 5-6 mm long, 2.5- 3 mm in diameter, ovoid or oblong, angular, dark, reddish brown, reticulate-pitted. Embryo ca. 5 mm long, straight; cotyledons 2-3 mm long, ovate, flat. DISTRIBUTION — Brazil (Parana, Rio de Janeiro, Santa Catarina, Sao Paulo). See Figure 41. SPECIMENS EXAMINED — BRAZIL. Parana: Ponta Grossa, 16 Oct. 1910, Dusen 10305 (s), 16 Oct. 1910, Dusen 10354 (F, GH, MICH, NY, s), 19 Oct. 1911, Dusen 8679 (s); Carvalho, 18 Nov. 1909, Dusen 8961 (F, GH, K, L, MICH, MO, NY, p, s), 5 Nov. 1911, Dusen 13298 (F, GH, K, L, MICH, MO, NY, P, s), 7 Nov. 1911, Dusen 13327 (F, GH, NY, s); Jaguariahyba, 10 Oct. 1911, Dusen 13180 (GH, s); Sena do Mar, Itupava, 450 m, 4 Nov. PLOWMAN: REVISION OF BRUNFELSIA 105 1915, Dusen 17301 (GH, NY, s); Mun. Sao Jose dos Pinhais, Campo Largo da Roseira, 13 Oct. 1946, Hatschbach 515 (G, us); Mun. Piraguara, Bauhado, 2 Nov. 1948, Hatschbach 1058 (us); Mun. Campina Grande do Sul: Rio Pardinho, Ver- tente Altantico, 29 Oct. 1965, Hatschbach 13055 (F, P, NY, us); Pico Caratuva, 1500-1600 m, 5 Oct. 1967, Hatschbach 17308 (c, uc); Mun. Morretes, Rio Ipiranga, 29 Nov. 1966, Hatschbach 15300 (NY, us); Mun. Quatro Barras, Rio Taquari, 8 Oct. 1968, Hatschbach 19937 (MICH, MO); km 40 da estrada Curitiba-Paranagua, 17 Oct. 1961, Pabst 5393/Pereira 6066 (F, w); *Quatro Barras, Morro Mae Catiro, Hatschbach 15071 (MBM); *Campina Grande do Sul, Serra Virgem Maria, Hatschbach 20304 (MBM); *Municipio Sao Mateus do Sul; Te- zoura, Hatschbach 22278 (MBM); *Municipio Campina Grande do Sul; Serra Capivari Grande, Hatschbach 22771 (MBM); *Municipio Cerro Asul; Barra do Turvo, Hatschbach 32637 (MBM); *Municipio Sao Jose dos Pinhaes; Guaricana, Hatschbach 34918 (MBM, MO); *Municipio Mor- retes; Estrada da Graciosa, Grota Funda, Hatsch- bach 40004 (F); *Quatro Barras, Arredores, Hatschbach 43273 (F); *Municipio Sao Jose dos Pinhaes; Purgatorio, Hatschbach 45295 (F); *Municipio Curitiba; Capao do Centre Politecni- co, Hatschbach & Cervi 46844 (F); *Municipio Quatro Barras; Morro do Anhangara, Kummrow 1575 (F); *Municipio Guaratuba; Serra do Ara- catuba, Kummrow 2420 (F). Rio de Janeiro: Alto Macahe de Nova Friburgo, 30 Oct. 1857, Glaziou 17168 (c, P, R); Serra do Mocoto, Santa Magda- lena, 1400 m, Aug. 1940, Santos Lima s.n. (RB); Rio de Janeiro, 1867, Glaziou 811 (BR, c), 1842, Raben 746 (BR). Santa Catarina: Mun. Brusque: Mata de Hoffman, 50 m, 19 Oct. 1950, Klein 319 (us); Azambuja, 41 m, 25 Nov. 1947, Reitz 2768 (uc, s); Brusque, 35 m, 25 Nov. 1947, Reitz C 1952 (s, us); Mato do Malucher, 40-50 m, 23 Feb. 1952, L. B. Smith 5786 (us). Mun. Itajai: Itajai, Morro da Fazenda, 150 m, 3 Nov. 1955, Klein 1751 (M, us), 17 May 1969, Plowman 2739 (GH); Itajai, Sep. 1870, Mueller 435 (K); Bra?© Joaquim, Luis Alves, 300 m, 30 Sep. 1954, Reitz & Klein 2160 (G, us); Una de Santa Catarina, Saco Grande, 300 m, 23 Nov. 1966, Klein & Bresolin 6864 (us). Mun. Bom Retire, Riozinho, 1000 m, 23 Dec. 1948, Reitz 2743 (us); pinheiral, 25 Nov. 1956, Smith, Reitz & Klein 7936 (K, NY, R, us); Mun. Ibirama, Horto Florestal I.N.P., 350 m, 2 Nov. 1953, Reitz & Klein 1158 (us), 450-700 m, 12 Nov. 1956, Smith & Klein 7532 (R, us); Blu- menau, Mata da Companhia Hering, 300 m, 22 Oct. 1959, Reitz & Klein 9205 (us). Mun. Rio do Sul, Serra do Matador, 550 m, 17 Oct. 1958, Reitz & Klein 7329 (us). Joinville, Estrada Dona Fran- ciscea, 450 m, 26 May 1957, Reitz & Klein 4238 (us), 600 m, 6 Nov. 1957, Reitz & Klein 5601 (NY, uc, us). Sao Paulo: Moaca, 22 Sep. 1912, Brade 6036 (s, SP); Mun. Iguape, Morro das Ped- ras, Sep. 1920, Brade 8088 (RB); Iguape, 20-100 m, Sep. 1901, Wettstein & Schiffner s.n. (w); cam- inho do Porto da Ribeira, 4 Oct. 1894, Lofgren & Edwall 2705 (SP); Bananal, Serra da Bocaina, Ser- tao do Rio Vermelho, 1200 m, 7 Oct. 1949, Brade & Duarte 20119 (GH, RB); Campos do Jordao, Oct. 1937, Campos Porto 3400 (GH, R), Oct. 1945, Lei- te 3660 (A); Mata da Boa Vista, 25 Nov. 1949, M. Kuhlmann 2176 (SP); Esta9ao Pilar do Sul, 28 Sep. 1898, Edwall 4329 (SP); Sao Paulo, Agua Funda, Mata do Institute de Botanica, 25 Mar. 1965, Rubens Faria 4 (SP); Jabaquara, 27 Sep. 1945, Handro s.n. (SP); Campinas, Heiner 209 (s), Sep. 1900, Campos Novaes 214 (us), 18 Aug. 1894, Campos Novaes 2960 (SP); Butantan, 25 Sep. 1917, Hoehne 582 (M, SP). Bosque de Saude, 9 Sep. 1922, Hoehne 7953 (SP, us); Hansa, 24 Oct. 1928, Hoehne 23169 (SP, us); Jaragua, 10 Oct. 1929, Hoehne 24405 (SP); Limeira, 30 Aug. 1951, M. Kuhlmann 2714 (SP); Cordeiropolis, 19 Nov. 1948, Levy 12 (SP); Araraquara, 28 Sep. 1888, Lofgren 948 (SP); Piruibe, 1 Nov. 1891, Lof- gren & Cowan 1660 (c, SP); Sao Bernardo, 18 Sep. 1908, Puttemans 6135 (SP); Semin. Espirito Santo-Santo Amaro, 18 Oct. 1942, Roth 356 (SP); Santa Branca, 21 Oct. 1901, Vert 6152 (SP). In his Flora Fluminensis of 1829,15 Vellozo in- cluded two species of Brunfelsia under the genus Besleria. One of these, Besleria inodora, was de- scribed 2 years later as Franciscea pauciflora by Chamisso and Schlechtendal from a form of the plant with a glandular-pubescent calyx. This name was transferred to Brunfelsia in 1846 by Bentham, who made the combination B. pauciflora. The ear- lier epithet inodora of Vellozo cannot be used for this species since it is preoccupied by B. inodora Mart. (1847) (= B. americana). In 1846 Bentham published B. calycina, which is essentially a glabrous form of B. pauciflora. He cited three syntypes. One of these (Lund 755) be- 15 As mentioned in the discussion of B. bonodora, there is evidence indicating that Vellozo's work was not offered for sale in 1825, which is the date that appears on the title page, but rather in September-November 1829, which becomes the effective date of publication (Carauta, 1973). 106 FIELDIANA: BOTANY longs to B. obovata; another, collected by Baron de Colchester in Ilha Santa Catarina, has been chosen as the lectotype. The third collection (Ba- de s.n.\ also from Santa Catarina, has not been located. Schmidt (1864) reduced B. calycina to a variety of B. pauciflora in recognition of their similarity. Owing to its elegant beauty as a conservatory ornamental, B. pauciflora was introduced at an early date into European horticulture and become very popular in the mid- 1800s. Several new "spe- cies" under the genus name Franciscea were meagerly described from cultivated forms, includ- ing F. eximia, F. macrantha, and F. lindeniana. These names continue to persist in the horticul- tural literature in various combinations and have created a great deal of confusion in nomenclature. These forms are here considered as cultivars. Brunfelsia pauciflora is relatively uniform in morphological characters, perhaps reflecting the uniformity of its habitat in coastal, mostly low- elevation, rain forest. Some variation does exist, however, in the number and size of the flowers and in the size of the leaves. The presence and nature of the indument in B. pauciflora has been used to distinguish varieties and to segregate species. The type specimen of B. pauciflora has a glandular pubescence in the calyx and pedicels. This form is found primarily inland in the states of Sao Paulo and Parana, where pre- cipitation is less than on the coast. Brunfelsia ca- lycina and B. pauciflora var. calycina were based on the more widespread form with a glabrous and perhaps more coriaceous calyx, occurring in coastal forests from Rio de Janeiro south to Santa Catarina. Closer examination of the nature of the indu- ment shows a continuous range of intermediate forms of pubescence and glaberrity. The calyx may bear small, scattered, punctate dots; very short, scattered, glandular hairs; a well-developed, glandular villosity; or may be completely gla- brous. Indeed, some variability in the develop- ment and density of the hairs can be found within a single population. Furthermore, the hairs may be shed from the calyx as the fruit develops. Be- cause of the instability of the calyx indument as a taxonomic character in this species, I have not segregated the populations with glabrous calyces as distinct forms or subspecies, as has been done in the past. Smith and Downs (1966) pointed out two col- lections (Reitz 2743; Smith, Reitz & Klein 7936) from the mixed Araucaria forests of Santa Catar- ina as "possible hybrids" (parents unknown). These specimens differ in having a nonglandular villosity on the lower surface of the leaves at the midrib, whereas the calyces are completely gla- brous. These forms suggest possible introgression of genes from other species, such as B. cuneifolia or B. brasiliensis. Both of these species contain a similar, nonglandular pubescence and occur in the planalto region of Santa Catarina. Brunfelsia pauciflora occurs primarily on the Atlantic-facing slope of the Serra do Mar of southeastern Brazil from the State of Rio de Ja- neiro to Santa Catarina, including the Serra dos Orgaos, Serra de Mantiqueira, Serra de Parana- piacaba, and several lesser planaltos and morros. Altitudinally, it ranges from near sea level on the coastal plain to about 1500 m at Pico Caratuvain, Parana. This is exclusively the zone of the mata pluvial, the wet tropical rain forest with up to 1600 mm of rainfall annually (Eiten, 1969). A few populations, as mentioned above, occur to the west in the mixed Araucaria forests of the plan- alto of Parana and Santa Catarina. This species is often found in shade along small streams and ravines, in humid primary and sec- ondary forests, and in other places where the soil is moist but well-drained. It apparently does not survive well when the forest is clear-cut. Brunfel- sia pauciflora flowers from September to Decem- ber, with the fruits appearing between January and March. Fruiting specimens are very scarce be- cause the large showy flowers are most often no- ticed by collectors. As an ornamental plant, B. pauciflora is one of the most important brunfelsias. It is commonly grown in conservatories in the temperate zone and frequently planted in tropical gardens. In limb di- ameter, the flowers are the largest of the genus, reaching 8 cm across. There is considerable vari- ation in color from deep, reddish purple to deep purple with a white spot or "eye" at the mouth. This spot is often surrounded by a blue-violet band that fades into the purple. The flowers fade rapidly with age to pale lavender or nearly white (but never pure white), with a concomitant in- crease of up to 1 cm in the diameter of the corolla. Several cultivars may now be recognized in cultivation. Cultivar "eximia" (B. eximia (Scheidw.) Bosse) has medium-sized leaves with an undulate margin and a densely glandular calyx with sharply acuminate teeth. Cultivar "macran- tha" (B. calycina var. macrantha (Lem.) L. H. Bailey and Raffill) is distinguished by large, leath- ery, plane leaves and by a large, glabrous calyx PLOWMAN: REVISION OF BRUNFELSIA 107 and a large corolla up to 8 cm across (Fig. 41). Cultivar "Lindeniana" (B. lindeniana (Planch.) G. Nichols) is a form with rosy-lilac flowers and slightly bullate leaves, according to Raffill (1928). Cultivar "floribunda" [B. calycina var. floribunda L. H. Bailey & Raffill] has smaller, violet flowers (to 3.7 cm across), shiny leaves, and is very free- flowering. Several of these cultivars represent very old clones, persisting in horticulture since their intro- duction in the last century. In an attempt to sta- bilize these names, I have designated as "repre- sentative collections" [for style of citation see Fig. 42 — Eds.] material at the Royal Botanic Gar- dens at Kew that agrees with the original descrip- tions of the horticultural forms. A living collec- tion of Brunfelsia cultivars is maintained in the Temperate House of Kew, some of which proba- bly represent the original clones of these forms. 22. Brunfelsia pilosa Plowman, Bot. Mus. Leafl. 24(2): 42. 1974. Type: BRAZIL. Santa Ca- tarina: Mun. Sao Miguel d'Oeste, forest above Rio Reperiguacu, Peperi, ca. 26°32'S, 53°44'W, 300-400 m, 21 Oct. 1964, Smith & Reitz 12777 (holotype, GH; isotypes, MO, R, uc). Figure 43. Shrub 0.5-2.0 m tall, diffusely branched from near base. Branches spreading and arching, slen- der, terete, leafy or naked. Bark rough, longitu- dinally cracked, shedding in thin, chartaceous flakes, yellowish brown. Branchlets slender, pi- lose to villous, grayish green, dark purple when young, the epidermis splitting lengthwise. Leaves appearing 2-ranked, scattered on branchlets, blade 3-7.5 cm long, 1.3-3 cm wide, narrowly elliptic, oblong-lanceolate or obovate, acuminate at apex, cuneate at base, pilose on both surfaces, primarily at midrib, dull dark green above, pale green be- neath, the young leaves dark purple, firmly mem- branaceous, lateral nerves 6-9, straight, often prominulous above; petiole short, 1-4 mm long, pilose to villous. Inflorescence terminal, sessile, usually with 1 flower, rarely 2-3. Flowers showy, deep violet fading to pure white with age, odor- less. Bracts 1-3, 1-8 mm long, linear-lanceolate, pilose to villous, caducous. Pedicel 1-3 mm long, stout, pilose; in fruit becoming corky, rugose-ver- rucose toward apex. Calyx 12-19 mm long, tu- bular-ventricose, appearing campanulate in exsic- cati, terete in cross section, sparsely to densely pilose with long weak hairs, rarely glabrous, pur- plish, drying reddish brown, membranaceous, teeth 4-10 mm long, subequal, lanceolate, acu- minate; calyx in fruit persistent, partially enclos- ing fruit, light green, becoming subcoriaceous, the sinuses becoming more deeply cut. Corolla tube 25-32 mm long, 1.5-3.0 mm in diameter, twice as long as calyx, glabrous; limb 30-47 mm in diameter, spreading, thickening at mouth promi- nent, round, white lobes 10-15 mm long, sub- equal, the uppermost posterior lobe somewhat larger, broadly rounded. Stamens included in up- permost part of corolla tube; filaments slender, glabrous, upper pair 3-4 mm long, lower pair 3- 5 mm long; anthers orbicular-reniform, 1-1.5 mm in diameter. Ovary 2-2.5 mm long, 1 mm in di- ameter, conical-ovoid; style 22-26 mm long, slen- der; stigma 1 mm long, briefly bifid, in the form of forceps. Fruit 12 mm long, 10 mm in diameter, ovoid to subglobose, apiculate at apex, smooth, dark green, shiny, pericarp thin-walled, dry at ma- turity, tardily dehiscent. Seeds ca. 10, 5-6 mm long, 2.5-4 mm in diameter, ellipsoid, reticulate- pitted. Embryo unknown. DISTRIBUTION — Brazil (Parana, Rio Grande do Sul, Santa Catarina, Sao Paulo), Paraguay, Argen- tina (Misiones). See Figure 37. SPECIMENS EXAMINED — BRAZIL. Parana: Curitiba, Bairro Sao Nicolau, 18 Nov. 1966, Ca- prilioni 1659 (us); Itaperussu, 17 Nov. 1908, Du- sen 7077 (GH, s); Jaguariahyva, 25 Oct. 1910, Du- sen 10443 (GH, MICH, NY, s); Tamandare, 4 Oct. 1914, G. Jonsson 1054a (F, GH, NY, s); Sao Ma- theus, 27 Feb. 1929, Gurgel s.n. (RB); Mun. Rio Branco do Sul, Sao Vicente, 27 Oct. 1967, Hatschbach 17610 (c, uc); Ponta Grossa, 2 Nov. 1928, Hoehne 23309 (SP, us); Foz do Iguacu, Parque Nacional de Igua9u, 8 Oct. 1946, /. G. Kuhlmann s.n. (RB); Ypiranga, 15 Sep. 1934, Reiss 99 (GH, NY); *Municipio Irati; Fazenda Al- eixo, Carvalho 83 (MBM); *Municipio Ivai; Bom Jardim, Hatschbach 22383 (NY); *Municipio Cur- itiba; Rio Atuba, Hatschbach 25613 (MBM); *Municipio Gal. Carneiro; Iratim, Hatschbach 28320 (MBM); *Municipio Pitanga; Borboletinha, FIG. 42. Representative specimen of Brunfelsia pauciflora cv. "Lindeniana" in the herbarium at Royal Botanic Gardens, Kew. 108 FIELDIANA: BOTANY - Revision of lirunl.-1-i.i (SoUnncne) Brunfelsia pauciflora (Cham. & Schlecht.) Benth. in DC. Prodr. JO: 199. 1846. cv* 'Lindeniana' Del. bv Timotlir 1'lowm.n I~1974 ""-"^^""i"' r.*i HERB. HORT. EOT. REG. KEW. '<**.' - Franciscea Lindeniana Planch. Belg. Hortio. 15:100. 1. 16. 1865. — PLOWMAN: REVISION OF BRUNFELSIA 109 BRUNFELSIA pilosa Bowman FIG. 43. Brunfelsia pilosa. Reproduced courtesy of the Botanical Museum of Harvard University. 110 FIELDIANA: BOTANY Hatschbach 32869 (GH); *Municipio Adrianopol- is; Rio Pardo, prox. a Barra, Hatschbach 37876 (MBM); *Municipio Balsa Nova; Campina da Cas- cavel, Hatschbach 39164 (GH); *Municipio Tiju- cas do Sul; Rincao, Hatschbach 40470 (F); *Mun- icipio Curitiba; Orto Guabirotuba, Hatschbach 45415 (F); *Municipio Campina Grande do Sul; Belizario, Hatschbach & Guimardes 15286 (MBM); *Municipio Jaguariaiava; Rio Samambaia, Hatschbach & Guimardes 25473 (MBM); *Muni- cipio Curitiba; Capanema, Kummrow 94 (MBM); *near Campo Largo, Lindeman & Haas 3248 (u, GH, MBM); *Parque Municipio do Iguacu (Muni- cipio Curitiba), Oliveira 100 (F). Rio Grande do Sul: Sao Leopoldo, 10 Sep. 1946, Henz 27246 (MO, NY), 20 m, 17 Sep. 1946, Henz 35305 (s); Mun. Rio Pardo, Fazenda Horticola, 70 m, Oct. 1923, Jurgens 19 (B); Colonia Santo Angelo, 4 Feb. 1893, C. Lindman A-597-b (s), Sep. 1900, Schwarzer 50 (L, s); Silveira Martins, 20 Mar. 1893, C. Lindman A-597-c (s); Rio dos Sinos, 8 Nov. 1949, Rambo 44295 (BR, L); Butterberg pro- pe Montenegro, 13 Nov. 1950, Rambo 49144 (GH, p, w); without locality, 1833, Gaudichaud 705 (P), 1835, Isabelle 6 (G); *Municipio Sta. Barbara; Cruz Alta, Lindeman et al. 8233 (u). Santa Ca- tarina: Mun. Ca£ador, Fazenda dos Carneiros, 1100 m, 7 Dec. 1962, Klein 3518 (us); Mun. Campos Novos, Palmares, 950 m, 28 Oct. 1963, Klein 4102 (us); Bituruna, Fazenda Etienne, 11 Feb 1948, Mello Filho 793 (R); Nova Teutonia, 25 Oct. 1943, Plaumann 164 (RB); Mun. Ararauja, Rodeio da Areia, 22 Nov. 1943, Reitz C-171 (RB); Santa Cecilia, 1,100 m, Reitz & Klein 14136 (NY); Mun. Sao Miguel d'Oeste, Canela Gaucha, 8 km NW of Sao Miguel d'Oeste, ca. 26°40'S, 53°34'W, 700-750 m, Smith & Reitz 12757 (NY, R, us); Mun. Abelardo Luz, N bank of Rio Chapeco at Abelardo Luz, ca. 26°35'S, 52°20'W, 900-1000 m, 23 Oct. 1964, Smith & Reitz 12870 (c, F, LE, R); Tubarao, Nov. 1889, Ule 1521 (P). Sao Paulo: Itapetininga, 17 Nov. 1887, Lofgren 389 (c, R, SP, us); Ypiranga, Nov. 1910, Luderwaldt 2117 (RB, SP); Campinas, Jundiahy, Mar. 1900, Campos No- vaes 216 (us); Villa de Serra Branca, 1 Nov. 1897, Puttemans 4328 (SP); Carandiru, Dec. 1912, Ta- mandare 244 (RB); Mandaquil, 23 Nov. 1906, Us- teri s.n. (G). Without state: Sellow 1573 (K, M, w). PARAGUAY. In regione fluminis Alto Pa- rana, Yaguarazapa, 1909-1910, Fiebrig 5460 (G, GH, us). ARGENTINA. Misiones: Dep. San Ja- vier, Acaragua, 220 m, 30 Sep. 1946, Bertoni 2974 (B, w); Dep. Candelaria: Bompland, 4 Oct. 1909, Jorgensen s.n. (BAB), Nov. 1910, Jorgensen s.n. (BAB); Santa Ana, Aug. 1901, Llamas 1530 (BAB, G). Dep. Iguazu: Cataratas de Iguazu, 13 May 1969, Plowman 2735 (GH); Delicia, camino a El Dorado, 4 Nov. 1949, Schwindt 2270 (c, LD, NY); Puerto Aguirre, Rio Iguazu, 19 Sep. 1922, I.N.T.A. 4435 (B). Brunfelsia pilosa is a distinct species related to B. uniflora and B. australis. Bentham saw a spec- imen (Sellow 1573) referable to the present spe- cies but included it within his concept of B. aus- tralis as a syntype. This specimen is rather ex- ceptional for B. pilosa in having a glabrous calyx and leaves. Brunfelsia pilosa is distinguished from related species in having very short, stout pedicels (1-3 mm long) and an inflated, tubular calyx with deeply cut teeth (4-- 10 mm long). The twigs, leaves, and calyx typically contain few to many long, weak trichomes. The flowers are large and showy with a corolla limb nearly 5 cm in diam- eter. Brunfelsia pilosa ranges from the Brazilian state of Sao Paulo south to Rio Grande do Sul and extreme northeastern Argentina (Misiones). It has a fairly wide altitudinal range, occurring from near sea level to 1100 m elevation. In the central part of its range, where it occurs most commonly, this species seems to be associated with the for- merly extensive Araucaria forests found in the south Brazilian planalto. In contrast to B. australis, which may become a small tree, B. pilosa grows as a weak shrub with slender, horizontal, arching branches. It occurs primarily in the understory of primary and sec- ondary woods, as well as in thickets and along watercourses. Flowering takes place from October to December with fruits appearing from February to May. Owing to its attractive flowers, this shrub is cul- tivated in tropical gardens, although not as exten- sively as B. australis. Brunfelsia pilosa appeared in the Florida nursery trade in the mid-1970s and is increasing in importance as an ornamental. In its native area, B. pilosa is known by a va- riety of common names: flor de primavera, pri- mavera, sempre triste, and manacd (Brazil); azuc- ena silvestre, azucena del monte, and Jazmin del Paraguay (Argentina). 23. Brunfelsia rupestris Plowman, Fieldiana, Bot. n. s. 8: 7. 1981. Type: BRAZIL. Minas Gerais: Estrada Diamantina a Corinto ate 20 km. Campo rupestre e cerrado. Arbusto ate 2 m, flores roxas. 12 Jan. 1976, Shepherd, de PLOWMAN: REVISION OF BRUNFELSIA 111 BRUNFELSIA rupest ris Plowman 5 cm. FIG. 44. Brunfelsia rupestris. Reproduced courtesy of the Field Museum of Natural History. Andrade, Konoshita & Tamashiro 3935 (ho- lotype, UEC 11912; isotypes, F 1877095, NY, UEC 11912A). Figure 44. Much-branched shrub 1-2 m tall. Bark on branches yellowish to reddish brown, cracked lon- gitudinally and transversely, exfoliating in thin flakes. Branch lets about 2 mm in diameter, gla- brous or puberulent. Leaves usually congested near branch tips, short petiolate, blade 10-40 mm long, 6-17 mm wide, lanceolate to oblong, api- cally acute, sometimes blunt or emarginate, strongly revolute at margin, basally cuneate, gla- brous above, puberulent beneath with short 112 FIELDIANA: BOTANY curved glandular or eglandular hairs, shiny, me- dium green above, dull beneath, drying ochra- ceous brown, thickly coriaceous, the lateral nerves 4-5, straight, scarcely distinct; petiole 1- 3 mm long, glabrous. Inflorescence terminal on current season's branchlets, short pedunculate, with 2-5(8) flowers. Flowers deep violet with a white "eye" at orifice, fading to lavender with age. Peduncle 1-5 mm long, short-branched, the branches articulating with the pedicels, persistent, puberulent. Bracts 16 mm long, linear-lanceolate or cymbiform, sparsely villous, ciliolate at mar- gin, caducous. Pedicel 4-12 mm long, 1-1.5 mm in diameter, glabrous. Calyx 10-15 mm long, 3- 6 mm in diameter, tubular-campanulate, terete, glabrous, light green, teeth 2-3 mm long, sub- equal, ovate to apically acute, the tip itself some- times blunt or truncate. Corolla tube orifice 2-3 mm in diameter; limb 15-22 mm in diameter, spreading, slightly thickened at mouth of tube, lobes 5-10 mm long, subequal, rounded. Stamens inserted in upper part of corolla tube; filaments strap-shaped, the anterior pair 2-3 mm long, api- cally incurved, included, the posterior pair 1-2 mm long; anthers 1.5 mm in diameter, orbicular- reniform. Ovary 1.5 mm long, ovoid; stigma 1 mm long, briefly bifid. Capsule enclosed by per- sistent calyx, 16 mm long, 14 mm in diameter, ovoid to globose, apiculate at apex, smooth, peri- carp thin, crustaceous, dry at maturity. Seeds 5 mm long, 2.5-3 mm in diameter, oblong-ellipsoid, somewhat angular, brown, reticulate-pitted. Em- bryo 4 mm long, straight; cotyledons 2 mm long, ovate. DISTRIBUTION — Restricted to the campo rupes- tre community in the Serra do Espinhaso, Minas Gerais, Brazil. See Figure 41. SPECIMENS EXAMINED — BRAZIL. Minas Ger- ais: Serra do Espinhaso, Municipio Diamantina: Guinda, 5 Nov. 1937, Mello Barreto 9479 (F, RB), 1300 m, 14 Nov. 1971, Hatschbach & Pelanda 27941 (GH); ca. 25 km SW of Diamantina on road to Gouveia, 1300 m, 16 Jan. 1969, Irwin et al. 22077 (F, MO, NY, UB); ca. 12 km SW of Diaman- tina, 1350 m, 23 Jan. 1969, Irwin et al. 22465 (NY, UB); Serra do Cruzeiro, 5 Nov. 1979, V.F. Ferreira et al. 887 (RB); *Municipio Gouveia; Da- tas, Hatschbach and Zelma 50296 (F). Brunfelsia rupestris is a very distinct species that is restricted to the compos rupestres at higher elevations in the Serra do Espinha?o of central Minas Gerais. It is most closely related to, and probably derived from, the widespread and poly- morphic species B. brasiliensis. Brunfelsia rupes- tris differs in having the leaves more congested, thickly coriaceous, strongly revolute at the mar- gins, sparsely pubescent, and with only four to five lateral nerves and in having glabrous pedicels and calyces. 24. Brunfelsia uniflora (Pohl) D. Don, Edin- burgh New Philos. J. 86. July. 1829. Figure 45. Franciscea uniflora Pohl, PI. Bras. Ic. 1: 2, t. 1. 1826. Type: BRAZIL. Rio de Janeiro: habitat in um- brosis inter frutices in via a Sumidorio ad Gov- erno et ad Rio Parahybuna, Sep.-Oct. 1818, Pohl [9] (holotype, w; isotype, PR). Franciscea hopeana Hook., Bot. Mag. 55: t. 2829. 1828. Type: Hooker, Bot. Mag. 55: t. 2829. 1828. Brunfelsia hopeana (Hook.) Benth. in DC., Prodr. 10: 200. 1846. Brunfelsia hopeana var. pubescens Benth. in DC., Prodr. 10: 200. 1846. Type: TRINIDAD. In in- sula Trinitatis montibus Monos Bocas, without date, Lockhart 197 (holotype, K; isotype, GOET). Brunfelsia uniflora var. pubescens (Benth.) R. E. D. Baker & N. W. Simmonds in R. O. Williams, Fl. Trinidad and Tobago 2(4): 270. 1953. Franciscea mutabilis H. Jacq., Ann. Fl. Pomone: 285. Jun. 1842. Type: Jacq., Ann. Fl. Pomone: 285. Jun. 1842. Brunfelsia mutabilis (Jacq.) Vilm., Rev. Hort. ser. 2, 1: 261. Aug. 1842. Shrub 0.5-3 m tall with diffuse, spreading branches, often branched from near base. Branch- es slender, terete, becoming somewhat knobby, naked beneath, glabrous. Bark grayish brown, shiny. Branchlets slender, glabrous or pubescent. Leaves scattered along the branchlets, highly variable in size and shape often in the same in- dividual, blade 2.5-8.0 cm long, 1.0-4.0 cm wide, oblong-elliptic, lanceolate or obovate, acute to acuminate at apex, narrowed or cuneate at base, sparsely glandular or variably pubescent beneath at the nerves, or glabrous, dark to light green above, paler green beneath, unusually dull, mem- branaceous to subcoriaceous, deciduous, lateral nerves 5-8, spreading, straight or arching; petiole short, 1-5 mm long, pubescent or glabrous. Inflo- rescence 1 -flowered, terminal, produced at tips of new shoots, sessile, rarely short pedunculate. Pe- duncle 1-6 mm long, slender, terete, glabrous or glandular pubescent, rarely villous, persistent. Bracts 1-3, 1-5 mm long, linear-lanceolate, cili- olate, pubescent at midrib, light green, caducous. Pedicel 1-6 mm long, erect, glabrous. Calyx 8- PLOWMAN: REVISION OF BRUNFELSIA 113 114 FIELDIANA: BOTANY 20 mm long, 3-7 mm in diameter, tubular to tu- bular-campanulate,16 narrowed at base, glabrous, rarely with sparse glandular hairs, pale green, membranaceous, teeth 1-3 mm, erect or incum- bent, triangular-ovate, acute to acuminate; calyx in fruit coriaceous, striately veined, dark green. Corolla tube 13-25 mm long, 1-2 mm diameter, 1-2 times as long as calyx, pentagonal in section, glabrous or sparsely glandular; limb 15-30 mm in diameter, spreading, lobes equal, broadly rounded, at times emarginate, overlapping at the lateral margins, narrowed at base. Stamens included in upper part of corolla tube; filaments slender, upper pair 3-5 mm long, lower pair 2-3 mm long; an- thers about 1 mm in diameter, orbicular-reniform. Ovary 2-3 mm long, 1-1.5 mm in diameter, nar- row conical to ovoid; style 15-23 mm long, slen- der, glabrous; stigma briefly bifid, lobes unequal, obtuse. Capsule nearly enclosed by persistent ca- lyx, 7-8 mm long, 8-18 mm in diameter, subglo- bose to ovoid, smooth, shiny, dark green, pericarp 0.5 mm thick, tardily dehiscent. Seeds 10-20, 3- 5 mm long, 2-3 mm in diameter, variable in shape, ovoid-ellipsoid, somewhat angulate, dark reddish brown to almost black, reticulate-pitted. Embryo 2-4 mm long, curved slightly; cotyle- dons ovate. DISTRIBUTION — Venezuela, Trinidad, Guyana, Brazil, Bolivia, Argentina. See Figure 46. SPECIMENS EXAMINED— VENEZUELA. Boli- var: Upata, 1839-1841, Otto 1003 (w). Cara- bobo: *Entre Trincheras y El Cambur, Guevara 2276 (F, NY). Isla Margarita: El Valle, 300 m, 3 Aug. 1903, Johnston 71 (GH), 2 Aug. 1901, John- ston & Miller 265 (GH, NY); without locality, *Fa- cultad Ciencias Forestales s.n. (NY). Guarico: Selvas de Tamanaco, cerca del Cementerio de Paso Real, 11-12 Jun. 1966, Aristeguieta 6153 (NY, YEN), 21 Sep. 1968, Plowman 1911 (ECON). Without state: Los Cedros, Aug. 1953, Gines 3605 (us). TRINIDAD. Monos Island, 100 ft, Apr. 1936, Graf 10 (NY). GUYANA. Western ex- 16 After completing his thesis, Powman described B. clandestine based on materials he had originally includ- ed in B. uniflora. The reference to the tubular-campan- ulate calyx in this description may derive from "fl. clan- destina." See discussion of B. clandestine! — N.H. tremity of Kanuku Mountains, in drainage of Tak- utu River, 200 m, 4-22 Mar. 1938, AC. Smith 3163 (A, B, F, G, LE, MO, NY, p, us, w). BRAZIL. Bahia: Espigao Mestre, 6 km S of Cocos, 520 m, 16 Mar. 1972, Anderson, Steiber & Kirkbride 37048 (wis); Itambe, Salobinho, basin of Rio Par- do, 14 Nov. 1942, Froes 12681/46; Morro do Chapeu, ca. 7 km S of town of Morro do Chapeu, 1,125 m, 17 Feb. 1971, Irwin, Harley & Smith 32406 (wis); Caatinga bei Tamburi, Oct. 1906, Ule 7061 (K, L); Serra Jacobina, St. Thome, Ponco d'Areia, 1843, Blanchet 3862 (BR, G, P); Blanchet 2616 (P); 1843, Blanchet s.n. (c, G, w); without locality, Jun. 1844, Blanchet s.n. (G), 1847, Blan- chet s.n. (G), Martius s.n. (L); *Monte Santo, Har- ley et al. 16432 (F, K); *Feira de Santana, Serra Sao Jose, Noblick 2038 (F); *Barra da Choca, Santos, T. Soares dos 2511 (F). Ceara: Horto Flo- restal de Ubajara, "Sitio Buriti," 5 Jun. 1942, Ai- res do Nascimento 19 (RB); Vi9osa de Ceara, 1860, Allemao 1249 (P, R); Mun. Sao Gonsalo, near Paracuru, 10 m, 7 Jan. 1945, Cutler 8271 (MO, us); Guaramiranga, Sao Salvador, 24 Aug. 1908, Ducke 1588 (MG); Fortaleza, tabuleiro alem de Aldeota, 5 Feb. 1957, Ducke 2954 (MG, R, RB); Varzea de Vaca, Feb. 1839, Gardner 2432 (BM, K); Serra do Bezouro, 20 Jan. 1958, Guedes 472 (MG); without locality, Allemao & Cysneiro 1250 (R), 1840, Gardner 1798 (CGE, F, G, L, NY, p, us, w); *Municipio de Sao Benedito; Sitio Castelo, Serra da Ipiapaba, Bezerra 334 (F); *Sitio Sao An- tonio, Serra de Maruoca, Fernandes, A., s.n. (F); *Camocim, Cafundo, S. J. Filho 8 (F). Espirito Santo: Bananal, 1924, Freire 14 (R, us); Gorto- cazes, 10 Nov. 1943, J. G. Kuhlmann s.n. (RB). Goias: Luziana, 18 Sep. 1967, Heringer 11549 (us). Minas Gerais: Vi9osa, state agricultural school, 20 Feb. 1959, Irwin 2674 (F, NY, R, us), 22 Feb. 1959, Irwin 2701 (F, NY, R, uc); 19 Nov. 1928, J. G. Kuhlmann s.n. (RB), 2 Apr. 1935, /. G. Kuhlmann s.n. (RB); oxcart road to Sao Miguel, about km 4, 685 m, 2 Apr. 1930, Mexia 4551 (GH, us); road to Barrosa, about km 6, 700 m, 15 Oct. 1930, Mexia 5182 (A, BH, BM, F, c, GB, GH, MICH, MO, NY, PH, s, uc, us); 1827, Pr. von Nevwied s.n. (BR). Caldas, 18 Oct. 1854, Lindberg 203 (BR, s); 25 Sep. 1873, Mosen 673 (c, LD, p, R, s), 1844- 1865, Regnell 1-374 (BR, c, GH, GOET, LE, M, NY, FIG. 45. Brunfelsia uniflora (originally as Franciscea uniflora). From Pohl, J. E., Plantarum Brasiliae Icones et Descriptiones (1826). PLOWMAN: REVISION OF BRUNFELSIA 115 FIG. 46. Distribution of Brunfelsia uniflora (solid circles). p, R, s, us); Cabo Verde, Oct. 1876, Barbosa Ro- drigues 261 (RB); Parahybuna, Oct. 1840, Gard- ner 5062 (EM, CGE, K, p, us, w); without locality, 1858, Weddell s.n. (G), 1845, Widgren s.n. (BR, LD, s). Pernambuco: Isla de Itamarica, Dec. 1839, Gardner 1095 (BM, CGE, G, NY, p, w); Per- nambuco, May 1838, Gardner s.n. (BM, K); Cabo Agostinho, 22 May 1952, D. de A. Lima 52-1039 (R); Olinda, 20 Apr. 1924, Picket 670 (SP); Tapera, 18 Dec. 1929, Pickel 2200 (us). Rio Grande do Norte: Monte Alegre, 26 Jan. 1961, Castellanos 23004 (R); near Natal, Mar. 1914, Dawe s.n. (K). Rio de Janeiro: Corcovado, Sep. 1833, Lus- chnath s.n. (LE), Nov. 1833, Luschnath s.n. (BR, LE), Oct.-Nov. 1832, Riedel 1112 (LE, NY, p, us, w); Restinga do Leblon, 5 Feb. 1948, Macedo s.n. (RB); Parahyba, Oct. 1840, Gardner 5566 (CGE, GH, K, w); Sobral prope Parahyba do Sul, Nov. 1881, Schwacke 3311 (GOET, R, us); Estrada de Minas, Miers 4721 (us); Serra d'Estrella, 1844, Weddell s.n. (NY, p); without locality, 1867, Gla- ziou 357 (BR); *Municipio Sao Sabastiao do Alto; Fazenda Bahia, Araujo 968 (F); *Municipio Silva Jardim; Reserva Biologica Po£o das Antas, ao Rio Aldeis Velha, Araujo 3405 (F); *Municipio Nit- teroi; Itaipu, Morro das Andorinhas, Araujo et al. 3202 (F); *entre Itacoatiara e Itaipuacu, Pedra do Alto Mourao, H. C. de Lima et al. 1637 (CORD, F, K, LA); *Rio de Janeiro, Bairro da Ur9a, Morro do Pao de Afucar, Martinelli 5135 (F); *Municipio Casemiro de Abreu; restinga proximo a Rio das Ostras, Martinelli et al. 5665 (F); *Itaipuacu; Pico Alto Mourao, Profice et al. 16 (F). Roraima: *Boa Vista, Rio Branco, Froes 23068 (IAN). Rio Branco, Serra de Carauma, Nov. 1908, Ule 7715 (K, MG); Road Boa Vista to Venezuela (BR 174); 116 FIELDIANA: BOTANY TABLE 7. Flowering and fruiting times of Brunfelsia uniflora in different parts of its geographical range. Region Flowering Fruiting Venezuela Northeastern Brazil Southeastern Brazil Bolivia April-August December- April October-November September-December August February-May February-April Unknown 20 km N of Boa Vista, 1 Feb. 1969, Prance et al. 9553 (wis). Sao Paulo: Botucatu, Edwall 6133 (SP); Campinas, Heiner s.n. (s); 2 May 1918, Campos Novaes 7756 (us); Sao Jose do Rio Par- do, 1 Oct. 1889, Lofgren & Edwall 1435 (SP, us); Xiririca, 16 Oct. 1894, Lofgren & Edwall 2742 (SP, us); BR-2a km 10 antes de Jacupiranga, 15 Oct. 1961, Pabst & Pereira 6023 (F, R, us); Ban- anal, Oct. 1833, Riedel s.n. (LE, us); pr. Sao Car- los, Jan. 1834, Riedel s.n. (LE); *SW de Jundiai, Serra de Japi, H. F. Leitdo Filho et al. 3219 (F). Without state: Pohl 264 (PR), Pohl s.n. (M). BO- LIVIA. Cochabamba: Prov. Ayopaya, Sailapata, 2,700 m, Dec. 1935, Cardenas 3380 (GH); Prov. Cochabamba, Siberia, 3,300 m, Jul. 1955, Car- denas 5216 (L, us); Carrasco, Jatum-Pino, 3300 m, Sep. 1961, Cardenas 5952 (K, us). Santa Cruz: Comarapa, 2800 m, 20 Oct. 1928, Stein- bach 8345 (A, BM, GH, K, MO, NY, s, us); *W of Comarapa on Carretera Fundamental 4, Davidson 3846 (F). Without department: Padcaya-Motovi, 2400 m, 24 Sep. 1927, Troll 240 (B, M). ARGEN- TINA. Jujuy: Dep. Ledesma, Serrama de Cali- legua, 1500 m, 18 Oct. 1963, Fabris 4502 (M). Salta: Oran, cerca del Achiral mas abajo de San Andres, 17-24 Sep. 1873, Lorentz & Hieronymus 258 (BR, G, LE, p). Brunfelsia uniflora was first described by Pohl in the genus Franciscea. Pohl collected the type in the state of Rio de Janeiro in Brazil. The ho- lotype, bearing the number 9, is preserved at Vi- enna (w); a probable isotype with the number 256 is found in Prague (PR). In 1828, Hooker de- scribed another species, F. hopeana, based on a living plant sent from Brazil, which, according to Hooker, differed from F. uniflora in having a shorter corolla tube. D. Don united Franciscea with the older genus Brunfelsia in 1829, making the correct combination B. uniflora. However, Bentham, in his monograph of 1846, used Hook- er's specific epithet in the combination B. ho- peana. This name is still widely, though incor- rectly, used for this species, particularly in the horticultural and pharmaceutical literature [see Plowman, 1977 — Eds.]. Brunfelsia uniflora is the most widespread and variable of the South American species of the ge- nus. It encompasses a heterogeneous complex of forms found throughout eastern Brazil with dis- junct populations in Venezuela and Bolivia. While showing some differentiation in response to dif- ferent habitats and as a result of geographical iso- lation, the variation is considered to fall within the normal range of a single polymorphic species. The characters and nature of the variability will be discussed according to geographical regions. Brunfelsia uniflora is distinguished from related species by its smaller, solitary flowers (tube 13- 25 mm long), the more or less tubular calyx, and the short (1-3 mm) calyx teeth. It is highly vari- able in the shape and size of the leaves, in the length of the calyx, and in degree and location of pubescence. This species grows under diverse environmen- tal conditions throughout its range, which is, in part, responsible for the different phenotypes that are encountered in different areas. It thrives in wet, montane rain forests in Rio de Janeiro and in the high Bolivian yungas. It has also become adapted to the drier regions of Minas Gerais, Sao Paulo, and Ceara, as well as the cerrado forma- tions of Goias and northeastern Brazil. It occurs sporadically as a component of the understory in primary and secondary forests and persists in cut- over areas (capoeiras). It grows in different types of soils, including red clay, sand, and moist hu- mus. Altitudinally, B. uniflora ranges from sea level to 3300 m in the Andes. Flowering and fruiting in B. uniflora occur at different times of the year in different parts of its range. These are indicated in Table 7. In the coastal rain forest of Rio de Janeiro and adjacent states, B. uniflora assumes its "typical" form, exemplified by the type collection and orig- inal description. These plants are very glabrous with obovate-elliptic leaves. The leaves tend to be larger and shinier than in other areas. PLOWMAN: REVISION OF BRUNFELSIA 117 Further inland, in the drier highlands of Minas Gerais, there is a tendency toward smaller and more pubescent leaves. In addition the flowers may be short-stalked and the calyx longer and narrower in relation to the corolla tube. Popula- tions showing this trend extend southward in the interior of Sao Paulo and northward to central Brazil and along the coast to the dry northeast of the country, stopping abruptly in the state of Ceara. Here the plant may become deciduous, flowering at the beginning of the wet season. Forms similar to those in drier areas of Brazil appear again along the coast of Venezuela, in Isla Margarita and Monos Island (Trinidad). In these northern populations, the young twigs and pedun- cle, if present, are frequently densely pubescent [perhaps because of environmental conditions; see Knapp, 1989 — Eds.]. Bentham tentatively named these plants B. hopeana var. pubescens pending observation of the corolla, which was missing in his specimen. When pubescence and calyx length in this form are considered across its entire area, the variation appears more or less continuous, often showing greater differences in adjacent populations than in those separated by thousands of kilometers. The development of pubescence, which may be glan- dular or nonglandular, is not a good character in this species. The relative length of the calyx is also variable, sometimes appearing tubular-cam- panulate, sometimes long tubular, with many in- termediates. [Later, Plowman differentiated B. claudestina, with tubular-campanulate calyx, from B. uniflora, with "narrowly tubular calyx." See discussion of B. claudestina. — N.H.] In contrast to past treatments of B. uniflora, these characters are not considered here for segregating varieties or subspecies. In southeastern Guyana and adjacent Brazil (Roraima) and southern Venezuela, there occurs another pubescent form of B. uniflora but with oblong-ovate leaves, slightly curved, tubular caly- ces, and hispidulous branchlets. Known only from four collections (A. C. Smith 3163, Ule 7715, Otto 1003, Prance et al. 9553), these distinctive pop- ulations skirt the eastern part of Guyana High- lands. Their uniformity suggests that they have been isolated from other populations of B. uniflo- ra for considerable time. A final component of B. uniflora occurs along the eastern edge of the Bolivian altiplano at about 2500 m altitude, extending south into northwest- ern Argentina. These populations closely resem- ble plants growing in the coastal rain forest of Rio de Janeiro. They differ in having shiny, coria- ceous leaves and longer pedicels. The Bolivian populations grow in very wet, montane cloud for- ests with Podocarpus and Alnus and are morpho- logically homogenous. In southern Brazil, from the state of Sao Paulo south to northeastern Argentina, several species occur that may superficially resemble, and have been confused with, B. uniflora. Two of these, B. australis and B. pilosa, are closely related to B. uniflora and probably have been derived from it through geographical isolation in the past. They form, in fact, a southern component of what may be referred to as the B. uniflora complex. Both B. pilosa and B. australis are distin- guished from B. uniflora by a number of constant morphological characters that are listed under those species. Few or no intermediates or putative hybrids have been found, although their ranges overlap somewhat in Sao Paulo (B. uniflora with B. pilosa) and in Rio Grande do Sul (B. pilosa with B. australis). Schmidt (1864) considered B. australis a variety of B. uniflora, but he did not see a full range of specimens of either species and included material of B. guianensis and B. pilosa. In the states of Parana and Santa Catarina, two additional species occur that may be confused with the B. uniflora complex. Smith and Downs (1966), in the Flora Ilustrada Catarinense, as- signed specimens of both B. pilosa and B. bras- iliensis subsp. macrocalyx to what they consid- ered B. uniflora. They further confused B. cunei- folia with B. brasiliensis . These workers appar- ently did not examine more typical material of B. uniflora and B. brasiliensis from farther north. The breakdown of sharp specific boundaries in these species (especially B. pilosa, B. cuneifolia, and B. brasiliensis) suggests a certain amount of hybridization among them in the past as they came together in the planalto of southern Brazil. Subsequent convergence in several characters may also be attributed to a relatively uniform en- vironment in the zone of the Araucaria forests. Similarities among these species are particularly noticeable in the size and shape of the leaves, the form and location of pubescence, and in the re- duction in the number (often to 1) of flowers per inflorescence. Field studies in which additional collections are correlated with local ecological conditions should prove very useful in under- standing their specific relationships. Brunfelsia uniflora is the source of manaca root, which is widely used in Brazil in the treat- ment of rheumatism and syphilis. It is known by 118 FIELDIANA: BOTANY a number of common names that, along with an account of the uses of the drug, are discussed in Plowman (1977). Dubious and Excluded Species Brunfelsia bahiensis Benth. in DC., Prodr. 10: 590. 1846. Type: BRAZIL. Bahia: without locality, 1834, Blanchet 1509 (holotype, c- DC, non vidi; photographs of holotype, F-645866, NY; isotypes, BM, G, p). Schmidt considered B. bahiensis a dubious spe- cies of Brunfelsia, and I am inclined to follow suit. It is known only from the type collection. One specimen from Minas Gerais (Pr. von Nev- wied s.n., 1816, BR) resembles the type but in fact seems better placed in B. bonodora. Another specimen from Bahia (Blanchet s.n., 1852, G) is also similar but is considered a form of B. hy- drangeiformis. Until additional material becomes available, the true affinities of this species must remain in doubt. Brunfelsia nyctaginoides Standl., Field Mus. Nat. Hist. Bot. ser. 22 (1): 47. 1940. Type: MEX- ICO. Chiapas: Monte Tacana, 1000-2000 m, Aug. 1938, Matuda 2411 (holotype, MICH; isotypes, G, K, MICH, MO, NY, us). This anomalous plant is known only from a few collections on high-elevation, volcanic peaks near the Guatemala-Mexico border. In the original de- scription, Standley (1940) expressed doubt as to the generic placement of this plant. He later rec- ognized its intermediate position between Bro- wallia, Streptosolen, and Brunfelsia (Standley and Steyermark, 1940). It must, however, be excluded from Brunfelsia on the basis of having a broadly lamellate stigma and plicate corolla lobes that are bright scarlet in color, resembling the corolla of Streptosolen ja- mesonii. Streptosolen has a distinctly twisted co- rolla tube that is lacking in the present plant. Its proper generic disposition must await a thorough evaluation of related genera of the Salpiglossi- deae. [It has now been placed in the monotypic genus Plowmania Hunz. (Hunziker and Subils, 1986).— Eds.] Nomina Nuda The following names, mostly of horticultural origin, have no taxonomic standing. Franciscea angusta Heynh., Nom. Bot. Hort. ed. 3. 551. 1850. Brunfelsia angusta Gentil., PI. Cult. Serres Jard. Bot. Brux. 40. 1907. Brunfelsia falcata Regel, Cat. PI. Hort. Aksakov. 21. 1860. Brunfelsia gracilis Heynh., Nom. Bot. Hort. ed. 2. 279. 1846. Franciscea gracilis W. Baxt. in Loud., Hort. Brit. Suppl. 3. 551. 1850. Brunfelsia latifolia Steud., Nom. Bot. ed. 2. 231. 1840. Non B. latifolia (Pohl) Benth. Brunfelsia lockhartii Heynh., Nom. Bot. Hort. ed. 2. 79. 1846. Brunfelsia magnifica Gentil., PI. Cult. Serres Jard. Bot. Brux. 40. 1907. Brunfelsia montana Lodd., Cat. PI. ed. 12. 4. 1820. Brunfelsia multiflora Regel, Cat. PI. Hort. Aksa- kov. 21. 1860. Brunfelsia schomburgkiana Klotsch ex Schom- burgk, Versuch. Fauna Fl. Br. Guy. 1155. 1848. Brunfelsia sieberi Regel., Cat. PI. Hort. Aksakov. 21. 1860. Brunfelsia spinosa Jacq., Enum. Syst. PI. 14. 1760. Brunfelsia spruceana J.A. Schmidt in Mart., Fl. Bras. 8(1): 258. 1864. A nomen nudum placed in the synonymy of B. maritima Benth. by Schmidt. Franciscea pohliana Heynh., Nom. Bot. Hort. ed. 2. 1846. Mania opifera Lacerda ex J. A. Schmidt in Mart., Fl. Bras. 8(1): 261. 1864. A nomen nudum placed in the synonymy of B. hopeana (Hook.) Benth. by Schmidt. XI. Editors' Acknowledgments It is impossible for us to acknowledge with cer- tainty all those whom Timothy Plowman would have wished to thank for their help. Because this work is based primarily on his Ph.D. dissertation, the original acknowledgments to individuals are reprinted here. We extend our apologies to anyone who should be mentioned but is not. As editors, we would like to add our own ac- knowledgments, especially to Dr. Michael Dillon of the Field Museum of Natural History, Chicago, who originally suggested taking on the task of preparing this revision for publication and who PLOWMAN: REVISION OF BRUNFELSIA 119 patiently answered all our queries and requests for advice. Nancy Hensold and Christine Niezgoda, also of the Field Museum, gave us invaluable aid in searching Plowman's notes and other archival material and provided additional data. Several in- dividuals and organizations kindly gave us per- mission to reproduce previously published illus- trations: Curtis' s Botanical Magazine; the Field Museum of Natural History, Chicago; the Botan- ical Museum, Harvard University; Christabel King; and the Linnean Society, London. The drawing of Brunfelsia amazonica was prepared by Margaret Tebbs, and all the distribution maps were prepared by Malcolm Penn. Kai Vollesen of the Royal Botanic Gardens, Kew, provided the photograph of B. obovata cv. Lindeniana, and Dr. Jim Mallet of University College, London, pro- vided the photograph of Methona larvae. The Photographic Unit at the Natural History Museum produced several of the figures published here, and Marian Short checked the manuscript. XII. Plowman's Acknowledgments The following acknowledgments are from Tim- othy Plowman 's dissertation: Many persons have generously given to me their time and assistance during the course of this study. I gratefully acknowledge their help, which has both enhanced the quality and facilitated the completion of this work. I would especially like to express my sincere gratitude to Prof. Richard Evans Schultes, who first suggested the genus Brunfelsia as a problem in ethnopharmacology and provided continued support and encouragement throughout the study. His profound interest in the plants and the people of South America demonstrated to me the impor- tance of field studies in ethnobotanical and taxo- nomic research. During the course of my field work, many in- dividuals graciously offered their facilities and hospitality, food, and lodging, and generally made my travels more pleasant and scientifically re- warding. I am particularly indebted to the follow- ing persons: Dr. Leandro Aristeguieta in Venezue- la; Drs. Jesus Idrobo, Eduardo Mora Osejo, and Alvaro Fernandez, Sr. Rafael Wandurraga, and Sr. Pedro Juajibioy Chindoy in Colombia; Dr. Ramon Ferreyra and Sr. Fernando Tina in Peru; Dr. Julian Camara Hernandez and Drs. Antonio and Carmen Krapovickas in Argentina; Dr. Roberto Klein and Sr. Dimitri Sucre in Brazil; Drs. George R. Proc- tor and C. D. Adams in Jamaica; and Dr. Roy Woodbury in Puerto Rico. The following made many useful suggestions and criticisms and assisted in solving taxonomic problems: Dr. Cleofe Calderon, Dr. Jose Cuatre- casas, Dr. William T Gillis, Dr. Leslie A. Garay, Mr. William Grime, Prof. Richard A. Howard, Mr. Tom E. Lockwood, Prof. Robert F. Raffauf, Prof. Reed C. Rollins, Dr. Velva E. Rudd, Dr. Elizabeth A. Shaw, Dr. Bernice G. Schubert, Dr. Lyman B. Smith, Dr. Thomas R. Soderstrom, Dr. William T. Stearn, Prof. Rolla M. Tryon, Prof. Carroll E. Wood, and Dr. John J. Wurdack. Technical assistance was provided by Drs. Alice F. Tryon, Umesh Banerjee, and Ihsan Al-Shehbaz and Mr. Gilbert Pierce. Dr. Fred Barkley and Mr. Eugene Courtney at the Botanical Research Station in Woburn, Mas- sachusetts, Mr. Al Fordham of the Arnold Arbo- retum, and Mr. Conrad Smith of Harvard Univer- sity provided much help with the care of living material. Bibliographic services and many kindnesses were provided by the staff of the libraries of the Gray Herbarium and Arnold Arboretum, particu- larly Mrs. Patricia Hall, and by Mrs. Mary Gau- det, Mrs. Lilian Hanscom, and Miss Esther Reyn- olds of the Botanical Museum. I would also like to thank Ms. Lynda Bates and Judith Gronim for the line drawings that appear in the text, and Mr. Coburn Bennett, Ms. Amina Anderson, and Mr. John Lupo, who were of in- valuable assistance with the photographic work. This study was supported in part by grants from the National Defense Education Act Title IV Fel- lowship and National Institutes of Health Training Grant (T-01 GM 00036). Field work was partly funded through the Evolutionary Biology Com- mittee, Harvard University (National Science Foundation Training Grant GB-7346), and the generosity of Mrs. Edward C. Sweeney. I am grateful to the curators of the herbaria mentioned in the text, who lent specimens of Brunfelsia used in this study. Lastly I would like to express my deep grati- tude to my parents, Dr. and Mrs. J. Wesley Plow- man, who have continuously supported and en- couraged me. XIII. Literature Cited ADAMS, C. D. 1972. Flowering Plants of Jamaica. Uni- versity of the West Indies, Mona, Jamaica, pp. 646- 648. 120 FIELDIANA: BOTANY ANDERSON, E. 1967. Plants, Man and Life. University of California Press, Berkeley, Calif., 251 pp. ARBER, A. 1912. Herbals, Their Origin And Evolution. Cambridge University Press, Cambridge, England, pp. 46-50, 171-172. BAEHNI, C. 1946. L'ouverture du bouton chez les fleurs de Solanees. Candollea, 10: 399-492. *BAILLON, H. E. 1888. Scrofulariacees. In Histoire des plantes, 9: 360-484. BENSON, L. 1962. Plant Taxonomy. Ronald Press, New York, N.Y., 494 pp. BENTHAM, G. 1835. Scrophularineae in Edward's Botan- ical Register, n. s., 8: sub pi. 1770. . 1846. Scrophulariaceae: Brunfelsia. In A. de Candolle, ed. Prodromus Systematis Naturalis Regni Vegetabilis, 10: 198-201, 590-591. BENTHAM, G., AND J. D. HOOKER. 1873. Genera Plantar- urn, vol. 2. Reeve and Company, London, England, pp. 882-913. BHADURI, P. N. 1933. Chromosome numbers of some solanaceous plants of Bengal. Journal of the Indian Botanical Society, 12(1): 56-64. BONNET, E. 1895. Lettres de Linne a David van Royen. Bulletin de L'Herbier Boissier, 3: 22. BRITTON, N. L., AND P. WILSON. 1925. Botany of Porto Rico and the Virgin Islands, pp. 178-179. In Scientific Survey of Porto Rico and the Virgin Islands, vol. 6, no. 1. New York Academy of Sciences, New York, NY. BROWN, K. S., JR. 1979. Ecologia geografica e evolu9ao nas florestas neotropicas. Thesis, Universidade Estad- ual de Campinas, Sao Paulo, Brazil. BRUNFELS, O. 1530. Herbarum Vivae Eicones. Johannes Schott, Strasbourg, France. CAIN, S. A. 1971. Foundations of Plant Geography. Haf- ner Publishing Company, New York, N.Y. CARAUTA, J. P. P. 1973. The text of Vellozo's Flora Flu- minensis and its effective date of publication. Taxon, 22(2/3):28 1-284. CHAMISSO, A., AND D. F. L. SCHLECHTENDAL. 1827. Fran- ciscea. In De plantis in expeditione speculatoria Ro- manzoffiana observatis. Linnaea, 2: 600-601. CONSTANCE, L. 1955. Systematics of angiosperms, pp. 430, 435-436. In A Century of Progress in the Natural Sciences. California Academy of Natural Sciences, San Francisco, Calif. CORNER, E. J. H. 1964. The Life of Plants. World Pub- lishing Company, New York, N.Y, pp. 207-226. CROIZAT, L. 1962. Space, Time, Form: The Biological Synthesis. Published by author, Caracas, Venezuela, pp. 138, 304-305. DANERT, S. 1958. Die Verzweigung der Solanaceen im Reproduktiven Bereich. Abhandlungen der Deutschen Akademie der Wissenschaften zu Berlin, 6: 1-183. D'ARCY, W. G. 1971. Solanaceae Studies I. Notes. An- nals of the Missouri Botanical Garden, 57: 259-263. * . 1974. Solanaceae. In Woodson, R. E., Jr., and R. W. Schery, eds., Flora of Panama. Annals of the Missouri Botanical Garden, 60(3): 573-780. -. 1989. (951) Proposal to conserve spelling * References marked with an asterisk did not appear in Plowman's original manuscript; they were added by the editors. 7450 Brunfelsia (Solanaceae). Taxon, 38: 510-511. DARLINGTON, C. D., AND E. K. JANAKI-AMMAL. 1945. Chromosome Atlas of Cultivated Plants. George Allen and Unwin Ltd., London, England, p. 253. DARLINGTON, C. D., AND A. P. WYLIE. 1955. Chromo- some Atlas of Flowering Plants. George Allen and Unwin Ltd., London, England. DAVIDSE, G. 1981. Chromosome numbers of miscella- neous angiosperms. Annals of the Missouri Botanical Garden, 68: 222-223. DAVIS, P. H., AND V. H. HEYWOOD. 1963. Principles of Angiosperm Taxonomy. Van Nostrand Co., Princeton, N.J., p. 34. DON, D. 1829. Observations on the characters and affin- ities of Darwinia, etc. Edinburgh New Philosophical Journal, July, p. 81. DON, G. 1837. A General System of Gardening and Bot- any, vol. 4. Rivington et al., London, England, pp. 476-477. *DRUMMOND, B. A., Ill, AND K. S. BROWN, JR. 1987. Ithomiinae (Lepidoptera: Nymphalidae): Summary of known larval food plants. Annals of the Missouri Bo- tanical Garden, 74: 341-358. *£DMONDS, J. M. 1983. Seed coat structure and devel- opment in Solanum section Solanum (Solanaceae). Botanical Journal of the Linnean Society, 87: 229- 246. EITEN, G. 1969. A vegeta£ao do Estado Sao Paulo. Bol- etim do Instituto de Botanica, 7: 1-147. ENDLICHER, S. 1839. Genera Plantarum. Fr. Beck, Vi- enna, Austria, p. 676. ERDTMAN, G. 1960. The acetolysis method. Svensk Bo- tanisk Tidskrift, 54: 561-564. FONT QUER, P. 1965. Diccionario de Botanica. Editorial Labor S. A., Barcelona, Spain, p. 220. *FOOTT, W. H. 1967. Occurrence of Frumenta nundi- nella (Lepidoptera: Gelechiidae) in Canada. Canadian Entomologist, 99: 443-444. FRANCEY, P. 1935. Monographic du Genre Cestrum L. Candollea, 6: 46-398. — . 1936. Monographic du Genre Cestrum L., Par- tie II. Candollea, 7: 1-132. GOODSPEED, T. H. 1 954. The Genus Nicotiana. Chronica Botanica Company, Waltham, Massachusetts. GOTTSCHALK, W. 1954. Die Chromosomenstruktur der Solanaceen und der Berucksichtigung phylogenetisch- er Fragestellungen. Chromosoma, 6: 539-626. GRISEBACH, A. H. R. 1861. Flora of the British West Indian Islands. Lovell Reeve and Company, London, England, pp. 431-432. GREUTER, W [CHAIRMAN OF THE EDITORIAL COMMITTEE], F R. BARRIE, H. M. BURDET, W. G. CHALONER, V. DEMOULIN, D. L. HAWKSWORTH, P. M. JORGENSEN, D. H. NICOLSON, P. C. SILVA, TREHANE, AND J. MCNEILL [SECRETARY TO THE EDITORIAL COMMITTEE]. 1994. In- ternational Code of Botanical Nomenclature. Regnum Veg. 131. Konigstein. *HAEGI, L. 1981. A conspectus of Solanaceae tribe An- thocercidae. Telopea, 2(2): 173-180. PLOWMAN: REVISION OF BRUNFELSIA 121 HASSLER, E. 1919. Solanacea paraguariensia vel minus cognita. II. Repert. Spec. Nov. Regni Veg., 15: 217- 245. HITCHCOCK, C. J. 1932. A monographic study of the ge- nus Lycium in the western hemisphere. Annals of the Missouri Botanical Garden, 19: 179-374. HOOKER, W. J. 1828. Franciscea Hopeana. Hooker's Bo- tanical Magazine, 55: t. 2829. HOPPE, H. A. 1958. Drogenkunde. Cram, De Gruyter and Company, Hamburg, Germany, pp. 396-397. *HuNT, D. R. 1978. Brunfelsia australis (Solanaceae). Curtis's Botanical Magazine, 182: 49-51, t. 760. *HUNZIKER, A. T., AND R. SuBiLS. 1986. Studies of So- lanaceae XXII: A new genus of the tribe Salpiglossi- deae. Kurtziana, 18: 121-132. HUTCHINSON, J. 1969. Evolution and Phylogeny of Flow- ering Plants. Academic Press, London, England, pp. 615-622, 631-634. *JARVIS, C. E., F. R. BARRIE, D. M. ALLAN, AND J. L. REVEAL. 1993. A list of Linnaean generic names and their types. In Regnum Vegetabile, vol. 127. Koeltz Scientific Books, Konigstein. JUSSIEU, A. L. DE. 1791. Genera Plantarum. Ziegler and Sons, Zurich, Switzerland, p. 142. *KNAPP, S. 1989. A revision of the Solarium nitidum species group (section Holophylla pro parte): Sola- naceae. Bulletin of the British Museum of Natural History, Botany, 19: 63-103. KRUG, C. W. L., AND I. URBAN. 1897. Diagnosen Neuer Arten. Notizblatt des Koniglichen Botanischen Gar- tens und Museums zu Berlin, 1(10): 324. LAMAS, G. 1973. Taxonomia e Evolu9ao dos Generos Ituna Doubleday (Danainae) e Paititia gen. n., Thyr- idia Hiibner e Methona Doubleday (Ithomiinae) (Nymphalidae, Lepidoptera). D.Sc. thesis, Universi- dade de Sao Paulo, Brazil. LAWRENCE, G. H. M. 1965. Taxonomy of Vascular Plants. Macmillan Company, New York, N.Y., 823 pp. LECOINTE, P. 1947. Arvores e Plantas Uteis, ed. 2. Ama- zonia Brasileira 3, sen 5. Brasiliana, 251: 279. LEENHOUTS, P. W. 1968. A Guide to the Practice of Her- barium Taxonomy. Regnum Vegetabile, vol. 58. In- ternational Bureau for Plant Taxonomy and Nomen- clature, Utrecht, Netherlands, p. 60. *LESTER, R. N., AND P. DURANDS. 1984. Enzyme treat- ment as an aid in the study of seed surface structures of Solarium species. Annals of Botany, 53: 129-131. LEWIS, W. H. 1971. High floristic endemism in low cloud forests of Panama. Biotropica, 3(1): 78-80. *LIEBHERR, J. K. 1988. General patterns in West Indian insects, and graphical biogeographic analysis of some circum-Caribbean Platynus beetles (Carabidae). Sys- tematic Zoology, 37: 385-409. LINNAEUS, C. 1742. Genera Plantarum, ed. 2. Leiden, Netherlands. . 1753. Species Plantarum. Stockholm, Sweden. — . 1754. Genera Plantarum, ed. 5. Stockholm, gio "De La Salle," no. 16. Vedado, La Habana, Cuba, pp. 362-365. *LLOYD, H. A., H. M. FALES, M. E. GOLDMAN, D. M. JERINA, T. PLOWMAN, AND R. E. SCHULTES. 1985. Brun- felsiamidine: A novel convulsant from the medicinal plant Brunfelsia grandiflora. Tetrahedron Letters, 26(22): 2623-2624. MACBRIDE, J. F. 1962. Flora of Peru: Solanaceae. Field Museum of Natural History, Botanical Series, 8(5-B, no. 1): 151-154. MADHAVADIAN, P. 1968. Chromosome numbers in South Indian Solanaceae. Caryologia, 21(4): 343-347. MALFAIT, B. T, AND M. G. DINKELMAN. 1972. Circum- Caribbean tectonic and igneous activity and the evo- lution of the Caribbean plate. Geological Society of America Bulletin, 83: 251-272. *MALLET, J. 1993. Speciation, raciation, and color pat- tern evolution in Heliconius butterflies: Evidence from hybrid zones. In Hybrid Zones and the Evolutionary Process. Oxford University Press, New York, N.Y, pp. 226-260. MARTIUS, C. F. P. VON. 1843. Systema Materiae Medicae Vegetabilis Brasiliensis. F. Fleischer, Leipzig, Ger- many, p. 67. MEISNER, C. F. 1840. Plantarum Vascularum Genera, vol. 1. Leipzig, Germany, p. 307. . 1840. Plantarum Vascularum Genera, vol. 2. Leipzig, Germany, pp. 218-219. MERCK AND COMPANY. 1930. Merck Index, ed. 4. Rah- way, N. J. METCALFE, C. R., AND L. CHALK. 1950. Anatomy of the Dicotyledons, vol. 2. Clarendon Press, Oxford, En- gland, p. 968. MIERS, J. 1849. Observations upon several genera hither to placed in Solanaceae, and upon others intermediate between that family and the Scrophulariaceae. Annals and Magazine of Natural History, Series 2, 3: 161- 179. . 1850. XXIII. Contributions to the botany of South America. Annals and Magazine of Natural His- tory, Series 2, 5: 247-250. -. 1855. Illustrations of South American Plants, vol. 1. H. Bailliere, London, England. -. 1857. Illustrations of South American Plants, Sweden. LIOGIER, E. E. 1957. Solanaceae. In Sauget, J. S., and E. E. Liogier, eds., Flora de Cuba, vol. 4, Contribuciones Ocasionales del Museu de Historia Natural del Cole- vol. 2. H. Bailliere, London, England. MONACHINO, J. V. 1953. Mire, a new species of Brun- felsia from Bolivia. Phytologia, 4(5): 342-347. MORI, S. A., AND L. A. MATTOS SILVA. 1979. The her- barium of the 'Centro de Pesquisas do Cacau' at Ita- buna, Brazil. Brittonia, 31: 177-196. *NELSON, B. W., C. A. C. FERREIRA, M. F. DA SILVA, AND M. L. KAWASAKI. 1990. Endemism centres, refugia and botanical collection density in Brazilian Amazo- nia. Nature, 345: 714-716. *OLMSTEAD, R. G., AND J. D. PALMER. 1992. A chloro- plast DNA phylogeny of the Solanaceae: Subfamilial relationships and character evolution. Annals of the Missouri Botanical Garden, 79: 346-360. *OLMSTEAD, R. G., AND J. A. SWEERE. 1994. Combining data in phylogenetic systematics: An empirical ap- proach using three molecular data sets in the Solana- ceae. Systematic Biology, 43(4): 467-481. 122 FIELDIANA: BOTANY *OLMSTEAD, R. G., J. A. SWEERE, R. E. SPANGLER, L. BOHS, AND J. D. PALMER. In press. Phylogeny and pro- visional classification of the Solanaceae based on chloroplast DNA. In Solanaceae IV. Royal Botanic Gardens, Kew, Richmond, Surrey, England. *PAGE, R. D. M., AND C. LYDEARD. 1994. Towards a cladistic biogeography of the Caribbean. Cladistics, 10: 21-41. PECKOLT, T. 1909. Heil-und Nutzpflanzen Brasiliens. Berichte der Deutschen Pharmazeutischen Gesells- chaft, 19: 307-315. Piso, W. 1648. De medicina Brasiliensi, p. 85. In Piso, W., and G. Marcgrave, Historia Naturalis Brasiliae. F. Hack, Leiden, Netherlands. . 1658. De Indiae Utriusque Re Naturali et Med- ica, F. Hack, Amsterdam, Netherlands, p. 184. PLOWMAN, T. 1973. Four new Brunfelsias from North- western South America. Botanical Museum Leaflets of Harvard University, 23(6): 245-272. * . 1977. Brunfelsia in ethnomedicine. Botanical Museum Leaflets of Harvard University, 25: 289-320. -. 1978. A new section of Brunfelsia: Section Forest: Medicinal and Toxic Plants of the Northwest Amazon. Dioscorides Press, Portland, Oregon. -. 1991. De plantis toxicariis e mundo novo trop- Guianenses. In Hawkes, J. G., ed., Systematic notes in the Solanaceae. Journal of the Linnean Society (Botany), 76: 294-295. 1979. The genus Brunfelsia: A conspectus of taxonomy and biogeography, pp. 475-491. In Haw- kes, J. G., R. N. Lester, and A. D Skelding, eds., The Biology and Taxonomy of the Solanaceae. Academic Press, London, England. -. 1981. Five new species of Brunfelsia from South America (Solanaceae). Fieldiana: Botany, n.s., 8: 1-16. PLUMIER, C. 1703. Catalogus Plantarum Americanarum, p. 12, t. 22. In Nova Plantarum Americanarum Gen- era. J. Boudot, Paris, France. . 1756. Brunfelsia americana. In Burman, J., ed., Plantarum Americanarum Fasc. 3. Amsterdam, Neth- erlands, p. 54, t. 65. POHL, J. E. 1826. Plantarum Brasiliae Icones et Descrip- tiones. Vienna, Austria, pp. 1-8, t. 1-7. PRANCE, G. T. 1973. Phytogeographic support for the theory of Pleistocene forest refuges in the Amazon basin, based on evidence from distribution patterns in Caryocaraceae, Chrysobalanaceae, Dichapetalaceae and Lecythidaceae. Acta Amazonica, 3: 5-27. RAFILL, C. P. 1928. Brunfelsias. Gardener's Chronicle and Gardening Illustrated, Series 3, 83: 370. RATERA, E. L. 1944. Numero de cromosomas de algunas Solanaceas argentinas. Instituto de Genetica, Univer- sidad de Buenos Aires, 2(9): 105-110. RICHARDS, P. W. 1952. The Tropical Rain Forest. Cam- bridge University Press, Cambridge, England, pp. 94- 95. RICKETT, H. W. 1944. The classification of inflores- cences. The Botanical Review, 10: 187-231. SCHMIDT, J. A. 1864. Scrophulariaceae. In Martius, C. P. F, ed., Flora Brasiliensis 8(1): 230-330. *SCHOLTZ, C. H. 1978. Notes on plant galls formed by Lepidoptera in the Transvaal. Journal of the Entomo- logical Society of South Africa, 41(1): 45-49. *SCHULTES, R. E., AND R. F. RAFFAUF. 1990. The Healing icale commentationes XXXVI: Phytochemical and ethnopharmacological notes on the Solanaceae of the northwest Amazon, pp. 25-49. In Hawkes, J. G., R. N. Lester, and N. Estrada R., eds., Solanaceae. III. Taxonomy, Chemistry, Evolution. Royal Botanic Gar- dens, Kew, Richmond, Surrey, England. SMITH, L. B. 1962. Origins of the flora of southern Bra- zil. Contributions from the United States National Herbarium, 35(3): 25-249. SMITH, L. B., AND R. J. DOWNS. 1966. In Reitz, P. R., ed. Flora Ilustrada Catarinense. Part 1 , Solanaceas, pp. 303-308. SMITH, L. B., AND R. C. SMITH. 1967. Itinerary of Wil- liams John Burchell in Brazil, 1825-1830. Phytologia, 14(8): 492-505. *SOLOMON, B. P., AND S. J. McNAUGHTON. 1979. Nu- merical and temporal relationships in a three-level food chain. Oecologia, 42: 47-56. *SOUEGES, R. 1907. Developpement et structure du teg- ument seminal chez les Solanacees. Annales des Sci- ences Naturelles, Botanique, Paris, 6:1-124. STAFLEU, F. A. 1967. Taxonomic Literature. International Bureau for Plant Taxonomy and Nomenclature, Utrecht, Netherlands, pp. 454-455. STAFLEU, F. A., AND R. S. COWAN. 1983. Taxonomic Lit- erature 2. Vol. 4, Sal-Ste. Bohn, Scheltema and Hol- kema, Utrecht, Netherlands. — . 1985. Taxonomic Literature 2. Vol. 5, P-Sak. Bohn, Scheltema and Holkema, Utrecht. STANDLEY, P. C. 1940. Studies of American plants IX. Publications of the Field Museum of Natural History, Botanical Series, 22(1): 47. STANDLEY, P. C. AND J. A. STEYERMARK. 1940. Studies of Central American plants II. Publications of the Field Museum of Natural History, Botanical Series, 22(5): 376. STEWARD, J. H. 1948. Tribes of the montana: An intro- duction, p. 531. In Steward, J. H., ed.. Handbook of South American Indians. Vol. 3, The Tropical Forest Tribes. Bulletin 143, Bureau of American Ethnology, Smithsonian Institution, Washington, DC. STEWARD, J. H., AND A. METRAUX. 1948. Tribes of the Peruvian and Ecuadorian montana, pp. 594, 605. In Steward, J. H., ed., Handbook of South American In- dians. Vol. 3, The Tropical Forest Tribes. Bulletin 143. Bureau of American Ethnology, Smithsonian Institu- tion, Washington, DC. STEYERMARK, J. A. 1968. Contribuciones a la flora de la Sierra de Imataca, Altiplanicie de Nuria y regi6n ad- yacente del Territorio Federal Delta Amacuro al sur del Rib Orinoco. Acta Botanica Venezuelica, 3: 49- 175. . 1979. Plant refuge and dispersal centres in Ven- ezuela— Their relict and endemic element, pp. 185- 221. In Larsen, K., and L. Holm-Nielsen, eds., Trop- ical Botany. Academic Press, London, England. SWANSON, C. P. 1957. Cytology and Cytogenetics. Pren- tice-Hall, Englewood Cliffs, N. J. p. 116. *SYMON, D. E. 1979. Fruit diversity and dispersal in PLOWMAN: REVISION OF BRUNFELSIA 123 Solatium in Australia. Journal of the Adelaide Botan- ical Garden, 1(6): 321-331. TASTEVIN, C. 1922. Nomes de plantas e animaes em lin- gua Tupy. Revista do Museu Paulista, 13: 688-763. TAUBERT, P. 1896. Beitrage zur Kenntnis der Flora des centralbrasilianischen Staates Goyaz. Botanische Jahr- biicher fur Systematik, Pflanzengeschichte und Pflan- zengeographie 21: 402-457. TRYON, R. M., JR. 1972. Endemic areas and geographic speciation in tropical American ferns. Biotropica, 4(3): 121-131. URBAN, I. 1906. Vitae, itineraque collectorum botani- corum. In Martius, C. P. F. von, ed., Flora Brasiliensis, 1(1): 138. VAVILOV, N. E. 1950. The Origin, Variation, Immunity and the Breeding of Cultivated Plants. Chronica Bo- tanica Company, Waltham, Massachusetts. VELLOZO, J. M. DA CONCEIQAO. 1829 ["1825"]. Flora Fluminensis. Archivos do Museu Nacional do Rio de Janeiro V. VUILLEUMIER, B. S. 1971. Pleistocene changes in the fau- na and flora of South America. Science, 173: 771- 779. WETTSTEIN, R. VON. 1895. Solanaceae. In A. Engler, A., and K. Prantl, eds., Die Naturlichen Pflanzenfamilien, 4(3b): 4-39. WURDACK, J. J. 1970. Erroneous data in Glaziou collec- tions of Melastomataceae. Taxon, 19(6): 911-913. Appendix I: List of Exsiccatae Acosta Soils 7576 (grandiflora subsp. grandiflora) Aguilar 1302 (australis), 841 (australis) Aires do Nascimento 19 (uniflora) Albuquerque et al. 1269 (grandiflora subsp. gran- diflora) Allard 21176 (grandiflora subsp. grandiflora), 22096 (grandiflora subsp. schultesii) Allemao 1249 (uniflora} Allemao & Cysneiro 1250 (uniflora) Almeida 1286 (brasiliensis subsp. brasiliensis) Alston & Lutz 138 (latifolid) Amaral, A. s.n. (brasiliensis subsp. brasiliensis) Amaral, I. L., et al. 803 (mire) Anderson 11129 (mire), 6751 (obovata var. cori- acea), 8949 (brasiliensis subsp. brasiliensis) Anderson et al. 37048 (uniflora) Andrade 1024 (brasiliensis subsp. brasiliensis) Angeli 250 (brasiliensis subsp. brasiliensis) Aparicio & Edmundo 868 (hydrangeiformis subsp. capitata) Araujo 3405 (uniflora), 954 (hydrangeiformis subsp. hydrangeiformis), 968 (uniflora) Araujo & Kennedy 24 Mar. 1971 (latifolia) Araujo et al. 3202 (uniflora) Aristeguieta 1669 (grandiflora subsp. schultesii), 6153 (uniflora) Assis 102 (brasiliensis subsp. brasiliensis), 137 (brasiliensis subsp. brasiliensis), 142 (bras- iliensis subsp. brasiliensis), 181 (brasiliensis subsp. brasiliensis) Atamp. (?) 6560 (brasiliensis subsp. brasiliensis) Bailey & Bailey 758 (brasiliensis subsp. bras- iliensis) Baker 61 (grandiflora subsp. grandiflora) Balansa 2240 (australis) Balgooy 1592 (australis) Bang 1398 (grandiflora subsp. schultesii), 2352 (mire) Barbosa Rodrigues 261 (uniflora) Barclay 911 (macrocarpa) Barreto 7787 (brasiliensis), 7790 (brasiliensis), 9749 (brasiliensis subsp. macrocalyx) Beck 13335 (grandiflora), 4937 (mire) Beck & Liberman 9379 (boliviano) Bedi s.n. (uniflora) Belshaw 3130 (grandiflora subsp. schultesii) Bernardi 18138 (australis), 20495 (australis), 3310 (grandiflora subsp. schultesii) Bertoni 2939 (australis), 2974 (pilosa), 3434 (australis) Besse et al. 513 (mire) Beyrich s.n. (brasiliensis subsp. brasiliensis) Bezerra 334 (uniflora) Black 48-2905 (amazonica) Blanchet 1455 (clandestina), 2616 (uniflora), 3352 (clandestina), 3354 (clandestina), 3862 (uniflora), Jun. 1844 (uniflora), s.n. (clandes- tina), s.n. (clandestina), s.n. 1843 (clandestina), s.n. 1847 (uniflora), s.n. 1857 (hydrangeiformis subsp. hydrangeiformis) Blanco 826 (imatacana) Boeke 1224 (grandiflora subsp. grandiflora), 1477 (mire) Boer 1047 (guianensis) Bonpland 528 (australis) Bornmuller 219 (australis) Bowie & Cunningham 100 (hydrangeiformis subsp. hydrangeiformis), 228 (brasiliensis subsp. brasiliensis) Brade 10569 (brasiliensis subsp. brasiliensis), 10583 (brasiliensis subsp. brasiliensis), 14074 (brasiliensis subsp. macrocalyx), 18643 (bon- odora), 18785 (brasiliensis subsp. brasiliensis), 20953 (brasiliensis subsp. macrocalyx), 21218 (brasiliensis subsp. brasiliensis), 6036 (pauci- flora), 6037 (brasiliensis subsp. macrocalyx), 7625 (brasiliensis subsp. macrocalyx), 8088 124 FIELDIANA: BOTANY (pauciflord), Oct. 1928 (hydrangeiformis subsp. capitatd) Brade & Duarte 20119 (pauciflord) Brade & Tamandare 6576 (brasiliensis subsp. macrocalyx) Braga 1169 (brasiliensis subsp. brasiliensis), 182 (brasiliensis subsp. brasiliensis), 203 (bras- iliensis subsp. brasiliensis) Branddo 2066 (brasiliensis subsp. brasiliensis) Bristan 1220 (chocoensis), 566 (chocoensis), 569 (chocoensis) British Guiana Forest Department 4422 (marti- ana) Brooke 5653 (boliviano) Brunei sp. nov. 1888 (hydrangeiformis subsp. capitatd) Buchtien 1299 (grandiflora subsp. schultesii), 1300 (grandiflora subsp. schultesii), 2029 (grandiflora subsp. schultesii), 5640 (mire), 7615 (mire) Bunting et al. 10351 (hydrangeiformis subsp. hy- drangeiformis) Burchell 1372 (brasiliensis subsp. brasiliensis), 8494 (burchellii), 8653 (burchellii), 8752 (bur- chellii) Buttura 157 (australis) Camargo 2123 (cuneifolia), 3 Oct. 1957 (cunei- folia) Campos Novaes 214 (pauciflord), 216 (pilosa), 2960 (pauciflord), 7756 (uniflora) Campos Porto 173 (brasiliensis subsp. macroca- lyx), 1879 (hydrangeiformis subsp. capitatd), 3381 (brasiliensis subsp. brasiliensis), 3400 (pauciflord), s.n. (brasiliensis subsp. bras- iliensis) Capanema s.n. (brasiliensis subsp. brasiliensis) Caprilioni 1659 (pilosa) Carauta 2420 (brasiliensis subsp. brasiliensis) Carauta & de Oliveira 1523 (brasiliensis) Cardenas 2813 (mire), 3380 (uniflora), 5216 (un- iflora), 5952 (uniflora) Carvalho 83 (pilosa) Carvalho & Gatti 795 (clandestina) Castellanos 23004 (uniflora), 23294 (obovata var. coriacea), 23973 (brasiliensis subsp. bras- iliensis) Cavalcante 2518 (guianensis) Chiar s.n. (australis) Claussen 146 (brasiliensis subsp. brasiliensis), 147 (brasiliensis subsp. brasiliensis), 157 (brasiliensis subsp. brasiliensis), 201 (hydran- geiformis subsp. hydrangeiformis), 334 (bras- iliensis subsp. brasiliensis), Nov. 1839 (hydran- geiformis subsp. hydrangeiformis), Nov. 1842 (hydrangeiformis subsp. capitatd), s.n. (bras- iliensis subsp. brasiliensis), s.n. (hydrangeifor- mis subsp. hydrangeiformis), s.n. 1841 (hy- drangeiformis subsp. hydrangeiformis) Constantino Nov. 1931 (brasiliensis subsp. bras- iliensis) Correa & Dressier 471 (dwyeri) Croat 13072 (dwyeri), 36856 (australis) Cuatrecasas 10611 (grandiflora subsp. schulte- sii), 10795 (grandiflora subsp. schultesii), 11275 (grandiflora subsp. schultesii) Curran 619 (brasiliensis subsp. brasiliensis), 623 (brasiliensis subsp. brasiliensis), s.n. (australis) Cutler 8271 (uniflora) D'Arcy 3959 (dwyeri) D'Arcy et al. 3948 (dwyeri) Davidse & Ramamoorthy 10709A (brasiliensis subsp. brasiliensis) Davidse et al. 11611 (brasiliensis) Davidson 3846 (uniflora) Davis & Marshall 1106 (mire) Davis & Yost 958 (grandiflora subsp. schultesii) Dawe 241 (grandiflora subsp. schultesii), Mar. 1914 (uniflora) Dobereiner & Tokarnia 732 (cuneifolia), 841 (australis) Dodson 2822 (grandiflora subsp. schultesii) Drummond 7330 (grandiflora subsp. schultesii) Duarte 2029 (brasiliensis subsp. brasiliensis), 5477 (bonodora), 631 (brasiliensis subsp. bras- iliensis), 8445 (hydrangeiformis subsp. hydran- geiformis), 8741 (brasiliensis subsp. bras- iliensis) Duarte & Pereira 22 Oct. 1964 (hydrangeiformis subsp. hydrangeiformis), 9 Dec. 1948 (hydran- geiformis subsp. capitatd) Duarte & Santos 102 (obovata var. coriacea) Duarte de Barros 1195 (brasiliensis subsp. bras- iliensis) Ducke 2954 (uniflora), 430 (amazonica), 865 (grandiflora subsp. schultesii), 29 Nov. 1928 (hydrangeiformis subsp. capitatd), 10 Jan. 1930 (mire), s.n. (amazonica), s.n. (guianensis), s.n. (martiana) Ducke & Kuhlmann s.n. (brasiliensis subsp. bras- iliensis) Duke 11351 (chocoensis), 13331 (chocoensis), 16047 (chiricaspi), 5300 (chocoensis), 9429 (dwyeri) Dupre s.n. 1842 (brasiliensis subsp. brasiliensis) Dusen 10305 (pauciflord), 10354 (pauciflord), 10443 (pilosa), 13180 (pauciflord), 13298 (pauciflord), 13327 (pauciflord), 13562 (bras- iliensis subsp. macrocalyx), 17301 (pauciflord), PLOWMAN: REVISION OF BRUNFELSIA 125 2023 (brasiliensis subsp. macrocalyx), 3388 (brasiliensis subsp. macrocalyx), 569 (bras- iliensis subsp. macrocalyx), 7077 (pilosa), 8679 (pauciflora), 8961 (pauciflora) Dwyer 7280 (dwyeri) Dwyer & Ganger 7349 (dwyeri) Dwyer et al. 5051 (dwyeri) Edwall, 15183 (hydrangeiformis subsp. capitata), 1981 p.p. (brasiliensis subsp. macrocalyx), 1981 p.p. (obovata var. obovatd), 4329 (pau- ciflora), 6133 (uniflora) Elburg 9819 (guianensis) Emmerich 985 (brasiliensis subsp. brasiliensis) Engler 1006 (brasiliensis subsp. brasiliensis), 1009 (brasiliensis subsp. brasiliensis), s.n. (brasiliensis subsp. brasiliensis) Escobar 359 (dwyeri) Fabris 4502 (uniflora) Facultad Ciencias For estates s.n. (uniflora) Farney et al. 584 (brasiliensis subsp. bras- iliensis), 597 (brasiliensis) Fernandes, A., s.n. (uniflora) Fernandez 365 (macrocarpa) Ferreira & Costa 16 (grandiflora subsp. grandi- flora) Ferreira, V.F., et al. 887 (rupestris) Ferreyra 4760 (grandiflora subsp. schultesii), 7778 (grandiflora subsp. schultesii), 985 (gran- diflora subsp. grandiflora) Fiebrig 49 (australis), 5460 (pilosa) Filho S. J. 8 (uniflora) Filho et al. 3219 (uniflora) Plaster 56 (hydrangeiformis subsp. capitata) Florschiitz & Maas 2924 (guianensis), 3094 (gui- anensis) Forestry Department of British Guiana 7495 (gui- anensis) Franklin & Glaziou 5319 (hydrangeiformis subsp. capitata) Frenzel 666 (cuneifolia) Froes 11620 (burchellii), 12681\46 (uniflora), 20067 (clandestina), 20210 (clandestina), 20535 (amazonica), 23068 (uniflora), 23974 (grandiflora subsp. schultesii), 24035 (grandi- flora subsp. schultesii), 25650 (martiana), 28393 (martiana) Garcia Barriga 4586 (grandiflora subsp. schul- tesii) Gardner, G. 1095 (uniflora), 1798 (uniflora), 2432 (uniflora), 248 (brasiliensis subsp. bras- iliensis), 5062 (uniflora), 5063 (brasiliensis subsp. brasiliensis) 5065 (hydrangeiformis subsp. hydrangeiformis), 5566 (uniflora), 563 (hydrangeiformis subsp. capitata), 564 (bras- iliensis subsp. brasiliensis) May 1838 (uniflora) Gaudichaud 442 (brasiliensis subsp. brasiliensis), 705 (pilosa) Gehrt 3667 (brasiliensis subsp. brasiliensis) Gentry 16249 (grandiflora subsp. schultesii) Gentry & Clewell 7138 (chocoensis) Gentry et al. 15707 (grandiflora subsp. schulte- sii), 18573 (grandiflora subsp. schultesii), 18664 (grandiflora subsp. schultesii) Germain 171 (brasiliensis subsp. brasiliensis) Gill 45 (grandiflora subsp. schultesii) Gines 3605 (uniflora) Ginzberger 93 (brasiliensis subsp. macrocalyx) Glaziou 11392 (hydrangeiformis subsp. hydran- geiformis), 11393 (hydrangeiformis subsp. hy- drangeiformis), 11394 (brasiliensis subsp. brasiliensis), 11395 (brasiliensis subsp. macro- calyx), 12110 (australis), 13478 (brasiliensis subsp. macrocalyx), 14171 (brasiliensis subsp. brasiliensis), 14172 (brasiliensis subsp. macro- calyx), 14173 (hydrangeiformis subsp. capita- ta), 14174 (hydrangeiformis subsp. capitata), 15132 (brasiliensis subsp. brasiliensis), 15311 (brasiliensis subsp. brasiliensis), 17168 (pau- ciflora), 17169 (brasiliensis subsp. macroca- lyx), 17170 (brasiliensis subsp. brasiliensis), 21920 (obovata var. coriacea), 357 (uniflora), 4909 (hydrangeiformis subsp. capitata), 5970 (latifolia), 6908 (brasiliensis subsp. macroca- lyx), 811 (pauciflora), 812 (brasiliensis subsp. brasiliensis), 9558 (hydrangeiformis subsp. capitata) Goes & Constantino 543 (brasiliensis subsp. brasiliensis), 883 (brasiliensis subsp. bras- iliensis) Graf 10 (uniflora) Granville 1378 (guianensis), 2204 (guianensis), 2224 (guianensis), B-4854 (guianensis) Guedes 265 (mire), 472 (uniflora) Guevara 2276 (uniflora) Guillemin 41 (brasiliensis subsp. brasiliensis), 171 (brasiliensis subsp. brasiliensis), 825 (brasiliensis subsp. brasiliensis), 954 (bras- iliensis subsp. brasiliensis) Gurgel 27 Feb. 1929 (pilosa) Handro 27 Sep. 1945 (pauciflora), 984 (obovata var. obovata) Harley 21898 (obovata var. coriacea) Harley et al. 16432 (uniflora) Harling & Andersson 11724 (grandiflora subsp. schultesii), 11771 (chiricaspi), 11804 (chiricas- pi), 13906 (grandiflora subsp. grandiflora), 13934 (grandiflora subsp. grandiflora) 126 FIELDIANA: BOTANY Marling et al. 7021 (chiricaspi), 7151 (chiricas- pi), 7474 (grandiflora subsp. schultesii), 7700 (grandiflora subsp. schultesii), 7772 (chiricas- pi) Hashimoto 1 (brasiliensis subsp. brasiliensis) Hassler 12559 (australis), 725 (australis) Hatschbach 1058 (pauciflora), 11878 (bras- iliensis subsp. macrocalyx), 13055 (pauciflora), 15071 (pauciflora), 15159 (cuneifolia), 15300 (pauciflora), 17308 (pauciflora), 17610 (pilo- sa), 18747 (brasiliensis subsp. macrocalyx), 19447 (brasiliensis subsp. macrocalyx), 19937 (pauciflora), 20304 (pauciflora), 20997 (bras- iliensis subsp. macrocalyx), 22278 (pauciflora), 22383 (pilosa), 22771 (pauciflora), 25613 (pi- losa), 26532 (brasiliensis subsp. macrocalyx), 28320 (pilosa), 32637 (pauciflora), 32869 (pi- losa), 34918 (pauciflora), 37876 (pilosa), 39164 (pilosa), 39209 (brasiliensis subsp. mac- rocalyx), 40004 (pauciflora), 40470 (pilosa), 42939 (brasiliensis subsp. brasiliensis), 43192 (australis), 43273 (pauciflora), 43398 (bras- iliensis subsp. brasiliensis), 43843 (obovata var. coriacea), 45295 (pauciflora), 45415 (pi- losa), 515 (pauciflora) Hatschbach & Ahumada 31333 (brasiliensis subsp. brasiliensis) Hatschbach & Cervi 46844 (pauciflora) Hatschbach & Guimaraes 15286 (pilosa), 25473 (pilosa), 30302 (cuneifolia), 45004 (brasiliensis subsp. brasiliensis) Hatschbach & Kummrow 37257 (obovata var. obovata), 45569 (brasiliensis subsp. bras- iliensis) Hatschbach & Pelanda 27941 (uniflora), 27993 (brasiliensis subsp. brasiliensis) Hatschbach & Zelma 50296 (rupestris) Heiner 209 (pauciflora), s.n. (uniflora) Heinrichs 496 (grandiflora subsp. schultesii) Henz 27246 (pilosa), 35305 (pilosa) Heringer 10556 (obovata var. coriacea), 10719 (obovata var. coriacea), 11549 (uniflora), 460 (hydrangeiformis subsp. capitata), 8662 (obo- vata var. coriacea), 8993/1197 (obovata var. coriacea), 9096/1290 (obovata var. coriacea) Heringer et al. 4281 (obovata var. coriacea), 6148 (brasiliensis subsp. brasiliensis) Hermann 11237 (grandiflora subsp. schultesii) Heyligers 438 (guianensis) Hoehne, F. C. 582 (pauciflora), 23169 (pauciflo- ra), 23309 (pilosa), 24405 (pauciflora), 4595 (brasiliensis subsp. macrocalyx), 4920 (bras- iliensis subsp. brasiliensis), 5481 (brasiliensis subsp. brasiliensis), 6129 (brasiliensis subsp. brasiliensis), 6846 (brasiliensis subsp. bras- iliensis), 7953 (pauciflora), 8250 (brasiliensis subsp. macrocalyx) Hoehne, W., 6154 (brasiliensis subsp. bras- iliensis), 6155 (brasiliensis subsp. brasiliensis) Huber 14990 (grandiflora subsp. grandiflora), 1556 (grandiflora subsp. grandiflora), 1562 (grandiflora subsp. grandiflora), 4204 (gran- diflora subsp. schultesii), 4377 (grandiflora subsp. schultesii), 4575 (grandiflora subsp. grandiflora), 4589 (grandiflora subsp. grandi- flora), 887 (guianensis) I.N.T.A. 4435 (pilosa) Irwin 2269 (brasiliensis subsp. brasiliensis), 2674 (uniflora), 2701 (uniflora), 48753 (grandiflora subsp. schultesii) Irwin et al. 13126 (obovata var. coriacea), 13194 (brasiliensis subsp. brasiliensis), 15799 (bras- iliensis subsp. brasiliensis), 19549 (brasiliensis subsp. brasiliensis), 20150 (brasiliensis subsp. brasiliensis), 22077 (rupestris), 22465 (rupes- tris), 26556 (brasiliensis subsp. brasiliensis), 29189 (brasiliensis subsp. brasiliensis), 29407 (brasiliensis subsp. brasiliensis), 30628 (bras- iliensis subsp. brasiliensis), 32406 (uniflora), 47359 (grandiflora subsp. schultesii), 47900 (guianensis), 55978 (guianensis), 8152 (obov- ata var. coriacea), 8514 (obovata var. coriacea) Isabelle 6 (pilosa) Jarenkow 132 (cuneifolia) Johnston 71 (uniflora) Johnston & Miller 265 (uniflora) Joly 1133 (brasiliensis subsp. brasiliensis) Jons son, G. 1054a (pilosa) Jorgensen 2098 (australis), 3661 (australis), 4 Oct. 1909 (pilosa), Nov. 1910 (pilosa) Juajibioy Chindoy 277 (grandiflora subsp. schul- tesii) Jurgen 19 (pilosa) Kennedy 1386 (chiricaspi), 316 (dwyeri) Kermann 270 (hydrangeiformis subsp. capitata) Kermes 1363 (australis) Killip & Smith 26844 (grandiflora subsp. schul- tesii), 27667 (grandiflora subsp. schultesii), 28644 (grandiflora subsp. schultesii) Klein 1751 (pauciflora), 319 (pauciflora), 3518 (pilosa), 4102 (pilosa), 4343 (cuneifolia) Klein & Bresolin 6864 (pauciflora) Klug 741 (grandiflora subsp. schultesii), 2880 (grandiflora subsp. schultesii), 3326 (grandiflo- ra subsp. schultesii) Krapovickas 29261 (australis) Krapovickas & Cristobal 11706 (australis), 15385 (australis) PLOWMAN: REVISION OF BRUNFELSIA 127 Krapovickas & Schinini 31476 (boliviano), 35155 (grandiflora subsp. schultesii) Krapovickas et al. 26092 (australis) Krukoff 1532 (mire), 5284 (grandiflora subsp. grandiflora) Kuhlmann, J.G., 1326 (mire), 1327-k (mire), 141 (obovata var. obovata), 1722 (guianensis), 2287 (mire), 809 (mire), 943 (amazonica), 8 Nov. 1922 (latifolia), 19 Nov. 1928 (uniflora), 3 Feb. 1930 (brasiliensis subsp. brasiliensis), 24 Oct. 1932 (hydrangeiformis subsp. hydran- geiformis), 2 Apr. 1935 (uniflora), 10 Nov. 1943 (uniflora), 8 Oct. 1946 (pilosa) Kuhlmann, M., 2176 (pauciflora), 2714 (pauciflo- ra), 2926 (obovata var. obovata), 4309 (bras- iliensis subsp. macrocalyx), 4359 (brasiliensis subsp. macrocalyx) Kummrow 1575 (pauciflora), 2420 (pauciflora), 2432 (cuneifolia), 94 (pilosa), 970 (brasiliensis subsp. macrocalyx) Lalandes s.n. (brasiliensis subsp. brasiliensis) Landrum et al. 4150 (brasiliensis subsp. bras- iliensis) Langsdorff 43 (hydrangeiformis subsp. hydran- geiformis), s.n. (brasiliensis subsp. bras- iliensis), s.n. (hydrangeiformis subsp. hydran- geiformis) Lanjouw 345 (guianensis) Lanjouw & Lindeman 2137 (guianensis), 2564 (guianensis), 2888 (guianensis) Leite 3660 (pauciflora), 4188 (hydrangeiformis subsp. hydrangeiformis) Lemme s.n. (guianensis) Levy 12 (pauciflora) Lima, A. S. 7859 (mire) Lima, D. de A., 52-1039 (uniflora) Lima, H. C. de, and Ramamoorthy 1224 (bras- iliensis subsp. brasiliensis) Lima, H. C. de, et al. 1637 (uniflora) Lima, H. C. de, et al. 2122 (grandiflora subsp. grandiflora) Lindberg 203 (uniflora) Lindeman 5889 (guianensis), 6069 (guianensis), 6162 (guianensis) Lindeman & Haas 1174 (brasiliensis subsp. mac- rocalyx), 3248 (pilosa), 5157 (brasiliensis subsp. macrocalyx) Lindeman et al. 8233 (pilosa) Lindeman, C. A-597-a (australis), A-597-b (pilo- sa), A-597-c (pilosa) Llamas 1530 (pilosa) Llamas sp. nov. (australis) Lobb 65 (grandiflora subsp. grandiflora), s.n. (brasiliensis subsp. brasiliensis), s.n. (hydran- geiformis subsp. capitata) Lofgren 86 (brasiliensis subsp. brasiliensis), 389 (pilosa), 3986 (brasiliensis subsp. brasiliensis), 948 (pauciflora) Lofgren & Cowan 1660 (pauciflora) Lofgren & Edwall 1435 (uniflora), 2705 (pauci- flora) Longfield 367 (macrocarpa) Lorentz & Hieronymus 258 (uniflora) Luderwaldt 1086 (brasiliensis subsp. bras- iliensis), 2117 (pilosa) Lugo 23 (grandiflora subsp. grandiflora), 2724 (chiricaspi), 2942 (chiricaspi), 2981 (chiricas- pi), 3476 (chiricaspi) Lund 754 (brasiliensis subsp. brasiliensis), 755 (obovata var. obovata), 756 (brasiliensis subsp. brasiliensis) Luschnath Sep. 1833 (uniflora), Nov. 1833 (uni- flora), s.n. 1832 (brasiliensis subsp. bras- iliensis), s.n. 1834 (brasiliensis subsp. bras- iliensis), s.n. 1834 (latifolia) Luteyn et al. 4769 (grandiflora subsp. grandiflo- ra) Lutz 1551 (hydrangeiformis subsp. capitata), Jan. 1925 (brasiliensis subsp. brasiliensis), Boc 14 (hydrangeiformis subsp. capitata) Liitzelburg 4019 (obovata var. coriacea), 6187 (brasiliensis subsp. macrocalyx) 6495 (hydran- geiformis subsp. capitata), 6559 (brasiliensis subsp. macrocalyx) Maas et al. 12963 (grandiflora subsp. grandiflo- ra), 3326 (hydrangeiformis subsp. hydrangei- formis), PI 2709 (grandiflora subsp. grandiflo- ra), P13228 (grandiflora subsp. schultesii) Macedo 3131 (brasiliensis subsp. brasiliensis), 5 Feb. 1948 (uniflora) Madison 986 (mire) Madison et al. 5390 (grandiflora subsp. schulte- sii) Magalhaes 1643 (hydrangeiformis subsp. hydran- geiformis) Maguire 23894 (guianensis) Markgraf 3495 (brasiliensis subsp. brasiliensis) Markgraf & Apparicio 10495 (hydrangeiformis subsp. capitata) Martinelli 1115 (hydrangeiformis subsp. hydran- geiformis), 5135 (uniflora) Martinelli et al. 5665 (uniflora) Martius 1292 (brasiliensis subsp. brasiliensis), 3300 (martiana), 541 (bonodora), Dec. 1817 (brasiliensis subsp. brasiliensis), s.n. (guianen- sis), s.n. (martiana), s.n. (uniflora) Matthews 1320 (grandiflora subsp. schultesii) 128 FIELDIANA: BOTANY Mattos Silva et al. 357 (clandestina) Mattos, J., & Mattos, N., 14284 (brasiliensis subsp. brasiliensis) McDaniel 15328 (grandiflora subsp. schultesii) Melin 186 (grandiflora subsp. schultesii) Mello Barreto 10530 (brasiliensis subsp. bras- iliensis), 7767 (brasiliensis subsp. brasiliensis), 7784 (brasiliensis subsp. brasiliensis), 7786 (brasiliensis subsp. brasiliensis), 7788 (bras- iliensis subsp. brasiliensis), 7789 (brasiliensis subsp. brasiliensis), 7791 (brasiliensis subsp. brasiliensis), 7793 (brasiliensis subsp. bras- iliensis), 7794 (brasiliensis subsp. brasiliensis), 9063 (brasiliensis subsp. brasiliensis), 9479 (rupestris), 9833 (brasiliensis subsp. bras- iliensis) Mello Filho 793 (pilosa) Mexia 4089-a (hydrangeiformis subsp. capitata), 4551 (uniflora), 5182 (uniflora), 5467 (bras- iliensis subsp. brasiliensis), 6444 (grandiflora subsp. schultesii), 7275 (grandiflora subsp. grandiflora) Meyer 12621 (australis), 166 (australis), 3076 (australis) Miers 4495 (brasiliensis subsp. brasiliensis), 4721 (uniflora), Dec. 1837 (hydrangeiformis subsp. capitata), Feb. 1838 (brasiliensis subsp. macrocalyx), s.n. 1838 (hydrangeiformis subsp. capitata) Mikan s.n. (latifolia) Mimura 53 (brasiliensis subsp. brasiliensis) Monies 27471 (australis), 4703 (australis) Mora 1040 (chiricaspi) Moreira 5775 (brasiliensis subsp. brasiliensis), s.n. (brasiliensis subsp. macrocalyx) Morel 3783 (australis), 4144 (australis), 8921 (australis) Mori 6828 (chocoensis) Mori & Bolten 8394 (guianensis), 8575 (guianen- sis) Mori & Boom 14726 (guianensis), 14834 (gui- anensis) Mori et al. 12082 (clandestina), 8767 (guianen- sis), 9775 (clandestina) Moura s.n., 1887 (hydrangeiformis subsp. hy- drangeiformis) Mosen 673 (uniflora) Mueller 435 (pauciflora) Muniez 99 (australis) Neill 6546 (grandiflora subsp. grandiflora) Neill & Palacios 7101 (grandiflora subsp. gran- diflora) Netto s.n. (brasiliensis subsp. brasiliensis) Netto et al. s.n. (brasiliensis subsp. brasiliensis) Neuwied, Pr. von s.n. 1827 (uniflora) Noblick 2038 (uniflora) Occhioni 1131 (brasiliensis subsp. macrocalyx), 9174 (brasiliensis subsp. brasiliensis) Oldeman B-1896 (guianensis) Oliveira 21 (brasiliensis subsp. brasiliensis) Oliveira 100 (pilosa) Otto 1003 (uniflora) Pabst 5393 (pauciflora), 7411 (brasiliensis subsp. brasiliensis), 8340 (brasiliensis subsp. bras- iliensis), 9115 (brasiliensis subsp. brasiliensis) Pabst & Brade 10283 (brasiliensis subsp. macro- calyx) Pabst & Pereira 6023 (uniflora) Paiva & Stehmann s.n. (australis) Palacios et al. 3467 (brasiliensis subsp. bras- iliensis) Palmer 155 (australis) Pearce 290 (grandiflora subsp. grandiflora), 295 (grandiflora subsp. grandiflora), Mar. 1868 (grandiflora subsp. grandiflora) Peckolt, O., Aug. 1934 (hydrangeiformis subsp. hydrangeiformis) Peckolt, T., 71 (hydrangeiformis subsp. capitata) Pedersen 418 (australis), 418a (australis) Pereira 6066 (pauciflora), 9241 (hydrangeiformis subsp. hydrangeiformis), 9451 (brasiliensis subsp. brasiliensis) Perez Arbelaez 688 (grandiflora subsp. schultesii) Perrottet s.n. (guianensis) Picket 2200 (uniflora), 670 (uniflora) Pierroti 6563 (australis) Pinheiro 1980 (clandestina), 2086 (clandestina) Pinkley 202 (grandiflora subsp. grandiflora), 420 (chiricaspi), 43 (grandiflora subsp. grandiflo- ra), 444 (grandiflora subsp. schultesii), 457 (grandiflora subsp. schultesii), 460 (grandiflora subsp. grandiflora), 563 (chiricaspi), 575 (grandiflora subsp. schultesii) Pires 51899 (guianensis) Pires & Silva 10218 (guianensis), 11436 (gui- anensis), 11811 (mire) Plaumann 164 (pilosa) Plowmann 10020 (obovata var. coriacea), 1911 (uniflora), 1919 (imatacana), 2039 (grandiflora subsp. schultesii), 2080 (chiricaspi), 2081 (chiricaspi), 2090 (grandiflora subsp. schulte- sii), 2092 (chiricaspi), 2183 (grandiflora subsp. schultesii), 2494 (grandiflora subsp. schultesii), 2713 (australis), 2723 (australis), 2735 (pilo- sa), 2739 (pauciflora), 3160 (dwyeri), 5825 (grandiflora subsp. schultesii), 5984 (grandiflo- ra subsp. schultesii) PLOWMAN: REVISION OF BRUNFELSIA 129 Plowman & Davis 4852 (mire}, 5000 (mire), 5036 (mire), 5172 (mire) Plowman & de Lima, H.C., 12901 (hydrangeifor- mis subsp. capitata) Plowman & Kennedy 2310 (grandiflora subsp. schultesii) Plowman & Sucre 2786 (brasiliensis subsp. bras- iliensis), 2891 (brasiliensis subsp. macrocalyx), 2906 (brasiliensis subsp. macrocalyx) Plowman et al. 10066 (clandestina), 10087 (clan- destina), 2404 (grandiflora subsp. schultesii), 2407 (grandiflora subsp. schultesii), 6455 (grandiflora subsp. schultesii), 6737 (grandiflo- ra subsp. schultesii), 6939 (grandiflora subsp. schultesii) Poeppig 2206 (grandiflora subsp. schultesii) Pohl 263 (brasiliensis subsp. brasiliensis), 264 (uniflord), 265 (brasiliensis subsp. bras- iliensis), 266 (brasiliensis subsp. brasiliensis), s.n. (uniflord) Poiteau s.n. (guianensis) Prance & Silva 59030 (obovata var. coriaced) Prance et al. 11611 (martiana), 11998 (grandiflo- ra subsp. grandiflora), 13471 (grandiflora subsp. grandiflora), 14551 (grandiflora subsp. schultesii), 3880 (amazonica), 4704 (martiana), 5854 (grandiflora subsp. schultesii), 6570 (grandiflora subsp. schultesii), 6933 (cuneifol- ia), 8265 (mire), 9553 (uniflord) Profice et al. 16 (uniflord) Puttemans 4328 (pilosa), 5890 (brasiliensis subsp. macrocalyx), 6135 (pauciflord) Raben 14 (hydrangeiformis subsp. hydrangeifor- mis), 703 (brasiliensis subsp. brasiliensis), 746 (pauciflord) Ragonese 3646 (australis) Rambo 29147 (australis}, 321 (australis), 37819 (australis), 44295 (pilosa), 49144 (pilosa) Regnell 173 (brasiliensis subsp. brasiliensis), I- 374 (uniflord), 1-375 (brasiliensis subsp. bras- iliensis) Reiss 99 (pilosa) Reitz 2743 (pauciflord), 2768 (pauciflord), 3985 (brasiliensis subsp. macrocalyx), 4375 (bras- iliensis subsp. macrocalyx), C-171 (pilosa), C- 1952 (pauciflord) Reitz & Klein 11250 (australis), 1158 (pauciflo- rd), 14136 (pilosa), 16178 (cuneifolid), 2160 (pauciflord), 4076 (cuneifolid), 4238 (pauciflo- rd), 5601 (pauciflord), 7329 (pauciflord), 7601 (brasiliensis subsp. macrocalyx), 9205 (pauci- flord) Revilla 1724 (grandiflora subsp. schultesii), 304 (grandiflora subsp. schultesii) Riedel 112 (bonodord), 18 (bonodora), 59 (bon- odora), s.n. (bonodora) 1112 (uniflord), 113 (hydrangeiformis subsp. capitata), 1190 (bras- iliensis subsp. brasiliensis), 1249 (brasiliensis subsp. brasiliensis), 1308 (latifolid), 1465 (brasiliensis subsp. brasiliensis), 1466 (bras- iliensis subsp. brasiliensis), 1467 (obovata var. obovata), 467 (brasiliensis subsp. brasiliensis), 11 Dec. 1823 (hydrangeiformis subsp. capita- ta), Oct. 1833 (uniflord), Jan. 1834 (uniflord), s.n. (brasiliensis subsp. brasiliensis), s.n. (lati- folid) Rizzini 7 Nov. 1972 (latifolid) Rodriguez 484 (australis) Rojas 12133 (australis) Rojas Acosta, N., 13004 (australis) Rosa 622 (grandiflora) Rose & Russel 20221 (brasiliensis subsp. bras- iliensis) Roth 356 (pauciflord) Rubens Faria 4 (pauciflord) Rusby 1030 (mire), 2122 (grandiflora subsp. schultesii), 2611 (mire), 621 (grandiflora subsp. schultesii), 862 (mire) Saldanha 8587 (brasiliensis subsp. brasiliensis) Sampaio 68 (brasiliensis subsp. brasiliensis), 74 (brasiliensis subsp. brasiliensis) Santos, N. 5790 (latifolid) Santos, T. Soares dos, 2511 (uniflord), 1964 (clandestina), 2015 (clandestina), 433 (clan- destina), 689 (clandestina) Santos, J. U. et al. 48 (grandiflora subsp. gran- diflora) Santos Lima Aug. 1940 (pauciflord) Santos-Lima 200 (hydrangeiformis subsp. hydran- geiformis) Sastre 1649 (guianensis), 4751 (guianensis) Schinini et al. 18580 (australis) Schott 5361 (martiana), s.n. (martiana) Schubert 2150 (guianensis) Schuch 591 (hydrangeiformis subsp. hydrangei- formis), s.n. (brasiliensis subsp. brasiliensis) Schultes 3340 (grandiflora subsp. schultesii), 3422 (chiricaspi) Schultes & Cabrera 19115 (grandiflora subsp. schultesii) Schultes & Rodrigues 26132A (martiana) Schulz 85 (australis), 8645 (guianensis) Schunke V., J. 8 (grandiflora subsp. schultesii), 1480 (grandiflora subsp. schultesii), 3480 (grandiflora subsp. grandiflora), 4034 (gran- diflora subsp. grandiflora), 4051 (grandiflora subsp. grandiflora) Schwacke 3311 (uniflord), s.n. (brasiliensis subsp. brasiliensis) 130 FIELDIANA: BOTANY Schwarz 139 (australis), 36 (australis), 48 (aus- tralis), 93 (australis), 8114 (australis), 8547 (australis), 8661 (australis), 8972 (australis) Schwarzer 50 (pilosa) Schwindt 2270 (pilosa) Scolnik 1495 (grandiflora subsp. grandiflora) Sellow 1573 (pilosa), 719 (hydrangeiformis subsp. capitata), s.n. (brasiliensis subsp. bras- iliensis), s.n. (hydrangeiformis subsp. capitata), s.n. (hydrangeiformis subsp. hydrangeiformis) Silva, N. T. & Santos, M.R., 4631 (guianensis) Silva, N. T. da 336 (guianensis) Silva, M. N. et al. 17 (mire) Smith, A.C., 3163 (uniflora) Smith, L.B., 1788 (brasiliensis subsp. macroca- lyx), 1989 (brasiliensis subsp. macrocalyx), 5786 (pauciflora) Smith, L.B., & Klein 7532 (pauciflora), 13044 (cuneifolia), 8567 (cuneifolia) Smith, L.B., & Reitz 12541 (cuneifolia), 12757 (pilosa), 12870 (pilosa) Smith, L.B., et al. 7936 (pauciflora) Solomon 7609 (grandiflora subsp. grandiflora), 7905 (grandiflora subsp. grandiflora) Solomon et al. 12022 (mire), 12071 (mire) Soukup 3041 (grandiflora subsp. schultesii), 446 (mire) Souza et al. 521 (brasiliensis subsp. brasiliensis) Spruce 1495 (amazonica), 3973 (grandiflora subsp. schultesii) St. Hilaire A-555 (hydrangeiformis subsp. hy- drangeiformis), A-746 (brasiliensis subsp. brasiliensis), B-2051 (obovata var. obovata), B- 58 (hydrangeiformis subsp. capitata), D-80 (brasiliensis subsp. brasiliensis), s.n. (bras- iliensis subsp. brasiliensis) Steinbach 1805 (grandiflora subsp. schultesii), 5487 (grandiflora subsp. schultesii), 7399 (grandiflora subsp. schultesii), 7505 (mire), 7936 (grandiflora subsp. schultesii), 8345 (un- iflora) Stephan s.n. (brasiliensis subsp. brasiliensis) Steyermark 86244 (imatacana), 86574 (imata- cana), 88290 (imatacana) Stork et al. 10461 (mire) Strang 172 (brasiliensis subsp. brasiliensis) Stuckert 16307 (australis) Sucre 2339 (brasiliensis subsp. brasiliensis), 2360 (brasiliensis subsp. brasiliensis), 4213 (bras- iliensis subsp. brasiliensis), 5314 (latifolia), 5806 (brasiliensis subsp. macrocalyx), 7446 (brasiliensis subsp. brasiliensis), 8002 (bras- iliensis) Sucre & Plowman 5086 (brasiliensis subsp. bras- iliensis), 5191 (brasiliensis subsp. macrocalyx), 5206 (brasiliensis subsp. macrocalyx) Sucre et al. 6428 (latifolia) Tamandare 244 (pilosa) Tastevin s.n. (grandiflora subsp. grandiflora) Taylor et al. 1084 (martiana) Tessman 3243 (grandiflora subsp. schultesii), 3541 (grandiflora subsp. grandiflora), 4921 (grandiflora subsp. schultesii), 5005 (grandiflo- ra subsp. schultesii), 5541 (grandiflora subsp. grandiflora) Torres 293 (grandiflora subsp. grandiflora) Traill 599 (grandiflora subsp. schultesii), 600 (grandiflora subsp. schultesii) Troll 240 (uniflora) Trujillo 3488 (imatacana) Tyson 5315 (d\vyeri) Tyson et al. 4335 (dwyeri) U.S. Exploring Expedition s.n. (hydrangeiformis subsp. capitata) U.S. South Pacific Exploring Expedition s.n. (brasiliensis subsp. brasiliensis) Ule 1521 (pilosa), 5453 (grandiflora subsp. schul- tesii), 6481 (grandiflora subsp. schultesii), 7061 (uniflora), 7715 (uniflora), 9746 (gran- diflora subsp. schultesii), 9747 (grandiflora subsp. schultesii), 9748 (grandiflora subsp. schultesii), 9749 (grandiflora subsp. schultesii), 9750 (mire), Nov. 1894 (hydrangeiformis subsp. hydrangeiformis), 23 Aug. 1896 (bras- iliensis subsp. brasiliensis), 10 Nov. 1896 (brasiliensis subsp. brasiliensis) Uribe, P., Dec. 1930 (grandiflora subsp. schulte- sii) Uribe Uribe 1353 (grandiflora subsp. schultesii) Usteri 23/306 (brasiliensis subsp. brasiliensis), 23 Nov. 1906 (pilosa) van Donselaar 3114 (guianensis) Vargas 15767 (mire), 16301 (grandiflora subsp. schultesii), 6311 (mire) Vasconcelos & Coelho 26 Feb. 1962 (grandiflora subsp. schultesii) Vauthier 42 (brasiliensis subsp. brasiliensis), 541 (brasiliensis subsp. brasiliensis) Venturi 4408 (australis) Vert 6152 (pauciflora) Vianna et al. 1574 (brasiliensis subsp. bras- iliensis) Vidal 311 (latifolia), 11-5435 (hydrangeiformis subsp. capitata), 11-5784 (brasiliensis subsp. brasiliensis), s.n. (brasiliensis subsp. bras- iliensis) Vieira et al. 641 (mire) PLOWMAN: REVISION OF BRUNFELSIA 131 Vincent s.n. (australis), s.n. (brasiliensis subsp. brasiliensis) Vogel 677 (brasiliensis subsp. macrocalyx) Warming 103 (brasiliensis subsp. brasiliensis) Wawra & Maly 426 (brasiliensis subsp. bras- iliensis) Weberbauer 6151 (grandiflora subsp. grandiflo- ra) Weddell 1648 (brasiliensis subsp. brasiliensis), 3621 (boliviano), 818 (hydrangeiformis subsp. capitatd), 878 (hydrangeiformis subsp. hydran- geiformis), s.n. 1844 (uniflord), s.n. 1858 (un- iflord) Weir 23 (brasiliensis subsp. brasiliensis) Wessels Boer 1195 (guianensis) Wettstein & Schiffner Sep. 1901 (pauciflord), s.n. (brasiliensis subsp. brasiliensis) White 1072 (mire], 1840 (grandiflora subsp. schultesii), 2386 (grandiflora subsp. schultesii) Widgren 486 (brasiliensis subsp. brasiliensis), 596 (hydrangeiformis subsp. capitatd), s.n. 1844 (hydrangeiformis subsp. capitatd), s.n. 1845 (uniflord) Wilbur & Weaver 11358 (dwyeri) Williams & Assis 6088 (brasiliensis subsp. bras- iliensis), 6859 (brasiliensis subsp. brasiliensis), 7928 (hydrangeiformis subsp. hydrangeifor- mis), 12 Feb. 45 (brasiliensis subsp. bras- iliensis) Williams, L. 5482 (grandiflora subsp. schultesii) Woolston 571 (australis) Woytkowksi 34536 (grandiflora subsp. grandiflo- ra), 35008 (grandiflora subsp. schultesii), 5026 (grandiflora subsp. grandiflora), 514 (grandi- flora subsp. grandiflora) 5525 (grandiflora subsp. schultesii), 6170 (grandiflora subsp. schultesii) Wullschlagel 1505 (guianensis) Ybarrola 2963 (australis), 3270 (australis), 3430 (australis) Zarucchi & Andrade 2316 (grandiflora subsp. grandiflora) Zaruma et al. 56 (chiricaspi) Zernys s.n. (brasiliensis subsp. macrocalyx) Appendix II: Brunfelsia sect. Brunfelsia Description Brunfelsia L. sect. Brunfelsia. Type species: B. americana L. Brunfelsia L., Sp. PL 191. 1753. Type species: B. americana L. Brunfelsia L. sect. Eubrunfelsia Griseb, Fl. Brit. W. Ind. 432. 1861. Type species: B. americana L. Brunfelsia L. subg. Brunfelsiopsis Urb., Symb. Ant. 1: 402. 1899. Type species: B. densifolia Krug & Urb. Brunfelsiopsis (Urb.) Kuntze in Post & Kuntze, Lex. 81. Dec. 1903 ("1904"). Type species: B. densifolia Krug & Urb. Brunfelsia sect. Brunfelsiopsis (Urb.) Dalla Torre & Harms, Gen. Siphon. 453. 1904. Type spe- cies: B. densifolia Krug & Urb. Shrubs or small trees to 10 m tall. Inflores- cence terminal or axillary. Flowers large, 1-many per inflorescence. Corolla white, turning yellow with age, or reddish purple; tube 6-24 times as long as calyx, gradually dilated toward apex, open, not constricted at mouth. Anthers oblong, weakly bilobate. Stigma subcapitate, convex, sub- entire or shallowly bilobate. West Indies (see Fig. 9). About 20 species.17 Names of Brunfelsia Sect. Brunfelsia Brunfelsia acunae Hadac, Fol. Geobot. Phytotax. 5: 430. 1970. Brunfelsia americana L., Sp. PI. 191. 1753. (B. terminalis Salisb., Prodr. Stirp. 109. 1796. B. violacea Lodd, Bot. Cab., t. 792. 1823. B. in- odora Mart., Del. Sem. Hort. Monac. 1847; Linnaea 24: 180. 1851. B. fallax Duchass. ex Griseb., Abh. Konigl. Ges. Wiss. Gottingen. 7: 242. 1857. B. americana var. pubescens Gri- seb., Fl. Brit. W. Ind. 432. 1864. B. abbottii Leonard, J. Wash. Acad. Sci. 17(3): 71. 1927.) Lesser Antilles, Puerto Rico, Dominican Re- public. Brunfelsia cestroides A. Rich., Fl. Cub. Fan. 3: 151. 1853. (B. vinciflora Griseb., Mem. Am. Acad. Arts 8(1): 523. 1860.) Cuba. Brunfelsia clarensis Britton & P. Wilson, Mem. Torrey Bot. Club 16(2): 102. 1920. Cuba. Brunfelsia densifolia Krug & Urb., Notizbl. Kon- igl. Bot. Gart. Berlin 1(10): 324. 1897. Puerto Rico. Brunfelsia grisebachii Amshoff, Contrib. Ocas. 17 This Caribbean group of brunfelsias is being revised at present by Victor Fuentes Fiallo of INIFAT, Havana, Cuba.— Eds. 132 FIELDIANA: BOTANY Mus. Hist. Nat. Col. "De la salle," Habana 15: 6. 1956. Cuba. Brunfelsia jamaicensis (Benth.) Griseb., Fl. Brit. W. Ind. 432. 1864. (B. nitida var. jamaicensis Benth. in DC., Prodr. 10: 201. 1846. B. harrisii Urb., Symb. Ant. 3: 373. 1903.) Jamaica. Brunfelsia lactea Krug & Urb., Notizbl. Bot. Gart. Berlin 1(10): 323. 1897. Puerto Rico. Brunfelsia linearis Ekman, Feddes Repert. Spec. Nov. Regni Veg. 21: 223. 1925. Cuba. Brunfelsia macroloba Urb., Symb. Ant. 9: 252. 1924. Cuba. Brunfelsia maliformis Urb., Symb. Ant. 3: 372. 1903. (B. fawcettii Urb, Symb. Ant. 3: 371. 1903.) Jamaica. Brunfelsia membranacea Urb., Symb. Ant. 5: 491. 1908. Jamaica. Brunfelsia nitida Benth. in DC., Prodr. 10: 201. 1846. (B. parvifolia A. Rich., Fl. Cub. Fan. 3: 151. 1853. B. longituba Lem., Jard. Fleur. 4: 61. 1854.) Cuba. Brunfelsia picardae Krug & Urb., Notizbl. Bot. Gart. Berlin 1(10): 321. 1897. Haiti. Brunfelsia plicata Urb., Symb. Ant. 6: 39. 1909. Jamaica. Brunfelsia pluriflora Urb., Symb. Ant. 9: 252. 1924. Cuba. Brunfelsia portoricensis Krug & Urb., Notizbl. Bot. Gart. Berlin 1(10): 322. 1897. Puerto Rico. Brunfelsia purpurea Griseb., Mem. Ann. Acad. Arts n. s. 8(1): 523. 1869. Cuba. Brunfelsia shaferi Britton & P. Wilson, Mem. Torrey Bot. Club 16(2): 102. 1920. Cuba. Brunfelsia sinuata A. Rich., Fl. Cub. Fan. 3: 151, t. 66. 1853. Cuba. Brunfelsia splendida Urb., Symb. Ant. 5: 491. 1908. Jamaica. Brunfelsia undulata Sw., Nov. Gen. Spec. PL 90. 1788. Jamaica. Appendix III: Additional References The following papers and accounts relevant to Brunfelsia were published after preparation of the original manuscript by Plowman. The list is by no means exhaustive. — Eds. BENNETT, B. C. 1992. Hallucinogenic plants of the Shuar and related indigenous groups in Amazonian Ecuador and Peru. Brittonia, 44(4): 483-493. BOAVENTURA, Y. M. S., AND L. H. PiEDADE. 1993. Floral biology and cytological analysis of Brunfelsia nitida Benth. Revista Brasileira de Genetica, 16(3): 785- 792. BRANTJES, N. B. M. 1978. Sensory responses to flowers in night-flying moths, pp. 13-19. In Richards, A. J., ed., The Pollination of Flowers by Insects. CARLQUIST, S. 1992. Wood anatomy of the Solanaceae: A survey. Allertonia, 6(4): 279-326. DAULATABAD, C. D., AND K. M. HOSAMANI. 1991. Un- usual fatty acids in Brunfelsia americana seed oil: A rich source of oil. Journal of the American Oil Chem- ist's Society, 68(8): 608-609. GASTIAZORO, M. T. C. DE. 1993. Estudio morfoanatomico de organos vegetativos en Cestroideae (Solanaceae) II: Tribu Salpiglossideae. Kurtziana, 22: 47-72. GUYOT, M. 1987. Les stomates du genre Brunfelsia (So- lanaceae). Bull. Mus. natn. Hist, nat., Paris, ser. 4, 9, section B, Adansonia, 3: 289-315. IYER, R. P., J. K. BROWN, M. G. CHAUBAL, AND M. H. MALONE. 1977. Brunfelsia hopeana I: Hippocratic screening and antiinflammatory evaluation. Lloydia, 40: 356-360. NYAWUAME, H. G. K., AND L. S. GILL. 1994. Epidermal studies of some species of family Solanaceae used in traditional medicine in West Africa. Feddes Reperto- rium, 105(1-2): 49-60. RUPPELT, B. M., E. F. R. PEREIRA, L. C. GON^ALVES, AND N. A. PEREIRA. 1991. Pharmacological screening of plants recommended by folk mesdicine as anti-snake venom — 1. Analgesic and anti-inflammatory activi- ties. Memorias do Institute Oswaldo Cruz, 86(suppl. II): 203-205. TANNER, E. V. J., AND V. KAPOS. 1982. Leaf structure of Jamaican upper montane rain-forest trees. Biotropica, 14(1): 16-24. THYR, B. D., M. J. SAMUEL, AND P. G. BROWN. 1975. New solanaceous host records for Corynebacterium michiganense. Plant Disease Reporter, 59(7): 595- 598. TOKARNIA, C. H., A. GAVA, L. STOLF, AND P. V. PEIXOTO. 1991. Experimental poisoning in cattle by Brunfelsia pauciflora (Solanaceae). Pesquisa Veterinaria Brasi- leira, ll(l-2):9-12. TRIGO, M. M. 1992. Contribution to the study of pollen in ornamental species. Solanaceae, Convolvulaceae and Hydrophyllaceae. Acta Botanica Malacitana, 17: 209-222. VICKERS, W. T, AND T. PLOWMAN. 1984. Useful plants of the Siona and Secoya Indians of Eastern Ecuador. Fieldiana, Botany, 15: 1-63. PLOWMAN: REVISION OF BRUNFELSIA 133 Index Synonyms and non-English common names are italicized. New names or combinations are in bold- face. Page numbers for illustrations are also in boldface. Anthocercideae 6 Atropaceae 4 Banisteriopsis caapi vii Besleria bonodora 59, 60 Besleria inodora 103 biogeography 29-34 Browallia 5 Brunfelsia 4, 34 Brunfelsia abbottii 132 Brunfelsia acuminata 62, 68 Brunfelsia acunae 132 Brunfelsia amazonica 38, 39 Brunfelsia americana 34, 132 var. pubescens 132 Brunfelsia angusta 1 19 Brunfelsia australis 53, 55 Brunfelsia bahiensis 119 Brunfelsia boliviana 57, 58 Brunfelsia bonodora 59, 60 Brunfelsia brasiliensis 61, 68 ssp. brasiliensis 62, 64 ssp. macrocalyx 66, 69 var. acuminata 62 Brunfelsia burchellii 40, 41 Brunfelsia calycina 103 f. grandiflora 78 var. eximia 103 var. floribunda 103 var. lindeniana 103 var. macrantha 103 Brunfelsia capitata 86 var. angustifolia 86 Brunfelsia cestroides 132 Brunfelsia chiricaspi vii, 70, 71 Brunfelsia chocoensis 40, 42 Brunfelsia clandestina 43, 44 Brunfelsia clarensis 132 Brunfelsia confertiflora 62 Brunfelsia cuneifolia 72, 73 Brunfelsia densiflora 34 Brunfelsia densifolia 132 Brunfelsia dwyeri 74, 75 Brunfelsia eximia 103 Brunfelsia falcata 1 1 9 Brunfelsia fallax 132 Brunfelsia fawcettii 133 Brunfelsia gracilis 119 Brunfelsia grandiflora 77 ssp. grandiflora 78, 79 ssp. schultesii vii, 53, 79, 80 Brunfelsia grisebachii 132 Brunfelsia guianensis 38, 46, 47 Brunfelsia harrisii 133 Brunfelsia hopeana 1 1 3 var. australis 53 var. macrocalyx 66 var. pubescens \ 1 3 Brunfelsia hydrangeiformis 84, 85 ssp. capitata 86, 87 ssp. hydrangeiformis 86, 85 var. glabriscula 86 Brunfelsia imatacana 89 Brunfelsia inodora 132 Brunfelsia jamaicensis 133 Brunfelsia lactea 133 Brunfelsia latifolia 91, 92, 119 Brunfelsia lindeniana 103 Brunfelsia linearis 133 Brunfelsia lockhartii 119 Brunfelsia longituba 133 Brunfelsia macrocarpa 93, 94 Brunfelsia macroloba 119 Brunfelsia macrophylla 86 Brunfelsia magnifica 119 Brunfelsia maliformis 133 Brunfelsia maritima 91 Brunfelsia martiana 49, 50 Brunfelsia membranacea 133 Brunfelsia mire 95, 96 Brunfelsia montana 119 Brunfelsia multiflora 119 Brunfelsia mutabilis 1 1 3 Brunfelsia nitida 133 \ar.jamaicensis 133 Brunfelsia nyctaginoides 1 1 9 Brunfelsia obovata 98, 100 var. coriacea 101 var. obovata 99 Brunfelsia paraguayensis 53 Brunfelsia parvifolia 133 Brunfelsia pauciflora 103, 104, 109 var. calycina 103 Brunfelsia picardae 133 Brunfelsia pilosa 108, 110 Brunfelsia plicata 133 Brunfelsia plurifolia 133 Brunfelsia portoricensis 133 Brunfelsia purpurea 133 Brunfelsia ramosissima 62, 68 var. confertiflora 62 var. laxiflora 62 var. parcifolia 62 Brunfelsia rupestris 111, 112 Brunfelsia schomburgkiana 119 Brunfelsia sieberi 1 1 9 Brunfelsia shaferi 133 Brunfelsia silvicola 101, 102 Brunfelsia sinuata 133 Brunfelsia spinosa 119 Brunfelsia splendida 133 Brunfelsia spruceana 38, 119 Brunfelsia tastevinii 78 Brunfelsia terminalis 132 Brunfelsia undulata 133 Brunfelsia uniflora 113, 114 f. intermedia 53 f. obovatifolia 53 f. typica 53 var. pubescens 1 1 3 Brunfelsia vinciflora 132 Brunfelsia violacea 132 Brunfelsiopsis 34 chiricaspi 72 Conejo, village vii cultivars 107 cytology 16 dispersal 23 ecology 20 edible fruits 43, 94 erva de macaco 69 evolutionary trends 26-30 Erythroxylum viii flor de Trovoada 74 flower morphology 10-15 Franciscea 4, 34, 51 Franciscea acuminata 62 Franciscea angusta 1 1 9 Franciscea australis 53 Franciscea calycina 103 Franciscea capitata 86 Franciscea confertiflora 62, 64 Franciscea divaricata 62 Franciscea eximia 103 Franciscea gracilis 119 Franciscea grandiflora 77 Franciscea hopeana 1 1 3 Franciscea hydrangeiformis 84 Franciscea latifolia 91 Franciscea lindeniana 103 Franciscea macrantha 103 Franciscea macrophylla 86 Franciscea maritima 91 Franciscea mutabilis 1 1 3 Franciscea obovata 98 Franciscea pauciflora 103 Franciscea pohliana 119 Franciscea ramosissima 62 Franciscea uniflora 34, 52 geographical distributions 29-34 Gerardia brasiliensis 61 guayabilla 95 Guianenses, section 38 hybridization 19-20, 83, 88, 98 134 FIELDIANA: BOTANY insect-plant relationships 23-26 Juajibioy, Pedro vii key to sections and species 35-37 Kofan customs vii, 72 lectotypes (new) 46, 51, 53, 59, 72, 86, 103 manacd 40, 49, 69 manacd assu 69 manacd da serra 69 manacd grande 82 manacd opifera 119 Mania opifera 119 medicinal use 77, 82, 84, 98, 118 mire 95 morphology 6-16, 26-30 ornamental 57, 82, 107, 111 Pinkley, Homer vii Plowman, Timothy vii-ix, xi Plowmania 6, 1 19 poisonous 69, 82 pollination biology 22 refugium hypothesis 33 Salpiglossideae 4, 5 sanango 82 sananguillo 82 Schultes, R. E. vii Schultesianthus megalandrus 76 section Franciscea 51 section Guianenses 38 Solanaceae 4 specialization, trends 26-30 Streptosolen 6 su-tim-ba-che vii thunderstorm flower 74 tsontinba"k"d vii tsontinb"k'o vii yage vii INDEX 135 UNIVERSITY OF ILLINOIS-URBANA